Premise of research. Early Cretaceous flowers recovered from Portugal and North America are some of the oldest and most informative angiosperm structures known. Analyses of this material provide a more direct basis for inferring floral structure and biology in early angiosperms than extrapolations based solely on extant taxa, and they have documented an unanticipated diversity of angiosperms, including the presence of many extinct forms, during the Early Cretaceous. The fossil flower described here from the Early Cretaceous of Portugal adds to the knowledge of this extinct diversity.

Methodology. Coalified fossil flowers were extracted from unconsolidated sediments and cleaned with HF, HCl, and water. Details were studied using scanning electron microscopy and synchrotron radiation X-ray tomographic microscopy. Phylogenetic relationships were evaluated by adding the fossil to a phylogenetic analysis of extant basal angiosperms based on morphological characters but constrained according to a topology based on molecular data.

Pivotal results. A new genus and species, Catanthus dolichostemon, are described on the basis of the fossil flowers. The perianth consists of three thick sepal-like tepals in an outer whorl and six thinner petallike tepals in two inner whorls. The androecium consists of several whorls of stamens with long, broad filaments and anthers with extrorse dehiscence. Pollen is trichotomocolpate and possibly monocolpate. The gynoecium is superior and apocarpous, consisting of six or, rarely, five carpels. Analysis of the phylogenetic position of Catanthus suggests a relationship to Canellales and Magnoliales.

Conclusions. Catanthus is a new Early Cretaceous magnoliid angiosperm for which the relationship to the two major clades of extant magnoliids (Laurales + Magnoliales or Canellales + Piperales) is not resolved securely. Catanthus adds to the evidence of substantial extinct diversity among early angiosperms, including the presence of extinct forms related to extant magnoliids, and it is consistent with a general pattern that angiosperm assemblages from the middle Albian and earlier are dominated by noneudicots.

Premise of research. The recognition of diverse and abundant chlamydospermous seeds from the Early Cretaceous of Denmark, Portugal, and eastern North America has been an unexpected outcome of studies of mesofloras that were initially focused on early angiosperms. These seeds provide structural information critical for understanding morphological and structural diversity in an important Mesozoic group of extinct gnetalean relatives.

Methodology. The fossil seeds were picked from Early Cretaceous mesofossil floras from localities in western Portugal and Virginia using a stereomicroscope. Selected seeds were studied in more detail for morphological and anatomical traits using SEM and synchrotron radiation X-ray tomographic microscopy.

Pivotal results. Six new species of chlamydospermous seeds are described that add substantially to the known diversity of Early Cretaceous chlamydosperms. In general seed organization, the fossils are similar to seeds of extant Gnetales, but none of the fossils can be assigned to any of the three living genera of Gnetales (Ephedra, Gnetum, and Welwitschia). All six species have similar closure of the micropylar canal but show considerable variation in the anatomy of the seed envelope. In micropylar closure, the fossils are most similar to extant Gnetum, but they differ in other respects from seeds of extant Gnetum, and one of the new seed taxa has polyplicate, ephedroid pollen in the micropyle. A well-preserved embryo with two cotyledons is preserved in seeds of Rothwellia foveata and provides the first information on the embryo in this Early Cretaceous chlamydospermous complex.

Conclusions. The chlamydospermous seeds described here show similarities to seeds of extant Gnetales. However, most of the fossils exhibit combinations of features that are unknown among extant species of Gnetales and clearly represent an extinct complex of plants that were important in Early Cretaceous vegetation, along with other extinct plant groups, including Bennettitales and Erdtmanithecales.

Premise of research. The fossil record of Aristolochiaceae (Piperales, magnoliids) is sparse, particularly from Cretaceous strata. Fossil seeds from the Early and mid-Cretaceous of Portugal and North America provide the earliest unequivocal documentation of the group.

Methodology. Detailed morphological and anatomical investigations of the fossil aristolochiaceous seeds were carried out using SEM and synchrotron radiation X-ray tomographic microscopy (SRXTM). Comparison with other seeds, extinct as well as extant, was based on published data and SRXTM analyses.

Pivotal results. Two new genera and two new species, Aristospermum huberi and Siratospermum mauldinense, are described based on fossil seeds from the Early and mid-Cretaceous of Portugal and North America. The seeds are anatropous and bitegmic, with the micropyle formed by the inner integument. The testa consists of an exotesta of varied thickness and an endotesta of crystalliferous cells. The tegmen is three cell layers thick and consists of an outermost layer of longitudinally aligned fibers, a middle layer of transversely aligned fibers perpendicular to the longitudinal fibers, and an inner layer of thin-walled cuboidal cells. In most seeds the exotesta is abraded, exposing the crystalliferous cells of the endotesta. Among extant plants, a similar seed coat with a crystalliferous endotesta and crossing fibers in the tegmen is unique to Aristolochiaceae.

Conclusion. The unique seed coat allows Aristospermum and Siratospermum to be assigned confidently to the lineage that today includes extant Aristolochiaceae. Aristospermum and Siratospermum provide the first unequivocal documentation of the Aristolochiaceae lineage of the Piperales during the Early Cretaceous diversification of angiosperms.

Premise of research. Small angiosperm fossils are diverse in Early Cretaceous mesofossil floras from Portugal and eastern North America. Investigations of these fossils have revealed an unexpectedly high diversity of extinct angiosperms related to lineages that are now species poor, such as Austrobaileyales, Nymphaeales, and Chloranthaceae. Here we analyze Early Cretaceous fruits and seeds from Portugal that are related to eumagnoliid angiosperms and that are also important for understanding extinct diversity in early angiosperms.

Methodology. The fossils were prepared by sieving in water; cleaned with HF, HCl, and water; and studied using scanning electron microscopy and synchrotron radiation X-ray tomographic microscopy. The systematic conclusion based on comparative studies was tested in a phylogenetic analysis.

Pivotal results. We recognize a new group of angiosperms based on fruits and seeds united by features that are otherwise unusual among angiosperms. Two genera, Serialis and Riaselis, are established and 10 species described. Both have unilocular fruiting units formed from a single carpel. Serialis has fruits with two or more seeds, while fruits of Riaselis are always one seeded. In Serialis, seeds are permanently attached to each other and dispersed as a unit. Both genera have anatropous and mesotestal-endotestal seeds with a tiny embryo and a distinctive vasculature in the testa extending from the hilum to the chalaza and then also on the antiraphal side to the micropyle. The fossils are most similar to seeds of Magnoliales but also share some features with seeds of Austrobaileya.

Conclusions. Serialis and Riaselis are the earliest fossils that can be assigned to the Magnoliales but are sufficiently different from those of all Magnoliales that they cannot be assigned to any extant family. Serialis and Riaselis provide further documentation of extensive extinct diversity among early angiosperms, and their abundance in the mesofossil floras suggests that they were common and widespread in early angiosperm communities.

A new microsporangiate cone, Renbernia zhoui, is described from the mesofossil flora extracted from exposures ofthe Potomac Group at Puddledock, Virginia, U.S.A., which are of Early Cretaceous (early-middle Albian) age. The cone consists of densely arranged dorsiventrally flattened laminar, flabelliform microsporangiophores that bear two elongate microsporangia on the presumed abaxial surface. The microsporangia are separated by sterile tissue that expands apically into a prominent hood-like sterile extension. The microsporangia have extrorse valvate dehiscence and both microsporangia and sterile apical expansion are covered by a short stiff hairs. In situ pollen resembles Decussosporites, elliptical with a long median colpus on the presumed distal surface flanked laterally by two equally long lateral colpi. A short transverse colpus on the presumed proximal surface links the two lateral colpi and divides the grain into two parts creating the appearance of two sacci. Renbernia zhoui is similar to Brenneria potomacensis described from the slightly older Drewry's Bluff and Dutch Gap mesofossil floras from the Potomac Group that also has Decussosporites-type pollen in situ. However, Renbernia microsporangiophores are more distinctly laminar and have sporangia that are more prominently elongated and with a hood-like apical extension of sterile tissue. The in situ pollen is also much smaller, the pollen wall is much more distinctly perforate-foveolate rather than more or less psilate, and in Renbernia the saccus-like structures are not inflated. The relationship of Brenneria and Renbernia, as well as the possible link between Decussosporites-type pollen and pollen of Eucommiidites (Erdtmanithecales) is discussed.

A new genus and species, Geminispermum virginiense, is described based on a well-preserved coalified cupulate reproductive unit recovered from the Early Cretaceous (early–middle Albian) Puddledock locality, Virginia, U.S.A. The reproductive unit is bisymmetrical and consists of an axis that bifurcates into two cupule-bearing stalks, each in the axil of a bract. Each cupule stalk bears a single non-valvate cupule recurved towards the center of the reproductive unit. The cupule opens distally by a short transverse slit with a distinct upper margin. Each cupule almost completely encloses a single orthotropous seed that is free from the cupule except at the base. The nucellus is also free from the integument except at the basal point of attachment. Geminispermum combines features of the ovulate structures of Caytoniales, Umkomasiales (= Corystospermales, including Doyleales) and Petriellales, but cannot be included in any of these existing orders as they are currently understood. The recurved, closed, non-valvate cupules are particularly similar to those of Caytonia, Petriellaea and Reymanownaea in external morphology, but differ in being one-seeded. The cupules of Geminispermum differ from the one-seeded cupules of Umkomasiales in being non-valvate and in having only a single cupule per bract. Geminispermum is perhaps most similar to the one- or two-seeded non-valvate cupules of Ktalenia from the Early Cretaceous of Argentina, but Ktalenia is poorly preserved, details of cupule architecture are uncertain, and the cupules appear to be associated with a single strongly dissected bract. Geminispermum is currently the only unequivocal seed plant cupule recovered from the Early Cretaceous Potomac Group and is distinct from all previously described cupulate reproductive structures.

Premise of research. Early Cretaceous Hedyosmum-like fossils are important because they provide information on the pistillate flowers and fruits of plants that produced Asteropollis pollen, which is common and widely distributed very early in the history of angiosperms. Hedyosmum (Chloranthaceae) is also the only extant genus for which there is a plausible fossil presence at such an early stage of angiosperm evolution.

Methodology. The fossils were sieved out of unconsolidated sediments and cleaned with HF, HCl, and water. External morphology and internal anatomy were studied using scanning electron microscopy and synchrotron radiation X-ray tomographic microscopy.

Pivotal results. New information on Hedyosmum-like fossils is provided based on pistillate flowers and fruits with adhering Asteropollis pollen from the Early Cretaceous of Portugal. The fossils are assigned to a new Early Cretaceous taxon, Hedyflora crystallifera, which in external morphology is closely similar to extant Hedyosmum. However, the fossils differ from the extant genus in having a crystalliferous endotesta with cells that have endoreticulate infillings, a feature characteristic of all extant Chloranthaceae except Hedyosmum. Extant Hedyosmum has a thin, unspecialized seed coat. This new discovery confirms earlier predictions that an endotestal seed coat is ancestral for Chloranthaceae as a whole but has been lost in the lineage leading to extant Hedyosmum.

Conclusions. Hedyflora confirms the divergence of the Hedyosmum lineage from other Chloranthaceae very early in the angiosperm radiation but refutes these early fossils as evidence of extant Hedyosmum in the Early Cretaceous.

A new taxon, Skyttegaardia galtieri, is described based on microsporangiophores with Monosulcites/Cycadopites pollen isolated from clays collected at the Skyttegård locality, island of Bornholm, Denmark, which are of earliest Cretaceous (Berriasian) age. Each microsporangiophore consists of a short, massive proximal fertile stalk-like portion with a truncate base, and a long sterile distal extension. A cavity on each side of the median line of the stalk-like portion, partially encloses a sporangium that dehisces by a longitudinal slit. The long distal extension tapers to a slender point and is curved toward the inferred adaxial side. The extension is irregularly angular in cross-section and the cuticle is thick with deep stomatal pits. The organization of the microsporangiophore, the in situ pollen and stomatal features suggest relationship with extant Cycadales. However, in all extant and fossil cycads there are usually many more sporangia per microsporangiophore, typically in groups of two to five, and they are borne on the surface of the proximal stalk-like portion rather than embedded in its tissues. These differences preclude secure inclusion of Skyttegaardia in Cycadales and open the possibility that these microsporangiophores were produced by a group of extinct plants, the other parts of which remain to be identified. The thick cuticle and sunken stomata of Skyttegaardia, together with the embedded sporangia, suggest adaptation to water stress, which is also consistent with the xeromorphic traits seen among the leaf fragments in the Skyttegaard flora and the arid conditions inferred from geological–geochemical proxies.

Three new genera, Atlantocarpus, Lambertiflora and Mugideiriflora, are described from the Early Cretaceous of North America and Portugal based on floral structures with multicarpellate and apocarpous gynoecia that have been studied using scanning electron microscopy and synchrotron radiation X-ray tomographic microscopy. Lambertiflora and Mugideiriflora have numerous perianth parts in several series, as well as many stamens and many carpels borne on a short conical receptacle. Atlantocarpus has many carpels arranged spirally on an elongated receptacle. Perianth and stamens are not preserved in Atlantocarpus, but scars of two sizes at the base of the receptacle indicate the presence of several series of tepals and stamens. Phylogenetic assessment of the three new genera indicates close relationships with members of extant Austrobaileyales, which is also favoured by the apparently ascidiate carpels of Atlantocarpus. However, the phylogenetic signal is not strong and the fossils also share many features with magnoliid angiosperms. Fully secure resolution of their relationships is hampered by lack of information of critical floral features in the fossil material, the constellation of likely plesiomorphic characters that they exhibit, and inadequate knowledge of character homologies and character evolution among extant taxa. There is also the broader concern about whether phylogenetic patterns can be reliably inferred for the earliest phases of angiosperm evolution based solely on a depauperate assemblage of extant taxa given extensive extinction over the last 100 million years. The new genera add to growing evidence of diverse extinct angiosperms with multiparted flowers during the Early Cretaceous that have a variety of relationships to extant ANA-grade angiosperms and magnoliids.

An Early Cretaceous mesofossil flora is described from the lower part of the Almargem Formation (late Barremian - early Aptian) from Torres Vedras (NE of Forte da Forca), Portugal. The flora is the oldest mesofossil flora containing angiosperm remains to be described in detail based on well-preserved flower, fruit and seed remains. In addition to angiosperms, the mesofossil flora also includes megaspores, sporangia and tiny leaves of spore-bearing plants. There are also twigs, cone fragments and seeds of conifers and seeds assigned to the BEG group. In total about 100 species have been distinguished. Most abundant in terms of plant fragments identified, are spore-bearing plants and conifers. Although only about 18% of the specimens can be attributed to angiosperms, angiosperm diversity is unexpected high considering the age of the flora. Angiosperms account for about 62 % of all species recognized. Angiosperm diversity is mainly at the level of ANA-grade angiosperms, eumagnoliids and in a few cases early diverging lineages of monocots. Eudicots are subordinate. Twenty new genera and 28 new species of angiosperms are established (Anaspermum operculatum gen. et sp. nov., Appofructus nudus gen. et sp. nov., Appomattoxia minuta sp. nov., Burgeria striata gen. et sp. nov., Canrightia elongata sp. nov., Choffaticarpus compactus gen. et sp. nov., Dejaxia brevicolpites gen. et sp. nov., Dinisia portugallica gen. et sp. nov., Eckhartia brevicolumella gen. et sp. nov., Eckhartia longicolumella sp. nov., Eckhartia intermedia sp. nov., Eckhartianthus lusitanicus gen. et sp. nov., Eckhartiopsis parva gen. et sp. nov., Gastonispermum antiquum sp. nov., Goczania rugosa gen. et sp. nov., Goczania inaequalis sp. nov., Goczania punctata sp. nov., Ibrahimia verminculata gen. et sp. nov., Juhaszia portugallica gen. et sp. nov., Kempia longicolpites gen. et sp. nov., Kvacekispermum costatum sp. nov., Mcdougallia irregularis gen. et sp. nov., Nicholsia brevicolpites gen. et sp. nov., Piercipollis simplex gen. et sp. nov., Reyanthus lusitanicus gen. et sp. nov., Samylinaea punctata gen. et sp. nov., Teebacia hughesii gen. et sp. nov., Vedresia elliptica gen. et sp. nov.). Comparison with results of a palynological study from the same horizon that yielded the mesofossil flora shows a marked underestimation of angiosperm diversity in the palynoflora, a pattern that has also been recognized elsewhere.

Angiosperm mesofossils are described from the Lower Cretaceous Almargem Formation exposed near the village of Catefica, Portugal, and are thought to be of Aptian-early Albian age. The mesofossil assemblage from Catefica is diverse and, in addition to the angiosperms described here, also contains a rich assemblage of non-angiosperm fossils, including leafy axes of bryophytes and lycopsids, lycopsid and salvinialean megaspores, and sporangia, sori and leaf fragments of ferns. Thereare also twigs, cones, cone scales, seeds and sporangia of several kinds of conifers. Other seed plants include 11 species of chlamydospermous seeds and vegetative axes related to the BEG group (Bennettiales-Erdtmanithecales-Gnetales). In terms of the number of plant fragments identified, angiosperms are most abundant in the Catefica assemblage and account for morethan half of all specimens. Angiosperms also dominate in number of species, but because the non-angiosperm fossils have notbeen studied in detail the total number of species in the flora is not yet established. Sixty-seven species of angiosperms arerecognized. Angiosperm diversity is mainly at the level of non-eudicots, including ANA-grade angiosperms, Chloranthaceae and magnoliids. Remains of chloranthoid angiosperms are especially common, both in the number of specimens and in numberof species recognized. About 40 % of the specimens, and more than 25 % of the species are chloranthoids. Remains of magnoliid angiosperms (Magnoliales, Laurales, Canellales, Piperales) are also prominent among the angiosperms. Eudicots are subordinate: only 3–4 % of all angiosperm specimens can be assigned confidently to eudicot angiosperms. Five new genera and six new species of angiosperms are established (Canrightia foveolata sp. nov., Elasmostemon paisii gen. et sp. nov., Endressistemon cateficensis gen. et sp. nov., Ibericarpus cuneiformis gen. et sp. nov., Proencistemon portugallicus gen. et sp. nov., Valvidistemon globiferus gen. et sp. nov.). Several other new taxa are also described, but not formally named.

A new fossil angiosperm, Paisia pantoporata, is described from the Early Cretaceous Catefica mesofossil flora, Portugal, based on coalified floral buds, flowers and isolated floral structures. The flowers are actinomorphic and structurally bisexual with a single whorl of five fleshy tepals, a single whorl of five stamens and a single whorl of five carpels. Tepals, stamens and carpels are opposite, arranged on the same radii and tepals are involute at the base clasping the stamens. Stamens have a massive filament that grades without a joint into the anther. The anthers are dithecate and tetrasporangiate with extensive connective tissue between the tiny pollen sacs. Pollen grains are pantoporate and spiny. The carpels are free, apparently plicate, with many ovules borne in two rows along the ventral margins. Paisia pantoporata is the oldest known flower with pantoporate pollen. Similar pantoporate pollen was also recognised in the associated dispersed palynoflora. Paisia is interpreted as a possibly insect pollinated, herbaceous plant with low pollen production and low dispersal potential of the pollen. The systematic position of Paisia is uncertain and Paisia pantoporata most likely belongs to an extinct lineage. Pantoporate pollen occurs scattered among all major groups of angiosperms and a close match to the fossils has not been identified. The pentamerous floral organisation together with structure of stamen, pollen and carpel suggests a phylogenetic position close to the early diverging eudicot lineages, probably in the Ranunculales.