A NEW SPECIES OF PICEA BASED ON SILICIFIED SEED CONES FROM THE OLIGOCENE OF WASHINGTON

Picea eichhornii n. sp. is described from anatomically preserved seed cones. The fossils are from the Early Oligocene Jansen Creek Member of the Makah Fm. which is exposed along the northern shore of the Olympic Peninsula, Washington. The cones are at least 5.5 cm long and up to 3.5 cm in diameter. The cone axis is 4–6 mm in diameter and contains a pith made up of thick-walled parenchyma cells. Resin canals occur in a single ring in the secondary xylem in some specimens but are absent in others. The cortex is mostly parenchymatous and contains numerous large axial resin canals that branch to supply the bract and scale. Vascular traces to each scale and its subtending bract diverge separately from the vascular cylinder of the cone axis. The bract is tongue-shaped and keeled at its base. It is 5 mm wide and up to 9 mm long. The bract trace fades out before entering the bract base while two resin canals extend into the bract base. The ovuliferous scale is about 2.3 cm long and has a thin, probably papery, apex. Resin canals of the scale occur abaxial to the vascular tissue in the scale base, but some bend around the margins of the vascular strand to become adaxial outward. About 20 resin canals occur in the abaxial scale sclerenchyma, and this is the main anatomical feature that distinguishes these cones as a new species. There are less than 14 such canals in cones in a reference collection of 15 modern species and in the two fossil species known from anatomically preserved material. While the new species adds to our knowledge of the diversity of Cenozoic Picea, its affinities within the genus remain undetermined.     

A NEW SCIADOPITYACEOUS SEED CONE FROM THE UPPER CRETACEOUS OF HOKKAIDO,JAPAN

A sciadopityaceous seed cone, Sciadopityostrobus kerae, gen. et sp. nov., is described on the basis of a permineralized specimen from the Upper Cretaceous (Cenomanian–Coniacian) of Hokkaido, Japan. The peel method was applied for anatomical observations. The seed cone consists of a cone axis receiving numerous cone scale complexes that are arranged helically. Each complex generally has five inverted ovules that are oriented adaxially. The cone is similar to those of living Sciadopitys verticillata with respect to its peltate cone scale complex, with free apices of both bract and ovuliferous scale, trichomes on the bract, and a trifurcated ovuliferous scale strand. In the fossil, the bract and ovuliferous scale strands fuse with each other in the basal part of the cone scale complex, while S. verticillata bract and ovuliferous scale strands are derived separately from the vascular cylinder and remain separate throughout their length. The present specimen is one of the oldest records of structurally preserved cones that can be assigned to the family Sciadopityaceae.     

Pinus MUTOI (Pinaceae), a new species of permineralized seed cone from the upper Cretaceous of Hokkaido,Japan

Pinus mutoi is described as a new species on the basis of a permineralized seed cone from the Upper Cretaceous of Hokkaido, Japan. The cone is at least 20 cm long and up to 6 cm in diameter, consisting of a cone axis and numerous cone-scale complexes that are arranged helically around the axis. Two winged seeds are borne on the adaxial surface of each ovuliferous scale. Each complex receives a single trace from the vascular cylinder of the cone axis. In the scale base, all the resin canals occur abaxially to the vascular strand. The spatulate bract of the fossil is unique to the specimen among the cones of both living and fossil Pinus. The central umbo, broad sclerotic cortex of cone axis, and absence of serotinous features of the fossil cone suggest affinity with the subsection Sylvestres of the section Pinus, subgenus Pinus. This is the first record of permineralized preserved Pinus cone from the Cretaceous of Eastern Eurasia.     

PINUS AVONENSIS,A NEW SPECIES OF PETRIFIED CONES FROM THE OLIGOCENE OF WESTERN MONTANA

Fossils from the Oligocene of western Montana described in this treatment are the first structurally preserved ovulate cones of Pinus to be reported from the Tertiary of North America. They are about 5.5 cm long and have a maximum diam of 2.5 cm. Numerous scales are arranged spirally around the axis and each scale bears two winged seeds. The bract subtending the ovuli-ferous scale is 3-4 mm long and is free from the scale throughout its length. The pith and cortex of the axis are constructed of thick-walled parenchyma cells and 18-21 resin canals occur at the inner edge of the cortex. Resin canals entering the base of the ovuliferous scale are restricted to the abaxial side with vascular tissues occupying the adaxial side. Vascular strands near the tip of the scale are strongly rounded on the adaxial or phloem side. At the abaxial side of the tip of the ovuliferous scale is a broadly rhomboidal apophysis with a raised umbo that terminates in a short spine. The fossils differ from the several Recent cones examined in having fewer resin canals and biseriate rays in the secondary xylem of the cone axis. The shape of the cone, its anatomical features, and the morphology of the tip of the cone scale indicate affinity with the subgenus Diploxylon.     

A NEW TAXODIACEOUS SEED CONE FROM THE OLIGOCENE OF WASHINGTON

Cunninghamiostrobus goedertii is described as a new species based on permineralized seed cones from the Early Oligocene of Washington. The fossils come from the Makah Formation and were found on the northern shore of the Olympic Peninsula. The cone consists of numerous bract-scale complexes arranged helically around a central axis. Each bract-scale complex has a large bract bearing a small flap of tissue adaxially that represents the ovuliferous scale. Up to three seeds were produced on each complex. The vascular trace to the bract-scale complex diverges from the vascular cylinder of the cone axis as a concentric strand. This divides in the outer cortex of the axis to form a large collateral bract tract abaxially and a minute scale trace adaxially. The latter continues outward to supply the ovuliferous scale. The bract trace divides, forming many strands outward which occur in a row with transfusion tissue between them. Many resin canals also occur in the bract. The new cone combines features found in modern cones of Athrolaxis, Cunninghamia, and Taiwania but is most similar to Cunninghamiostrobus yubariensis from the Late Cretaceous of Japan.     

PICEA DIETTERTIANA,A NEW SPECIES OF PETRIFIED CONES FROM THE OLIGOCENE OF WESTERN MONTANA

Fossils described in this treatment are the first structurally-preserved ovulate cones of Picea to be reported from the Tertiary. They are 5.0-5.8 cm long and 1.6-1.8 cm at their widest diameter. Numerous ovuliferous scales are arranged spirally around the axis and each bore two winged seeds. The bract subtending the scale is 4.0-7.0 mm long and is fused to the scale for 1.0 mm. The base of the bract is inflated on the abaxial side extending for about 1.0 mm between the seeds of the adjacent scales. Both the scales and their subtending bracts are recurved at their point of divergence from the axis. The ovuliferous scales taper gradually to a point, and the thickness of the tissues at the scale apex indicates that they were woody. Anatomically, the silicified cones are very similar to those of the Recent species and indicate that all important features of the latter had evolved by Oligocene time.     

SILICIFIED PINUS REMAINS FROM THE MIOCENE OF WASHINGTON

Silicified leaves, dwarf shoots, pollen cones, and seed cones of Pinus from a Late Miocene chert bed within the Yakima Basalt Formation near Yakima, Washington are interpreted as coming from a single new species, P. foisyi. The needles and dwarf shoots are those of a three-needle pine. The needles contain two to four medial resin canals, a biform hypodermis, and endodermal cells with uniformly thickened walls. The pollen cones are ellipsoidal and about 1 cm long, and many contain bisaccate pollen grains. The seed cones are at least 6 cm long and are slightly asymmetrical. The cone axis has a broad sclerotic outer cortex, and the seed wing extends from a thick parenchymatous base. The scale apex bears a conspicuously swollen projection. The foliage and seed cones are identifiable with the Subgenus Pinus, Section Pinus, Subsection Oocarpae independently of one another, and together indicate a fossil species related to the modem Californian closed cone pines. Pinus foisyi represents one of the earliest occurrences of cone asymmetry associated with this group. However, cone serotiny characteristic of the modem species appears to have evolved after the Late Miocene.     

PETRIFIED CONES AND NEEDLE-BEARING TWIGS OF A NEW TAXODIACEOUS CONIFER FROM THE EARLY CRETACEOUS OF CALIFORNIA

Ovulate cones identified as Abietites ellipticus Fontaine, from the Early Cretaceous of northern California, have been reinvestigated. Rather than being preserved as imprints as originally described, the fossils are petrified. Two cones are attached to needle-bearing twigs. The organs are similar to those of the living Cunninghamia lanceolata. However, the scale portion of the bract-scale complex of the fossil cones is situated close to the bract apex, while that in modern Cunninghamia cones is midway between the bract base and apex. This plus other structural differences warrant treatment of the material as a new species of Cunninghamiostrobus Stopes and Fujii.     

PINUS WOLFEI,A NEW PETRIFIED CONE FROM THE EOCENE OF WASHINGTON

A silicified cone from the Late Eocene of Washington is described as a new fossil species of Pinus. The cone was probably 9–10 cm long and 3–5 cm at its widest diam in the living condition and is peculiar in having abundant resin canals in the secondary xylem of the axis arranged in three concentric rings near the cone base. The bract of the fossil is also unusual in having resin canals of distinctly unequal sizes and a vascular strand that is adaxially concave. In the absence of external features of the scale tips, these anatomical conditions along with the construction of the outer cortex of the axis of thick-walled cells suggest closest affinity of the new species with the subsections Contortae, Oocarpae, and Sylvestres of the section Pinus, subgenus Pinus.     

PITYOSTROBUS PUBESCENS,A NEW SPECIES OF PINACEOUS CONES FROM THE LATE CRETACEOUS OF NEW JERSEY

This new species is based on a single semifusinized cone from the Late Cretaceous (Cenomanian) of New Jersey. The cone is flattened but essentially complete. It is 55 mm long and 8 by 13 mm in diam. Scale apices are thin, rounded, and entire, lacking evidence of an umbo or spine. The vascular cylinder of the cone axis is organized as a series of separate strands. The scale is made up of a basal portion that stands out nearly perpendicular to the cone axis and a sharply upturned distal portion. The bract base has a pronounced abaxial keel. Bract and scale traces diverge from the vascular cylinder of the cone axis separately from one another. A poorly developed interseminal ridge is present at the chalazal end of the seeds. An unusual feature of the cone is the presence of a trichome-bearing epidermis on the cone axis, bract-scale complex, and near the scale apex. Resin canals diverge into the bract-scale complex abaxial to the scale trace with branches becoming adaxial to the scale trace outward. A number of features of the new species occur in cones of Abies, Cedrus, Keteleeria, Larix, Picea, Pseudolarix, Pseudotsuga, and Tsuga as well as in the extinct genus Pseudoaraucaria. Features of Pinus are absent. This suggests that Pseudoaraucaria may have served as an ancestral source for modern genera other than Pinus with Pityostrobus pubescens representing an evolutionary intermediate.     

THE MOTION OF WINDBORNE POLLEN GRAINS AROUND CONIFER OVULATE CONES: IMPLICATIONS ON WIND POLLINATION

Comparisons are presented between the three-dimensional airflow patterns created around and by a scale model of a conifer ovulate cone and the trajectories of windborne pollen grains around Picea, Larix, and Pinus ovulate cones. Three general components of the airflow pattern around an ovulate cone model are 1) doldrum-like eddies, rotating over the adaxial surfaces of cone scales and directed toward attached ovules, 2) airflow spiralling around the cone axis along cone scale orthostichies and parastichies, and 3) a complex pattern of vortices (“umbilicus”) directed toward the leeward surface of the ovulate cone. The observed trajectories of pollen grains around cones of Picea, Larix, and Pinus conform to two of these three airflow components: 1) pollen grains are seen to roll along cone scales toward the distal scale margin and to become reentrained in airflow directed backward toward attached ovules, and 2) pollen grains passing around the cone are deflected into the “umbilicus” airflow pattern, where they either settle on or impact with cone scales (approach trajectories), or where they approach the leeward cone surface but are deflected away by airflow passing under the cone (Z-shaped trajectories). Vectoral analyses of pollen grain motion reveal a complex pattern of trajectories influenced by boundary layer conditions defined by ovulate cone geometry and ambient airflow speed. Wind tunnel studies of ovulate cones subtended by leaves and stem indicate that leaves circumscribing the cone act as a snowfence, deflecting windborne pollen toward the cone. Vectoral analyses of airflow patterns and pollen grain trajectories close to ovulate cones indicate that wind pollination in conifers is a non-stochastic aerodynamic process influenced by cone-leaf morphology and the behavior of pollen grains as windborne particles.     

Great spotted woodpeckers Dendrocopos major exert multiple forms of phenotypic selection on Scots pine Pinus sylvestris

Relatively few animal species extract seeds from closed conifer cones because of the forces required to spread apart or penetrate the woody scales. Those species that forage on seeds in closed cones tend to forage selectively, and therefore act as selective agents on cone structure. However, little is known about the foraging preferences and thus phenotypic selection that is exerted on conifers by many species that forage extensively on seeds in closed cones, including especially woodpeckers (Picidae). Great spotted woodpeckers Dendrocopos major are one of the main predators of seeds in closed cones of Scots pine Pinus sylvestris in central and eastern Europe. To estimate the cone preferences of these woodpeckers foraging on Scots pine, we contrasted traits of cones that were and were not foraged on by woodpeckers. Woodpeckers preferred to forage on shorter cones when scales were thin (smaller apophyses) but preferred cones of intermediate length when scales were thicker, providing evidence for correlational selection. The preference for intermediate‐sized cones indicates that woodpeckers exert disruptive selection on cone length when cones have thicker scales, but the overall selection on cone length across all scale types indicates directional favoring the evolution of longer cones. Woodpeckers avoided cones with thicker scales, which would lead to directional selection favoring the evolution of thicker scales. Preferences for intermediate‐sized cones have been found in tree squirrels and directional selection favoring the evolution of cones with thicker scales may be a common outcome of the foraging behavior of birds.     

PINUS BUCHANANII,A NEW SPECIES BASED ON A PETRIFIED CONE FROM THE OLIGOCENE OF WASHINGTON

A silicified seed cone of Oligocene age from the Olympic Peninsula of Washington represents a new species of Pinus. The cone was about 8 cm long and 3 cm at its widest diameter in the living condition. Its scale apices are thickened and each has a dorsal umbo. Internal cone construction confirms the assignment of the new species to Pinus and suggests affinity with the subsections Australes and Ponderosae of the section Pinus, subgenus Pinus. The cone is peculiar in having a stout bract trace that is slightly concave on its adaxial side and in having resin canals that diverge from the axial secretory system toward the bract but constrict markedly and terminate before entering the bract.     

Reconstruction of the Jurassic conifer Sewardiodendron laxum (Taxodiaceae)

Compressed seed cones and pollen cones of Sewardiodendron laxum are described from the Middle Jurassic of Yima, Henan, central China. They are either organically attached to or associated with leafy shoots. Seed cones are terminally borne. Each cone is ovate to elongated, up to 6.5 cm long and 3.5 cm wide, and consists of a stout axis and numerous helically arranged bract-scale complexes. The bract protrudes beyond and is partially fused with the reduced ovuliferous scale. The ovuliferous scale bears approximately six inverted, small, and flattened seeds. Pollen cones are borne in terminal clusters. Microsporophylls are helically arranged, each bearing three abaxial, basally fused pollen sacs. Pollen is assaccate, rounded, and with an inconspicuous pore. Morphological, structural, and cuticular features of seed cones, pollen cones, and leafy shoots of S. laxum are compared with those of fossil and extant conifers. S. laxum is included in Taxodiaceae and believed to have its closest affinities with a Mesozoic conifer Elatides and a group of Cunninghamia-like conifers. It is reconstructed as a half-evergreen tree that lived in a humid, warm-temperate climate.     

濒危植物梵净山冷杉的球果与种子性状研究北大核心CSCD

梵净山冷杉(Abies fanjingshanensis)为国家一级保护濒危植物。为了揭示梵净山冷杉球果发育和成熟过程的性状特征,以确定成熟球果的适宜采种期。该研究对梵净山冷杉自然生长区内成年结实母树的球果进行跟踪观测和定期采集,通过物理解剖和形态学参数测定,比较分析不同采种期梵净山冷杉的球果、种鳞和种子的性状差异。结果显示:(1)梵净山冷杉的球果每年7月中旬开始形成,10月中旬开始成熟,发育早期至发育后期球果的长度和宽度显著增加,发育后期至成熟期形态和颜色均无明显变化。(2)成熟球果平均长、宽、鲜重、干重和相对含水量分别为7.18 cm、3.84 cm、36.98 g、20.33 g和45.06%。(3)成熟球果的平均种鳞层数为30.76层,种鳞总数为250.67片,平均出种量436.67粒,平均种子饱满率82.49%。(4)成熟球果基部、中部、上部的种子性状不同,且球果中部的种子性状参数最大,饱满率最高,种子平均长、宽、厚分别为9.14、2.30、2.37 mm,千粒重11.44 g。研究表明,梵净山冷杉的球果从形成至成熟过程约3个月,成熟球果的最佳采种时间为10月下旬,发育后期和成熟球果的形态和颜色差异不明显,不能以颜色和形态作为球果成熟的判定依据;种鳞与球果的发育和成熟同步,球果中部的种鳞能够完全发育,且形状参数最大,孕育的种子最饱满;梵净山冷杉成熟球果出种量大、饱满率高,但是种子小、重量轻。