Barremian Angiosperm Pollen and Associated Palynomorphs from the Dakhla Oasis Area, Egypt

Pollen grains characteristic of the local pre-tricolpate, pre-Aptian phase of angiosperm evolution have been recovered from the upper part of the Six Hills Formation in the Dakhla Oasis area, Egypt. Highest abundance (up to 16.5 per cent) and diversity of angiosperm pollen is attained in samples from the Tineida 2 borehole, which also display a remarkable variety of different aperture types for a palynoflora of late Barremian age. Monosulcate, columellate tectates such as Retimonocolpites are most important, but a single trichotomosulcate grain as well as monoporate and inaperturate types that have not been reported from many coeval palynofloras, are also present. The following new taxa are described using same-grain SEM/LM techniques: Tucanopollis annulatus Schrank, sp. nov., Retimonocolpites pennyi Schrank and Mahmoud, sp. nov. and Retiacolpites columellatus Schrank, gen. et sp. nov. Phytogeographic links are closest with palynofloras previously described from the late Barremian of West Africa (Gabon, Congo), that is within the Northern Gondwana or Dicheiropollis etruscus/Afropollis Province.     

Profile of a flower: How rates of morphological evolution drive floral diversification in Ericales and angiosperms

Premise

Recent studies of floral disparity in the asterid order Ericales have shown that flowers vary strongly among families and that disparity is unequally distributed between the three flower modules (perianth, androecium, gynoecium). However, it remains unknown whether these patterns are driven by heterogeneous rates of morphological evolution or other factors.

Methods

Here, we compiled a data set of 33 floral characters scored for 414 species of Ericales sampled from 346 genera and all 22 families. We conducted ancestral state reconstructions using an equal-rates Markov model for each character. We estimated rates of morphological evolution for Ericales and for a separate angiosperm-wide data set of 19 characters and 792 species, creating “rate profiles” for Ericales, angiosperms, and major angiosperm subclades. We compared morphological rates among flower modules within each data set separately and between data sets, and we compared rates among angiosperm subclades using the angiosperm data set.

Results

The androecium exhibits the highest evolutionary rates across most characters, whereas most perianth and gynoecium characters evolve more slowly in both Ericales and angiosperms. Both high and low rates of morphological evolution can result in high floral disparity in Ericales. Analyses of an angiosperm-wide floral data set reveal that this pattern appears to be conserved across most major angiosperm clades.

Conclusions

Elevated rates of morphological evolution in the androecium of Ericales may explain the higher disparity reported for this floral module. Comparing rates of morphological evolution through rate profiles proves to be a powerful tool in understanding floral evolution.     

Sterols and fatty acids of some non-photosynthetic angiosperms

Sterols and fatty acids were extracted and identified from three parasitic angiosperms, Cuscuta campestris, Monotropa uniflora and M. hypopitys. Each plant contained the typical 16 and 18-carbon fatty acids of angiosperms, but the partially-photosynthetic Cuscuta contained much larger quantities of linolenic acid than the non-green Monotropa species which had smaller amounts of linolenic acid characteristic of non-photosynthetic tissue. Sterol quantity was three times higher in Cuscuta than in the Monotropa species. Sitosterol was the major sterol in all species with smaller amounts of campesterol and cholesterol.     

Carpels in Brasenia (Cabombaceae) are completely ascidiate despite a long stigmatic crest

BACKGROUND AND AIMS: The morphological structure of anthetic carpels of Brasenia (Cabombaceae), a member of the phylogenetically basal ANITA grade, has not been studied before. The carpel has a long stigmatic crest on the ventral side and could give the impression of a conduplicate structure. This is in contrast to the carpel structure in other genera of the ANITA grade. Therefore, a study of carpel development and carpel structure at anthesis was carried out. METHODS: Carpels of Brasenia schreberi were studied at different developmental stages up to anthesis by means of microtome section series and SEM to analyse and reconstruct the outer and inner carpel morphology. KEY RESULTS: Carpels of Brasenia are extremely ascidiate up to anthesis. The elongate stigma originates around the mouth of the young carpel, which is slightly curved toward the centre of the flower. Subsequently, the stigmatic zone below the mouth expands by massive intercalary elongation. CONCLUSIONS: In their ascidiate shape, carpels of Brasenia are similar to carpels of Cabomba, the other genus of Cabombaceae, which, in contrast, has a short stigma restricted to the tip of the carpel. Thus, the morphological structure is independent of the extent (and one-sidedness) of the stigma. The outer shape of carpels at anthesis does not allow the inference of the inner morphological surface. If an angiosperm carpel has a one-sided stigma it can be extremely conduplicate or extremely ascidiate. Therefore, caution has to be used in the interpretation of the structure of fossil carpels.     

Analysis of Acorus calamus chloroplast genome and its phylogenetic implications

Determining the phylogenetic relationships among the major lines of angiosperms is a long-standing problem, yet the uncertainty as to the phylogenetic affinity of these lines persists. While a number of studies have suggested that the ANITA (Amborella-Nymphaeales-Illiciales-Trimeniales-Aristolochiales) grade is basal within angiosperms, studies of complete chloroplast genome sequences also suggested an alternative tree, wherein the line leading to the grasses branches first among the angiosperms. To improve taxon sampling in the existing chloroplast genome data, we sequenced the chloroplast genome of the monocot Acorus calamus. We generated a concatenated alignment (89,436 positions for 15 taxa), encompassing almost all sequences usable for phylogeny reconstruction within spermatophytes. The data still contain support for both the ANITA-basal and grasses-basal hypotheses. Using simulations we can show that were the ANITA-basal hypothesis true, parsimony (and distance-based methods with many models) would be expected to fail to recover it. The self-evident explanation for this failure appears to be a long-branch attraction (LBA) between the clade of grasses and the out-group. However, this LBA cannot explain the discrepancies observed between tree topology recovered using the maximum likelihood (ML) method and the topologies recovered using the parsimony and distance-based methods when grasses are deleted. Furthermore, the fact that neither maximum parsimony nor distance methods consistently recover the ML tree, when according to the simulations they would be expected to, when the out-group (Pinus) is deleted, suggests that either the generating tree is not correct or the best symmetric model is misspecified (or both). We demonstrate that the tree recovered under ML is extremely sensitive to model specification and that the best symmetric model is misspecified. Hence, we remain agnostic regarding phylogenetic relationships among basal angiosperm lineages.     

Stomatal patterning in angiosperms

My thesis is that understanding stomatal patterning requires a holistic perspective. Since stomata are structures critical to the survival of terrestrial plants, they need to be viewed in relation to their function and their interface with other structural components. With this outlook, I begin by discussing pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views. I suggest areas where I believe profitable studies might enable us to better understand stomatal patterning. The final sections of the paper review stomatal patterning on angiosperm leaves and present a theory of patterning. With the abundance of molecular information, and coming genomic sequences and new tools, an opportunity exists to dissect the process of how cells are selected to become different from their neighbors and assume a fate critical to plant survival. Understanding this biological process at the molecular level requires comprehending the broad base on which stomatal patterning rests.     

Phylogenetic trait conservation in the partner choice of a group of ectomycorrhizal trees

Ecological interactions are frequently conserved across evolutionary time. In the case of mutualisms, these conserved interactions may play a large role in structuring mutualist communities. We hypothesized that phylogenetic trait conservation could play a key role in determining patterns of association in the ectomycorrhizal symbiosis, a globally important trophic mutualism. We used the association between members of the pantropical plant tribe Pisonieae and its fungal mutualist partners as a model system to test the prediction that Pisonieae‐associating ectomycorrhizal fungi will be more closely related than expected by chance, reflecting a conserved trait. We tested this prediction using previously published and newly generated sequences in a Bayesian framework incorporating phylogenetic uncertainty. We report that phylogenetic trait conservation does exist in this association. We generated a five‐marker phylogeny of members of the Pisonieae and used this phylogeny in a Bayesian relaxed molecular clock analysis. We established that the most recent common ancestors of Pisonieae species and Pisonieae‐associating fungi sharing phylogenetic conservation of their patterns of ectomycorrhizal association occurred no more recently than 14.2 Ma. We therefore suggest that phylogenetic trait conservation in the Pisonieae ectomycorrhizal mutualism association represents an inherited syndrome which has existed for at least 14 Myr.     

Leaf energy balance modelling as a tool to infer habitat preference in the early angiosperms

Despite more than a century of research, some key aspects of habitat preference and ecology of the earliest angiosperms remain poorly constrained. Proposed growth ecology has varied from opportunistic weedy species growing in full sun to slow-growing species limited to the shaded understorey of gymnosperm forests. Evidence suggests that the earliest angiosperms possessed low transpiration rates: gas exchange rates for extant basal angiosperms are low, as are the reconstructed gas exchange rates for the oldest known angiosperm leaf fossils. Leaves with low transpirational capacity are vulnerable to overheating in full sun, favouring the hypothesis that early angiosperms were limited to the shaded understorey. Here, modelled leaf temperatures are used to examine the thermal tolerance of some of the earliest angiosperms. Our results indicate that small leaf size could have mitigated the low transpirational cooling capacity of many early angiosperms, enabling many species to survive in full sun. We propose that during the earliest phases of the angiosperm leaf record, angiosperms may not have been limited to the understorey, and that some species were able to compete with ferns and gymnosperms in both shaded and sunny habitats, especially in the absence of competition from more rapidly growing and transpiring advanced lineages of angiosperms.     

A synthesis of radial growth patterns preceding tree mortality

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan‐continental tree‐ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1–100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long‐lasting declines were found for gymnosperms, shade‐ and drought‐tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark‐beetles) typically showed relatively small and short‐term growth reductions. Our analysis did not highlight any universal trade‐off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark‐beetle attack, while long‐term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth‐based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark‐beetle outbreaks.