Sublepidodendron grabaui comb. nov., a lycopsid from the Upper Devonian of China

Sublepidodendron wusihense (Sze) Sze and Lepidostrobus grabaui Sze were based on compressions from the upper part of the Wutung Formation (Famennian) of Jiangsu, South China. After studying the morphology and anatomy of abundant well-preserved specimens from two localities, Sublepidodendron wusihense and Lepidostrobus grabaui are reconsidered and viewed as Sublepidodendron grabaui (Sze) comb. nov. This plant is an arborescent, heterosporous lycopsid known from trunk, branches and cones. Leaf bases are spirally arranged, fusiform in outline, with a vascular bundle scar and keel. One specimen is known with a cone attached at the tip of a distal branch. The trunk has an intrastelar parenchyma concentration (pith), exarch primary xylem and secondary xylem. The branch anatomy varies from exarch primary xylem with a small, centrally located pith, to a solid exarch primary xylem strand. Based on the morphology and anatomy of both vegetative and reproductive organs, Sublepidodendron grabaui is placed into Sublepidodendraceae ( sensu Kräusel & Weyland, 1949), and Isoëtales ( sensu DiMichele & Bateman, 1996).  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 299–311.     

Hsüa deflexa sp. nov. from the Xujiachong Formation (Lower Devonian) of eastern Yunnan, China

A new vascular plant, Hsüa deflexa sp. nov., is documented from the Lower Devonian ((upper) Pragian-lower Emsian) Xujiachong Formation, near Xujiachong village, Qujing District, eastern Yunnan, South China. In three dimensions, the branching system comprises a robust creeping main axis and comparatively slender erect lateral axes inserted oppositely or alternately. The lateral axes depart at right angles from the main axis. Towards the apex, the main axis is perhaps erect with the lateral axes attached at smaller angles. The lateral axes dichotomize equally one to three times in different planes and decrease in diameter and length acropetally. They bear sparse and irregular tiny spines. Apices of a pair of distal lateral axes curve in opposite directions and each terminates in a rounded to reniform sporangium. The sporangium dehisces into two equal valves along an indistinct convex marginal thickening. The xylem is possibly centrarch. This plant fits Hsüa in branching pattern, sporangial characters and xylem maturation. Hsüa deflexa sp. nov. differs mainly from the type species H.  robusta in the presence of axial spines, perpendicular extension of lateral axes from the main axis and curving of distal lateral axes. Based on the possibly centrarch xylem and terminal sporangium, this plant relates to the rhyniophytes ( sensu Banks, 1968). In view of the equal sporangial valves with marginal thickenings it resembles the zosterophyllophytes ( sensu Banks, 1968). Hsüa is now treated as incertae sedis .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 255−271.     

A new late Westphalian fossil marattialean fern from Nova Scotia

Sydneia manleyi gen. et sp. nov. is based on part of a fertile frond from the upper Westphalian D of the Sydney Coalfield, Nova Scotia, Canada. It has small synangia composed of laterally fused sporangia that are elongate and with a circular cross-section. The sporangia yielded variably sized monolete and trilete spores with laevigate and microspinate ornamentation; intermediate forms were also observed. The spores can be correlated with the sporae dispersae species Latosporites minutus , Punctatosporites oculus and Laevigatosporites minimus . Size distribution of the spores is variable and highly skewed, suggesting heterogeneity of the spores within the sporangium. Spore ultrastructure indicates that the fossil is part of a fern, and the morphology of the spores and synangia indicate marattialean affinities.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 199–212.     

Estinnophyton wahnbachense (Kräusel et Weyland) comb. nov., une plante remarquable du siegenien d'allemagne

Specimens collected from the type locality of Protolepidodendron wahnbachense Kräusel et Weyland 1932 (Germany) are studied. The narrow axes, with persistent leaves which are bifurcated and spirally arranged, resemble those of P. wahnbachense. However, the vegetative and fertile leaves bifurcate twice and the four resulting segments are spread in various planes; the fertile leaves support two pairs of stalked elongate sporangia which are directed towards the axis. Estinnophyton wahnbachense (Kräusel et Weyland) comb. nov. is thus proposed as a name for these Siegenian plant remains, the questionable affinities of which are discussed.     

A new species of Estinnophyton from the Lower Devonian Posongchong Formation, Yunnan, China; its phylogenetic and palaeophytogeographical significance

A new species of early vascular land plant, Estinnophyton yunnanense , is described from the Lower Devonian (upper Pragian) Posongchong Formation at Zhichang village, Gumu Town, Wenshan District, Yunnan Province, China. This plant possesses aerial stems with isotomous branching. The leaves are slender, once- or twice-bifurcated, and arranged in low spirals or pseudowhorls. Fertile leaves are morphologically identical to the vegetative ones, clustered on the axis, forming a loose strobilus-like structure. There are three to six fertile leaves in each gyre. Two ovoid-elongate sporangia are singly attached to the adaxial surface each subtended by a recurved stalk, either at the base of each segment in a once-bifurcate fertile leaf or below the points of successive divisions in a twice-bifurcate fertile leaf. Estinnophyton yunnanense differs from two other species described previously from Europe, E .  gracile and E .  wahnbachense , mainly in possessing two single stalked sporangia on each fertile leaf rather than two pairs of sporangia, as well as in the dimensions of its leaves. In view of the complex taxonomy and nomenclature surrounding the type material of Estinnophyton , we considered possible emendation of the generic diagnosis based on the new Chinese material. Following comparison with typical members of the Protolepidodendrales and other taxa, Estinnophyton is considered to have had a closer affinity with primitive sphenopsids than the protolepidodendralean lycopsids. A comparison between Chinese and European floras containing Estinnophyton indicates that South China and Western Europe had floristic elements in common although they were in different phytogeographical units during the Early Devonian epoch.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 201–216.     

The pollen cones of Ginkgo from the Early Cretaceous of China, and their bearing on the evolutionary significance

A well-preserved pollen cone of the genus Ginkgo was found in the Yixian Formation (Early Cretaceous) of Liaoning Province, China, and is described as the new species Ginkgo liaoningensis . The pollen cone, catkin-like, consists of a main axis bearing scales at its base and spirally arranged sporophylls bearing 3–4 (−2) oval or elliptical sporangia. The sporangia are pendulous and dehiscent by a longitudinal slit. The mature pollen grains are monocolpate and elongate-elliptical, and the juvenile pollen grains are found in the tetrad stage. The characteristics of G. liaoningensis are very similar to those of the living Ginkgo biloba , but the sporophylls of G. liaoningensis bear 3–4 (−2) sporangia, whereas those of G. biloba bear mostly two sporangia. G. liaoningensis differs from fossil cones of Ginkgo huttoni from the Jurassic of Yorkshire, UK, and an unnamed specimen of Ginkgo from the Upper Cretaceous of Alberta, Canada, in the size and number of sporangia. The comparison between G. liaoningensis (fossil) and G. biloba (living) supports the reduction hypothesis of ovule organs in Ginkgo , with the number of sporangia having experienced the process of reduction from three or four to two since the Early Cretaceous.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 133–144.     

The Upper Permian articulate Phyllotheca australis from South Africa

RAYNER, R. J., 1992. The Upper Permian articulate Phyllotheca australis from South Africa. The taxonomic position of two fossil articulates, which form an important element of the Permian Gondwana flora, is examined in the light of recent collections from South Africa. Vegetative and fertile axes, dispersed and attached sporangia, in-situ spores, isolated and attached whorls of leaves, rhizomes bearing roots and branches, naked axes and a permineralized axis are all assigned to Phyllotheca australis. The large numbers of axes with attached sporangia allow a reinterpretation of the fertile structure and the specific diagnosis is emended. Although it is recognized that the fossils assigned to this taxon may represent a species complex, there exist no clear morphological discontinuities between the material previously assigned to P. australis and P. indica from widely separated Gondwanan localities. Phyllotheca australis was common throughout Gondwana, and is a reliable index fossil of the Upper Permian.     

Gumuia gyzzata A New Plant from the Lower Devonian of Yunnan,China

A new genus and species, Gumuia zyzzata is found in the Posonchong Formation of the Lower Devonian (Siegenian) from Wenshan district of Yunnan, China. It has both lateral and terminal sporangia. Successive sporangia develop on alternate sides of short axes near the hases of preceding sporangia, producing a zigzag fertile organization. The author supposes that it is a sympodium Opposite sporangia are suppositionally referred to the result from condensation of the axis between alternate sporangia. The new plant is tentatively referred to a putative zosterophyll.     

古木蕨——云南早泥盆世一植物新属

本文报道了滇东南文山地区下泥盆统坡松冲组(西根阶)地层中一陆生植物新属、种——曲轴古木蕨(Gumuia zyzzata gen.et sp.nov.)。植物矮小,匍匐或直立生长,二叉分枝。能育轴上孢子囊侧出,居于顶端或侧部位置,后者呈亚对生、交互对生或互生在曲折能育轴的折点上。孢子囊圆至横椭圆形,无柄或极短柄,成熟时通常切线垂直茎轴着生,并沿远端开裂。在系统分类上,依据新植物孢子囊侧出的性状,它暂被归人工蕨类,这有待于解剖结构的进一步证实。文中还对合轴分枝这一重要性状的发生做了探讨。     

贵州早泥盆世刺镰蕨的新观察

刺镰蕨在早泥盆世是一种广布于世界的植物,尽管报道自许多地点,但远未完全地了解它,其主要原因是能育材料罕见。我们观察的标本采自贵州早泥盆世的丹林组。它显示具柄的孢子囊侧生于靠近叶腋的轴上,我们的标本近似球形的孢子囊和它的形态略小于雷诺的肾形孢子囊外,其它方面如：带状的营养轴上具有疏螺旋排列的镰刀状叶,具短柄的孢子囊侧生于轴上以及管胞具穿孔壁等性状与雷诺修订后的刺镰蕨完全一致。我们同意雷诺的观点,将镰蕨目升为纲,与工蕨纲和石松纲并列。它的系统位置是介于工蕨纲和石松纲之间。其下包含一目、一科三个属——镰蕨属、巴拉格瓦纳彻属和星木属。这种植物的生殖部分在中国尚属首次报道。     

RELLIMIA THOMSONII FROM THE GIVETIAN OF NEW YORK: SECONDARY GROWTH IN THREE ORDERS OF BRANCHING

Pyritized axes of the vegetative branching system of Rellimia thomsonii, from the Givetian of New York were described with emphasis on the presence and position of the secondary tissues in relation to the known axis orders. The morphology and anatomy of four axis orders (N, N + 1, N + 2, N + 3) was described using two well-preserved specimens. One specimen showed the largest known (N) axis order, with spirally attached N + 1 axes. The other specimen, from a more distal portion of the plant, showed the N + 2 axis order with spirally arranged N + 3 axes. The shape of the primary xylem, discernible in three of the axis orders (N + 1, N + 2, N + 3), was a three-lobed protostele with mesarch maturation of the primary xylem. The protostele of the N + 3 axis order, with only primary growth, was identical to that of fertile N + 3 axes. The other axis orders (N + 2, N + 1, N) had wood surrounding the primary xylem. The wood of the N+2 axis was limited, and occurred only in the basal portion of the axis, but that of the N+1 and N axes was extensively developed with two and three growth layers, respectively. This wood was pycnoxylic with narrow rays similar to other aneurophytalean progymnosperms. Surrounding the secondary xylem was secondary phloem indicating the vascular cambium in Rellimia was bifacial. The presence of growth layers was discussed in relation to the climatic interpretation placed on their presence and how it correlates with paleogeographic reconstructions of the Devonian.     

New Observations on Drepanophycus spinaeformis from the Lower Devonian of Guizhou,China

Drepanophycus spinaeformis Goeppert is a plant of world-wide distribution in the Lower Devonian. Although this species has been reported from numerous localities, it is far from being understood well, because the fertile material is rare. Observation on specimens from the Danlin Formation of Guizhou showed that the sporangia were borne on short stalks attached laterally to the axes near the leaf axil, nearly sphaeroid, and smaller than Rayner’ kidney sporangia. Howerer, other character, e. g. banded nutritive axis bearing spiral drepanoid leaves, perforated tracheid wall, are the same as those described by Rayner. We agree with Rayner in raising Drepanophycales to the status of class, Drepanophycopsida, parallel with Zosterophyllopsida and Lycopsida. The systematic position of Drepanophycopsida is in between Zosterophyllopsida and Lycopsida. The class includes one order, one family and three genera (Drepanophycus, Baragwanathia and Asteroxylon).The fertile parts of the plant are reported for the first time in China.     

Primary growth and morphological markers of interannual growth limits in Cupressaceae from Patagonia

The present study shows experimentally that primary growth of the native Patagonian Cupressaceae Austrocedrus chilensis (D. Don) Pic.-Serm et Bizz., Fitzroya cupressoides (Molina) I. M. Johnst. and Pilgerodendron uviferum (D. Don) Florin is potentially continuous. External morphological markers of interannual growth limits have been identified for the main axes of these species after several years of observation. Such limits correspond with axis portions in which internodes are relatively short and leaves have a small distal free end forming an acute angle with the axis. The causality of these results and their utility for studying the productivity, adaptation to different environments, population management and conservation of these threatened species are discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146 , 285–293.     

FERTILE CONIFEROPHYTE REMAINS FROM THE LATE TRIASSIC DEEP RIVER BASIN,NORTH CAROLINA

Seed cones (Compsostrobus neotericus gen. et sp. nov.), pollen cones, and vegetative remains of coniferophytes occur in Upper Triassic rocks of the Deep River Basin (Pekin Formation) of Central North Carolina. Seed cones have spatulate ovuliferous scales, each with two ovules and subtended by an elongated bract with an attenuate tip. Cuticle of seed cones resembles that of leaves on vegetative axes. Slender leaves are borne along two sides of the axis. Pollen cones have helically arranged microsporophylls, each with two abaxial sporangia bearing pollen grains of the Alisporites type. Seed cones, pollen cones, and vegetative remains suggest a coniferophyte very modern in aspect.     

CRENATICAULIS,A NEW GENUS OF DEVONIAN PLANTS ALLIED TO ZOSTEROPHYLLUM,AND ITS BEARING ON THE CLASSIFICATION OF EARLY LAND PLANTS

The name Crenaticaulis verruculosus is proposed for slender, pseudomonopodially and dichoto-mously branching plants that bore opposite to subopposite sporangia along the stem and two rows of prominent, multicellular teeth on opposite sides of the terete axis. Epidermal cells were either narrow and elongate parallel to the stem or short with a papillate outer tangential wall. Axillary tubercles were present on one side of the stem near lateral branches. Some tubercles bore remnants of branches. Sporangia dehisced along their distal margins into two unequal halves. No spores were found. Occasional short lengths of stem were petrified by iron pyrite. Sections revealed a cortex consisting of four to six rows of thick-walled cells and a xylem strand. The strand was elliptical in transverse section and maturation was exarch. Tracheids were chiefly scalariform. It is suggested that the so-called axillary tubercles, known in several plants of Devonian age, were scars of rhizophores like those in the modern genus Selaginella. The plant is referred to the subdivision Zosterophyllophytina.     

Two new fossil matoniaceous stem genera from Tasmania, Australia

Two new genera, Tasmanopteris and Heweria, from Tasmania are the first definite permineralized rhizomes assignable to the Matoniaceae. Tasmanopteris richmondii sp. nov., of mide-Mesozoic age, is from the Lune River site in southwestern Tasmania and is composed of four or more annular solenostelos separated by cortices with an inner mesarch xylem ring and outer exarch xylem rings. Secretory canals occur in the cortices between the protoxylem clusters of its outer xylem rings. The specimen of Heweria kempii sp. nov., which is very similar to extant Matonia, was collected from a Lower Tertiary conglomerate in central Tasmania but was reworked from older sediments. Anatomically, the rhizomes of Heweria consist of 3–6 annular solenosteles with the inner and middle xylem rings being mesarch, whereas the outer ring is exarch. Roots arise opposite the protoxylem clusters of the outermost ring. Epidermal hairs on the stems of this genus are like those of living Matonia in being attenuate, uniseriate, and multicellular.

Associated venation pattern of partially preserved leaves, isolated sporangia, annuli, and spores in this specimen provide additional support for the relationships of this fossil to modern Matonia.     

The relationship of Sophora sect. Edwardsia (Fabaceae) to Sophora tomentosa, the type species of the genus Sophora, observed from DNA sequence data and morphological characters

Sophora tomentosa , the type species of the genus Sophora , is shown by phylogenetic analyses of rbc L and ITS sequence data to be sister to Sophora sect. Edwardsia . S. tomentosa and most of the species from sect. Edwardsia share hypogeal germination, exstipulate leaves, and terete filaments. These species have buoyant seeds, and are distributed by ocean currents throughout the pantropics ( S. tomentosa ) and around southern temperate oceanic islands (sect. Edwardsia ). S. tomentosa differs from the species of sect. Edwardsia by its frutescent growth habit, terminal elongate inflorescence and smooth-walled legume. S. macrocarpa is unusual in sect. Edwardsia as its leaves have stipules, the filaments are winged, and the legume is smooth-walled.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 439–446.     

Observations on Nothia aphylla Lyon ex Høeg

Naked branched axes associated with sporangia, originally described by Kidston and Lang (1920) as the probable fertile region of Asteroxylon mackiei, were later shown by Lyon (1964) to belong to an unknown plant which he named Nothia aphylla. The name was validated by Høeg in 1967. Nothia aphylla is described here in greater detail from new material collected from the Rhynie locality. The sporangia are lateral in terminal branched spikes. Their arrangement ranges from spiral to semiverticillate. The fertile axes terminate in clusters of 2–5 sporangia. The stalked sporangia are rather reniform with a distal transversely-extended dehiscence slit. Systematically Nothia has similarities to both Zosterophyllophytina and Rhyniophytina.     

The primary vascular system of Callixylon

The primary vascular system of Callixylon (main stem axes of Archaeopteris) is composed of a discontinous cylinder of longitudinal bundles from some of which traces diverge radially in a regular, helical pattern. Tangential divisions and fusions of primary xylem strands occur in approximately equal frequency and, thus, the number of bundles in the system remains constant in the specimen studied. Following divergence, traces increase in size and become ribbed distally. The available evidence suggests that all of the traces observed in this study are branch traces. Branch bases, embedded in secondary xylem, have the anatomical characteristics of penultimate axes of lateral branch systems of Archaeopteris. It is suggested that some regions of main axes of Archaeopteris might have borne lateral branch systems only, lacking entirely simple leaves. Lateral branch systems, therefore, might have been more similar to compound leaves than previously supposed.     

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.