Cretaceous diversification of angiosperms in the western part of the Iberian Peninsula

The classic leaf fossil floras from the Cretaceous of the Lusitanian Basin, Portugal, which were first described more than one hundred years ago, have played an important role in the development of ideas on the early evolution of angiosperms. Insights into the nature of vegetational change in the Lusitanian Basin through the Cretaceous have also come from studies of fossil pollen and spores, but the discovery of a series of mesofossil floras containing well-preserved angiosperm reproductive structures has provided a new basis for understanding the systematic relationships and biology of angiosperms at several stratigraphic levels through the Cretaceous. In the earliest mesofossil floras from the Torres Vedras locality, which are of probable Late Barremian-Early Aptian age, angiosperms are surprisingly diverse with about 50 different taxa. In slightly later mesofossil floras, which are of probable Late Aptian-Early Albian age, the diversity of angiosperms is still more substantial with more than hundred different kinds of angiosperm reproductive structures recognized from the Famalicão locality alone. However, this early diversity is largely among angiosperm lineages that produced monoaperturate pollen (e.g., Chloranthaceae, Nymphaeales) and early diverging monocots (Alismatales). Eudicots are rare in these Early Cretaceous mesofossil floras, but already by the Late Cenomanian the vegetation of the western Iberian Peninsula is dominated by angiosperms belonging to various groups of core eudicots. The Normapolles complex is a particularly conspicuous element in both mesofossil floras and in palynological assemblages. In the Late Cretaceous mesofossil floras from Esgueira and Mira, which are of Campanian-Maastrichtian age, core eudicots are also floristically dominant and flowers show great organisational similarity to fossil flowers from other Late Cretaceous floras described from other localities in Asia, Europe and North America.     

Preliminary palaeobotanical notes on the Early Cretaceous Cintura Formation (Sonora,Mexico)

The Cintura Formation of Albian–Cenomanian age in the Cabullona Basin yielded an important fossil flora, including palynomorphs, leaf impressions and fossil trunks. At the base of the Marquechi Member a poorly preserved palynomorph assemblage is recognised. The palynological assemblage is dominated by gymnosperms, mainly by cheirolepidiacean genus Classopollis. A noteworthy feature is the presence of angiosperm pollen grains of the genera Clavatipollenites, Retimonocolpites and Tucanopollis. A rich, diverse and well-preserved macroflora of leaf impressions has been observed in the uppermost part of this formation belonging to the San Juan Member. There are at least eight morphospecies of leaves including an abundance of taxa with possible botanical affinities to the family Sapindaceae. The angiosperm pollen grains together with the sapindalean leaves constitute the oldest record of such remains in Mexico.     

Fossil pollen records of extant angiosperms

The fossil record for angiosperm pollen types which are comparable to recent taxa is evaluated, following a similar survey published in 1970. Special attention is paid to the dating of the sediments. Evidence for 139 families is considered to be reliable, for others the records are cited as provisional, pending the accumulation of more evidence. Some published records are shown to be erroneous. In the early Cretaceous only types occur indicating the presence of plants ancestral to Magnoliidae and Liliatae. In the Turonian increased differentiation is evident, culminating in the Maestrichtian when evidence for the presence of most major taxa of angiosperms is available. In the Tertiary diversification at lower taxonomic level continues and identification with living taxa becomes more certain. Herbaceous groups tend to appear later in the record than woody ones.     

The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence

Gymnosperms, and conifers in particular, are sometimes very productive trees yet angiosperms dominate most temperate and tropical vegetation. Current explanations for angiosperm success emphasize the advantages of insect pollination and seed dispersal by animals for the colonization of isolated habitats. Differences between gymnosperm and angiosperm reproductive and vegetative growth rates have been largely ignored. Gymnosperms are all woody, perennial and usually have long reproductive cycles. Their leaves are not as fully vascularized as those of angiosperms and are more stereotyped in shape and size. Gymnosperm tracheids are generally more resistant to solute flow than angiosperm vessels. A consequence of the less efficient transport system is that maximum growth rates of gymnosperms are lower than maximum growth rates of angiosperms in well lit, well watered habitats. Gymnosperm seedlings may be particularly uncompetitive since their growth depends on a single cohort of relatively inefficient leaves. Later, some gymnosperms attain a higher productivity than co-occurring angiosperm trees by accumulating several cohorts of leaves with a higher total leaf area. These functional constraints on gymnosperm growth rates suggest that gymnosperms will be restricted to areas where growth of angiosperm competitors is reduced, for example, by cold or nutrient shortages. Biogeographic evidence supports this prediction since conifers are largely confined to high latitudes and elevations or nutrient-poor soils. Experimental studies show that competition in the regeneration niche (between conifer seedlings and angiosperm herbs and shrubs) is common and significantly affects conifer growth and survival, Fast-growing angiosperms, especially herbs and shrubs, may also change the frequency of disturbance regimes thereby excluding slower-growing gymnosperms. Shade-tolerant and early successional conifers share similar characteristics of slow initial growth and low plasticity to a change in resources. Shade-tolerant gymnosperms would be expected to occur only where forest openings are small or otherwise unsuitable for rapid filling by fast-growing angiosperm trees, lianas or shrubs. The limited evidence available suggests that shade-tolerant conifers are confined to forests with small gap sizes where large disturbances are very rare. The regeneration hypothesis for gymnosperm exclusion by angiosperms is consistent with several aspects of the fossil record such as the early disappearance of gymnosperms from early successional environments where competition with angiosperms would have been most severe. However there are unresolved difficulties in interpreting process from paleoecological pattern which prevent the testing of alternative hypotheses.     

The evolution of embryo size in angiosperms and other seed plants: implications for the evolution of seed dormancy

Abstract.— Seed dormancy plays an important role in germination ecology and seed plant evolution. Morphological seed dormancy is caused by an underdeveloped embryo that must mature prior to germination. It has been suggested that the presence of an underdeveloped embryo is plesiomorphic among seed plants and that parallel directional change in embryo morphology has occurred separately in gymnosperms and in angiosperms. We test these hypotheses using original data on embryo morphology of key basal taxa, a published dataset, and the generalized least squares (GLS) method of ancestral character state reconstruction. Reconstructions for embryo to seed ratio (E:S) using family means for 179 families showed that E:S has increased between the ancestral angiosperm and almost all extant angiosperm taxa. Species in the rosid clade have particularly large embryos relative to the angiosperm ancestor. Results for the gymnosperms show a similar but smaller increase. There were no statistically significant differences in E:S between basal taxa and any derived group due to extremely large standard errors produced by GLS models. However, differences between reconstructed values for the angiosperm ancestor and more highly nested nodes are large and these results are robust to topological and branch-length manipulations. Our analysis supports the idea that the underdeveloped embryo is primitive among seed plants and that there has been a directional change in E:S within both angiosperms and gymnosperms. Our analysis suggests that dormancy enforced by an underdeveloped embryo is plesiomorphic among angiosperms and that nondormancy and other dormancy types probably evolved within the angiosperms. The shift in E:S was likely a heterochronic change, and has important implications for the life history of seed plants.     

Characterization of the expression patterns of LEAFY/FLORICAULA and NEEDLY orthologs in female and male cones of the conifer genera Picea, Podocarpus, and Taxus: implications for current evo-devo hypotheses for gymnosperms

The identity of genes causally implicated in the development and evolutionary origin of reproductive characters in gymnosperms is largely unknown. Working within the framework of plant evolutionary developmental biology, here we have cloned, sequenced, performed phylogenetic analyses upon and tested the expression patterns of LEAFY/FLORICAULA and NEEDLY orthologs in reproductive structures from selected species of the conifer genera Picea, Podocarpus, and Taxus. Contrary to expectations based on previous assessments, expression of LFY/FLO and NLY in cones of these taxa was found to occur simultaneously in a single reproductive axis, initially overlapping but later in mutually exclusive primordia and/or groups of developing cells in both female and male structures. These observations directly affect the status of the "mostly male theory" for the origin of the angiosperm flower. On the other hand, comparative spatiotemporal patterns of the expression of these genes suggest a complex genetic regulatory network of cone development, as well as a scheme of functional divergence for LFY/FLO with respect to NLY homologs in gymnosperms, both with clear heterochronic aspects. Results presented in this study contribute to the understanding of the molecular-genetic basis of morphological evolution in conifer cones, and may aid in establishing a foundation for gymnosperm-specific, testable evo-devo hypotheses.     

MADS-box genes active in developing pollen cones of Norway spruce (Picea abies) are homologous to the B-class floral homeotic genes in angiosperms.

The reproductive organs of conifers, the pollen cones and seed cones, differ in morphology from the angiosperm flower in several fundamental respects. In this report we present evidence to suggest that the two plant groups, in spite of these morphological differences and the long evolutionary distance between them, share important features in regulating the development of the reproductive organs. We present the cloning of three genes, DAL11, DAL12, and DAL13, from Norway spruce, all of which are related to the angiosperm B-class of homeotic genes. The B-class genes determine the identities of petals and stamens. They are members of a family of MADS-box genes, which also includes C-class genes that act to determine the identity of carpels and, in concert with B genes specify stamens in the angiosperm flower. Phylogenetic analyses and the presence of B-class specific C-terminal motifs in the DAL protein sequences imply homology to the B-class genes. Specific expression of all three genes in developing pollen cones suggests that the genes are involved in one aspect of B function, the regulation of development of the pollen-bearing organs. The different temporal and spatial expression patterns of the three DAL genes in the developing pollen cones indicate that the genes have attained at least in part distinct functions. The DAL11, DAL12, and 13 expression patterns in the pollen cone partly overlap with that of the previously identified DAL2 gene, which is structurally and functionally related to the angiosperm C-class genes. This result supports the hypothesis that an interaction between B- and C-type genes is required for male organ development in conifers like in the angiosperms. Taken together, our data suggests that central components in the regulatory mechanisms for reproductive organ development are conserved between conifers and angiosperms and, thus, among all seed plants.     

Occurrence of endodermis with a casparian strip in stem and leaf

It is well known that an endodermis with casparian strip always occurs in roots, but few people are aware that it also occurs in stems and leaves of some vascular plants. The rather sparse literature on endodermis in aerial organs was last included in a review in 1943. The present compilation, which does not consider hydathodes, nectaries, or other secretory structures, emphasizes distribution of cauline and foliar endodermis with casparian strip. It occurs unevenly among major taxa: quite common in rhizomes and leaves among pteridophyte groups, with exceptions; absent in gymnosperm stems but found in leaves at least among some conifers; in stems of at least 30 mostly herbaceous angiosperm families, but far less common in leaves, where it is mostly reported from petioles. Etiolation can induce casparian strips in stems and petioles of some herbaceous plants, but results from leaf blades are questionable. There are recent reports of an endodermis with casparian strip in leaves of both woody and herbaceous taxa. The physiological function, if any, of a casparian strip in aerial organs remains unknown.     

Diversification of myco-heterotrophic angiosperms: Evidence from Burmanniaceae

Background  

Myco-heterotrophy evolved independently several times during angiosperm evolution. Although many species of myco-heterotrophic plants are highly endemic and long-distance dispersal seems unlikely, some genera are widely dispersed and have pantropical distributions, often with large disjunctions. Traditionally this has been interpreted as evidence for an old age of these taxa. However, due to their scarcity and highly reduced plastid genomes our understanding about the evolutionary histories of the angiosperm myco-heterotrophic groups is poor.     

Fossil evidence and phylogeny: the age of major angiosperm clades based on mesofossil and macrofossil evidence from Cretaceous deposits

The fossil record has played an important role in the history of evolutionary thought, has aided the determination of key relationships through mosaics, and has allowed an assessment of a number of ecological hypotheses. Nonetheless, expectations that it might accurately and precisely mirror the progression of taxa through time seem optimistic in light of the many factors potentially interfering with uniform preservation. In view of these limitations, attempts to use the fossil record to corroborate phylogenetic hypotheses based on extensive comparisons among extant taxa may be misplaced. Instead we suggest a method-minimum age node mapping-for combining reliable fossil evidence with hypotheses of phylogeny. We use this methodology in conjunction with a phylogeny for angiosperms to assess timing in the history of major angiosperm clades. This method places many clades both with and without fossil records in temporal perspective, reveals discrepancies among clades in propensities for preservation, and raises some interesting questions about angiosperm evolution. By providing a context for understanding the gaps in the angiosperm fossil record this technique lends credibility and support to the remainder of the angiosperm record and to its applications in understanding a variety of aspects of angiosperm history. In effect, this methodology empowers the fossil record.     

Timing in the evolution of derived floral characters: upper cretaceous (turonian) taxa with tricolpate and tricolpate-derived pollen

Various hypotheses that seek to explain the rich species diversity of angiosperms relative to other seed plants are briefly mentioned or reviewed. Of these, the subset that relates angiosperm diversity in some way to the relationship between angiosperms and insects, particularly anthophilous insects, is here the object of attention. Specifically, I address and reject the possibility that the relationship between angiosperm diversification and insects, particularly those demonstrating a preference for flowers with derived floral characteristics associated with insect pollination, may be ruled out because of asynchronous patterns of diversification in the fossil record. New data on floral structure from the Turonian of the Atlantic Coastal Plain reveal a surprising diversity of floral characters in taxa bearing tricolpate and tricolporate-derived pollen. The characters and taxa that appear in these Turonian sediments suggest that rather specific modes of insect pollination, perhaps involving highly derived insect pollinators, already existed at 90 Ma. Given the observed rate of diversification of angiosperms during that time and the pattern of evolution in insects, including what can be inferred about the history of the Apidae, these new floral data suggest that hypotheses relating angiosperm diversity to highly specific pollinators are still valid in the context of fossil evidence. Even so, consistency with fossil evidence is not necessarily proof of these relationships. In any case, there may well be multiple causes of relatively high angiosperm species diversity and understanding the relative importance of each of these requires neontological as well as paleontological investigations. One promising approach is to work within the context of phylogenetic patterns with more fossil data.     

Sexing pollen reveals female bias in a dioecious plant

* Information on angiosperm sex ratios has largely been restricted to surveys of flowering individuals. These often deviate from equality, with male bias more commonly reported. Female-biased sex ratios are concentrated in a few taxa and have been linked to the possession of heteromorphic sex chromosomes and bias introduced during the gametophytic stage of the life cycle. It has been proposed that differences in gamete quantity and quality could give rise to female bias, although there is no direct evidence with which to evaluate this possibility. * Here, we use flow cytometry to investigate microgametophytic 'sex ratios' in a flowering plant. We demonstrate that differences in DNA content between the sexes in Rumex nivalis, a species with heteromorphic sex chromosomes, make it possible to distinguish female- vs male-determining pollen nuclei. * We found a small but significant female bias in microgametophytes produced by males (mean 0.515) with significant variation among family means (range 0.463-0.586), and 18 of 22 families averaging > 0.50. * The observed female bias at the gametophytic stage of the life cycle is consistent with the direction of bias previously reported for seeds and vegetative and reproductive plants in wild populations of R. nivalis, but is insufficient to fully explain the degree of bias.     

SEM analysis of leaf epicuticular waxes of Sedum section Gormania (Crassulaceae)

Epicuticular leaf waxes in the 12 taxa classified inSedum, sect.Gormania show variation in size, shape, and frequency. All of the taxa exhibit wax platelets, but the platelets in the three putative relicts (S. albomarginatum, S. moranii, S. oblanceolatum) are larger than those in related taxa, and those inS. paradisum are poorly developed. Interspecific hybrids tend to show wax patterns intermediate between those of their respective parents. Hypothetical hybrid origins for the four polyploid taxa in the section have been supported previously by morphological and reproductive data and are further clarified by leaf wax data.     

Three loblolly pine CesA genes expressed in developing xylem are orthologous to secondary cell wall CesA genes of angiosperms

Specific plant cellulose synthases (CesA), encoded by a multigene family, are necessary for secondary wall synthesis in vascular tissues and are critical to wood production. We obtained full-length clones for the three CesAs that are highly expressed in developing xylem and examined their phylogenetic relationships and expression patterns in loblolly pine tissues. Full-length CesA clones were isolated from cDNA of developing loblolly pine (Pinus taeda) xylem and phylogenetic inferences made from plant CesA protein sequences. Expression of the three genes was examined by Northern blot analysis and semiquantitative RT-PCR. Each of three PtCesA genes is orthologous to one of the three angiosperm secondary cell wall CesAs. The PtCesAs are coexpressed in tissues of loblolly pine with tissues undergoing secondary cell wall biosynthesis showing the highest levels of expression. Phylogenetic and expression analyses suggest that functional roles for these loblolly pine CesAs are analogous to those of orthologs in angiosperm taxa. Based upon evidence from this and other studies, we suggest division of seed plant CesA genes into six major paralogous groups, each containing orthologs from various taxa. Available evidence suggests that paralogous CesA genes and their distinct functional roles evolved before the divergence of gymnosperm and angiosperm lineages.     

Recurrent violations of invariant rules for offspring size: evidence from turtles and the implications for small clutch size models

Smith and Fretwell’s classic model predicts that parents can maximize fitness by dividing the energy available for reproduction into offspring of an optimal size. However, this model breaks down when clutch size is small (~1–10 offspring). Invariant rules are an extension of the Smith–Fretwell model, and these rules predict how offspring size will vary among and within individuals that produce small clutch sizes. Here, we provide a narrow test of invariant rules using three turtle species, then we synthesize and re-analyze existing data from 18 different species (comprising five Orders) to evaluate whether invariant rules are followed across broad taxa. We do not find support for most invariant rules in turtles, and our re-analysis demonstrates a general mismatch between observed and expected values across all taxa evaluated, suggesting that invariant rules fail to predict reproductive patterns in nature. Morphological constraints on offspring size and reproductive effort may be important reasons for disparities between theory and observation both in turtles and other taxa. Paradoxically, morphological constraints are most common in small-bodied species and individuals, but these same candidates are also those which produce the small clutch sizes that are necessary to test invariant rules, such that a fair test of invariant rules will often be challenging. Mismatches between theory and observation might also occur because theory assumes that mothers exert control over resource allocation to offspring. In fact, there is evidence of widespread genetic correlations among investment per offspring and reproductive effort, such that these traits are not independent.     

果实类型多样性的形成机制和进化

果实是被子植物特有的繁殖器官,果实的类型影响种子传播的模式、有效性和距离。果实类型的多样性在一定程度上造成了不同植物类群在生态上的差异,并对被子植物的进化速度和分化模式产生重要影响。本文对果实类型多样性形成中起关键作用的通路及其进化机制进行了综述,讨论了影响果实类型进化的各种因素,介绍了有关果实类型进化方向和模式的研究进展,为研究被子植物果实多样性的形成和演化提供新的思路。     

Is the anthophyte hypothesis alive and well? New evidence from the reproductive structures of Bennettitales

Bennettitales is an extinct group of seed plants with reproductive structures that are similar in some respects to both Gnetales and angiosperms, but systematic relationships among the three clades remain controversial. This study summarizes characters of bennettitalean plants and presents new evidence for the structure of cones and seeds that help clarify relationships of Bennettitales to flowering plants, Gnetales, and other potential angiosperm sister groups. Bennettitales have simple mono- or bisporangiate cones. Seeds are borne terminally on sporophylls. They have a unique structure that includes a nucellus with a solid apex, no pollen chamber, and a single integument, and they are clearly not enclosed by a cupule or other specialized structures. Such features differ substantially from Gnetales, flowering plants, and the seed fern Caytonia, providing no compelling evidence for the origin of the angiospermous carpel. Cladistic tests were performed to assess the strength of the "anthophyte hypothesis" and possible relationships of Bennettitales, Gnetales, and Caytonia to flowering plants. Our results do not support the anthophyte hypothesis for the origin of angiosperms by a transformation of fertile organs that were already aggregated into a cone or flower-like structure. However, the anthophyte topology of the seed plant tree continues to be supported by morphological analyses of living and extinct taxa.     

Evolution of a unique anatomical precision in angiosperm leaf venation lifts constraints on vascular plant ecology

The main role of leaf venation is to supply water across the photosynthetic surface to keep stomata open and allow access to atmospheric CO₂ despite evaporative demand. The optimal uniform delivery of water occurs when the distance between veins equals the depth of vein placement within the leaf away from the evaporative surface. As presented here, only angiosperms maintain this anatomical optimum across all leaf thicknesses and different habitats, including sheltered environments where this optimization need not be required. Intriguingly, basal angiosperm lineages tend to be underinvested hydraulically; uniformly high optimization is derived independently in the magnoliids, monocots and core eudicots. Gymnosperms and ferns, including available fossils, are limited by their inability to produce high vein densities. The common association of ferns with shaded humid environments may, in part, be a direct evolutionary consequence of their inability to produce hydraulically optimized leaves. Some gymnosperms do approach optimal vein placement, but only by virtue of their ability to produce thick leaves most appropriate in environments requiring water conservation. Thus, this simple anatomical metric presents an important perspective on the evolution and phylogenetic distribution of plant ecologies and further evidence that the vegetative biology of flowering plants—not just their reproductive biology—is unique.     

Testing reticulate evolution of four Vitis species from East Asia using restriction‐site associated DNA sequencing

Reticulate evolution is an important driving force of angiosperm evolution. It has been proposed as an important evolutionary process in Vitis L. subgenus Vitis. Events of natural hybridization and introgression of several taxa native to North America have been hypothesized and discussed. However, there is no convincing evidence of reticulate evolution reported for closely related Vitis taxa from East Asia. We aim to explore natural hybridization and introgression among four closely related Vitis taxa from East Asia (V. amurensis Ruprecht, V. romanetii Romanet du Caillaud, V. shenxiensis C. L. Li, and V. piasezkii Maximowicz) with the restriction‐site associated DNA sequencing technique. A total of 46 accessions, covering the potential morphological and geographic variation of each species, are sequenced. Our results show a complex evolutionary pattern of the four Vitis species. The phylogenetic inference suggests that V. amurensis is monophyletic, however, V. romanetii, V. shenxiensis, and V. piasezkii do not appear to be monophyletic. Significant signals of introgression in some accessions have been detected by population structure analyses. D‐statistics analysis and population structure analyses support the presence of introgression between V. shenxiensis/V. piasezkii and V. romanetii in sympatric populations, but a strong signal of admixture has not been recognized between distantly located populations. Our results provide strong evidence of reticulate evolution among V. romanetii, V. shenxiensis, and V. piasezkii.     

The biology of rarity: Patterns, causes and consequences

There is a growing body of evidence suggesting that locally rare and geographically restricted species may have characteristics that differ from those of taxa that are more common. Several studies show that rare taxa have lower levels of self-incompatibility, a tendency toward asexual reproductive pathways, lower overall reproductive effort and poorer dispersal abilities. There are several mechanisms that could be responsible for such differences, but they may in practice be difficult to differentiate. Nonetheless, the documentation of recurrent rare-common differences is of vital importance because it may allow us to compensate partially for the bias of the published literature toward studies of common taxa.