Studies of the earliest Cretaceous angiosperms in the 1970s made only broad comparisons with living taxa, but discoveries of fossil flowers and increasingly robust molecular phylogenies of living angiosperms allow more secure recognition of extant clades. The middle to late Albian rise of tricolpate pollen and the first local dominance of angiosperm leaves mark the influx of near-basal lines of eudicots. Associated flowers indicate that palmately lobed ‘platanoids’ and Sapindopsis are both stem relatives of Platanus, while Nelumbites was related to Nelumbo (also Proteales) and Spanomera to Buxaceae. Monocots are attested by Aptian Liliacidites pollen and Acaciaephyllum leaves and Albian araceous inflorescences. Several Albian–Cenomanian fossils belong to Magnoliidae in the revised monophyletic sense, including Archaeanthus in Magnoliales and Virginianthus and Mauldinia in Laurales, while late Barremian pollen tetrads (Walkeripollis) are related to Winteraceae. In the basal ANITA grade, Nymphaeales are represented by Aptian and Albian flowers and whole plants (Monetianthus, Carpestella and Pluricarpellatia). Epidermal similarities of lower Potomac leaves to woody members of the ANITA grade are consistent with Albian flowers assignable to Austrobaileyales (Anacostia). Aptian to Cenomanian mesofossils represent both crown group Chloranthaceae (Asteropollis plant) and stem relatives of Chloranthaceae and/or Ceratophyllum (Canrightia, Zlatkocarpus, Pennipollis plant and possibly Appomattoxia).

New morphological and phylogenetic data prompt us to present an updated review of floral morphology and its evolution in the basal ANITA grade of living angiosperms, Chloranthaceae, and Ceratophyllum. Floral phyllotaxis is complex whorled in Nymphaeales and spiral in Amborella and Austrobaileyales. It is unresolved whether phyllotaxis was ancestrally whorled or spiral, but if it was whorled, the whorls were trimerous. The flowers are probably ancestrally bisexual because in most families with unisexual flowers these flowers exhibit rudiments of the opposite sex. Carpels are largely ascidiate and the closure line is short, either transverse or longitudinal. A style is usually absent or, if present, generally short and not plicate. Angiospermy (carpel sealing) is by secretion, rather than postgenital fusion, except in large-flowered Nymphaeales and in Illicium, correlated with unusual fruits. Carpels with a single, median, pendent ovule are probably plesiomorphic. Chloranthaceae and Ceratophyllaceae have an unsettled phylogenetic position, but in some phylogenetic analyses they form a clade, which may be sister to the remaining mesangiosperms (Magnoliidae, monocots, eudicots). This position is supported by their carpel characters, which are similar to those of the ANITA grade and different from those of most other mesangiosperms.

Premise of research. Discoveries of fossil flowers in Cretaceous rocks offer improved evidence for relationships with living clades, but for more secure inferences formal phylogenetic analyses are desirable. We extend previous analyses of magnoliids, monocots, and basal eudicots to Aptian, Albian, and Cenomanian fossils related to the basal "ANITA" lines and Chloranthaceae. Methodology. We performed parsimony analyses of a morphological data set of Recent angiosperms and published fossils, with the arrangement of Recent taxa constrained to backbone trees based primarily on molecular data. Pivotal results. Not only Monetianthus (as previously inferred) but also Carpestella is nested within Nymphaeaceae, while Pluricarpellatia may be a stem relative of Cabombaceae or Nymphaeaceae. Anacostia (with Similipollis pollen) is nested within Austrobaileyales. The position of Couperites (with Clavatipollenites pollen) is ambiguous: it may be on the stem lineage of Chloranthaceae (and Ceratophyllum, if this extant aquatic is related to Chloranthaceae), nested in Chloranthaceae, or more basal. Plants with Asteropollis pollen and reduced tepals are related to the chloranthaceous genus Hedyosmum. Zlatkocarpus, which also has a reduced perianth, may be either a stem relative or a crown group member of Chloranthaceae. Plants that produced loosely reticulate Pennipollis pollen are more likely related to Chloranthaceae and/or Ceratophyllum than to monocots. We confirm that Canrightia, with bisexual flowers and a reduced perianth, is a stem relative of Chloranthaceae. Despite similarities to Piperales, Appomattoxia (with Tucanopollis pollen) is more likely near the base of the ANITA grade or related to Chloranthaceae and/or Ceratophyllum. Conclusions. The Cretaceous rise of angiosperms involved the radiation not only of magnoliids, eudicots, and monocots but also of basal ANITA lines, including both aquatic Nymphaeales and woody groups. Our results reaffirm the early diversity of Chloranthaceae and clarify their floral evolution, in which a shift to unisexual flowers preceded loss of the perianth. © 2014 by The University of Chicago. All rights reserved.

Chloranthaceae were one of the first common lines during the early radiation of angiosperms, possibly reflecting adaptation to more open habitats. Phylogenetic analyses clarify the position of Cretaceous mesofossils in molecular trees of Recent taxa. Plants that produced Asteropollis pollen, with tepals adnate to a single carpel, are nested in crown group Chloranthaceae with Hedyosmum; Canrightiopsis, with three stamens and no perianth, is sister to Sarcandra and Chloranthus; and Canrightia is a stem relative that illustrates a still bisexual stage in floral reduction. Plants that produced Pennipollis pollen are related to Chloranthaceae and/or Ceratophyllum rather than monocots. Appomattoxia, which produced Tucanopollis pollen, has equivocal affinities, but Pseudoasterophyllites, with similar pollen and stems with reduced leaves, may be a link between Chloranthaceae and Ceratophyllum. These results imply that flowers became unisexual before losing the perianth, while bisexual flowers in Canrightiopsis, Sarcandra, and Chloranthus are secondarily derived from unisexual flowers.

Because the Gulf Coast Lower Cretaceous is dated by ammonites, it has great potential as a reference section for correlating continental sequences such as the Potomac Group and dating events in the rise of angiosperms. Middle Albian terrestrial palynofloras from Oklahoma have been described in detail, but the early Albian interval, represented in the Glen Rose Formation of Texas, is less well known. Samples from two localities on the Paluxy River, which correlate with horizons containing late early Albian ammonites, are dominated by Classopollis and Exesipollenites, but angiosperm pollen is the next most common terrestrial element, and there are several index spore species for Zone II in the Potomac Group. Among the angiosperms, reticulate monosulcates are most common and diverse, but there are also several tricolpate species. Stratigraphically important angiosperms include the Clavatipollenites rotundus group and reticulate tricolpates, which appear in the upper part of Potomac Zone I and the dated earliest Albian of England and Portugal. However, there are also tricolpates with striate-reticulate sculpture, a pollen type that is not known from upper Zone I but appears in the late early Albian of Portugal. This assemblage contrasts with floras from Potomac Zone II and the middle Albian (lower Fredericksburg Group) of Oklahoma, where tricolpates overtook monosulcate angiosperms in species diversity. These results confirm arguments based on the Portuguese section that there is a significant hiatus between Potomac Zones I and II, and that this gap is at least partly late early Albian. The dominance of Classopollis and Exesipollenites and the occurrence of isolated Northern Gondwanan elements (Sergipea, Tucanopollis) suggest that Texas lay in a transition zone between Southern Laurasia and the hotter and drier Northern Gondwana province, but regional studies are needed to disentangle geographic and climatic factors from effects of the lagoonal local environment.

Phylogenetic analyses imply that monocots were a key group in the early radiation of angiosperms, yet they are much rarer than other major clades in the Early Cretaceous macrofossil record. Here we describe a well-preserved leaf and several inflorescences related to the near-basal monocot family Araceae and abundant monocot leaves of uncertain affinities from two latest Albian localities in north-eastern Spain. Orontiophyllum ferreri sp. nov. has a multistranded midrib, several orders of parallel-pinnate veins, two orders of transverse veins, and paracytic-oblique stomata. This suite of characters (but with both anomocytic and paracytic-oblique stomata) is characteristic today of Orontium in the near-basal araceous subfamily Orontioideae, and later Cretaceous and early Cenozoic leaves assigned to Orontiophyllum have similar architecture. Sedimentology and anatomy suggest a (semi)aquatic ecology. Other monocot leaves at the same locality are linear and parallel-veined but have similar stomata. Although anomocytic stomata have been proposed as ancestral in monocots, O. ferreri, the associated linear leaves, Albian–Cenomanian cuticles from Australia and Portugal, and extant data are consistent with the hypothesis that variable paracytic-oblique stomata are ancestral. Turolospadix bogneri gen. et sp. nov., from the other locality, includes spadices of ebracteate flowers with four tepals, a central gynoecium, and a long stipe (vs a spathe attached just below the fertile zone as in most Araceae). Phylogenetic analyses indicate that the character combinations seen in O. ferreri and T. bogneri are ancestral for Araceae, and they could be either sister to Araceae or nested within a basal grade of the family. Together with fossils from the Aptian–Albian of Brazil and Portugal, the Spanish fossils indicate that Araceae are among the oldest extant monocot families, but they were associated with diverse linear-leaved monocots of uncertain affinities.

Recent advances in molecular phylogenetics and a series of important palaeobotanical discoveries have revolutionized our understanding of angiosperm diversification. Yet, the origin and early evolution of their most characteristic feature, the flower, remains poorly understood. In particular, the structure of the ancestral flower of all living angiosperms is still uncertain. Here we report model-based reconstructions for ancestral flowers at the deepest nodes in the phylogeny of angiosperms, using the largest data set of floral traits ever assembled. We reconstruct the ancestral angiosperm flower as bisexual and radially symmetric, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels. Although uncertainty remains for some of the characters, our reconstruction allows us to propose a new plausible scenario for the early diversification of flowers, leading to new testable hypotheses for future research on angiosperms.