Giant cordaitalean trees in early Permian riparian canopies in North China: Evidence from anatomically preserved trunks in Yangquan,Shanxi Province

Cordaitaleans, as close relatives of modern conifers, had a long geological history in the Cathaysia from the Visean (Mississippian, lower Carboniferous) to the end of Permian. They became prominent since the late Pennsylvanian, and best developed during the Cisuralian (early Permian) in North China, serving as the volumetrically dominant to subdominant elements of wetland floras. Architecture and ecology of the Cathaysian cordaitaleans from non-peat-forming environments are poorly known. Here, we report giant cordaitalean trunks and describe their morphology and brief anatomical features from the Cisuralian Taiyuan Formation in Yangquan, Shanxi Province, North China. These trunks are characterized by the Artisia-like pith and pycnoxylic xylem. Absence of growth rings in the logs suggests they grew under non-seasonal humid tropical conditions. They are preserved in sandstone bodies interpreted as deposits of distributary river channels on the delta plain. Several trunks with attached rooting systems indicate that these trees may have been growing on channel levees or delta plains, and brought into the channels by lateral bank erosion. Allometric estimates of tree height suggests that the largest trees were up to approximately 43.5 m tall. Mature cordaitaleans with straight trunks were probably the tallest trees and formed the canopy of the riparian forest in North China during the Cisuralian.     

The Late Triassic Schilderia Adamanica and Woodworthia Arizonica Trees of the Petrified Forest National Park, Arizona, Usa

Two fossil tree species, both with unusual characteristics, occur in the Upper Triassic of the Petrified Forest National Park, Arizona, USA and adjacent areas. The first, Schilderia adamanica , has a highly idiosyncratic secondary xylem structure which contains normal uniseriate and broad complex multiseriate 'herring-bone' rays. The trunk cross-section of the secondary xylem may be either of a normal pycnoxylic type with a central pith and rays radiating from that or it may consist of appressed 'xylem masses' with rays curving towards one another at their extremities. The second, Woodworthia arizonica , has narrow, horizontal vascular traces traversing the entire radial width of the secondary xylem. By analogy with extant tree species, these traces would have terminated on preventitious buds deeply embedded in the bark which, in the case of these fossil trunks, have failed to be silicified. Such buds have the capacity to develop into epicormic shoots when the crown foliage of the tree is damaged. A further specimen of W. arizonica is recorded for the first time from the Permian of southern Brazil. Reconstruction drawings of both trees are produced.     

Sclerospiroxylon xinjiangensis nov. sp., a gymnospermous wood from the Kungurian (lower Permian) southern Bogda Mountains,northwestern China: Systematics and palaeoecology

A new silicified wood, Sclerospiroxylon xinjiangensis Wan, Yang et Wang nov. sp., is described from the Cisuralian (lower Permian) Hongyanchi Formation in southeast Tarlong section, Turpan City, Xinjiang Uygur Autonomous Region, northwestern China. The fossil wood is composed of pith, primary xylem and Prototaxoxylon-type secondary xylem. The pith is solid, circular, heterocellular, with sclerenchyma and parenchyma. The primary xylem is endarch to mesarch, with scalariform thickenings on tracheid walls. The secondary xylem is pycnoxylic, composed of tracheids and parenchymatous rays. Growth rings are distinct. Tracheids have mostly uniseriate, partially biseriate araucarian pitting on their radial walls. Helical thickenings are always present on both the radial and the tangential walls. Rays are 2–14 cells high, with smooth walls. There are 2 to 7, commonly 2 to 4 cupressoid pits in each cross-field. Leaf traces suggest that S. xinjiangensis nov. sp. was evergreen with a leaf retention time of at least 15 years. Based on the sedimentological evidence, growth rings within the S. xinjiangensis nov. sp. could have been caused by seasonal climatic variations, with unfavorable seasons of drought or low temperature. Low percentage of latewood in each growth ring is probably due to the intensity of climatic seasonality and/or long leaf longevity.     

Structure of a corystosperm fossil forest from the Late Triassic of Argentina

A palaeoecological study of a standing Late Triassic forest containing 150 silicified stumps from the Río Blanco Formation of Mendoza province, Argentina is described. A mapped portion of the forest floor provides quantitative data — tree density, mean separation of trees and basal area per stump — which, along with taxonomic and sedimentologic information, allowed the reconstruction of a plant community that grew along river banks and within proximal floodplain environments. Analysis of architectural and phenological data from monotypic forest indicates an evergreen community composed of a corystosperm genus with a canopy height of c. 13–21 m. The corystosperm taxon: Elchaxylon, like Rhexoxylon, has polyxylic axes with centripetal secondary xylem but does not generate perimedullar bundles and the centrifugal secondary xylem produces an undivided solid pycnoxylic cylinder. Vegetation analysis shows that the forest has a clustered distribution pattern with high density. Forest density ranges between 727 and 1504 trees/ha but there are first order clusters with elevated density (mean nearest neighbour distance of 1.85 m). The histogram of diameter classes based on 131 stumps suggests an earlier colonization by few older pioneers (the largest ones) followed by establishment of a large younger cohort of coeval trees. Based on 9 series and 139 rings, the mean ring width and mean sensitivity (MS) were 3.47 mm and 0.31 respectively. MS values and the presence of false rings indicate the forest community responded to stressed ecosystems. The growth rings are very erratic and range from 0.27 to 8.94 mm. For fossil growth analysis it was assumed that wider rings would suggest a warmer climate and the considerable range in growth rates would indicate variability in the limiting factors among subsequent cycles.     

The occurrence of vertical resin canals in Keteleeria, with reference to its systematic position in Pinaceae

The variation in the number of resin canals in the secondary xylem of Keteleeria fortunei (A. Murray bis) Carrière is determined. At all levels from base to top of a tree, resin canals are absent rearer the pith, but they (sporadically) appear after a certain cambial age. In addition, the occurrence of resin canals in seven named species of Keteleeria was surveyed. It was confirmed that normal vertical resin canals do occur in the mature parts of secondary xylem, but that they may be absent in the juvenile parts of secondary xylem of stems, as well as in branches and in seedlings or saplings. Earlier investigations reporting the absence of resin canals altogether in this genus may have been misled by the use of younger (immature) specimens instead of mature stems of trees. The distribution pattern of resin canals found in Keteleeria is systematically significant because it supports a new system of classification of Pinaceae, in which Keteleeria is believed to have a close relationship with Nothotsuga.     

A new anatomically preserved ginkgoalean genus from the Upper Cretaceous (Cenomanian) of the Czech Republic

A new genus of ginkgoalean woody branch, Pecínovicladus kvaceki gen. et sp. nov., is described from the Upper Cretaceous (mid to late Cenomanian) Peruc-Korycany Formation at Pecínov Quarry, near Prague, Czech Republic. Eighteen branch specimens, anatomically preserved as charcoal, the product of wildfire, occur as allochthonous assemblages in intertidal facies and as parauthochthonous assemblages in supratidal salt marsh facies. Primary branches range from 7–13 mm in diameter, and are composed of a central, parenchymatous pith, a medial pycnoxylic xylem layer, and an outer periderm layer. Closely spaced, helically arranged leaf bases and rare secondary branch scars characterize the exterior. The branches are confidently referred to the Ginkgoales on the basis of a variety of diagnostic xylem features, most notably the presence of inflated axial parenchyma containing moulds of crystals, and by the precise anatomical correspondence of the leaf bases to those of detached, but facies-associated Nehvizdya obtusa leaves of known ginkgoalean affinity. The newly described ginkgoalean, together with cheirolepidiaceous conifers, formed fire-prone vegetation in halophytic salt marsh environments under a seasonal, subtropical climate.     

Vertical and radial profiles in tracheid characteristics along the trunk of Douglas-fir trees with implications for water transport

The main stems of three young Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirbel) Franco) trees were dissected to obtain samples of secondary xylem from internodes axially along the trunk and radially within each internode. From these samples, measurements were obtained of tracheid diameter, length, the number of inter-tracheid pits per tracheid, and the diameter of the pit membranes. In addition, samples were obtained along the trunks of three old growth trees and also a small sample of roots for measurement of tracheid diameter. A gradient was apparent in all measured anatomical characters vertically along a sequence among the outer growth rings. These gradients arose not because of a gradient vertically along the internodes, but because of the strong gradients present at each internode among growth rings out from the pith. Tracheid characteristics were correlated: wider and longer tracheids had more numerous pits and wider pits, such that total pit area was about 6% of tracheid wall area independent of tracheid size. A stem model combining growth rings in parallel and internodes in series allowed for estimates of whole trunk conductance as a function of tree age. Conductance of the stem (xylem area specific conductivity) declined during the early growth of the trees, but appeared to approach a stable value as the trees aged.     

Ontogeny of the proventitious epicormic buds in Quercus petraea.

In the present work, we described the fate of proventitious epicormic buds on the trunks of 40-year-old Quercus petraea trees and in parallel the vascular trace they produced in the wood. Our results show that small and large individual epicormic buds can survive as buds for 40 years and that both are composed of a terminal meristem and scales. Meristematic areas are detected in the scale axils of small buds; in addition to these meristems the large buds also have secondary bud primordia. The small buds are connected to the pith of the main stem by a unique trace, whereas the large buds are connected by one or multiple traces. A single trace might imply that the whole bud is still alive and multiple traces might indicate that the terminal meristem has died. In the latter case, each trace is connected to a secondary bud of the large bud. The buds found in a cluster are composed of a terminal meristem and scales with axillary meristems in the scale axils. A cluster is connected to the pith of a stem either by a unique trace when it seems to be the result of partial abscission of an epicormic shoot or multiple traces when it might have originated from an epicormic bud in which the terminal meristem has died. Whatever the type of the bud, the vascular trace in the bark is composed of a cambium, secondary xylem and parenchyma cells and the trace present in the wood had parenchyma cells with vestiges of secondary xylem. Each year, the vascular trace should be produced in the bark by the cambium of the tree but not by the bud itself. On 40-year-old Q. petraea, we observed a proliferation of epicormic buds and in parallel a multiplication of the number of vascular traces in the trunk, but the knots caused by the traces of epicormic buds in the wood, either as individuals or in clusters, are minor since their colours are only slightly darker than those of woody rays and they are less than 2 mm in diameter. The knots will appear when epicormic buds develop into shoots. Received: 30 March 1999 / Accepted: 09 June 1999     

扁圆封印木（相似种）茎干的解剖特征

贵州省水城矿区晚二叠世煤核中扁圆封印木（相似种Ｓｉｇｉｌｌａｒｉａ ｃｆ．ｂｒａｒｄｉｉＢｒｏｎｇｎ．）茎干的主要解剖特征如下：管状中柱,具多边形薄壁细胞组成的髓。初生木质部成环带状,外缘呈规则的齿槽状,向心式发育。次生木质部显束状特征,横切面管胞为方圆至长方形,纵切面为梯状壁增厚,并具流苏纹。射线１—２列细胞宽,数个至十余个细胞高。叶迹起源于初生木质部外缘的槽中,中始式,但以向心发育为主。     

Anatomical Characteristics of the Stem of Sigillaria cf. brardii from Coal Balls in Guizhou Province,China

The stem specimens of Sigillaria cf. brardii were collected from the coal balls of Upper Permian in Shuicheng Coal Mines in Guizhou Province. The main anatomical characteristics of Sigillaria cf. brardii are described as follows: The stem is siphonostelic, with pith composed entirely of polygonal parenchyma cells, there are secondary walls in some pith cell cavities these secondary walls show the characters of cell division. Surrounding the pith is the continuous cylindrical primary xylem which consists entirely of tracheids. The outermost, and part are the protoxylem elements show spiral secondary thickenings. In cross section, the outer edge of exarch primary xylem appears regularly sinuous, with trace of mesarch leaf originating from the furrows. The centripetal metaxylem is characterized by scalariform wall thickenings on the tracheids, and delicated strands of secondary wall materials extending between abjacent bars, these structures are called fimbris, or williamson striations, and are characteristic in lepidodendrids. The secondary xylem consists of tracheids and vascular rays. The tracheids, too, have scalariform wall thickenings and fimbris. The rays are one-to twocell width and several to more than ten cells in height.     

Laurel forests in Tenerife, Canary Islands

The general wood structure, vessel size and distribution along the stem xylem radius and in petioles were studied in Laurus azorica trees living in a Tenerife laurel forest. The fractions of volume occupied by dry matter, water and air in percentage of wood fresh volume were also studied. The wood showed a diffuse-porous structure, with solitary vessels or vessels somewhat clustered in small radially oriented groups. Vessels had a diameter ranging from 20 to 130 µm. This diameter was minimal close to the pith, increased more than 2-fold with age, and reached its maximum width close to the cambium. Vessel density decreased from 36 vessels mm^-2 near the pith to about 13 vessels mm^-2 near the cambium. Accordingly, the lumen area was small in young xylem close to the pith (0.0015 mm²), reaching a value 5 times larger (0.007 mm²) near the cambium than in the centre of the stem. Lumen area of vessels in petioles was about 1.5% of petiole cross-sectional area and thus much lower than in stems. Mean hydraulic diameter of these vessels was about 20 µm, and mean vessel density about 136 per petiole. There were only small differences in proportions of dry matter, water and air along stem radius. The relevance of each one of these fractions in the wood is discussed as evidence of the possible existence of a number of embolized vessels dispersed in the total functional cross-sectional area of the xylem.     

Palaeoginkgoxylon zhoui, a new ginkgophyte wood from the Guadalupian (Permian) of China and its evolutionary implications

A new taxon of ginkgophyte affinity Palaeoginkgoxylon zhoui gen. nov. et sp. nov. is described from the Guadalupian Lower Shihhotse Formation of the Hulstai coalfield, Inner Mongolia Autonomous Region (Nei Mongol), northern China, on the basis of the anatomical structures of the broad eustele and pycnoxylic secondary xylem. The anatomical structure of the new woody tree trunk resembles both the early gymnosperms of Eristophyton-Pitus types and the modern Ginkgo. Therefore, the new tree trunk is interpreted as representing a transitional stage in the evolution of Ginkgo from early arborescent lignophytes since the Early Carboniferous.     

On Paleozoic plants from marine strata: Trivena arkansana (Lyginopteridaceae) gen. et sp. nov., a lyginopterid from the Fayetteville Formation (middle Chesterian/Upper Mississippian) of Arkansas, USA

Five permineralized seed fern stems from the Fayetteville Formation (middle Chesterian/Upper Mississippian) of Arkansas conform to the concept of lyginopterid seed ferns. However, these specimens are unlike all previously reported lyginopterids, and the name Trivena arkansana (Lyginopteridaceae) gen. et sp. nov. is proposed. The stems are up to 30 by 19 mm in diameter and have pentagonal pith and eustele of five cryptic sympodia. Secondary tissues include abundant xylem with numerous wide rays and phloem surrounded by a periderm. The cortex is parenchymatous with abundant sclerotic clusters: some clusters are randomly dispersed and some are in discontinuous rows. Sclerenchyma bands form the "Dictyoxylon"-type outer cortex. Leaf traces diverge in a 2/5 phyllotaxy. Traces, accompanied by concentric secondary xylem, increase in size as they extend through the secondary xylem of the stem. The trace assumes a squat C shape at the outer margin of the secondary xylem and in the cortex divides into three discrete bundles, each surrounded by secondary xylem. Galleries within the phloem contain arthropod coprolites and exhibit wound response, suggesting plant-arthropod coevolution. The discovery of this new lyginopterid stem adds to the growing list of unique taxa described from the Fayetteville Formation and further solidifies its reputation as one of the most important Upper Mississippian plant fossil sites in North America.     

STRUCTURALLY PRESERVED FOSSIL PLANTS FROM ANTARCTICA. I. ANTARCTICYCAS,GEN. NOV., A TRIASSIC CYCAD STEM FROM THE BEARDMORE GLACIER AREA

Silicified stems with typical cycadalean anatomy are described from specimens collected from the Fremouw Formation (Triassic) in the Transantarctic Mountains of Antarctica. Axes are slender with a large parenchymatous pith and cortex separated by a narrow ring of vascular tissue. Mucilage canals are present in both pith and cortex. Vascular tissue consists of endarch primary xylem, a narrow band of secondary xylem tracheids, cambial zone, and region of secondary phloem. Vascular bundles contain uni- to triseriate rays with larger rays up to 2 mm wide separating the individual bundles. Pitting on primary xylem elements ranges from helical to scalariform; secondary xylem tracheids exhibit alternate circular bordered pits. Traces, often accompanied by a mucilage canal, extend out through the large rays into the cortex where some assume a girdling configuration. A zone of periderm is present at the periphery of the stem. Large and small roots are attached to the stem and are conspicuous in the surrounding matrix. The anatomy of the Antarctic cycad is compared with that of other fossil and extant cycadalean stems.     

New records of Early Cretaceous petrified wood in Yumen,northwestern Gansu Province,China and their palaeoclimatic implications

Several fossil woods from Early Cretaceous sediments in Yumen City in northwestern North China Block, China, have been described. They belong to two fossil wood taxa, Piceoxylon yumeniense Zhou, Peng, Deng, Zhang and Yang n. sp. and Protophyllocladoxylon chijinense Zhou, Peng, Deng, Zhang and Yang n. sp. The well-preserved specimens yield secondary xylem with distinct growth rings. Piceoxylon yumeniense exhibits cross-field with taxodioid pits as well as two distinct xylem ray types. The bi- to triseriate rays are characterised by one or two horizontal resin canals with unequal uniseriate ends. Protophyllocladoxylon chijinense Zhou, Peng, Deng, Zhang and Yang n. sp. shows window-like cross-field pits, which are occasionally cupressoid and have uniseriate xylem rays. These fossil wood records improve our understanding of the fossil diversity, floral composition and palaeoclimate of the Xiagou Formation. Palaeoclimatic analysis of the palaeoxylogical assemblage indicates that the northwestern Gansu region predominantly exhibited a warm and wet climate condition, while a brief cooling event may have occurred in the region during the Early Cretaceous.     

Periodicity of Wood Formation in Some Trees of Lowland Rainforest in Nigeria

Freshly prepared chlor-zinc-iodide was used to determine theperiodicity of wood formation at breast height (144 cm fromthe ground) in the trunks of some trees growing in the LowlandRainforest around Ibadan, Nigeria. Wood formation shows seasonal periodicity in the plants studied.The cambial derivates on the xylem side differentiate into woodcells, which at a certain stage of differentiation have abundantcellulose in their secondary walls. The cellulose stains deepblue in chlor-zinc-iodide. This has been used as a criterionfor deciding that wood formation has started. When no cellswith deep blue staining secondary walls are found the cambiumis known to be dormant or quiscent. The resumption of cambial activity is correlated positivelywith bud break and unfolding of new leaves. In Bombax buonopozenseP. Beauv. the relationship may be obscured by local cambialactivity induced by injury. Wood formation stops in the trunks either towards the end ofthe rainy season or at the early part of the dry season. Itstarts either during the dry season or at the beginning of therainy season; but the bulk of the wood is formed during therainy season. Cambial activity stops in most cases before leaffall. At the cessation of wood formation the fully lignifiedxylem elements abut on the xylem mother cells or on xylem cellswith incompletely thickened cell walls. Presence or absence of a starch-free-zone and the noding ofthe vascular rays also give indications of seasonal periodicityin wood formation. Growth rings are periodic and one growthring is generally formed each year.     

Gymnosperm trees from the Permian of Antarctica: An anatomically preserved trunk of Kaokoxylon sp.

Anatomically preserved gymnosperm axes are relatively abundant in Permian localities of Antarctica, but their anatomy has rarely been studied in detail, which limits comparison with other Gondwanan morphotaxa. Here we describe a silicified trunk collected from the Upper Permian Buckley Formation at Coalsack Bluff, in the central Transantarctic Mountains. The trunk has a small heterogeneous pith approximately 4 mm in diameter containing conspicuous sclerotic nests, endarch primary xylem maturation, paired leaf traces, and secondary xylem of the Araucarioxylon type. Comparison with contemporaneous gymnosperm axes from Antarctica indicates that the Coalsack Bluff trunk represents a new Permian morphotaxon for the region. The anatomical characters of the pith and secondary xylem suggest an affinity with the genus Kaokoxylon Kräusel, previously reported from Permian and Triassic localities of Southern Africa, South America, India, and Australia.     

New petrified gymnosperms from the Permian of Maranhão (Pedra de Fogo Formation), Brazil: Ductolobatopitys nov. gen. and Kaokoxylon

Continuing the study of petrified gymnosperm trunks recovered from the Pedra de Fogo Formation, we identify here two new taxa from the Permian deposits of the Parnaíba Basin, northeastern Brazil. One taxon is an endemic form named Ductolobatopitys mussae Conceição, Neregato et Iannuzzi, nov. gen., nov. sp., characterized by solenoid, lobed and non-septate heterocellular pith, cauline bundles with endarch maturation, and secondary xylem with araucarian radial pitting on the tracheid walls. The other form is assigned to the genus Kaokoxylon, which has been recorded from most of Gondwana, including the Parnaíba Basin, but is recorded for the first time from the Pedra de Fogo Formation with the new species Kaokoxylon brasiliensis. It is characterized by solid, non-septate heterocellular pith with sclerenchyma cells, endarch cauline bundles, and uni-to triseriate radial pitting on the walls of the tracheids. The sedimentological interpretations of the outcrops where the fossils were collected indicate that these plants lived on the shores of large continental lakes, with relatively high humidity but possibly periods of drought. These inferences are supported by growth interruptions in the secondary xylem, the presence of calamitalean and tree-fern stems, and microbialites that crop in the same area. These new finds not only increase the known diversity of the flora in the Pedra de Fogo Formation, but also provide more accurate information for understanding the floristic elements that formed the subtropical flora during the Cisuralian in this basin in Western Gondwana.     

Radial secondary growth and formation of successive cambia and their products in Ipomoea hederifolia L. (Convolvulaceae)

Ipomoea hederifolia stems increase in thickness using a combination of different types of cambial variant, such as the discontinuous concentric rings of cambia, the development of included phloem, the reverse orientation of discontinuous cambial segments, the internal phloem, the formation of secondary xylem and phloem from the internal cambium, and differentiation of cork in the pith. After primary growth, the first ring of cambium arises between the external primary phloem and primary xylem, producing secondary phloem centrifugally and secondary xylem centripetally. The stem becomes lobed, flat, undulating, or irregular in shape as a result of the formation of both discontinuous and continuous concentric rings of cambia. As the formation of secondary xylem is greater in one region than in another, this results in the formation of a grooved stem. Successive cambia formed after the first ring are of two distinct functional types: (1) functionally normal successive cambia that divide to form secondary xylem centripetally and secondary phloem centrifugally, like other dicotyledons that show successive rings, and (2) abnormal cambia with reverse orientation. The former type of successive rings originates from the parenchyma cells located outside the phloem produced by previous cambium. The latter type of cambium develops from the conjunctive tissue located at the base of the secondary xylem formed by functionally normal cambia. This cambium is functionally inverted, producing secondary xylem centrifugally and secondary phloem centripetally. In later secondary growth, xylem parenchyma situated deep inside the secondary xylem undergoes de‐differentiation, and re‐differentiates into included phloem islands in secondary xylem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 30–40.     

The Late Triassic Araucarioxylon Arizonicum trees of the Petrified Forest National Park, Arizona, USA

Examination and measurement of many of the trunks attributed to Araucarioxylon arizonicum Knowlton eroded from the Late Triassic Chinle Formation in the Petrified Forest National Park, Arizona demonstrate that the living tree did not closely resemble any of the present-day Araucaria trees of the southern hemisphere as postulated in past reconstructions. The research indicates that it was a tall monopodial tree with branches occurring in a disordered manner on the trunk from the base to the crown. Calculations using the allometric method of Niklas indicate that the trees were of considerable size. The largest recorded trunk has a basal diameter of nearly 3 m and may represent a tree 59 m high, when living. The root system of the A. arizonicum tree consisted of a ring of four to six steeply inclined lateral roots and a massive, vertically directed tap root. Many of the trunks still have their root systems attached, a circumstance that indicates their felling by the cut-bank operations of the local river system. The massive roots of these trunks, particularly the large tap root, are consistent with growth in soft, deep, alluvial soil, and the thin scale bark is to be expected in a tropical climate free from frost.