Systematics of fossil platanoids and hamamelids

The data on fossil platanoids and hamamelids are generalized, their morphological diversity and probable patterns of the establishment of the extant families Platanaceae and Hamamelidaceae are analyzed. It is shown that morphological and epidermal characters of polymorphic leaves of typical platanoid appearance were formed in the Late Albian and remained essentially invariable to the present time, indicating the morphological stasis of these leaves combined with a wide variation range. In view of association with essentially different reproductive structures, it is proposed to classify these leaves by the morphological system irrespective of the natural system of angiosperms. A new system of extinct platanoids and hamamelids, which is based on reproductive structures and includes two orders, Hamamelidales and Sarbaicarpales ordo. nov., is proposed. Hamamelidales comprises two extant families, Platanaceae (with the subfamilies Platanoideae subfam. nov. and Gynoplatananthoideae subfam. nov.) and Hamamelidaceae, and the extinct family Bogutchanthaceae fam. nov.; the new extinct order Sarbaicarpales ordo. nov. consists of two new families, Sarbaicarpaceae fam. nov. and Kasicarpaceae fam. nov. In a system of flowering plants that is based on molecular data, the families Platanaceae and Hamamelidaceae are assigned to remote orders, excluding close relationship (APG, 2003). At the same time, the system of APG II often contradicts morphological and paleontological data, while traditional ideas of morphologists concerning the common origin of these families have recently been supported by paleobotanic evidence. Probable origin of the families Platanaceae and Hamamelidaceae from a common polymorphic ancestral group is discussed.     

Infructescences of Friisicarpus sarbaensis sp. nov. (Platanaceae) from the Cenomanian-Turonian of western Kazakhstan

Platanoid capitate infructescences are described as a new species Friisicarpus sarbaensis. Infructescences of Friisicarpus N. Maslova et Herman have been first found in the Cretaceous of Kazakhstan. A joint burial of Friisicarpus infructescences and other platanoid reproductive structures as well as Platanus-like leaves are discussed. This indicates a considerable role of the group in the Cretaceous flora of Kazakhstan.     

Phylogenetic and growth form variation in the scaling of nitrogen and phosphorus in the seed plants

Plant biomass and nutrient allocation explicitly links the evolved strategies of plant species to the material and energy cycles of ecosystems. Allocation of nitrogen (N) and phosphorus (P) is of particular interest because N and P play pivotal roles in many aspects of plant biology, and their availability frequently limits plant growth. Here we present a comparative scaling analysis of a global data compilation detailing the N and P contents of leaves, stems, roots, and reproductive structures of 1,287 species in 152 seed plant families. We find that P and N contents (as well as N : P) are generally highly correlated both within and across organs and that differences exist between woody and herbaceous taxa. Between plant organs, the quantitative form of the scaling relationship changes systematically, depending on whether the organs considered are primarily structural (i.e., stems, roots) or metabolically active (i.e., leaves, reproductive structures). While we find significant phylogenetic signals in the data, similar scaling relationships occur in independently evolving plant lineages, which implies that both the contingencies of evolutionary history and some degree of environmental convergence have led to a common set of rules that constrain the partitioning of nutrients among plant organs.     

Reproductive structures of an extinct platanoid from the early Cretaceous (latest Albian) of eastern North America

Two new species of platanoid reproductive structure are described from the Bull Mountain locality in the Patapsco Formation (Potomac Group) of northeastern Maryland, USA. Pistillate inflorescences and infructescences (Platanocarpus elkneckensis sp. nov.) consist of flowers and fruits in sessile globose heads that are borne on an elongate axis. Individual pistillate flowers consist of five free carpels surrounded by prominent tepals. Staminate inflorescences, flowers and isolated stamens are assigned to Hamatia elkneckensis gen. et sp. nov. Staminate flowers are borne in a globose head with a small number of stamens (five?) per flower. Stamens consist of very short filaments, long anthers with strongly valvate dehiscence and an apically expanded connective. The connective expansion is frequently very well-developed, hook-like and extends down the ventral surface of the stamen. Anthers contain small, tricolporate, reticulate pollen. Association evidence, similarity of inflorescence structure and the occurrence of Hamatia-type pollen on flowers, carpels and fruits of Platanocarpus elkneckensis suggests that the staminate and pistillate material was produced by a single species of plant. The “Hamatia-plant” provides further evidence of pentamerous floral structure in mid-Cretaceous platanoids and documents the occurrence of unequivocal tricolporate pollen in the platanoid complex.     

Variability in growth and reproduction in F1-families of an Erophila verna population

Summary Seeds from two morphologically different groups in an Erophila verna dune population were used in a growth experiment to analyse and quantify flexibility and plasticity of growth and reproductive characteristics.A strong correlation between plant morphology and individual seed weight was shown in parents and progeny. Seeds of plants with narrow leaves were twice as heavy (H-plants) as those from plants with broad leaves (L-plants) Families of H-plants allocated ca. 10% more dry matter to the roots. These characters were fixed, presumably due to selfing. Plants of both types are characterized by a graded control function between the vegetative and the reproductive phase. Most of the families showed a single switch from vegetative to reproductive growth, two families showed even two switch periods.Between-family differentiation is interpreted as a result of multiniche selection in populations of Erophila verna. Within-family variation is disadvantageous due to low fecundity.     

Leaf Architecture and Systematics of the Hamamelidaceae Sensu Lato

Hamamelids have a long fossil history and an important fossil record. Their interesting biogeographic relationships indicate a great age. There exist good surveys of the pollen and floral organs of this family whereas it is so far poorly known from leaf architecture. The leaf architecture of all 29 genera with more than 60 among the total of 140 species of the family was surveyed in this work using clearified leaves. It is found that leaf architecture analysis may shed light on the relationships within the family and the conclusion of evolution based on leaf architecture basically accords with that based on others. The major categories of leaf architecture of Hamamelids observed in this work are as follows: leaf form, leaf margin, tooth type, venation, marginal ultimate venation, areolation and trichome. It must be emphasized that of all these characters the tooth type is the most stable and useful for systematics. In this work a new tooth type is recognized under the name altingioid. Teeth of this type are obviously asymmetrical, with a persistent transparent gland on the top, and with their lateral veinlets free, not reaching the medial vein. All three genera of the subfamily Liquidambaroideae have this tooth type, whereas most leaves of the rest genera of this family have fothergilloid teeth, which are basically symmetrical, without glands. The venation in the fothergilloid tooth is almost the same as that in the altingioid tooth, the only difference being that the lateral veins on the abaxial side of the altingioid teeth are usually absent or very weak and short if present. The present authors consider that the subfamily Liquidambaroideae has to be separated from the family Hamamelidaceae sensu lato and treated as an independent family, Altingiaceae, on the basis of the special tooth type. different pollen morphology and flower structure. The stability of tooth type may serve classification not only of order and family level, but also of tribe, genus and species level with the help of characters of teeth, such as shape, size, density, distribution, single or double, with or without glands. By comparison of Hamamelidaceae and Altingiaceae with some primitive families of subclass Hamamelidae, namely, Trochodendraceae, Tetracentraceae, Cercidiphyllaceae, Eupteleaceae and Platanaceae, the putative evolutionary trend of tooth types is outlined as follows: ↑ altingioid Chloranthoid → Cercidiphylloid →platanoid → fothergilloid In general evolutionarytrend of teeth within these families is reduction and simplification in structure.     

Allelopathic potential of Tagetes minuta terpenes by a chemical, anatomical and phytotoxic approach

The dominant terpenes in the essential oils of the leaves and reproductive structures of Tagetes minuta L. (Asteraceae) were studied throughout its life cycle in a natural population. The anatomy of the secretory cavities was described in order to correlate the changes in terpene content with structural changes. Finally, the phytotoxic effect of ocimenones on germination was also evaluated. Monoterpenes increased in both green leaves and reproductive structures throughout the plant's life cycle, whereas the opposite occurred in senescent leaves. Spathulenol was the main component in senescing leaves. The highest content of ocimenones, the bioactive components of the essential oil, was found in the reproductive structures. Bioassays showed that both pure ocimenones and fruit material from T. minuta delayed and inhibited the germination of cohabitant species. A relationship between allelopathy, biosynthesis, catabolism and terpene release is proposed for the chemical ecological effect of T. minuta.     

中生代银杏类植物系统发育、分类和演化趋向

长期以来银杏类植物化石分类都依据营养叶形态为基础。由于叶形态的多型性和异源性,导致分类和系统发育解释的紊乱。根据对保存完好的繁殖器官(胚珠器官)系统发育分析结果所作的银杏目分类表明中生代除了银杏和银杏科以外,至少还存在着3．5个已灭绝的科级单元。此方案把已知其繁殖器官的成员和仅仅根据营养器官建立起来的分类位置不明的属严格地区分开来,并注明各科的限定性特征和已知成员的地质地理分布。银杏目植物自古生代起源,至早中生代以后朝着不同的方向辐射,呈现出丰富的多样性并经历了错综复杂的演化过程,其总的演化趋向是退缩：叶片扁化、蹊化和融合;胚珠器官简化,胚珠增大、数目减少,珠柄趋于消失。     

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.     

Paleobotanical evidence on the early radiation of nonmagnoliid dicotyledons

Paleobotanical studies indicate that several isolated and systematically depauperate groups of extant woody dicotyledons originated in the Mid Cretaceous. TheChloranthaceae had probably differentiated into insect-pollinated (Chloranthus andSarcandra) and wind-pollinated (Ascarina andHedyosmum) forms by the end of the Albian, and leaves referable to theTrochodendrales are known from the Albian and Cenomanian. In the latest Cretaceous and Early Tertiary, extinct representatives of theTrochodendrales includedNordenskioldia and theJoffrea-Nyssidium complex. ThePlatanaceae also differentiated before the end of the Albian and initially had insect-pollinated, unisexual flowers with five carpels or stamens. Some of these features persisted in the platanoid lineage until the Early Tertiary, and during the Paleocene and Eocene thePlatanaceae included forms with elliptical, palmate and pinnate foliage. The history of thePlatanaceae suggests that several features of the reproductive morphology of extant taxa may have arisen in association with a trend toward wind pollination. In the Mid Cretaceous, platanoid foliage partially intergrades with pinnateSapindopsis and pedateDebeya-Dewalquea leaves suggesting a close relationship betweenPlatanaceae andRosidae andFagaceae respectively. TheChloranthaceae, Trochodendrales, andPlatanaceae all occupy a somewhat intermediate position between theMagnoliidae andHamamelidae and are of considerable interest with respect to their role in the initial radiation of nonmagnoliid (higher) dicotyledons.     

Distribution and utilization of assimilated carbon in red clover during the first year of vegetation

Summary The pattern of distribution of¹⁴C labelled assimilates and translocation with time was measured in red clover during one reproductive cycle. Measurements were made on whole plants grown outdoors in pots by exposing the aerial parts to¹⁴CO₂ during one photoperiod. Simultaneously, root respiration and N₂ fixation were recorded.At the beginning of the vegetative period, 2/3 of the assimilates remained in the leaves (basal leaves), and 1/3 were directed to the root system. Then the development of branches required as much as 40% of the C and the root allocation decreased. Reproductive structures diverted 17% of the current photosynthates. Nitrogen fixation was optimal during the maximum extension of the basal leaves and decreased during the development of branches. During this period, C allocation to the nodulated roots was high with an estimated amount of 3.2 mg of C per mg of N fixed.With time, translocation occured within the foliage, from basal leaves to the leaves of the branches and to the new basal leaves developed after senescence of the branches. Remobilization to the reproductive structures remained minimal indicating that flower and seed growth was supported by current photosynthesis.     

Correlative controls of senescence and plant death in Arabidopsis thaliana (Brassicaceae)

Like most monocarpic plants, longevity of Arabidopsis thaliana plants is controlled by the reproductive structures; however, they appear to work differently from most dicots studied. Neither male- and female-sterility mutations (ms1-1 and bell1, respectively) nor surgical removal of the stems with inflorescences (bolts) at various stages significantly increased the longevity of individual rosette leaves, yet the mutants and treated plants lived 20-50 d longer, measured by the death of the last rosette and/or the last cauline leaf. A series of growth mutations (clv2-4, clv3-2, det3, vam1 enh, and dark green) also increased plant longevity by 20-30 d but did not delay the overall development of the plants. The mutations prolonged plant life through the production of new leaves and stems with inflorescences (bolts) rather than by extending leaf longevity. In growing stems, the newly-formed leaves may induce senescence in the older leaves; however, removal of the younger leaves did not significantly increase the life of the older leaves on the compressed stems of Arabidopsis. Since plants that produce more bolts also live longer, the reproductive load (dry weight) of the bolts did not seem to drive leaf or whole plant senescence here. The developing reproductive structures caused the death of the plant by preventing regeneration of leaves and bolts, which are green and presumably photosynthetic. They also exerted a correlative control (repression) on the development of additional reproductive structures.     

Photoreceptors and signals in the photoperiodic control of development

Many plant species are sensitive to changes in the seasons, especially with regard to their reproductive behaviour. Sexual or vegetative reproductive structures are often only formed at times of the year when days are sufficiently long, or short. Plants perceive daylength in the leaves, but reproductive changes occur in shoot apices in response to the movement of signals throughout the plant. There is good evidence that phytochrome A is an essential component of the daylength-sensing mechanism in long-day plants, and preliminary evidence suggests a potential interaction between phytochrome C and daylength sensitivity in short-day plants. The sensitivity of reproductive processes to photoperiodic control is directly altered by photoreceptor action, particularly in the case of phytochrome B, which has a major influence on whether flowering or tuberization occurs under non-inductive conditions in both long- and short-day plants, but is not involved in daylength measurement. The signals which move from leaves to the sites of reproductive development are not known, but there is good evidence that gibberellins may be important and some preliminary indication that brassinosteroids might also be involved in photoperiodic signalling.     

Morphogenesis of proximal branch leaves in mosses

The formation of deeply dissected and compound leaves at the bases of branches, their homology between different groups of mosses, and probable factors responsible for their development are considered. Previous authors differ in the interpretation of such leaves and in most cases describe them as special morphological structures named pseudoparaphyllia. It is shown, however, that this term has been applied both to whole leaves and to separate leaf parts. Among the patterns of leaf formation deviating from the basic type, a special place belongs to the Hampeella variant, where deeply dissected and compound leaves are formed due to the delayed development of branch primordia. The families representing this variant occupy a basal position in the phylogenetic tree of pleurocarpous mosses. The Leucodon variant, where splitting of leaves into lobes is apparently explained by strong stem extension, is not specific for any definite phylogenetic group and manifests itself in different families. The Hypnum variant is also not associated with certain phylogenetic lineages, but it provides an example of more profound specialization.     

A new species of the genus <Emphasis Type="Italic">Platimeliphyllum</Emphasis> N. Maslova from the Paleocene of the Amur Region,Russia

A new species, Platimeliphyllum valentinii, is described from the Paleocene of the Amur Region on the basis of macromorphological and epidermal characters. It was shown that leaves of Platimeliphyllum N. Maslova associate with two fundamentally different types of reproductive structures: platanaceous Archaranthus N. Maslova et Kodrul and Bogutchanthus N. Maslova, Kodrul et Tekleva, which combines characters of the Platanaceae and Hamamelidaceae. The higher evolutionary stability of leaf structures in comparison with reproductive organs is discussed.     

Climatic changes in eastern Asia as indicated by fossil floras. II. Late cretaceous and Danian

Four Late Cretaceous phytoclimatic zones — subtropical, warm-temperature, temparature and boreal — are recognized in the Northern Hemisphere. Warm-temperature vegetation terminates at North Sakhalin and Vancouver Island. Floras of various phytoclimatic zones display parallel evolution in response to climatic changes, i.e., a temperature rise up to the Campanian interrupted by minor Coniacian cooling, and subsequent deterioration of climate culminating in the Late Danian. Cooling episodes were accompanied by expansions of dicotyledons with platanoid leaves, whereas the entire-margined leaf proportion increased during climatic optima. The floristic succession was also influenced by tectonic events, such as orogenic and volcanic activity which commenced in Late Cenomanian-Turonian times. Major replacements of ecological dominants occurred at the Maastrichtian/Danian and Early/Late Danian boundaries.     

Biomass allocation patterns in terrestrial, epiphytic and aquatic species of Utricularia (Lentibulariaceae)

Utricularia forms the largest genus of carnivorous plants and is characterized by the possession of typical traps (“bladders”). Total biomass allocation was examined in three aquatic, six terrestrial and one epiphytic species of Utricularia from natural habitats in West Africa and from the Botanical Gardens, Bonn. Total biomass of aquatic species was considerably higher than that of terrestrial or epiphytic species. Epiphytic Utricularia accumulate about 35% of their biomass in green leaves, in contrast to 65% of nearly chlorophylless reproductive structures and traps. Aquatic species allocated more than 85% of their total biomass to stolons, leaves and traps, but only 10–13% to reproductive structures. This is in stark contrast to the allocation patterns of terrestrial bladderworts. These species allocate nearly 90% of their total biomass in reproductive structures, and only about 10% to stolons, leaves and traps. This reduction of photosynthetically active plant tissue strongly suggests that as a consequence of the alternative resource of chemical energy, the carnivorous habit might have partly replaced autotrophy in certain terrestrial Utricularia species, especially in some smaller ones.     

Density-dependent reproductive and vegetative allocation in the aquatic plant Pistia stratiotes (Araceae)

Pistia stratiotes is an aquatic macrophyte that grows in temporary-ponds in the southern Pantanal, Brazil. It reproduces both sexually and asexually and is usually observed forming dense mats on the water surface, a condition favored by the plant's vegetative reproduction coupled with an ability for rapid growth. In this study we examined the effect of densely crowded conditions on the production of reproductive and vegetative structures. In addition, we verified whether there is a trade-off between clonal growth and investment in sexual reproductive structures, and whether there is an allocation pattern with plant size. Individual plant biomass and the number of the rosettes producing sexual reproductive structures and vegetative growth structures both increased with density. Increase in plant size resulted in increased proportional allocation to sexual reproductive structures and vegetative growth structures. Allocation of biomass to reproduction did not occur at the expense of clonal growth. Thus, the density response appears as a increase of rosettes producing sexual reproductive structures and vegetative growth structures. Therefore, long leaves and stolons may be adaptive under densely crowded conditions where competition for light is intense. An important aspect in the study of trade-offs is the size-dependency of the allocation patterns .Usually, larger plants produce more biomass. Therefore, larger plants can allocate more biomass to both vegetative and sexual reproduction than smaller plants and thus show a positive correlation between both traits rather than the expected negative one.     

Resource partitioning to male and female flowers of Spinacia oleracea L. in relation to whole-plant monocarpic senescence

Male plants of spinach (Spinacea oleracea L.) senesce following flowering. It has been suggested that nutrient drain by male flowers is insufficient to trigger senescence. The partitioning of radiolabelled photosynthate between vegetative and reproductive tissue was compared in male (staminate) versus female (pistillate) plants. After the start of flowering staminate plants senesce 3 weeks earlier than pistillate plants. Soon after the start of flowering, staminate plants allocated several times as much photosynthate to flowering structures as did pistillate plants. The buds of staminate flowers with developing pollen had the greatest draw of photosynthate. When the staminate plants begin to show senescence 68% of fixed C was allocated to the staminate reproductive structures. In the pistillate plants, export to the developing fruits and young flowers remained near 10% until mid-reproductive development, when it increased to 40%, declining to 27% as the plants started to senesce. These differences were also present on a sink-mass corrected basis. Flowers on staminate spinach plants develop faster than pistillate flowers and have a greater draw of photosynthate than do pistillate flowers and fruits, although for a shorter period. Pistillate plants also produce more leaf area within the inflorescence to sustain the developing fruits. The (14)C in the staminate flowers declined due to respiration, especially during pollen maturation; no such loss occurred in pistillate reproductive structures. The partitioning to the reproductive structures correlates with the greater production of floral versus vegetative tissue in staminate plants and their more rapid senescence. As at senescence the leaves still had adequate carbohydrate, the resources are clearly phloem-transported compounds other than carbohydrates. The extent of the resource redistribution to reproductive structures and away from the development of new vegetative sinks, starting very early in the reproductive phase, is sufficient to account for the triggering of senescence in the rest of the plant.