Global patterns and ecological drivers of taxonomic and phylogenetic endemism in angiosperm genera

Endemism of lineages lies at the core of understanding variation in community composition among geographic regions because it reflects how speciation,extinction,and dispersal have influenced current distributions.Here,we investigated geographic patterns and ecological drivers of taxonomic and phylogenetic endemism of angiosperm genera across the world.We identify centers of paleo-endemism and neo-endemism of angiosperm genera,and show that they are mostly located in the Southern Hemisphere in tropical and subtropical regions,particularly in Asia and Australia.Different categories of phylogenetic endemism centers can be differentiated using current climate conditions.Current climate,historical climate change,and geographic variables together explained~80%of global variation in taxonomic and phylogenetic endemism,while 42-46%,1%,and 15%were independently explained by these three types of variables,respectively.Thus our findings show that past climate change,current climate,and geography act together in shaping endemism,which are consistent with the findings of previous studies that higher temperature and topographic heterogeneity promote endemism.Our study showed that many centers of phylogenetic endemism of angiosperms,including regions in Amazonia,Venezuela,and west-central tropical Africa that have not previously been identified as biodiversity hotspots,are missed by taxon-based measures of endemism,indicating the importance of including evolutionary history in biodiversity assessment.

The Protective Role of Xanthophyll Cycle in Resurrection Angiosperm Boea hygrometrica During Dehydration and Rehydration

The protective role of xanthophyll cycle in resurrection angiosperm Boea hygrometrica (Bunge) R.Br. was investigated by analysis of the changes of chlorophyll fluorescence and xanthophyll cycle components in response to dehydration and rehydration in detached leaves under very weak light condition (3 mumol photons.m(-2).s(-1)) and in the dark. With declines in the values of PSII photochemical efficiency (Fv/Fm), PSII actual quantum yield (Phi(PSII)), photochemical quenching (qP) and non-photochemical quenching (NPQ) during dehydration, zeaxanthin significantly increased in control Boea leaves under very weak light condition, while no zeaxanthin accumulation was detected in Boea leaves treated with dithiothreitol (DTT) and Boea leaves in the dark, and after 3 d rehydration, the parameters Fv/Fm, Phi(PSII), qP and NPQ showed full recovery in control Boea leaves under very weak light condition, but the parameters only underwent partial recovery in Boea leaves treated with DTT and Boea leaves in the dark, suggesting that the recovery of photosystem II (PSII) photochemical activities in Boea leaves was obviously affected by treatments with DTT and darkness, therefore, zeaxanthin may play an important protective role in desiccated Boea leaves even under very weak light conditions.

Photosynthesis of Resurrection Angiosperms

Resurrection plants which are able to quickly reactivate after falling into a period of anabiosis caused by dehydration have been very rare among angiosperms, especially among dicotyledons whose chlorophyll content and chloroplast structure little changed in the course of desiccation, therefore has been called homoiochlorophyllous desiccation-tolerant plants (HDTs). Another type of resurrection angiosperms that lost its chlorophyll dining desiccation is called poikilochlorophyllous desiccation-tolerant plants (PDTs). HDTs have been received more attention because of simplicity of protection mechanism which is much easy to the study and utilization of the desiccation tolerance of resurrection angiosperms. Recent advances in studies of photosynthesis of resurrection angiosperms indicate that photochemical activities are sensitive indicators for the study of physiological state of resurrection angiosperms during desiccation and rehydration. Photochemical activities of resurrection angiosperms are inhibited with loss of water similar to those of general plants, however, the magic thing is that they could reactivate rapidly during rehydration even losing more than 95% water. Up-regulations in xanthophyll cycle and antioxidative systems as well as preservation in integrity and stability of photosynthetic membranes during desiccation may be very important to desiccation tolerance of resurrection angiosperms. The fact that phosphate treatment in rehydration stage also strongly influences resurrection indicated importance of studies on rehydration stages of resurrection angiosperms.

In Vitro Fertilization of Angiosperms-10-Year Effort in China

This review gives a brief retrospect to the development on in vitro fertilization (IVF) of angiosperms in China. During the last decade Chinese scientists put great enthusiasm and efforts on IVF system construction and built up notable contributions to the flourish of this field. Keeping pace with international development and participating international cooperation in the field of IVF, Chinese scientists have now focused on the investigation of basic mechanism relevant to possible gamete interaction, egg cell activation and early embryogenesis by IVF. In vitro manipulation techniques are combined with cytological and molecular biological approaches to unveil the double fertilization mysteries.

Xingxueanthus: An Enigmatic Jurassic Seed Plant and Its Implications for the Origin of Angiospermy

The origin of angiosperms has been tantalizing botanists for centuries. Despite the efforts of palaeobotanists, most of the pre-Cretaceous angiosperms are regarded either non-convincing or misdated. The applications of SEM and LM （light microscope） enable us to recognize a coalified fossil plant, Xingxueanthus sinensis gen. et sp. nov., from the Haifanggou Formation （Middle Jurassic, 〉160 Ma） in western Liaoning, China. Xingxueanthus is an ＂inflorescence＂ with more than 20 female units spirally arranged. Each female unit is situated in the axii of a bract. The female unit is composed of an ovule-container and a style-like projection at the top. There is a vertical column bearing several ovules in the ovule-container. The general morphology and the internal structure of Xingxueanthus distinguish itself from any known fossil and extant gymnosperms, and its structures are more comparable to those of angiosperms. Xingxueanthus, if taken as a gymnosperm, would represent a new class, demonstrate an evolutionarily advanced status of ovule-protection in gymnosperms never seen before, and provide new insights into the origin of angiospermy. Alternatively, if taken as an angiosperm, together with Schmeissneria, it would increase the diversity of Jurassic angiosperms, which has been underestimated for a long time, and suggest a much earlier origin of angiospermy than currently accepted.

An Undercover Angiosperm from the Jurassic of China

Searching for early angiosperms is a riveting activity in botany because it helps to resolve the phyiogenetic relationships among seed plants and among angiosperms themselves. One of the challenges for this job is what the target fossils look like. Most possibly early angiosperms may elude our scrutiny with gymnospermous appearances. This possibility becomes a reality in a Jurassic plant, Solaranthus gen. nov, which bears a peltaspermalean appearance and enclosed ovules. According to knowledge available hitherto, the latter feature makes it an angiosperm. However, such a feature is more likely to be eclipsed by its gymnospermous appearance. The early age and unexpected character assemblage of Solaranthus urge for a fresh look on the assumed-simple relationship between angiosperms and gymnosperms. Its resemblance to the order Peltaspermales favors the Mostly Male Theory.

Late Cretaceous Aquatic Angiosperms from Jiayin,Heilongjiang,Northeast China

Three taxa of Late Cretaceous aquatic angiosperms, Queruexia angulata （Lesq.） Krysht., Cobbania corrugata （Lesq.） Stockey et al. and Nelumbites cf. extenuinervis Upchurch et al. from Jiayin of Heilongjiang, NE China, are described in detail. Among them, Cobbania and Nelumbites from the Upper Cretaceous in China are reported for the first time. The aquatic angiosperm assemblage of Queruexia-Cobbania-Nelumbites appears to imply a seasonal, warm and moist environment in the Jiayin area during the Santonian-Campanian time.

A Dichocarpum-like Angiosperm from the Early Cretaceous of China

The Yixian Formation（the Early Cretaceous） of China has yielded a diverse group of early angiosperms, revealing the charm of angiosperms from the formation is far from finishing. Here we report a fossil angiosperm, Nothodichocarpum gen. et sp. nov, with physically connected female parts, male parts, branches, and leaves with pinnate reticulate venation from the Yixian Formation of Liaoning, China. In addition to the previous reports, this report underscores the diversity of angiosperms in the Early Cretaceous Yixian Formation and earlier origin of angiosperms. The morphological variations in the reproductive parts in different stages allow us to infer the development of this early angiosperm. This fossil plant demonstrates a unique character assemblage that sheds new light on the origin and evolution of carpels.

Mid-Cretaceous Hothouse Climate and the Expansion of Early Angiosperms

The remarkable transition of early angiosperms from a small to a dominant group characterized the terrestrial ecosystem of the Cretaceous. This transition was instigated and promoted by environmental changes. Mid-Cretaceous is characterized by major geological events that affected the global environment. δ^18O, palaeothermometer TEX86, and other climatic indices from marine sediments suggest rapid temperature increase during mid-Cretaceous despite occasional short cooling events. Simultaneously, terrestrial deposits in East Asia changed from coal-bearing to shale, then to red beds and evaporites. Plant assemblages and other paleoclimate indicators point to rapid aridification for midCretaceous terrestrial environments. In addition, the wildfires were frequently spread all over the earth by the numerous charcoal evidence during the Mid-Cretaceous. Thus, we speculate that the seasonally dry and hot conditions of mid-Cretaceous created a fiery hothouse world. Early angiosperms increased in abundance and diversity and evolved from a few aquatic species to terrestrial herbaceous and then to the diversified flora of today. Angiosperms showed rapid physiological evolution in vein density and leaf area that improved photosynthesis and water absorption. These ecophysiological changes made early angiosperms well adapted to the hot and dry environment in mid-Cretaceous. Moreover, these physiological changes facilitated the fire–angiosperm cycles in mid-Cretaceous that likely further stimulated the early angiosperm evolution.

A Whole Plant Herbaceous Angiosperm from the Middle Jurassic of China

In contrast to woody habit with secondary growth, truthful herbaceous habit lacking secondary growth is restricted to angiosperms among seed plants. Although angiosperms might have occurred as early as in the Triassic and herbaceous habit theoretically may have been well adopted by pioneer angiosperms, pre-Cretaceous herbs are missing hitherto, leaving the origin of herbs and evolution of herbaceous angiosperms mysterious. Here we report Juraherba bodae gen. et sp. nov, a whole plant herbaceous angiosperm, from the Middle Jurassic （〉164 Ma） at Daohugou Village, Inner Mongolia, China, a fossil Lagerst^itten that is worldwide famous for various fossil finds. The angiospermous affinity of Juraherba is ensured by its enclosed ovules/seeds. The plant is small but complete, with physically connected hairy root, stem, leaves, and fructifications. The Middle Jurassic age recommends Juraherba as the earliest record of herbaceous seed plants, demanding a refresh look at the evolutionary history of angiosperms.

A Novel Angiosperm from the Early Cretaceous and Its Implications for Carpel-Deriving

The Yixian Formation （Lower Cretaceous） of China is famous for its fossils of early angiosperms, and these fossils have shed otherwise unavailable light on the evolution of angiosperms. The seeds in Archaefructus and Nothodichocarpum are inserted along the dorsal of the fruits while those in Sinocarpus are along the ventral of the fruits, suggesting different Bau-plans and pathways deriving the carpels in these plants of the Early Cretaceous. Adding further fossil evidence, here we report a novel fossil angiosperm, Neofructus lingyuanensis gen. et sp. nov, with its seeds inserted on both ventral and dorsal sides of the fruits documented through light microscopic and SEM observations, suggesting a novel way forming gynoecium in the Yixian Formation. These fossil angiosperms indicate that angiosperm gynoecia in the Yixian Formation （the Barremian to Aptian） are derived in obviously different ways, implying an evolutionary scenario for angiosperms quite different from formerly assumed.

Geographic patterns of taxonomic and phylogenetic β-diversity of angiosperm genera in regional floras across the world

Beta diversity(β-diversity)is the scalar between local(α)and regional(γ)diversity.Understanding geographic patterns ofβ-diversity is central to ecology,biogeography,and conservation biology.A full understanding of the origin and maintenance of geographic patterns ofβ-diversity requires exploring both taxonomic and phylogeneticβ-diversity,as well as their respective turnover and nestedness components,and exploring phylogenetic p-diversity at different evolutionary depths.In this study,we explore and map geographic patterns ofβ-diversity for angiosperm genera in regional floras across the world.We examine both taxonomic and phylogeneticβ-diversity and their constituent components,and both tip-weighted and basal-weighted phylogeneticβ-diversity,and relate them to latitude.On the one hand,our study found that the global distribution ofβ-diversity is highly heterogeneous.This is the case for both taxonomic and phylogeneticβ-diversity,and for both tip-weighted and basal-weighted phylogeneticβ-diversity.On the other hand,our study found that there are highly consistent geographic patterns among different metrics ofβ-diversity.In most cases,metrics ofβ-diversity are negatively associated with latitude,particularly in the Northern Hemisphere.Different metrics of taxonomicβ-diversity are strongly and positively correlated with their counterparts of phylogeneticβ-diversity.

A Biased, Misleading Review on Early Angiosperms

A recently published review by Herendeen?et al.?is misleading, self-centered, self-praising, and self-conflicting. They excluded the famous early angiosperm?Archaefructus?from their list of exemplar angiosperms, which contained only fossil plants they published themselves, leaving the impression that they were only authoritative on the origin and early history of angiosperms. Their 57-year-old “No Angiosperms Until the Cretaceous” conception does not reflect the truth about the origin and early history of angiosperms. Reinforcing such vapidly repeated statement does not help resolving any problem in science but leads to no solution for the origin of angiosperms. The authors tried to establish a criterion identifying a fossil angiosperm but their own exemplar angiosperm?Monetianthus?overturns their own criterion. Apparently, such a review does not positively contribute much to science.

Comparative Studies of Tracheary Element Structure of Some Gymnosperms with Angiosperms

Tissues of the pinna and rachis of Cycas diannaensis and pinna, rachis, and root of Cycas taiwaniana, rachis of Cycas szechuanensis, stem of Metasequoia glyptostroboides (Taxodiaceae), stems of Chamaecyparis obtusa (Sieb.et Zucc.) Endl cv. Tetragona (Cupressaceae), and leaves and stems of Michelia alba and Michelia figo and stems of Amygdalus persica (angiosperms) were compared using the scanning electron microscopy. In all species of these gymnosperms, their many tracheary elemnts have perforations in end walls and lateral walls. These structures are the same as vessels of angiosperms;therefore, these tracheary elements are vessel elements. Many types of vessels were found in cycads: pitted vessels in M. glyptostroboides, spiral and pitted vessels in Chamaecyparis obtusa cv. Tetragona. The development and structural characteristics of vessels of cycads, the two other gymnosperms, and the angiosperms were identical. Some characters such as extent of incline of perforation plate in the end wall showed that vessel characters of some angiosperms were more primitive than the cycads or M. glyptostroboides and C. obtusa cv. Tetragona. Many of the vessel elements of the angiosperms were band shaped, without end walls, and had only two lateral walls and other two margins;the end was acuate or with an arc margin;the end of some vessel elements was acute with no perforations, many perforations located only in lateral wall. Such results are rarely reported in previous work. In all species, perforations were seen only in tracheae, and the walls of parenchyma cells only had a thin primary wall and without perforation. Analysis and discussion to the experimental measures which were observed and research of the structure of vessel or tracheid, pointed out that several measures all could be used and the Jeffrey’ method effects were better. Comparing the vessels of cycads, Taxodiaceae and Cupressaceae helps us to understand the mechanism in which these most primitive or more primitive extant gymnosperms were adapted to harsh environments and to understand these species’ evolutionary extent, and has the significance to the studies of plant anatomy, plant systematics and plant evolution.

Origin of Angiosperms and Their Diversification in the Cretaceous

The dominating hypothesis stated that angiosperms originated in the Early Cretaceous, there were no pre-Cretaceous angiosperms, and carpels (the basic units of gynoecium) were derived from former megasporophylls bearing ovules/seeds along their margins through longitudinally folding and enrolling. However, there are increasing evidences of pre-Cretaceous angiosperms, the assumed megasporophyll actually does not exist, and the Cretaceous-only history of angiosperms appears much shorter than suggested by molecular clocks. Here I will integrate new knowledge of living and fossil plants to give a plausible explanation for the origin and early evolution of angiosperms. Several lines of evidence indicate that the ancestor of angiosperms may well have been present in the Triassic. The former gap between angiosperms and gymnosperms is artificial. Some Triassic fossils playing a role intermediate between angiosperms and gymnosperms seem to favor the Unifying Theory.

Nanjinganthus is an angiosperm,isn't it?

Nanjinganthus is an Early Jurassic angiosperm recognized based on the study of over 200 specimens.However,some other authors have misinterpreted these fossils.Here the authors try to remedy the problems,by pointing out the logical pitfalls in these publications and underscoring a long-used,workable criterion for early angiosperms.The paper explains the cons and pros of this criterion,hoping to bring palaeobotany and plant taxonomy back to a consistent and practical track.Nanjinganthus is an angiosperm.

Androecium of Archaefructus, the Late Jurassic Angiosperms from Western Liaoning, China

Androecium of the earliest known flowering plant Archaefructus liaoningensis was found from the Upper Jurassic Jianshangou Formation of western Liaoning, China. The androecium consists of numerous stamens bearing in pair on the reproductive axes below conduplicate carpels. The stamens are composed of a short filament and basifixed anther for each. Monosulcate pollen in situ are found from the anthers. The characters of the androecium reveals that Archaefructus are probably protandrous, and the paired stamens and monosulcate pollen appear to indicate that Archaefructus, as primitive angiosperms,might be derived from extinct seed -ferns during the Older Mesozoic. Archaefructus is considered Late Jurassic in age.

Inference of phylogenetic relationships among key angiosperm lineages using a compatibility method on a molecular data set

Phylogenetic relationships among the five key angiosperm lineages, Ceratophyllum, Chloranthaceae, eudicots, magnoliids, and monocots, have resisted resolution despite several large-scale analyses sampling taxa and characters extensively and using various analytical methods. Meanwhile, compatibility methods, which were explored together with parsimony and likelihood methods during the early development stage of phylogenetics, have been greatly under-appreciated and not been used to analyze the massive amount of sequence data to recon-struct the basal angiosperm phylogeny. In this study, we used a compatibility method on a data set of eight genes (mitochondrial atp1, matR, and nad5, plastid atpB, matK, rbcL, and rpoC2, and nuclear 18S rDNA) gathered in an earlier study. We selected two sets of characters that are compatible with more of the other characters than a random character would be with at probabilities of pM<0.1 and pM<0.5 respectively. The resulting data matrices were subjected to parsimony and likelihood bootstrap analyses. Our unrooted parsimony analyses showed that Cerato-phyllum was immediately related to eudicots, this larger lineage was immediately related to magnoliids, and monocots were closely related to Chloranthaceae. All these relationships received 76%–96% bootstrap support. A likelihood analysis of the 8 gene pM<0.5 compatible site matrix recovered the same topology but with low support. Likelihood analyses of other compatible site matrices produced different topologies that were all weakly sup-ported. The topology reconstructed in the parsimony analyses agrees with the one recovered in the previous study using both parsimony and likelihood methods when no character was eliminated. Parts of this topology have also been recovered in several earlier studies. Hence, this topology plausibly reflects the true relationships among the five key angiosperm lineages.

Nuclear phylogenomics of angiosperms and insights into their relationships and evolution

Angiosperms(flowering plants) are by far the most diverse land plant group with over 300,000 species. The sudden appearance of diverse angiosperms in the fossil record was referred to by Darwin as the “abominable mystery,”hence contributing to the heightened interest in angiosperm evolution. Angiosperms display wide ranges of morphological, physiological,and ecological characters, some of which have probably influenced their species richness. The evolutionary analyses of these characteristics help to address questions of angiosperm diversification and require well resolved phylogeny. Following the great successes of phylogenetic analyses using plastid sequences,dozens to thousands of nuclear genes from next-generation sequencing have been used in angiosperm phylogenomic analyses, providing well resolved phylogenies and new insights into the evolution of angiosperms. In this review we focus on recent nuclear phylogenomic analyses of large angiosperm clades, orders, families,and subdivisions of some families and provide a summarized Nuclear Phylogenetic Tree of Angiosperm Families. The newly established nuclear phylogenetic relationships are highlighted and compared with previous phylogenetic results. The sequenced genomes of Amborella,Nymphaea, Chloranthus, Ceratophyllum, and species of monocots, Magnoliids, and basal eudicots, have facilitated the phylogenomics of relationships among five major angiosperms clades. All but one of the 64 angiosperm orders were included in nuclear phylogenomics with well resolved relationships except the placements of several orders. Most families have been included with robust and highly supported placements, especially for relationships within several large and important orders and families.Additionally, we examine the divergence time estimation and biogeographic analyses of angiosperm on the basis of the nuclear phylogenomic frameworks and discuss the differences compared with previous analyses. Furthermore,we discuss the implications of nuclear phylogenomic analyses on ancestral reconstruction of morphological, physiological, and ecological characters of angiosperm groups, limitations of current nuclear phylogenomic studies, and the taxa that require future attention.

Angiosperm-wide analysis of fruit and ovary evolution aided by a new nuclear phylogeny supports association of the same ovary type with both dry and fleshy fruits

Fruit functions in seed protection and dispersal and belongs to many dry and fleshy types,yet their evolutionary pattern remains unclear in part due to uncertainties in the phylogenetic relationships among several orders and families.Thus we used nuclear genes of 502 angiosperm species representing 231 families to reconstruct a well supported phylogeny,with resolved relationships for orders and families with previously uncertain placements.Using this phylogeny as a framework,molecular dating supports a Triassic origin of the crown angiosperms,followed by the emergence of most orders in the Jurassic and Cretaceous and their rise to ecological dominance during the Cretaceous Terrestrial Revolution.The robust phylogeny allowed an examination of the evolutionary pattern of fruit and ovary types,revealing a trend of parallel carpel fusions during early diversifications in eudicots,monocots,and magnoliids.Moreover,taxa in the same order or family with the same ovary type can develop either dry or fleshy fruits with strong correlations between specific types of dry and fleshy fruits;such associations of ovary,dry and fleshy fruits define several ovaryfruit"modules"each found in multiple families.One of the frequent modules has an ovary containing multiple ovules,capsules and berries,and another with an ovary having one or two ovules,achenes(or other single-seeded dry fruits)and drupes.This new perspective of relationships among fruit types highlights the closeness of specific dry and fleshy fruit types,such as capsule and berry,that develop from the same ovary type and belong to the same module relative to dry and fleshy fruits of other modules(such as achenes and drupes).Further analyses of gene families containing known genes for ovary and fruit development identified phylogenetic nodes with multiple gene duplications,supporting a possible role of whole-genome duplications,in combination with climate changes and animal behaviors,in angiosperm fruit and ovary diversification.