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Text-fig. 36. Synchrotron radiation X-ray tomographic microscopy (SRXTM, a–e) images of "Paisia-like follicle"; Catefica locality, Portugal. a, b) Volume rendering of follicle in lateral (a) and ventral (b) views showing the decurrent stigmatic region that extends from the follicle base to the apex but lacks a distinct papillate zone; c) Longitudinal section (volume rendering cut at orthoslice yz0341) of follicle showing under-developed ovules towards the base and numerous well-developed ovules/seeds in the upper part suggesting that the follicle is probably mature; d) Dorsal view of follicle apex showing the cleft in the presumed stigmatic apical region; e) Transverse section (orthoslice xy1294) of follicle with one dorsal and two ventral bundles and two placentae bulging into the locule, one on either side of the ventral suture; note the strongly compressed outer epidermis and the homogenized cells of the mesocarp. Specimen, Catefica 49-S174915 (a–e). Scale bars = 300 Μm (a–d), 100 Μm (e). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 36. Synchrotron radiation X-ray tomographic microscopy (SRXTM, a–e) images of "Paisia-like follicle"; Catefica locality, Portugal. a, b) Volume rendering of follicle in lateral (a) and ventral (b) views showing the decurrent stigmatic region that extends from the follicle base to the apex but lacks a distinct papillate zone; c) Longitudinal section (volume rendering cut at orthoslice yz0341) of follicle showing under-developed ovules towards the base and numerous well-developed ovules/seeds in the upper part suggesting that the follicle is probably mature; d) Dorsal view of follicle apex showing the cleft in the presumed stigmatic apical region; e) Transverse section (orthoslice xy1294) of follicle with one dorsal and two ventral bundles and two placentae bulging into the locule, one on either side of the ventral suture; note the strongly compressed outer epidermis and the homogenized cells of the mesocarp. Specimen, Catefica 49-S174915 (a–e). Scale bars = 300 Μm (a–d), 100 Μm (e).
Text-fig. 35. Scanning electron microscope (SEM, a, b, d) and synchrotron radiation X-ray tomographic microscopy (SRXTM, c, e) images of "Paisia-like follicle"; Catefica locality, Portugal. a) Lateral view of slender follicle with an almost straight ventral margin and a slightly curved dorsal margin; b) Lateral view of small, broad follicle with slightly sinuous ventral margin and rounded dorsal margin; c) Lateral view of small, broad follicle with slightly sinuous ventral margin and rounded dorsal margin; d) Apical part of follicle in (a) showing the slight apical cleft in the probable stigmatic region; e) Transverse section (orthoslice xy0407) of follicle showing ovules and distinct follicle wall with small, thin-walled cells of the outer epidermis (arrow), larger, isodiametric cells of the mesocarp and an inner layer of smaller, thin-walled cells. Specimens, Catefica 50-S171523 (a, d), Catefica 343-S171515 (b), Catefica 49-S174929 (c, e). Scale bars = 300 Μm (a–c), 100 Μm (d, e). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 35. Scanning electron microscope (SEM, a, b, d) and synchrotron radiation X-ray tomographic microscopy (SRXTM, c, e) images of "Paisia-like follicle"; Catefica locality, Portugal. a) Lateral view of slender follicle with an almost straight ventral margin and a slightly curved dorsal margin; b) Lateral view of small, broad follicle with slightly sinuous ventral margin and rounded dorsal margin; c) Lateral view of small, broad follicle with slightly sinuous ventral margin and rounded dorsal margin; d) Apical part of follicle in (a) showing the slight apical cleft in the probable stigmatic region; e) Transverse section (orthoslice xy0407) of follicle showing ovules and distinct follicle wall with small, thin-walled cells of the outer epidermis (arrow), larger, isodiametric cells of the mesocarp and an inner layer of smaller, thin-walled cells. Specimens, Catefica 50-S171523 (a, d), Catefica 343-S171515 (b), Catefica 49-S174929 (c, e). Scale bars = 300 Μm (a–c), 100 Μm (d, e).
Text-fig. 34. Synchrotron radiation X-ray tomographic microscopy (SRXTM, a–h) images of "Paisia-like follicle"; Catefica locality, Portugal. a–d) Volume rendering of follicles in lateral (a, c) and ventral (b, d) views showing the decurrent stigmatic region that extends from base of the follicle to the apex; note papillate zone forming a probable stigma along the full length of the ventral suture; e) Longitudinal section (volume rendering cut at orthoslice yz0326) near the base of the follicle showing two ovules with a striate-reticulate surface (asterisks); note transverse fibers lining the inner follicle wall and large cells of the mesocarp; f) Transverse section (volume rendering cut at orthoslice xy2475) of follicle showing two rows of ovules borne on placentae on either side of the ventral suture (asterisks); note also the pronounced and densely-spaced papillae around the ventral suture; g) Transverse section (orthoslice xy1988) of follicle showing two ventral vascular bundles and one dorsal bundle (arrows) and ovules/seeds borne on two placentae, one on either side of the ventral suture; note the remains of the small thin-walled cells of the outer epidermis that cover the thicker-walled cells of the mesocarp; h) Transverse section (orthoslice xy2860) of follicle showing two ventral bundles and one dorsal bundle (arrows) and ovules/seeds in two rows on the placentae, one on either side of the ventral suture; note the remains of small epidermal cells and the large rounded cells of the mesocarp with thicker walls. Specimens, Catefica 49-S174916 (a, b), Catefica 49-S174917 (c–f, h), Catefica 50-S171525 (g). Scale bars = 300 Μm (a–d), 100 Μm (e–h). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 34. Synchrotron radiation X-ray tomographic microscopy (SRXTM, a–h) images of "Paisia-like follicle"; Catefica locality, Portugal. a–d) Volume rendering of follicles in lateral (a, c) and ventral (b, d) views showing the decurrent stigmatic region that extends from base of the follicle to the apex; note papillate zone forming a probable stigma along the full length of the ventral suture; e) Longitudinal section (volume rendering cut at orthoslice yz0326) near the base of the follicle showing two ovules with a striate-reticulate surface (asterisks); note transverse fibers lining the inner follicle wall and large cells of the mesocarp; f) Transverse section (volume rendering cut at orthoslice xy2475) of follicle showing two rows of ovules borne on placentae on either side of the ventral suture (asterisks); note also the pronounced and densely-spaced papillae around the ventral suture; g) Transverse section (orthoslice xy1988) of follicle showing two ventral vascular bundles and one dorsal bundle (arrows) and ovules/seeds borne on two placentae, one on either side of the ventral suture; note the remains of the small thin-walled cells of the outer epidermis that cover the thicker-walled cells of the mesocarp; h) Transverse section (orthoslice xy2860) of follicle showing two ventral bundles and one dorsal bundle (arrows) and ovules/seeds in two rows on the placentae, one on either side of the ventral suture; note the remains of small epidermal cells and the large rounded cells of the mesocarp with thicker walls. Specimens, Catefica 49-S174916 (a, b), Catefica 49-S174917 (c–f, h), Catefica 50-S171525 (g). Scale bars = 300 Μm (a–d), 100 Μm (e–h).
Text-fig. 16. Scanning electron microscope (SEM) images of "Stamen fragments with in situ Clavatipollenites- or Asteropollis-type pollen" (sp. 1: a–c; sp. 2: d–f; sp. 3: g–i); Catefica locality, Portugal. a) Stamen fragment showing pollen sacs; b) Distal view of pollen grain from (a) showing semitectate-reticulate tectum; c) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered, columellae supporting muri with fine pits and rounded supratectal ornamentation; note orbiculae with a finely spiny surface (arrows); d) Stamen showing very short filament, lateral pollen sacs and short apical extension of the narrow connective; e) Folded pollen grain from (d) showing semitectate-reticulate tectum; f) Detail of pollen wall from (d) showing the semitectatereticulate tectum and muri with fine rounded ornamentation; g) Stamen fragment; h, i) Detail of pollen grains from (g) showing the semitectate-reticulate tectum with smooth muri, long scattered columellae and tiny scattered orbicules (arrow). Specimens, Catefica 50-S170395 (a–c), Catefica 49-S172561 (d–f), Catefica 50-S170390 (g–i). Scale bars = 600 Μm (a, d, g), 6 Μm (b, e, h), 1.5 Μm (c, f, i). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 16. Scanning electron microscope (SEM) images of "Stamen fragments with in situ Clavatipollenites- or Asteropollis-type pollen" (sp. 1: a–c; sp. 2: d–f; sp. 3: g–i); Catefica locality, Portugal. a) Stamen fragment showing pollen sacs; b) Distal view of pollen grain from (a) showing semitectate-reticulate tectum; c) Detail of pollen wall showing the semitectate-reticulate tectum and long, scattered, columellae supporting muri with fine pits and rounded supratectal ornamentation; note orbiculae with a finely spiny surface (arrows); d) Stamen showing very short filament, lateral pollen sacs and short apical extension of the narrow connective; e) Folded pollen grain from (d) showing semitectate-reticulate tectum; f) Detail of pollen wall from (d) showing the semitectatereticulate tectum and muri with fine rounded ornamentation; g) Stamen fragment; h, i) Detail of pollen grains from (g) showing the semitectate-reticulate tectum with smooth muri, long scattered columellae and tiny scattered orbicules (arrow). Specimens, Catefica 50-S170395 (a–c), Catefica 49-S172561 (d–f), Catefica 50-S170390 (g–i). Scale bars = 600 Μm (a, d, g), 6 Μm (b, e, h), 1.5 Μm (c, f, i).
Text-fig. 4. Synchrotron radiation X-ray tomographic microscopy (SRXTM) images of seeds of Canrightia foveolata sp. nov.; Catefica locality, Portugal. a–d) Volume renderings of abraded seeds in ventral (a), lateral (b, d) and apical (c) views showing the slightly protruding chalaza (arrows) and dense longitudinal grooves with shallow pits in the surface of the endotesta; e) Transverse section of seed (orthoslice xy0665) showing the irregular grooved surface of the endotesta (oi-en) and the tegmen comprised of two layers of thick-walled cells that surround the cells of the prominent endothelium (asterisk); f) Longitudinal section (orthoslice xz1195) through seed showing the thin-walled endothelium cells (asterisk) surrounded by the thicker cells of the outer tegmen and endotesta; g) Longitudinal section (orthoslice yz0727) through seed showing outlines of angular crystals evenly distributed in cells of the endotesta (oi-en); note the outer epidermis of the tegmen (ii-o) composed of thick-walled cells; h) Longitudinal section (orthoslice xz0940) of seed showing details of the chalazal region with course of the vascular bundle (vb), cells of the prominent endothelium (asterisk), crystalliferous endotesta of the outer integument (oi-en) and the distinct thick walled cells of the outer cells of the tegmen (ii-o); i) Longitudinal and tangential section (orthoslice yz0542) through the endotesta (oi-en) showing the outlines of densely spaced crystals. Specimens, Catefica 242-S175178 (a–c, e–h), Catefica 49-S175179 (d, i). Scale bars = 300 Μm (a–d), 100 Μm (e, f, h, i), 50 Μm (g). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 4. Synchrotron radiation X-ray tomographic microscopy (SRXTM) images of seeds of Canrightia foveolata sp. nov.; Catefica locality, Portugal. a–d) Volume renderings of abraded seeds in ventral (a), lateral (b, d) and apical (c) views showing the slightly protruding chalaza (arrows) and dense longitudinal grooves with shallow pits in the surface of the endotesta; e) Transverse section of seed (orthoslice xy0665) showing the irregular grooved surface of the endotesta (oi-en) and the tegmen comprised of two layers of thick-walled cells that surround the cells of the prominent endothelium (asterisk); f) Longitudinal section (orthoslice xz1195) through seed showing the thin-walled endothelium cells (asterisk) surrounded by the thicker cells of the outer tegmen and endotesta; g) Longitudinal section (orthoslice yz0727) through seed showing outlines of angular crystals evenly distributed in cells of the endotesta (oi-en); note the outer epidermis of the tegmen (ii-o) composed of thick-walled cells; h) Longitudinal section (orthoslice xz0940) of seed showing details of the chalazal region with course of the vascular bundle (vb), cells of the prominent endothelium (asterisk), crystalliferous endotesta of the outer integument (oi-en) and the distinct thick walled cells of the outer cells of the tegmen (ii-o); i) Longitudinal and tangential section (orthoslice yz0542) through the endotesta (oi-en) showing the outlines of densely spaced crystals. Specimens, Catefica 242-S175178 (a–c, e–h), Catefica 49-S175179 (d, i). Scale bars = 300 Μm (a–d), 100 Μm (e, f, h, i), 50 Μm (g).
Text-fig. 15. Scanning electron microscope (SEM) images of stamen with "Asteropollis-type pollen sp. 2"; Catefica locality, Portugal. a) Stamen with pollen in situ showing the dome-shaped extension of the connective with hairs; b–d) Pollen in situ in stamen in (a) showing poorly defined pentachotomocolpate (b, d) and possibly tetrachotomocolpate (c) apertures and semitectate-reticulate tectum; e) Detail of broken pollen grains showing the loosely attached reticulum, long scattered columellae and thick foot layer. Specimen, Catefica 49-S101209 (a–e). Scale bars = 600 Μm (a), 6 Μm (b–e). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 15. Scanning electron microscope (SEM) images of stamen with "Asteropollis-type pollen sp. 2"; Catefica locality, Portugal. a) Stamen with pollen in situ showing the dome-shaped extension of the connective with hairs; b–d) Pollen in situ in stamen in (a) showing poorly defined pentachotomocolpate (b, d) and possibly tetrachotomocolpate (c) apertures and semitectate-reticulate tectum; e) Detail of broken pollen grains showing the loosely attached reticulum, long scattered columellae and thick foot layer. Specimen, Catefica 49-S101209 (a–e). Scale bars = 600 Μm (a), 6 Μm (b–e).
Text-fig. 23. Synchrotron radiation X-ray tomographic microscopy (SRXTM) images of Ibericarpus cuneiformis gen. et sp. nov.; Catefica locality, Portugal. a) Longitudinal section (orthoslice yz0827) showing fruitlet with apical stigmatic region and a single seed enclosed within the fruitlet wall that has a thick epidermal cuticle; note the tiny embryo (emb) internal to the micropyle (mi) and oriented toward the base of the fruitlet; b) Transverse section through apical part of two fruitlets (orthoslice xy0810) showing the fruitlet wall (fr) composed of small thin-walled cells covered by an epidermis of isodiametric cells (ep) with a thick outer cuticle (cu); c) Longitudinal section (orthoslice xz0370) through basal part of fruitlet perpendicular to section in (a) showing the micropyle (mi), embryo (emb) composed of tiny cells, and the thick cuticle (cu) covering the bulging cells of the fruitlet epidermis. Specimen, Catefica 50-S174907 (a–c). Scale bars = 300 Μm (a–c). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 23. Synchrotron radiation X-ray tomographic microscopy (SRXTM) images of Ibericarpus cuneiformis gen. et sp. nov.; Catefica locality, Portugal. a) Longitudinal section (orthoslice yz0827) showing fruitlet with apical stigmatic region and a single seed enclosed within the fruitlet wall that has a thick epidermal cuticle; note the tiny embryo (emb) internal to the micropyle (mi) and oriented toward the base of the fruitlet; b) Transverse section through apical part of two fruitlets (orthoslice xy0810) showing the fruitlet wall (fr) composed of small thin-walled cells covered by an epidermis of isodiametric cells (ep) with a thick outer cuticle (cu); c) Longitudinal section (orthoslice xz0370) through basal part of fruitlet perpendicular to section in (a) showing the micropyle (mi), embryo (emb) composed of tiny cells, and the thick cuticle (cu) covering the bulging cells of the fruitlet epidermis. Specimen, Catefica 50-S174907 (a–c). Scale bars = 300 Μm (a–c).
Text-fig. 31. Scanning electron microscope (SEM) images of isolated pollen of Piercipollis sp. (a, b) and Teebacia sp. (c–e); Catefica locality, Portugal. a) Isolated pollen grain (arrow) adhering to the much larger pollen of Araucariacites sp. in fragment of a conifer cone; note the size difference between the angiosperm and conifer pollen that is typical in Early Cretaceous floras; b) Pollen grain in (a) enlarged showing the extended aperture and homobrochate reticulum with smooth muri supported by long, scattered, columellae; c) Isolated pollen grains adhering to the outer surface of a Saportanthus parvus flower; d) Pollen grain in (c) enlarged showing the open reticulum and muri ornamented by fine transverse ribs; e) Detail of pollen grain in (d) showing the muri supported by long, scattered columellae; note the fine transverse ribs on the muri. Specimens, Catefica 49-S170139-01 (a, b), Catefica 361-S174322-01 (c–e). Scale bars = 50 Μm (a), 20 Μm (c), 6 Μm (b, d), 1.5 Μm (e). in The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms
Text-fig. 31. Scanning electron microscope (SEM) images of isolated pollen of Piercipollis sp. (a, b) and Teebacia sp. (c–e); Catefica locality, Portugal. a) Isolated pollen grain (arrow) adhering to the much larger pollen of Araucariacites sp. in fragment of a conifer cone; note the size difference between the angiosperm and conifer pollen that is typical in Early Cretaceous floras; b) Pollen grain in (a) enlarged showing the extended aperture and homobrochate reticulum with smooth muri supported by long, scattered, columellae; c) Isolated pollen grains adhering to the outer surface of a Saportanthus parvus flower; d) Pollen grain in (c) enlarged showing the open reticulum and muri ornamented by fine transverse ribs; e) Detail of pollen grain in (d) showing the muri supported by long, scattered columellae; note the fine transverse ribs on the muri. Specimens, Catefica 49-S170139-01 (a, b), Catefica 361-S174322-01 (c–e). Scale bars = 50 Μm (a), 20 Μm (c), 6 Μm (b, d), 1.5 Μm (e).
Integrating fossil flowers into the angiosperm phylogeny using molecular and morphological evidence
<p><span>F</span><span>ossils are essential to infer past evolutionary processes. The assignment of fossils to extant clades has traditionally relied on morphological similarity and on apomorphies shared with extant taxa. The use of explicit phylogenetic analyses to establish fossil affinities has so far remained limited. In this study, we built a comprehensive framework to investigate the phylogenetic placement of 24 exceptionally preserved fossil flowers. For this, we assembled a new species-level dataset of 30 floral traits for 1,201 extant species that were sampled to capture the stem and crown nodes of all angiosperm families. We explored multiple analytical approaches to integrate the fossils into the phylogeny, including different phylogenetic estimation methods, topological-constrained analyses, and combining molecular and morphological data of extant and fossil species. Our results were widely consistent across approaches and showed minor differences in the support of fossils at different phylogenetic positions. The placement of some fossils agrees with previously suggested relationships, but for others, a new placement is indicated. We also identified fossils that are well supported within particular extant families, whereas others showed high phylogenetic uncertainty. Finally, we present recommendations for future analyses combining molecular and morphological evidence, regarding the selection of fossils and appropriate methodologies, and provide some perspectives on how to integrate fossils into the investigation of divergence times and the temporal evolution of morphological traits.</span></p>
Data from: A single evolutionary innovation drives the deep evolution of symbiotic N2-fixation in angiosperms
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Data from: Another look at the root of the angiosperms reveals a familiar tale
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Data from: The relationship of recombination rate, genome structure, and patterns of molecular evolution across angiosperms
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Data from: Positive selection drives adaptive diversification of the 4-coumarate: CoA ligase (4CL) gene in angiosperms
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Data from: Evaluating the impact of genomic data and priors on Bayesian estimates of the angiosperm evolutionary timescale
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Data from: Non-equilibrium dynamics and floral trait interactions shape extant angiosperm diversity
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Data from: Mating systems and selection efficacy: a test using chloroplastic sequence data in Angiosperms
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Data from: Heterogeneous rates of molecular evolution and diversification could explain the Triassic age estimate for angiosperms
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Data from: Inferring complex phylogenies using parsimony: an empirical approach using three large DNA data sets for angiosperms
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Data from: Phylogeny of Saxifragales (angiosperms, eudicots): analysis of a rapid, ancient radiation
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Data from: Dioecy does not consistently accelerate or slow lineage diversification across multiple genera of angiosperms
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