BARINOPHYTON CITRULLIFORME (BARINOPHYTALES INCERTAE SEDIS,BARINOPHYTACEAE) FROM THE UPPER DEVONIAN OF PENNSYLVANIA

Barinophyton citrulliforme is definitely proven to be a tracheophyte. The vascular cylinder of the main axis is an exarch protostele composed of tracheids having a continuous secondary wall folded into protrusions into the cell lumen. These protrusions delineate the position of annular thickenings which were deposited earlier than the continuous secondary wall. Between successive protrusions, the later-deposited secondary wall is interrupted by minute pitlike structures. It is suggested that the secondary walls of the tracheids were laid down in a two-phase depositional sequence. The fertile system of B. citrulliforme consists of a main axis bearing spirally arranged strobili. The strobilus consists of an axis that bears two alternate rows of sporangiferous appendages. The sporangiferous appendages are borne laterally along the strobilar axis and recurve abaxially around the axis. The sporangia are attached along the inside curve of the appendages, one sporangium per appendage, each containing both microspores and megaspores. This species thus exhibits sporangial heterospory which is considered to be an adaptation to an aquatic habit and the sporangia are considered to be functional analogs of the sporocarps of Marsilea. The interpretation of the strobilus is morphologically identical to Ananiev's interpretation of the strobilus of Pectinophyton bipectinatum; consequently P. bipectinatum is here transferred to Barinophyton as B. robustius comb. nov.     

An anatomically preserved lycopsid from the Upper Devonian of South China

An anatomically preserved lycopsid, Lobodendron fanwanensegen. et sp. nov. Liu, Wang, Xue & Meng, is described from the Upper Devonian (Famennian) Wutong Formation of Changxing County, Zhejiang Province, China. The fossil plant bears slender, dichotomously branched axes. The vascular strand consists of solid terete primary xylem and lobed secondary xylem, which implies the result from the activity of possibly discontinuous cambium. The new plant has character combinations that do not conform to any branches in the canopy of the tree-lycopsids known previously, but resemble those of the basal part of some pseudoherbaceous lycopsids. This new plant may exemplify a Late Devonian lycopsid with a pseudoherbaceous growth habit.     

A FERTILE SPECIES OF TRIPHYLLOPTERIS FROM THE EARLY CARBONIFEROUS (MISSISSIPPIAN) OF SOUTHWESTERN VIRGINIA

The first occurrence of attached fertile structures to foliage assignable to the form genus Triphyllopteris Schimper is reported from the early Carboniferous Price Formation of southwestern Virginia. Sporangium-bearing branch systems consisting of a basal undivided foliar region and a distal much-divided fertile region are described as a new species T. uberis Skog and Gensel. Ovoid fusiform sporangia, 2.0 mm long and 0.5 mm wide and containing trilete spores, terminate ultimate divisions of the branch systems and collectively form masses up to 4 cm in all dimensions. T. uberis is compared to previously known sporangiate organs of Late Devonian and early Carboniferous age, many of which lack attached foliage. Similarities in the overall branching pattern of many of these non-synangiate, sporangiate organs of early Carboniferous age is noted. The possible mode of dispersal and affinities of the plant bearing T. uberis fertile foliage is discussed.     

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.     

Vegetative Characters,Growth Habit and Microsporangiate Strobilus of Lycopsid Minostrobus chaohuensis

Late Devonian Minostrobus chaohuensis is one of the earliest monosporangiate-strobilate isoetaleans. Based on new material of this plant, the vegetative axis and microsporangiate strobilus are studied in detail, and the whole plant knowledge is summarized. The vegetative axis is isotomously branched. The stem is up to 55 mm in diameter with helically arranged leaf cushions. Stems and thick branches bear long fusiform leaf cushions and interareas with vertical linear ornamentations. A ligule pit, oblanceolate leaf scar, and vascular bundle scar appear on the leaf cushion. Distal axes have persistent lanceolate leaves and rhombic leaf bases. The microsporangiate strobilus is cylindrical in shape, possesses sporophyll with alate pedicel and long triangular lamina, uniseriate sporangial wall, subarchesporial pad inside the sporangium, and microspore with cingulum. Based on comparisons with other isoetaleans, the usage of the terms “leaf cushion” and “leaf base” is discussed, and Minostrobus chaohuensis is considered as a tree-like lycopsid. It suggests that arborescent isoetaleans with monosporangiate strobili had appeared and diversified in the Late Devonian. The multi-dichotomous branching system of Minostrobus provides new data on the evolution of growth architecture in rhizomorphic lycopsids.     

Re-study of Minostrobus chaohuensis Wang (Lycopsida) from the Upper Devonian of Anhui,South China

Minostrobus chaohuensis Wang was previously known only as a lycopsid megasporangiate strobilus from the Upper Devonian of Anhui Province, South China. Our comprehensive study of the newly collected materials reveals the details of its morphology, anatomy, and reproduction, which allow us to emend its generic and specific diagnoses. M. chaohuensis is reconstructed as a plant with multi-dichotomous branching system, helically arranged leaves, and monoecious and monosporangiate strobili (i.e., separate megasporangiate and microsporangiate strobili in one individual). The anatomy of both fertile and sterile portions of Minostrobus indicates that the exarch primary xylem strand is the solid protostele, with the peripheral protoxylem ridges and Williamson's striations in metaxylem tracheids. The key reproductive and anatomical characters suggest that Minostrobus chaohuensis is far more likely to represent a distal shoot of pseudo-herbaceous or arborescent lycopsids within the order Isoëtales sensu lato. It is suggested that the monosporangiate-strobilus clade in the Isoëtales may include primitive, monoecious taxa in the Late Devonian and advanced, dioecious ones in the Carboniferous. The hypothesis that the more phylogenetically advanced monosporangiate-strobilus clade might have well diverged from the basal bisporangiate-strobilus clade of arborescent lycopsids by the Late Devonian is further supported.     

STENOKOLEOS BIFIDUS SP. N. IN THE UPPER DEVONIAN OF NEW YORK STATE

Stenokoleos is a genus for petrified axes from the Mississippian New Albany Shale to which an Upper Devonian occurrence in New York is added. Two orders of branching were known and the plant was thought to be related to coenopterid ferns. The new petrified axes from New York reveal three orders of branching. A pair of rachides emerges from one side of the stem at each node. Their position alternates at successive nodes (distichous). Each rachis bears alternately arranged pinnae. The shape of the xylem strand and the number of protoxylem areas are variable. Traces to the pairs of rachides arise either as two separate strands or as a single strand that is presumed to divide while still within the cortex of the stem. Traces to pinnae are ellipsoid or clepsydroid. Tracheids are scalariform and uni- or biseriate, circular-bordered pitted. Peripheral loops are present in all orders of branches. Protoxylem strands are numerous and maturation is mesarch. Cortex is parenchymatous where it is preserved but outer cortex is missing. Stenokoleos and Reimanniopsis are placed in a new family, Stenokoleaceae. This is classified as Incertae Sedis among Pterophytina in Tracheophyta. It is suggested that the plant is related more closely to the Mississippian pteridosperms Tristichia and Tetrastichia than to the coenopterid ferns.     

A Silurian 'marrellomorph' arthropod

Xylokorys chledophilia, a new arthropod with three-dimensionally preserved soft tissues, is described from the Herefordshire (Silurian) Lagerstätte of England. The head and trunk are covered by a relatively featureless ovoid carapace, which comprises a domed central part and a flange-like border. The head bears five pairs of appendages. The first is uniramous, with dorsal and ventral projections distally. Appendages two to four are biramous and each endopod terminates in two projections. Appendage five is possibly biramous. The hypostome is very long and subrectangular in outline. There are approximately 35 pairs of biramous trunk appendages. Each exopod comprises a long slender shaft bearing numerous fine filaments; each endopod comprises a ribbon-like shaft bearing paddle-like endites. Morphological comparisons and cladistic analyses of X. chledophilia indicate affinity with Vachonisia rogeri from the Lower Devonian Hunsrück Slate, within the marrellomorphs, but assignment to Marrellomorpha is provisional pending revision of other members of this clade. Xylokorys is the first ‘marrellomorph’ to be reported from the Silurian. It is interpreted as a benthic particle filter feeder, which may also have consumed prey items.     

MORE EVIDENCE FOR CONIFER DIVERSITY IN THE UPPER TRIASSIC OF NORTH CAROLINA

An ovulate strobilus from the Upper Triassic Deep River Basin, North Carolina, has helically arranged, loosely aggregated, elongated, spatulate bracts with axillary ovule-bearing appendages with about 8–10 ovules attached in two lateral rows, with outwardly directed micropyles. The axillary ovuliferous appendage is homologous with the voltzialean fertile dwarf shoot, but probably not directly evolved from it. More credible is a suggested origin from a completely fertile axillary appendage such as that of the Lower Permian Trichopitys. The occurrence of this cone, Metridiostrobus palissyaeoides, gen. and sp. nov., along with Compsostrobus neotericus and Voltzia andrewsii, reflects considerable diversity among conifer ovulate cones during the Upper Triassic.     

A new species of Lepidostrobus from the Upper Devonian of Xinjiang, China and its bearing on the phylogenetic significance of the order Isoëtales

Lepidostrobus xinjiangensis sp. nov. is described from Upper Devonian rocks of the eastern Junggar Basin, Xinjiang, north-west China. It provides new insights into the reproductive diversification and phylogenetic relationships of lycopsids in the Late Devonian. The strobilus shares more characteristics with arborescent lycopsids than any herbaceous forms, and falls within the diagnosis of Lepidostrobus . Each sporophyll consists of a deltoid pedicel and a triangular lamina. Sporophylls are horizontally inserted into the strobilus axis in low spirals. The pedicel, with lateral alations and an abaxial keel, extends distally into the upturned lamina and downturned heel, producing a peltate appearance. A single sporangium with terminal longitudinal dehiscence is a radially elongate, dorsiventrally flattened ovoid and is attached along its length to the adaxial surface of the pedicel. A column-like subarchesporial pad is found in the sporangium. A possible ligule occurs on the adaxial surface of the pedicel distal to the sporangium. The strobilus is microsporangiate, containing Lycospora -type spores with granulate ornamentation and an equatorial flange. Based on this new species, the reproductive diversification and evolutionary pattern in arborescent lycopsids from the Devonian through the Carboniferous are discussed in a phylogenetic framework. We suggest that the reproductive strategies represented by bisporangiate- and monosporangiate-strobili had proliferated by the Late Devonian, which implies that phylogenetically advanced arborescent lycopsids bearing Lepidostrobus strobili had an earlier origin than previously thought.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143 , 55−67.     

New Middle Cambrian bivalved arthropods from the Burgess Shale (British Columbia,Canada)

The morphology of two new bivalved arthropods, Loricicaris spinocaudatus gen. et sp. nov. and Nereocaris briggsi sp. nov. from the middle Cambrian (Series 3, Stage 5) Burgess Shale Formation (Collins Quarry locality on Mount Stephen, Yoho National Park, British Columbia, Canada), is described. The material was originally assigned to the genus Branchiocaris, but exhibits distinctive character combinations meriting its assignment to other taxa. Loricicaris spinocaudatus possesses an elongate and spinose abdomen comparable to the contemporaneous Perspicaris and Canadaspis, as well as chelate second head appendages and subtriangular exopods, comparable to Branchiocaris. Nereocaris briggsi possesses a laterally compressed carapace, elongate and delicate appendages and a medial eye located between a pair of lateral eyes on a rhomboidal eye stalk. Although undoubtedly congeneric with Nereocaris exilis from a slightly younger horizon of the Burgess Shale Formation, N. briggsi differs in overall proportions and segment number, warranting assignment to a new species. The newly described taxa were coded into an extensive cladistic analysis of 755 characters, and 312 extinct and extant panarthropods, including a variety of Cambrian bivalved arthropods from both the Burgess Shale and the Chengjiang Lagerstätten. Cambrian bivalved arthropods consistently resolved as a paraphyletic assemblage at the base of Arthropoda. Important innovations in arthropod history such as the specialization of the deutocerebral head appendages and a shift from a nekton‐benthic deposit feeding habit to a benthic scavenging/predatory habit, the symplesiomorphic feeding condition of Euarthropoda (crown‐group arthropods), were found to have occurred among basal bivalved arthropods.     

湖北长阳上泥盆统一种石松植物

描述一种采自湖北上泥盆统弗拉阶黄家蹬组中的石松植物。该植物茎轴纤细。叶基纺锤形,螺旋排列。叶线形,叶缘具刺。具顶生的孢子叶球。其孢子叶匙状或披针形,边缘具刺。孢子囊呈圆形或椭圆形。植物茎具原生中柱。原生木质部呈小脊状位于中柱边缘。后生木质部管胞由梯纹分子组成,在加厚棒之间没有类似“威廉姆逊纹”的连接物。该植物与采自湖南中泥盆统基维特阶的Longostachys(Zhu,Huand Feng) Caiand Chen可比较。它们在茎轴、线形和具刺的叶、纺锤形和螺旋排列的叶基、匙状披针形的孢子叶,以及叶、叶基和孢子叶的度量等特征方面均非常相似。两者在解剖特征上存有差别,即当前植物不具次生木质部,不具髓,后生木质部加厚棒之间不具连接物。考虑到现有特征并不足以建立新属种,暂归入cf.Longostachyssp.     

New Sphenophyllum plant from the Upper Devonian of Zhejiang Province,China

Sphenophyllum was an important and long-surviving sphenopsid genus in the Paleozoic floras, with a worldwide distribution. A new species, Sphenophyllum changxingense sp. nov., is described from the Upper Devonian Wutong Formation of Changxing County, Zhejiang Province, China. This plant is characterized by two orders of slender axes and wedge-shaped leaves borne in whorls. The axes bear short spines and show longitudinal ridges and furrows on surface. Three to eight isophyllous leaves, with one, two, or no second-order axes, are attached at each node of first-order axes. Leaves bear spines and show a bilobate morphology; the two leaf lobes divide distally to form several marginal segments, each segment with a leaf vein. Sphenophyllum changxingense represents an early and primitive species within the genus, in light of the absence of heterophylly and specialized hook-like leaves. Like some Carboniferous and Permian species, it appears to have formed dense mats with mutually supportive axes. This plant adds to the known diversity of early sphenopsids in the Late Devonian.     

TRILOBOXYLON ASHLANDICUM GEN. AND SP. N. FROM THE UPPER DEVONIAN OF NEW YORK

A new taxon is described from the Upper Devonian, Oneonta formation in the northern Cats-kills of New York. Triloboxylon gen. n. is represented by petrifactions showing two orders of branching. The main axis bears branches spirally and the latter bear the ultimate appendages spirally. Ultimate appendages branch dichotomously twice, in one plane. Primary xylem of the main axis and branches is three-armed with mesarch protoxylem extending in a continuous band within each arm. Primary xylem of the ultimate appendage is terete and dichotomizes twice. Metaxylem elements are characterized by scalariform and circular-bordered pitting on all walls. The cortex is composed entirely of isodiametrical parenchyma cells. Triloboxylon is compared with other genera with three-lobed protosteles. Its possible affinity with the Aneurophytales is shown. The morphological nature of the “frond” of the Aneurophytales and the possibility that the group possesses the morphological equivalent of simple leaves are discussed.     

A pyritized polychaete from the Devonian of Ontario

A polychaete from the Middle Devonian Arkona Shale at Hungry Hollow, Arkona, Ontario is preserved in three dimensions in pyrite. The prostomium bears a single median antenna, a pair of lateral antennae and a pair of ventral palps. It is assumed to be fused to a reduced peristomium. The anteriormost three pairs of trunk appendages are modified as tentacular cirri, the third long and biramous. The remainder of the finely annulated trunk bears at least 21 similar biramous parapodia, some of which preserve evidence of chaetae. The postsegmental pygidium is very small and may bear up to two pairs of cirri. The polychaete, Arkonips topororum, falls within the Palpata, Aciculata, among the crown group Phyllodocida. Its remarkable preservation highlights the potential of the Arkona Shale to yield other examples of soft-tissue preservation.     

A new homosporous,arborescent lycopsid from the Middle Devonian of Xinjiang,Northwest China

Abstract: A new arborescent lycopsid, Hoxtolgaya robusta gen. et sp. nov., is described from the Middle Devonian of Xinjiang, Northwest China. It has stems up to 90 mm wide with fusiform leaf bases and long linear microphylls. Sporophylls are not aggregated into strobili and are isomorphic, with sporangia homosporous and bearing Acinosporites‐type microspores. A syndrome of characters in Hoxtolgaya, including the arborescence and the homospory, implies that the arborescent habit is not necessarily correlated with the heterospory in the early evolution of arborescent lycopsids. Homosporous, arborescent lycopsids probably represent one of the transient forms between the Devonian herbaceous protolepidodendrids and the Devonian–Carboniferous heterosporous arborescent lycopsids.     

The origin of the ovule

Of several theories for the origin of the ovule advanced in this century and based largely on fossil evidence, the telomic concept is widely considered the most plausible. Its principal tenet is the evolution of the integument through fusion of sterile branches or telomes around a terminal megasporangium. The only point of agreement in these theories is that the entire nucellus is a megasporangium that retains a single megaspore and the endosporic female gametophyte. Their differences center on the origin of the integument. A new concept offered here on evidence from ovules of both Paleozoic and extant seed plants significantly alters the telomic theory. It proposes that the nucellus is a sporangiophore of stem origin that bears a terminal megasporangium; that at least some of the fused integumentary telomes were fertile; and that among all features cited to characterize ovules, the unique nature of the retained megaspore alone defines the seed habit. Changes in the seed plant megaspore that extended the period of nutrient absorption over the whole course of female gametophyte development, along with complex physiological changes in the nucellus, were probably achieved along a single phylogenetic line beginning in a Late Devonian population of progymnosperms. For such a combination of events to have occurred more than once is highly unlikely, and, therefore, a monophyletic origin for seed plants is proposed. Several primitive features in ovule structure, some not evidenced since the Lower Carboniferous Period, occur in a mutant form of Arabidopsis thaliana isolated from genetically transformed plants. Their recurrence provides additional support for the proposed concept of ovule origin and also suggests that the genetic mechanisms for expression of primitive features in advanced taxa could be initiated in each case by mutation of a single homeotic gene.     

The first trees. The Archaeopteris model

The earliest self-supporting organisms exceeding 2 m in height evolved about 370 million years ago, approximately 100 million years after the rise of the first land plants. Evidence for the tree habit is usually indirect and assessed from the diameter of the available stem fragments. Four systematic groups of Devonian plants evolved the tree habit independantly: the Lycopsida, Cladoxylopsida, and progymnosperms in the Middle Devonian, the Equisetopsida in the Late Devonian. All share a free-sporing life cycle which limits their habitats to wet areas. Their branching pattern involves the strict division of their apices, whether equally or unequally. The progymnosperm genus Archaeopteris was widespread worldwide and evolved the highest trees of the Devonian (maximum height estimated at 40 m). Besides it ecological significance as the dominant component of the earliest forests, Archaeopteris currently represents the closest known relative to the seed plants with which it shares two derived characters, the heterosporous life cycle, and the possession of leaves. Another distinctive feature of Archaeopteris trees is represented by the double function of their wood for both support and conduction. New analyses involving vascular trace analysis in anatomically preserved specimens have demonstrated that Archaeopteris is not the simple tree reconstructed by Beck (1962). In this fate model, Archaeopteris consisted of an erect trunk bearing short-lived, flattened, leaf-like branch systems forming a terminal crown. New evidence indicates that laterally to these appendages of apical origin, a new type of branches, of adventitious origin, evolved which development compares to that of the axillary branches of the seed plants. These branches which were large and long-lived represent major architectural components of the tree. Evidence for vascular structures comparable to those produced on stem cuttings in modern plants suggest that Archaeopteris may have evolved vegetative strategies for propagation. The set of "modern" characters of Archaeopteris may explain its success until the Devonian/Carboniferous boundary when its extinction is correlated to the radiation of the earliest seed plants.     

“Lycostrobus” chinleana,an equisetalean cone from the Upper Triassic of the southwestern United States and its phylogenetic implications

Detailed study of the cone Lycostrobus chinleana Daugherty shows that the fossil was incorrectly attributed to the Lycopodiales by the author and to the quillworts by Retallack and that it actually should be assigned to the Equisetales. The cone, which occurs in the Upper Triassic Chinle Formation at several localities in the southwestern United States, is ∼2.5 cm wide and nearly 6 cm long and consists of a stout axis bearing whorls of peltate sporangiophores. Each sporangiophore is composed of a slender stalk and a hexagonal disk, which typically bears a single, generally long, lanceolate, forward-directed leaf-like umbo tip on the outer surface and several recurrent sporangia on the inner surface. Small round to oval trilete spores occur in the sporangia. Since the leaf-like umbo tip is similar to the sterile leaves found in certain calamite cones and the recurrent sporangia are equisetalean-like, it appears that the cone may represent a intermediate stage between Calamites and modern Equisetum. According to this hypothesis, the nonbracteate Equisetum cone could have developed from a bracteate calamite cone, through reduction and fusion of the bracts and the sporangiophores, rather than by the loss of whorls of bracts of the Calamites cone as suggested earlier by others. As a result of this study the cone is assigned to the new Equisetalean genus Equicalastrobus and redescribed under the name E. chinleana (Daugherty) Grauvogel-Stamm and Ash, n. comb.     

PROTEOKALON A NEW GENUS OF PROGYMNOSPERMS FROM THE DEVONIAN OF NEW YORK STATE AND ITS BEARING ON PHYLOGENETIC TRENDS IN THE GROUP

Proteokalon gen. nov. is described from the Upper Devonian Catskill deposits of New York. Two orders of branching and ultimate appendages are preserved' by petrifaction and by compression. The first order bears branches decussately and has a skewed four-armed protostele that occasionally dichotomizes. Second-order branches dichotomize rarely and most have T-shaped or three-armed protosteles. They bear ultimate appendages alternately, either in lateral pairs, or singly from the abaxial side. These appendages divide several times in one plane. Their vascular strand is terete. Maturation of the primary xylem is mesarch, and it consists of tracheids and parenchyma. Secondary xylem and phloem and a periderm are present. The outer cortex has a system of hypodermal fibers. Proteokalon is most similar to Tetraxylopteris and Triloboxylon of the Aneurophytales. A comparison of the stratigraphic occurrence of Protopteridium, Aneurophyton, Tetraxylopterism, Sphenoxylon, Triloboxylon, and Proteokalon suggests some evolutionary trends among the Aneurophytales.