Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

Rodeites Sahni reinvestigated—I

A detailed investigation of Rodeites Sahni from nearly 100 specimens collected from the Deccan Intertrappean beds of India has brought to light some new important facts: (1) The sporocarp wall is thicker and the spores within it are larger than hitherto described. (2) The mode of sorus attachment, observed for the first time, is similar to that of Marsilea rather than Regnellidium. (3) The presence of a distinct cellular gametophyte inside some megaspores is an entirely new finding for the family Marsileaceae. The presence of a male prothallus inside smaller microspores is also newly observed. (4) The occurrence of sporocarps in groups and their attachment are unique. (5) The prismatic layer is distinct in the sporocarp wall of Rodeites The above observations help in reconsidering the position of Rodeites in the Marsileaceae.     

The Correlation of Physiological Differences and Varions Leaf Forms of an Aqnatic Plant

An analysis of the celluar constituents of the leaves of an aquatic fern, Marsilea vestita, was done after and during the formation of different leaf forms. Land leaves of Marsilea produced almost twice as much dry material, soluble carbohydrate, insoluble protein and soluble peptides compared to submerged leaves on a fresh weight basis, but on a dry weight basis there is tittle difference between the two types of leaves. the submerged leaf soluble nitrogen fraction consisted mostly of ammonia and free amino acids, while that of the land leaf probably con tained mostly peptides. Free arginine was present in large quantities in megaspores and as the plant matured, at the expense of the megaspore, the amount of arginine left in the pool diminished. A correlation was found between soluble sugar increase and leaf elongation, Just prior to the production of submerged leaves the developing leaves contained low levels of protein and soluble sugar, but high levels of soluble nitrogen and an increased amount of starch. A theory which involves metabolism was proposed to account for the leaf form changes in Marsilea.     

Ethylene biosynthesis in Regnellidium diphyllum and Marsilea quadrifolia

The pathway of ethylene biosynthesis was examined in two lower plants, the semi-aquatic ferns Regnellidium diphyllum Lindm. and Marsilea quadrifolia L. As a positive control for the ethylene-biosynthetic pathway of higher plants, leaves of Arabidopsis thaliana (L.) Heynh. were included in each experiment. Ethylene production by Regnellidium and Marsilea was not increased by treatment of leaflets with 1-aminocyclopropane-1-carboxylic acid (ACC), the precursor of ethylene in higher plants. Similarly, ethylene production was not inhibited by application of aminoethoxyvinylglycine and -aminoisobutyric acid, inhibitors of the ethylene biosynthetic enzymes ACC synthase and ACC oxidase, respectively. However, ACC was present in both ferns, as was ACC synthase. Compared to leaves of Arabidopsis, leaflets of Regnellidium and Marsilea incorporated little [¹⁴C]ACC and [¹⁴C]methionine into [¹⁴C]ethylene. From these data, it appears that the formation of ethylene in both ferns occurs mainly, if not only, via an ACC-independent route, even though the capacity to synthesize ACC is present in these lower plants.Abbreviations ACC 1-aminocyclopropane-1-carboxylic acid - AdoMet S-adenosyl-l-methionine - AIB -aminoisobutyric acid - AVG aminoethoxyvinylglycine This research was supported by the U.S. Department of Energy through grant No. DE-FG02-91ER20021 and, in part, by a fellowship of the National Engineering and Research Council of Canada to Jacqueline Chernys.     

A new marsilealean fern species from the Early Cretaceous of Jordan

Leaflets of Marsileaceae are described from the Albian (Early Cretaceous) strata of Jordan. The fossils are from the Jarash Formation (Kurnub Group) and are found in fluvial sediments along with water lily leaves. The small wedge-shaped leaves have dichotomous veins that anastomose and form a marginal vein. Based on comparisons to living genera, Marsileaceaephyllum mahisensis Hu, Taylor, Brenner et Basha, n. sp., is most similar to Marsilea, in particular, with terrestrial leaflet forms; yet, it is distinct from living and fossil species by its small size and the few dichotomously branched middle veins that have a monopodial course. In addition, a single similar-veined smaller leaf with a retuse apex is thought to be a juvenile leaf of the same species. This is the first megafossil evidence of the family from Africa/Arabian Peninsula.     

Plant expansins are a complex multigene family with an ancient evolutionary origin

Expansins are a group of extracellular proteins that directly modify the mechanical properties of plant cell walls, leading to turgor-driven cell extension. Within the completely sequenced Arabidopsis genome, we identified 38 expansin sequences that fall into three discrete subfamilies. Based on phylogenetic analysis and shared intron patterns, we propose a new, systematic nomenclature of Arabidopsis expansins. Further phylogenetic analysis, including expansin sequences found here in monocots, pine (Pinus radiata, Pinus taeda), fern (Regnellidium diphyllum, Marsilea quadrifolia), and moss (Physcomitrella patens) indicate that the three plant expansin subfamilies arose and began diversifying very early in, if not before, colonization of land by plants. Closely related "expansin-like" sequences were also identified in the social amoeba, Dictyostelium discoidium, suggesting that these wall-modifying proteins have a very deep evolutionary origin.     

云南20种蕨类植物叶表皮微形态特征研究

利用光学显微镜对云南20种不同科、属的蕨类植物的叶表皮形态进行了观察和研究。结果表明：它们的叶片表面存在不同的附属物,而有的种类无附属物;20种蕨类植物共具8种气孔器类型(极细胞型、腋下细胞型、辐射状细胞型、双不等细胞型、无规则四细胞型、共环极细胞型、聚腋下细胞型和不规则型), 它们在气孔器组成上具多型现象,气孔均为下生型,分布方式及大小多样;叶表皮细胞的微形态在科属间表现出一定的差异,主要有不规则型、多边型或规则条状,表皮细胞垂周壁呈波纹状、深波状或波浪状。上述研究结果为蕨类植物的系统分类及演化提供依据。     

MORPHOLOGY OF MARSILEA VESTITA. III. MORPHOGENESIS OF THE LEAVES OF ETIOLATED PLANTS

The laminae of etiolated Marsilea vestita leaves develop by means of marginal meristems. Unlike light-grown plants, the form of the etiolated plant is not affected by growth on a solid medium. All of the young leaves isolated from light-grown submerged plants will elongate in darkness. The smallest, etiolated, uncoiled leaves develop into land leaves when they are placed in light, and this development occurs regardless of whether the leaves remain on the plant or are isolated on nutrient agar. Only these smallest leaves (2.3 mm average length) are actually capable of being converted from a submerged leaf form to a land leaf form by darkness.     

α-Expansins in the semiaquatic ferns Marsilea quadrifolia and Regnellidium diphyllum: evolutionary aspects and physiological role in rachis elongation

To investigate the evolutionary history of expansins and their role in cell elongation in early land plants, we isolated two α-expansin genes, Mq-EXP1 and Rd-EXP1, respectively, from the semiaquatic ferns Marsilea quadrifolia L. and Regnellidium diphyllum Lindm. The deduced amino acid sequences of the fern expansins exhibit a high degree of identity to those of seed plants, showing that expansin genes were conserved during the evolution of vascular plants. Gel-blot analysis of M. quadrifolia and R. diphyllum genomic DNA indicated that, in both ferns, α-expansins are encoded by multigene families. Expression of α-expansin genes probed with Mq-EXP1 was confined to the elongating region of the Marsilea rachis. Cell-wall proteins of M.␣quadrifolia induced in-vitro extension of acidified cucumber cell walls. In R. diphyllum, expression of Rd-EXP1 increased when elongation of the rachis was enhanced by submergence or ethylene. These results indicate that α-expansins act as wall-loosening proteins in ferns, as has been proposed for angiosperms. In addition, Rd-EXP1 may play a role in mediating elongation of the rachis in submerged plants. Received: 7 March 2000 / Accepted: 29 April 2000     

Architectural mutation and leaf form, for the palmate series

Palmate leaf form occurs in both the ferns and angiosperms. The palmate leaf form, and its variants, is present in distantly separated clades within both ferns and angiosperms. There tend not to be intermediate forms which link these palmate leaves to other leaf forms within the taxonomic groups in question. The recurrence of homoplasious leaf forms in separate taxonomic groups could be a consequence of the algorithmic like mode of leaf growth. Leaves develop through the reiteration of modular units. It is probable that the homoplasious leaf forms in different taxa are derived independently through re-combinations of the parameters in the basic leaf form development algorithm.     

Evolutionary patterns in the terebratulid genus Caryona Cooper (Brachiopoda,Jurassic of France)

The study of ontogenetic and morphological changes in different species of the brachiopod genus Caryona COOPER (Terebratulidae) from the Lower Callovian‐Lower Oxfordian of northern France shows a succession (and an interference) of processes which vary according to the characters studied. This is an example of “mosaic evolution”;. The ontogenetic changes result from heterochrony, especially acceleration and hypermorphism. The main phylogenetic tendency is peramorphosis in most characters. This evolutionary pattern agrees with phyletic gradualism.

Moreover, analysis of the internal characters demonstrates the limited value of these in discrimination of the various genera.     

ON THE STEM APEX,LEAF INITIATION AND EARLY LEAF ONTOGENY IN FILICALEAN FERNS

A general account of the stem apex organization in ferns is presented in support of the classical single apical cell concept. The range in variation of apical cells and of their modes of division are described. Evidence is brought out to indicate probable directing effects of the apical cell on modes of division of surrounding cells and on the leaf mother cell. Initiation of and eventual establishment of a stabilized apex in fern leaves is described. Of the more than 50 genera studied, the leaves of all are traceable to a single mother cell from which the leaf apical cell is cut out. Apical dichotomies are described in a number of genera as well as their effect on early leaf development. Results are discussed in a phylogenetic and morphogenetic context of leaf appendicularization.     

Ultrastructural study of Arcellites humilis Villar de Seoane and Archangelsky, 2008, from the Cretaceous of Patagonia,Argentina

A scanning and transmission electron microscopy-based ultrastructural study of Arcellites humilis Villar de Seoane and Archangelsky, 2008, from the Kachaike Formation and Piedra Clavada Formation (Albian-Cenomanian) from several localities in Patagonia (Argentina), is presented in this paper. TEM-photographs of thin-sectioned specimens show a thick spore body wall composed of three layers: a thin and dense exine, an inner granular perine sublayer with irregularly disposed small channels and lacunae, and an outer alveolate perine sublayer with channels perpendicular to the external surface. Transverse sections of the A. humilis wall are compared with fossil and extant species of Marsileaceae. We show that the wall ultrastructure is similar in A. humilis, A. santacrucensis, A. disciformis, A. stellatus and Regnellidium upatoiensis. However, the body wall and acrolamella of A. humilis are more similar to those of R. diphyllum Lindman than to those of the Marsilea L. or Pilularia L. species. Water ferns such as members of the Marsileaceae played an important role in aquatic or semi-aquatic niches in Patagonian Cretaceous floras, suggesting that high humidity and temperature prevailed during the Albian-Cenomanian in this region of Argentina.     

STIPE ANATOMY,WATER POTENTIALS,AND XYLEM CONDUCTANCES IN SEVEN SPECIES OF FERNS (FILICOPSIDA)

Anatomy and water relations were studied for the desert fern Notholaena parryi, as well as six other ferns representing three different orders which occupied xeric as well as mesic habitats. Tracheid number and diameter, and total xylem cross sectional area increased during leaf development for N. parryi; the whole plant conductance (volume flow of water through a stipe divided by the rhizome-to-leaf water potential drop) increased but tended to level off as the leaves matured. The reported occurrences of very steep water potential gradients (about 25 MPa m^–1) in stipes of N. parryi were confirmed. The ferns with the highest whole plant conductances (Alsophila australis, Botrychium dissectum, and Adiantum capillus-veneris) had the largest or greatest number of tracheids. Numerous tracheids in Botrychium dissectum offset a low tracheary conductivity, whereas Marsilea vestita had few tracheids resulting in a low whole plant conductance. Whole plant conductances for the ferns were 2 to 3 orders of magnitude less than those generally observed for angiosperms and 6 orders less than for gymnosperms. However, the relative conductivity (whole plant conductance times stipe xylem length divided by xylem area) was only 5- to 10-fold less than for angiosperms and about the same as for the gymnosperms. Stipe water relations in these ferns are discussed in relation to the evolution of xylem anatomy.     

The ontogenetic approach to chlorophyll fluorescence studies of plant photosynthetic apparatus under stressful conditions

Based on the analysis of reasons limiting the application of the method of chlorophyll fluorescence induction for estimating the state of the leaf photosynthetic apparatus under prolonged stress, the necessity of the ontogenetic approach consisting in a more exact determination of leaf age was substantiated. A comparison of the calendar and ontogenetic ways of determination of age of cucumber leaves under controlled conditions revealed essential distinctions in the estimation of plant leaf photosynthetic apparatus by the method of chlorophyll fluorescence induction for two variants distinguishing by the cultivation light regime ("white", 400-700 nm, and "red", 600-700 nm). It was shown that, in the case of prolonged effect of the stress factor on the plant, the unambiguity of the interpretation of chlorophyll fluorescence induction parameters in the estimation of the state of their photosynthetic apparatus depends essentially on the choice of the ontogenetic period of leaves of plants being compared and the accuracy of determination of leaf age.     

A quantitative method for inferring locomotory shifts in amniotes during ontogeny,its application to dinosaurs and its bearing on the evolution of posture

Evolutionary transitions between quadrupedal and bipedal postures are pivotal to the diversification of amniotes on land, including in our own lineage (Hominini). Heterochrony is suggested as a macroevolutionary mechanism for postural transitions but understanding postural evolution in deep time is hindered by a lack of methods for inferring posture in extinct species. Dinosaurs are an excellent natural laboratory for understanding postural transitions because they demonstrate at least four instances of quadrupedality evolving from bipedality, and heterochronic processes have been put forward as an explanatory model for these transitions. We extend a quantitative method for reliably inferring posture in tetrapods to the study of ontogenetic postural transitions using measurements of proportional limb robusticity. We apply this to ontogenetic series of living and extinct amniotes, focusing on dinosaurs. Our method correctly predicts the general pattern of ontogenetic conservation of quadrupedal and bipedal postures in many living amniote species and infers the same pattern in some dinosaurs. Furthermore, it correctly predicts the ontogenetic postural shift from quadrupedal crawling to bipedal walking in humans. We also infer a transition from early ontogenetic quadrupedality to late-ontogenetic bipedality in the transitional sauropodomorph dinosaur Mussaurus patagonicus and possibly in the early branching ceratopsian Psittacosaurus lujiatunensis but not in the sauropodomorph Massospondylus carinatus. The phylogenetic positions of these ontogenetic shifts suggest that heterochrony may play a role in the macroevolution of posture, at least in dinosaurs. Our method has substantial potential for testing evolutionary transitions between locomotor modes, especially in elucidating the role of evolutionary mechanisms like heterochrony.     

Heterochrony: Developmental mechanisms and evolutionary results

The concept of heterochrony, that the relative timing of ontogenetic events can shift during evolution, has been a major paradigm for understanding the role of developmental processes in evolution. In this paper we consider heterochrony from the perspective of developmental biology. Our objective is to redefine heterochrony more broadly so that the concept becomes readily applicable to the evolution of the full range of ontogenetic processes, from embryogenesis through the adult. Throughout, we stress the importance of considering heterochrony from a hierarchical perspective. Thus, we recognize that a heterochronic change at one level of organization may be the result of non-heterochronic events at an underlying level. As such, heterochrony must be studied using a combination of genetic, molecular, cellular, and morphological approaches.     

The ontogeny of sexual dimorphism in the cranium of Bornean orang-utans (Pongo pygmaeus pygmaeus): II. Allometry and heterochrony

Based on a homogeneous sample of 212 individuals spanning all postnatal periods, we examine the ontogeny of cranial sexual dimorphism in Bornean orang-utans (Pongo pygmaeus pygmaeus) by means of allometric analysis and in terms of heterochrony. The bivariate growth allometries of 20 cranial dimensions against basicranial length yield two major patterns. Confirming the null hypothesis, strong ontogenetic scaling, where growth regressions of both sexes fall along a single ontogenetic continuum, and where shape differences between adult males and females result from the extension of relative growth in the smaller females to larger size in males, is found in 10 cases. Ontogenetic scaling is particularly strong in proportions of (1) the neurocranium directly associated with brain size, (2) the orbital region, and (3) the dental arcade. In terms of heterochrony such a pattern most likely is the result of a process termed "time hypermorphosis", i.e. an extension of the growth period in time in males. The second major pattern seen in the remaining 10 cases shows a departure from ontogenetic scaling, with males exhibiting a significantly steeper slope than females. Departures from ontogenetic scaling, where size and shape are dissociated with adult males being disproportionately larger than adult females, are found in proportions of cranial regions directly associated with secondary sexual character development: prognathism, canine size, and cheek pad area. In terms of heterochrony such a pattern most likely is the result of a process termed "acceleration", i.e. the rate of shape change is increased in males.     

Roles of the af and tl genes in pea leaf morphogenesis: characterization of the double mutant (afaftltl)

The pleiofila phenotype (afaftltl double mutant) of Pisum sativum arises from two single-gene, recessive mutations known to affect the identity of leaf pinnae, afila (af), and acacia (tl). The wild-type leaf consists of proximal leaflets and distal tendrils, whereas the pleiofila leaf consists of branched pinnae terminating in small leaflets. Using morphological measurements, histology, and SEM, we characterized the variation in leaf form along the plant axis, in leaflet anatomy, and in leaf development in embryonic, early postembryonic, and late postembryonic leaves of aftl and wild-type plants. Leaves on aftl plants increase in complexity more rapidly during shoot ontogeny than those on wild-type plants. Leaflets of aftl plants have identical histology to wild-type leaflets although they have smaller and fewer cells. Pinna initiation is acropetal in early postembryonic leaves of aftl plants and in all leaves of wild-type plants, whereas in late postembryonic leaves of aftl plants pinna initiation is bidirectional. Most phenotypic differences between these genotypes can be attributed to differential timing (heterochrony) of major developmental events.