Rotafolia songziensis gen. et comb. nov., a sphenopsid from the Late Devonian of Hubei, China

A sphenopsid from the Upper Devonian (Famennian) Xiejingsi Formation, south-western Hubei Province, China, previously named as various species in Sphenophyllum , Hamatophyton , Bowmanites and Sphenophyllostachys , is now reinvestigated and assigned to a new taxon, Rotafolia songziensis gen. et comb. nov. Its ribbed axes are anisotomous and possess slightly expanded nodes. Lateral axes are inserted at nodes on main axes. Whorls of much divided vegetative leaves are attached at nearly right angles to nodes of basal axes, and at acute angles to nodes of terminal axes. There are six leaves per whorl. The terminal strobilus includes a central axis and verticils of fertile units. Each fertile unit consists of a bract and numerous sporangia. The margin of the elongate-cuneate bract bears a distal and many lateral elongate segments. Clusters of elongate sporangia are abaxially attached to the base of the bract at the same level. The axis has an actinostele, composed of a three-ribbed, exarch primary xylem and radial secondary xylem. Although Rotafolia songziensis closely resembles Hamatophyton verticillatum in axis character, leaf morphology and primary xylem type, they are quite different in strobilar structure. Taxonomically, Rotafolia is placed in the order Sphenophyllales by three well-defined characters: 1) whorled appendages; 2) ribbed protosteles; 3) exarch primary xylem maturation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 21–37.     

A new genus of Early Devonian plants with novel strobilar structures and vegetative appendages from the Posongchong Formation of Yunnan,China

A new plant, Dibracophyton acrovatum gen. et sp. nov., is described from the Lower Devonian (Pragian) Posongchong Formation of Wenshan District, southeastern Yunnan, China. The plant has creeping axes from which arise vegetative and fertile axes. The vegetative axes helically bear lateral dichotomous appendages with curved or round tips. The fertile axes possess terminal strobili with numerous fertile units arranged in irregular helices. Each fertile unit consists of a stalked long-elliptical sporangium, with dehiscence into two equal valves, and two discrete long-ovate bracts covering sporangium from above–below directions. A new genus and species is thus established based on these characters and is temporarily regarded as incertae sedis of Tracheophyta, although it perhaps closes to the barinophytes in affinity. Detailed comparisons with other plants having a similar architecture, i.e., sporangia closely associated with modified vegetative structures, are made. The discovery of D. acrovatum further enriches the composition of the Posongchong flora and demonstrates great morphological disparity of the Early Devonian vascular plants.     

Axis differentiation in two South American Nothofagus species (Nothofagaceae)

An analysis was carried out on the length, diameter and number of leaves, and the ratios between these variables for current-year growth units (sibling growth units) derived from different nodes of previous-year growth units (parent growth units) of young Nothofagus dombeyi and Nothofagus pumilio trees. Changes in sibling growth unit length, diameter, and number of leaves with position on the parent growth unit were assessed. In both species, sibling-growth unit morphology varied according to both the axis type of the parent growth unit and the position of the sibling growth unit on its parent growth unit. For the largest parent growth units, the length, diameter and number of leaves of their sibling growth units decreased from distal to proximal positions on the parent growth unit. Distal sibling growth units had a more slender stem and longer internodes than proximal sibling growth units. Sibling growth units in equivalent positions tended to have a more slender stem for N. dombeyi than for N. pumilio. Long main-branch growth units of N. pumilio had longer internodes than those of N. dombeyi; the converse was true for shorter growth units. The growth unit diameter/leaf number ratio was consistently higher for N. pumilio than for N. dombeyi. Nothofagus pumilio axes would go through a faster transition from an 'exploring' morphology to an 'exploiting' morphology than N. dombeyi axes. Within- and between-species variations in growth unit morphology should be considered when assessing the adaptive value of the branching pattern of plants.     

A FERTILE AXIS OF TRILOBOXYLON ASHLANDICUM,A PROGYMNOSPERM FROM THE UPPER DEVONIAN OF NEW YORK

A partially petrified impression of Triloboxylon ashlandicum (Aneurophytales) is the first recognized fertile axis of the genus. Identification of the fertile axis rests on the similarity of its anatomy with that of previously described vegetative specimens. Fertile organs replace some vegetative branches along part of the main axis. Fertile organs are twice dichotomized in one plane and bear elongate sporangia arranged pinnately. Vegetative branches differ in that they bear the ultimate appendages of the plant helically. The latter organs dichotomize many times in one plane. Although similar in size and morphology to the ultimate appendages, the fertile organs are homologous by position and vascular supply to the vegetative branches which they replace. Sporangia of T. ashlandicum dehisce longitudinally and terminate in an apiculate tip. Spores are unknown. Fertile organs of T. ashlandicum resemble those of other Aneurophytales and support the earlier placement of Triloboxylon in the order on anatomical grounds. T. ashlandicum differs from other Aneurophytales, however, by bearing vegetative organs at the distal end of its fertile axis.     

A new euphyllophyte Kunia venusta gen. et sp. nov. from the Middle Devonian of Yunnan,South China

Abstract Kunia venusta gen. et sp. nov. is reported from the late Middle Devonian (Givetian) Haikou Formation near Kunming City, Yunnan Province, China. This plant has three orders of naked axes that divide pseudomonopodially. The second‐ and third‐order axes occur in a helix. Fertile appendages are distantly spaced and helically inserted to the third‐order axis; they comprise equally dichotomous branches terminated by two clusters of paired and fusiform sporangia. Sterile appendages are dichotomous and distally recurved. A comparison is made with the basal euphyllophytes including the trimerophytes, cladoxylopsids, zygopterids, stauropterids, and some relevant genera of uncertain affinity. The new plant resembles them in dichotomous appendages with terminal elongated sporangia, but differs mainly in the three orders of pseudomonopodial axes bearing helical laterals. It is thus placed in the Euphyllophytina as incertae sedis. It is suggested that an evolutionary divergence in the branching pattern and appendage morphology might have occurred in the Middle Devonian euphyllophytes, that is, maintaining three dimensions versus yielding more or less planation.     

Avian skin development and the evolutionary origin of feathers

The discovery of several dinosaurs with filamentous integumentary appendages of different morphologies has stimulated models for the evolutionary origin of feathers. In order to understand these models, knowledge of the development of the avian integument must be put into an evolutionary context. Thus, we present a review of avian scale and feather development, which summarizes the morphogenetic events involved, as well as the expression of the beta (beta) keratin multigene family that characterizes the epidermal appendages of reptiles and birds. First we review information on the evolution of the ectodermal epidermis and its beta (beta) keratins. Then we examine the morphogenesis of scutate scales and feathers including studies in which the extraembryonic ectoderm of the chorion is used to examine dermal induction. We also present studies on the scaleless (sc) mutant, and, because of the recent discovery of "four-winged" dinosaurs, we review earlier studies of a chicken strain, Silkie, that expresses ptilopody (pti), "feathered feet." We conclude that the ability of the ectodermal epidermis to generate discrete cell populations capable of forming functional structural elements consisting of specific members of the beta keratin multigene family was a plesiomorphic feature of the archosaurian ancestor of crocodilians and birds. Evidence suggests that the discrete epidermal lineages that make up the embryonic feather filament of extant birds are homologous with similar embryonic lineages of the developing scutate scales of birds and the scales of alligators. We believe that the early expression of conserved signaling modules in the embryonic skin of the avian ancestor led to the early morphogenesis of the embryonic feather filament, with its periderm, sheath, and barb ridge lineages forming the first protofeather. Invagination of the epidermis of the protofeather led to formation of the follicle providing for feather renewal and diversification. The observations that scale formation in birds involves an inhibition of feather formation coupled with observations on the feathered feet of the scaleless (High-line) and Silkie strains support the view that the ancestor of modern birds may have had feathered hind limbs similar to those recently discovered in nonavian dromaeosaurids. And finally, our recent observation on the bristles of the wild turkey beard raises the possibility that similar integumentary appendages may have adorned nonavian dinosaurs, and thus all filamentous integumentary appendages may not be homologous to modern feathers.     

A novel lycopsid from the Upper Devonian of Jiangsu, China

A new lycopsid, Monilistrobus yixingensis gen. et sp. nov., is described from the Wutung Formation (Famennian, Upper Devonian) of Jiangsu, China. The plant has many features typical of other Upper Devonian lycopsids, including dichotomous branching, helically arranged obovate expanded leaf bases, linear leaves with spiny appendages along the lateral margins, sporophylls widened proximally, and one elliptical sporangium attached to the adaxial surface. The most distinctive novel feature of the plant is that the modified sporophylls are arranged tightly into fertile zones or cone-like structures that are separated by lengths of axes with more lax sterile microphylls only: therefore the cone-like structures are strung along the branches like beads on a necklace.     

Comparative morphology of the gametophytes of Equisetum subgenus Hippochaete and the sexual behaviour of E. ramosissimum subsp. debile, (Roxb.) Hauke, E. hyemale var. affine (Engelm.) A.A., and E. laevigatum A. Br.

Gametophytes of Equisetum , subgenus Hippochaete , species E. hyemale, E. ramosissimum, E. laevigatum, E. variegatum and E. scirpoides are morphologically distinct. Lamellae from female branches and the archegonial neck cells provide a range of diagnostic features but antheridia are uniform throughout this subgenus, unlike the subgenus Equisetum , in which antheridial morphology is the most reliable criterion for specific identification of gametophytes. The classification of Equisetum into two subgenera based on sporophytic features is supported by gametophyte morphology: column lamellae and sunken antheridia with two opercular cells occur in Hippochaete whereas plate lamellae and projecting antheridia prevail in subgenus Equisetum. The absence of well-marked discontinuities in gametophyte morphology in Hippochaete reinforces cytologica! and hybridization data indicating that the taxa are more closely interrelated man in the subgenus Equisetum. No differences in gametophyte morphology were detected at the intraspecific level. Neither gametophyte morphology nor sexuality provide any definitive data to support me theory that Hippochaete contains the most primitive extant horsetails.
Sexuality in E. ramosissimum subsp. debile, E. hyemale var. affine and E. laevigatum is far more labile than in other taxa of Equisetum. Both male and female secondary gametophytes may be derived from initially male or female individuals as a result of lamellar proliferation.     

Phylogenetic diversification of Equisetum (Equisetales) as inferred from Lower Cretaceous species of British Columbia, Canada

Three types of anatomically preserved vegetative shoots with features that characterize crown group Equisetum have been discovered in Lower Cretaceous deposits (≈136 Ma) of British Columbia, Canada, suggesting the genus is much older than currently believed. Specimens include two types of aerial shoots described as E. haukeanum sp. nov. and E. vancouverense sp. nov. and one type of subterranean rhizome. Shoots are 1-2 mm in diameter, jointed, and in cross section have fluted stems with a hollow pith. Distinctive patterns of cortical sclerenchyma and different ridge morphologies characterize each shoot morphotype. Nodes display irregular branching, highly fused leaf sheaths, and a nodal diaphragm. The aerial stem morphospecies have vallecular canals on alternating radii with carinal canals of an equisetostele surrounded by only a few tracheids. No secondary tissues are produced. Bands of surficial stomata flank the furrows of one morphospecies. Rhizomes and aerial shoots are of a similar size, suggesting that the plants were equivalent in stature to the smallest living Equisetum species. These fossils augment our understanding of evolutionary transformations that led from Paleozoic Archaeocalamitaceae and Calamitaceae to crown group Equisetaceae, suggesting that the initial diversification of Equisetum began far earlier than suggested by molecular-clock-based estimates.     

Epidermal keratinocyte stem cells: their maintenance and regulation

The epidermis is a stratified epithelium consisting of inter follicular regions and appendages (hair follicles, sweat glands, sebaceous glands). The dominant cell type (the keratinocyte) is arranged in groups of cells termed epidermal proliferative units (EPUs), and one centrally-located clonogenic stem cell is ultimately responsible for replacing the remainder of the cells in the unit. Evidence is reviewed which indicates that the epidermal Langerhan's cell (ELC), and the cells comprising the dermis, may modify the keratinocyte microenvironment to create stem cell ‘niches’ and cellular diversity within the basal layer.     

The origin,patterning and evolution of insect appendages

The appendages of the adult fruit fly and other insects and Arthropods develop from secondary embryonic fields that form after the primary anterior/posterior and dorsal/ventral axes of the embryo have been determined. In Drosophila, the position and fate of the different fields formed within each segment are determined by genes acting along both embryonic axes, within individual segments, and within specific fields. Since the major architectural differences between most Arthropod classes and orders involve variations in the number, type and morphology of body appendages, the elucidation of the embryology and molecular genetics of the origin and patterning of insect limb fields may help to facilitate an understanding of both the mechanism of appendage formation and some of the major steps in the morphological evolution of the Arthropods. In this review, we will discuss recent studies that have advanced our understanding of both the origin and patterning of Drosophila leg and wing secondary fields. These results provide fresh insights into potentially general mechanisms of how body parts develop and evolve.     

On Occurring of Sporangia of Protolepidodendron Krejci and Its Significance

This paper deals with the characteristics and occurring state of the sporangia of Protolepidodendron and its significance. The authors discovered that the sporangia with a short sporangiophore spread out from the axis of the fertile branch. There is a distance from the base of sporophyllary leaves to the setting point of the sporangiophore. Some sporangia divideinto suspended and symmetrical sporangia at the tip of the sporangiophore. But the most of sporangia do not divide. According to the characteristics of the sporangia of Protolepidodendron Krejci, which can be distinguished from genera Barrandeina (Krejci) Stur and Longos-tachys Zhu, Hu et Feng. Protolepidodendron lixianense sp. nov. is also described in this paper.     

Periodicity of reproductive growth in lycopsids: An example from the Upper Devonian of Zhejiang Province,China

Monilistrobus Wang et Berry is a fossil lycopsid reported from the Upper Devonian Wutong Formation of Jiangsu Province, China, and includes only the type species Monilistrobus yixingensis Wang et Berry. Along a single fertile axis, Monilistrobus has cone-like structures separated by and alternated with a length of vegetative region, which is an important character here termed as periodicity of reproductive growth. The alternation of fertile and vegetative growth occurs regularly and produces repeated intercalary (non-terminal) fertile zones. New specimens, identified as Monilistrobus cf. yixingensis, from the same formation but of Zhejiang Province extend the geographic distribution of this genus. These new specimens demonstrate morphological variations within this genus, such as the lack of spiny appendages on the leaves, different measurements of leaf bases, and different shapes and positions of cone-like structures. The distribution of intercalary fertile zones, as summarized for the first time in this paper, occurs in the Drepanophycales, the Protolepidodendrales, the Isoëtales, the Lycopodiales, the Selaginellales, and Wuxia Berry, Wang et Cai and Monilistrobus of uncertain affinities. Thus, this peculiar trait might have evolved in the very early evolutionary stage of the lycopsids, as indicated in the basal group Drepanophycales. However, the presence of repeated, two or more intercalary fertile zones along a single fertile axis is demonstrated only in Monilistrobus and the extant Huperzia Bernhardi.     

Isolated clusters of paired tandemly repeated sequences in the Xenopus laevis genome.

There exist in the Xenopus laevis genome clusters of tandemly repeated DNA sequences, consisting of two types of 393-base-pair repeating unit. Each such cluster contains several units of one of these paired tandem repeats (PTR-1), followed by several units of the other repeat (PTR-2). The number of repeats of each type is variable from cluster to cluster and averages about seven of each type per cluster. Every cluster has ca. 1,000 base pairs of common left flanking sequence (adjacent to the PTR-1 repeats) and 1,000 base pairs of common right flanking sequence (adjacent to the PTR-2 repeats). Beyond these common flanks, the DNA sequences are different in the eight cloned genomic fragments we have studied. Thus, the hundreds of PTR clusters in the genome are dispersed at apparently unrelated sites. Nucleotide sequences of representative PTR-1 and PTR-2 repeats are 64% homologous. These sequences do not reveal an obvious function. However, the related species X. mulleri and X. borealis have sequences homologous to PTR-1 and PTR-2, which show the same repeat lengths and genomic organization. This evolutionary conservation suggests positive selection for the clusters. Maintenance of these sequences at dispersed sites imposes constraints on possible mechanisms of concerted evolution.     

Fossil Araceae from a Paleocene neotropical rainforest in Colombia

Both the fossil record and molecular data support a long evolutionary history for the Araceae. Although the family is diverse in tropical America today, most araceous fossils, however, have been recorded from middle and high latitudes. Here, we report fossil leaves of Araceae from the middle-late Paleocene of northern Colombia, and review fossil araceous pollen grains from the same interval. Two of the fossil leaf species are placed in the new fossil morphogenus Petrocardium Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov.; these fossils are very similar in leaf morphology to extant Anthurium; however, their relationship to the genus is still unresolved. A third fossil leaf type from Cerrejón is recognized as a species of the extant genus Montrichardia, the first fossil record for this genus. These fossils inhabited a coastal rainforest ～60-58 million years ago with broadly similar habitat preferences to modern Araceae.     

The problem of the skin derivatives' origin in the amniote evolution. The skin appendages--scale, feather, and hair

Overview of modern data on morphology of the skin derivatives in the higher vertebrates is given. Analysis of convergent similarities between the hair and feathers themselves as well as between their follicles makes it possible to forward a "generative" concept of the evolutionary origin of various ecto-mesodermal derivatives, such as keratinized dermal appendages (scales, feathers, hair). This concept appeared as a result of the author's studies on the skin derivatives, as well as of the data on molecular biology and the tissue engineering showing similar mechanisms of morphogenesis of the dermal appendages. Recurrently published ideas on various heterochronies in generations of the skin derivatives both in the onto- and the phylogeneses are also taken into acount. Various dermal appendages have appeared in the evolution of the higher vertebrates as independent generations of the ecto- and mesodermal tissues. Their parallel origin was caused by similar changes in the metabolism and molecular regulation of morphogenesis.     

海南七仙岭7种卷柏属植物的微观形态与生境间的关系分析

为探讨生境对卷柏属(Selaginella)植物微观形态的影响,利用扫描电子显微镜对海南七仙岭采集的7种卷柏属植物的侧叶、中叶、孢子叶的叶表皮形态以及小孢子形态进行全面观察分析,并计算气孔器大小、气孔密度、孢子大小等,比较分析其微观形态的区别及微形态与生境间的关系,为卷柏属植物的分类提供依据。结果显示:(1)同种卷柏属植物的侧叶、中叶与孢子叶在叶表皮形态上具明显差异,尤其是孢子叶上的气孔与营养叶相对较小且稀疏,与孢子叶的繁殖功能相符合。(2)不同种卷柏属植物的叶表皮特征也明显不同,主要表现在叶缘刺、气孔和瘤状突起上,表明这些特征可以作为卷柏属植物种间区分的依据。(3)卷柏属植物的小孢子形态稳定,纹饰多样;部分种间的小孢子形态相似,但可通过纹饰类型、裂缝的曲直进行区分。(4)琼海卷柏的小孢子具有独特的网状纹饰,暗示其具有独特的演化途径。(5)卷柏属植物叶表皮的气孔特征、瘤状突起特征,小孢子的颜色、纹饰,与海拔、生境的湿度有一定的相关性,但其形成机理有待进一步研究。     

An evolutionary treatment of the morphology and physiology of circulatory organs in insects

An overview from an evolutionary perspective is presented on the research of the past 2 decades on insect circulatory organs. Based on various functional morphology it is clear that the flow mode of the dorsal vessel ('heart') has changed during the evolution of hexapods. In all apterygotes and mayflies the flow is bidirectional. In most pterygote insects, however, it is unidirectional. In some endopterygote insects, the direction of the flow alternates. This is achieved by heartbeat reversal, which may have various physiological functions and is a derived condition that probably occurred several times during the course of insect evolution. Special attention is given to the hemolymph flow in body appendages. In ancestral hexapods, they are supplied by arteries, whereas circulation in appendages of higher insects is accomplished by accessory pulsatile organs. These auxiliary hearts are autonomous pumps and exhibit a great diversity in their functional morphology. They represent evolutionary innovations which evolved by recruitment of building blocks from various organ systems and were assembled into new functional units. Almost all pulsatile circulatory organs in insects investigated exhibit a myogenic automatism with a superimposed neuronal control. The neuroanatomy of insect circulatory organs has been investigated only in a small number of species but in considerable detail. Numerous potential peptidergic and a few aminergic mediators could be demonstrated by immunocytochemical and biochemical methods. The cardiotropic effectiveness of these mediators may vary among species and it can be stated that there is no uniform picture of the control of the various circulatory organs in insects. A possible explanation for the differences may lie in the different evolutionary origins of the muscular components. Furthermore, insect circulatory organs may represent important neurohemal releasing sites.