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.     

STEMS AND LEAVES OF CUNNINGHAMIOSTROBUS GOEDERTII FROM THE OLIGOCENE OF WASHINGTON

This report is based on nine specimens of fossil conifer stems and leaves from the Early Oligocene Jansen Creek Member of the Makah Formation. They were collected along the northern shore of the Olympic Peninsula of Washington. The fossils are preserved as siliceous permineralizations and were exposed in surface view along rock fractures. Details of leaf morphology and epidermal construction appear on fracture surfaces of certain specimens while the cellular construction of the leaves and twigs is visible in thin sections. Leaves are dorsiventrally flattened, attached to twigs that contain up to four growth increments of secondary xylem, up to 2.3 cm long, 3.5 mm wide, and have parallel margins with minute teeth. The leaves are about 0.5 mm thick and have a central vascular strand surrounded by transfusion tissue. A large resin canal occurs abaxial to the vascular strand, and generally two additional resin canals occur in the mesophyll near each leaf margin. Leaves are mostly hypostomatic, with sunken stomata in two longitudinal bands, one to each side of the midline of the leaf and each containing eight to 13 longitudinal rows of stomata. Several unusual anatomical features in the stems also occur in the peduncle and cone axis of seed cones described as Cunninghamiostrobus goedertii, which occurs at the same locality. Thus, the leafy twigs belong to the same species as produced the cones. The cones, leaves, and shoots of this Tertiary conifer are similar to those of modern Cunninghamia but differ from the living species in several respects.     

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.     

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.     

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.     

PETRIFIED PLANTS FROM THE UPPER MISSISSIPPIAN OF NORTH AMERICA. II. LEPIDOSTROBUS FAYETTEVILLENSE SP. N.

Structurally preserved arborescent lycopsid fructifications are described from the Fayetteville Shale (Upper Mississippian) of northwestern Arkansas. Specimens of Lepidostrobus fayettevillense sp. n. range from complete cones 22.5 cm long and approximately 1.0 cm in diameter to smaller water-worn fragments. The cones consist of a central axis bearing closely spaced, spirally arranged sporophylls which extend from the cone axis at right angles. Each sporophyll consists of a pedicel which is turned up at its end to form a distal lamina. Sporangia are large and attached to the adaxial surface of each sporophyll. The vascular cylinder consists of a centrally located exarch protostele at least 1.1 mm in diameter. The new species is compared with morphologically similar lycopsid fructifications of equivalent age.     

A NEW JURASSIC ISOETITES (ISOETALES) FROM THE WALLOWA TERRANE IN HELLS CANYON,OREGON AND IDAHO

A new species of the quillwort Isoetites, I. rolandii sp. nov. Ash and Pigg, is described from the Middle Jurassic Coon Hollow Formation in the Wallowa terrane in Hells Canyon, Oregon and Idaho. The new species is based on coalified impressions and mold-casts of an isoetaceous corm with narrow, elongate leaves, and isolated masses of leaves that have been studied using light microscopy. Plants are about 10 cm in maximum height and consist of cormose plant bases from 1.5 to 2.5 cm wide and 0.6 cm high bearing incomplete microphyllous leaves up to 8.4 cm long and 3.0 mm wide, that taper to about 2.0 mm wide distally. Associated casts of sporophyll bases and sporangia contain spherical structures about 440 μm in diameter that may represent megaspores. Based on the occurrence of complete plants, some specimens of I. rolandii are thought to have been preserved in situ. The new Isoetites provides the first floristic evidence for the occurrence of an aquatic or semiaquatic paleoenvironment in the Wallowa terrane. It is also the first documented occurrence of Isoetes-like plants in the Jurassic of western North America.     

Reinvestigation of Nystroemia pectiniformis Halle, an enigmatic seed plant from the Upper Permian of China

Reinvestigation of Nystroemia pectiniformis Halle from the Upper Shihhotze Formation of Shanxi Province, China, has led to the identification of new and important features of this enigmatic Late Permian seed plant, permitting its typification and diagnosis. After reassembling several of the previously studied specimens to form a single articulated branching system comprising at least four orders of branching, previously unknown features of its branching pattern and morphology have been characterized. First–order axes are wide and branch to one side only, bearing second–order branches either singly or in pairs and of two kinds: one fertile and bearing characteristic ovulate branching systems and the other presumably vegetative. Ovulate second–order axes are narrow and branch to one side only, producing numerous, closely spaced lateral branches in two alternate to sub–opposite rows. Lateral branches are slender and produce numerous ovulate branching systems to one side of the axis only. Ovulate branching systems divide unequally to produce 3–15 ultimate axes of different lengths that are planated. Each ultimate axis bears a single terminal ovule with 180 degree rotational symmetry and two horn–like integumentary projections distally. The other kind of second–order axes are distinct from those bearing ovules; they are wider and longer and branches occur on both sides of the secondary axis, lacking divisions in close proximity to the first–order axis. These have only been observed incomplete although their distinct morphology indicates they are unlikely to be ovulate branches from which ovules/seeds have been shed. Additional organs of the Nystroemia plant are considered, including pollen organs previously assigned by Halle to the same species (displaying its characteristic branching style), and also leaves of Chiropteris reniformis Halle that were probably borne on the larger kind of second–order branches. Implications of Nystroemia on seed plant evolution and distribution are discussed, and it is concluded that this most likely represents a late stratigraphic occurrence of a plesiomorphic hydrasperman–type seed plant with affinities closely allied to members of the Lyginopteridales.     

A new species of the extinct genus Austrohamia (Cupressaceae s.l.) in the Daohugou Jurassic flora of China and its phytogeographical implications

Abstract A new conifer, Austrohamia acanthobractea, sp. nov., is described from the Jurassic Daohugou flora, Inner Mongolia Autonomous Region, China. The material consists of impressions represented by well‐preserved leafy twigs and branches as well as ovulate cones. Leafy shoots with at least two orders of branching; ultimate branchlets alternate or sub‐opposite with helically arranged leaves, decurrent at base with distal rounded tip; dorsiventrally flattened and univeined. Ovuliferous cones elliptical, less than 1 cm long, terminally borne on ultimate and penultimate branches, composed of helically arranged bracts with ovules disposed on their adaxial surfaces. The presence of similar, if not identical taxa, on both sides of the Pacific indicates the cosmopolitan distribution of primitive Cupressaceae between East Asia (Eurasia) and South America in the Pangaea.     

HALLETHECA RETICULATUS GEN. ET SP. N.: A SYNANGIATE PENNSYLVANIAN PTERIDOSPERM POLLEN ORGAN

A new structurally preserved synangiate pollen organ is described from the upper Pennsylvanian (Mattoon Formation) of southeastern Illinois. The specimen of Halletheca reticulatus gen. et sp. nov. measures approximately 1.5 cm long by approximately 5.0 mm in diam, and consists of five elongate sporangial tubes equidistantly arranged around a solid central column, and embedded in a thin-walled ground tissue. The vascular system consists of five terete bundles characterized by scalariform tracheids. Sporangia are thick-walled and contain pollen of the Monoletes-type. Ultrastructural studies of the pollen grain wall show it to consist of a uniform network of muri which branch and re-unite to form small lumina. A comparison of the ultrastructure of the pollen grain wall of the new fructification is made with the wall organization found in similar pollen of Dolerotheca. Relationships between the new taxon and other presumed seed fern pollen organs differing in preservation mode are discussed.     

Permineralized seed fern cupules from the Triassic of Antarctica: implications for cupule and carpel evolution

In this paper we describe the first anatomically preserved Mesozoic seed fern cupule–Petriellaea. The multiovulate cupules were produced singly at the end of a short dichotomizing axis. Cupules are bilateral with a dorsal groove and transverse narrow ventral opening. The vascular system of the cupule consists of a series of traces that extend up the dorsal surface of the cupule and down the ventral face. Ovules are orthotropus, sessile, and borne on the adaxial surface of the leaflike cupule either singly or in multiple rows. They are up to 1.5 mm long, triangular in transverse section, and characterized by a multilayered integument. Nucellus and integument are fused throughout their length, but no pollen chamber is present. In the chalaza is a small vascular disc of transfusion tracheids that represents the extent of the ovule vascular system. Ovules are interpreted as being fossilized at a prepollination stage, although a few possess some evidence of a cellularized megagametophyte. These permineralized cupules indicate that in at least one Mesozoic seed fern group, ovule enclosure resulted from the transverse folding (tip to petiole) of a megasporophyll bearing adaxial ovules. Cupule morphology and ovule enclosure in other Late Paleozoic and Mesozoic seed ferns is discussed.     

Elatides sandaolingensis n. sp. (Cupressaceae sensu lato) — A new fossil conifer with cones from the Middle Jurassic of Xinjiang,northwestern China

Here we describe a new conifer fossil, Elatides sandaolingensis Z.X. Wang and B.N. Sun n. sp., from the Middle Jurassic Xishanyao Formation in the Turpan-Hami Basin, Xinjiang Uygur Autonomous Region, northwestern China. The materials consist of compressions represented by well-preserved leafy shoots, pollen cones, and seed cone. Leaves are characterized by long triangular shapes, with straight apex and entire margins, and two stomatal bands on the abaxial surface. Pollen cones are terminally disposed on the ultimate leafy shoots, borne singly or in clusters. Pollen sacs are long-oval shaped, with three pollen sacs fused together. Pollen grains are spherical and have small germinal papilla and few wrinkles. The seed cone is oblong, with more than 35 helically arranged bract-scale complexes, which are characterized by long-oval shape and triangular apex. Compared with the extant nine genera of Taxodiaceous Cupressaceae in the morphology of seed cone and pollen cones, the present fossil consistently shares many characteristics with the extant genus Cunninghamia, but differs in other aspects. After being compared with the reported fossil records of the fossil plants, the current species is found to be different from any known species; thus, the present fossil is referred to as a new species of Elatides. From the similarity between the present fossil and Cunninghamia, it can be inferred that there may be a genetic relationship between these two genera. Additionally, the new species has thin cuticles and slightly sunken stomata, which can provide evidence indicating that the climate of the Turpan-Hami Basin in Middle Jurassic might have been warm and humid. By studying the geological history and geographical distribution of Elatides, it can be inferred that this genus may have originated in Switzerland, and it was migrated from Xinjiang to North and South China through the Ural Mountains.     

PERMINERALIZED FRUITS AND SEEDS FROM THE PRINCETON CHERT (MIDDLE EOCENE) OF BRITISH COLUMBIA: ARACEAE

Some of the most conspicuous fruit and seed remains from the Middle Eocene Princeton chert locality (Allenby Formation) are assignable to the Araceae. Several thousand campylotropous, reniform, spiny seeds 2.5–3.2 mm long x 1.8–2.3 mm wide have been found dispersed in the chert. Two fruits, each with a single locule and at least eight ovules, show three distinct wall layers and appear to have been fleshy. The two layers of seed integument have pitted isodiametric sclereids aligned in radial rows, the outermost being thinner walled. Spines are borne in three rows on the dorsal seed surface while the ventral side is flattened. Idioblasts that probably contained raphides are present in the outer integument. Nucellar tissue is attached to the integument for most of its length and is well-preserved in some seeds near the chalaza and in the large conical shaped area beneath the micropyle. Endosperm cells with dark contents and curved, linear, monocotyledonous embryos are present in some seeds. Sections of seeds of living araceous taxa of the subfamilies Monsteroideae (Epipremnum, Rhodospatha) and Lasioideae (Urospatha, Cyrtosperma) were made for comparison. The fossil seeds represent a new taxon most closely related to Cyrtosperma, Keratosperma allenbyensis Cevallos-Ferriz et Stockey gen. et sp. nov., Family: Araceae, Subfamily: Lasioideae, Tribe: Lasieae. The Princeton remains are the oldest described seeds of this tribe known to date and add to our knowledge of the subtropical elements of the Princeton flora.     

POROSTROBUS NATHORSTII SP. NOV.: A NEW SPECIES OF LYCOPSID CONE FROM THE EARLY PENNSYLVANIAN OF ILLINOIS

Thirty-one specimens of a small megasporangiate lycopsid cone referable to the genus Porostrobus Nathorst and abundant associated dispersed megaspores have been collected from Early Pennsylvanian strata in the Allied Stone Company quarry, Milan, Illinois. Based on other elements in the flora, the deposit is considered to be part of the Morrowan Caseyville Formation and probably of Namurian age. This is the first reported occurrence of Porostrobus in North America and the cones are recognized as a new species, P. nathorstii. The environment of deposition indicates that the cones may have been transported from the parent plant prior to preservation. Cones are preserved as coalified compressions measuring 15–36 mm long by 2.5–7 mm wide, and are characterized by an apical tuft of leaves up to 20 mm long. Sporophylls are spirally arranged on a narrow cone axis, lack a heel or keel, and have a long distal lamina. Sporangia contain a single functional megaspore tetrad. Mature megaspores are 750–1, 150 μm in diameter, have prominent trilete sutures raised to form a gula, and have numerous branched hairs confined to an equatorial band. Megaspores correspond to the dispersed form Setosisporites praetextus (Zerndt) Potonie and Kremp. Porostrobus nathorstii is the only species of the genus described to date that is monosporangiate.     

HETEROGENEITY IN POLLEN ALLELE FREQUENCIES AMONG CONES,WHORLS, AND TREES OF TABLE MOUNTAIN PINE (PINUS PUNGENS)

Pinus pungens Lambert (Table Mountain pine) is characterized by serotinous cones that are borne several (two to five) per whorl. Sporophytic and gametophytic tissues from whorls collected in different canopy zones of eight trees were electrophoresed and surveyed for six allozyme loci. Pollen allele frequencies received by individual trees, whorls on trees, and cones in whorls were estimated and used to estimate outcrossing rates and the distribution of genetic diversity within and among each level. Significant differences in pollen allele frequencies were detected at all levels, indicating that the genetic composition of the pollen pool available to different female strobili is heterogeneous. The fine-scale genetic structure in the seed pool is a record of past reproductive events and is the precursor to genetic structure in the next generation.     

MITROSPERMUM VINCULUM SP. NOV., A CARDIOCARPALEAN OVULE FROM THE UPPER PENNSYLVANIAN OF OHIO

Numerous anatomically preserved ovules assignable to the genus Mitrospermum have been discovered in Upper Pennsylvanian sediments of Eastern Ohio. Although basically similar to Mitrospermum compressum, the newly discovered specimens exhibit several consistent differences. Ovules are strongly platyspermic, up to 4.2 mm long, 4.0 mm wide, and 0.6 mm thick. In the minor plane, ovules are broadest at the base and taper toward the micropyle. The integument exhibits three topographic regions: endotesta, sclerotesta, and sarcotesta. The sarcotesta is extremely broad in the major plane, where it forms two membranous wings. A single terete vascular bundle enters the base of the ovule, traverses the integument, and divides to form two integumentary bundles and a conspicuous nucellar platform. Integumentary bundles extend toward the tip of the ovule at the margin of the sarcotesta and sclerotesta. A pollen chamber with a prominent nucellar beak is delimited at the tip of the nucellus. Consistent differences in vascularization, size, nature of the seed base, features of the pollen chamber, and the Late Pennsylvanian age demonstrate that the specimens represent a distinct species. The discovery of these ovules extends the stratigraphic range of Mitrospermum to the Upper Pennsylvanian of Ohio.     

The actin microfilament network within elongating pollen tubes of the gymnospermPicea abies (Norway spruce)

Summary Actin microfilaments form a dense network within pollen tubes of the gymnosperm Norway spruce (Picea abies). Microfilaments emanate from within the pollen grain and form long, branching arrays passing through the aperture and down the length of the pollen tube to the tip. Pollen tubes are densely packed with large amyloplasts, which are surrounded by branching microfilament bundles. The vegetative nucleus is suspended within the elongating pollen tube within a complex array of microfilaments oriented both parallel to and perpendicular with the growing axis. Microfilament bundles branch out along the nuclear surface, and some filaments terminate on or emanate from the surface. Microfilaments in the pollen tube tip form a 6 m thick, dense, uniform layer beneath the plasma membrane. This layer ensheathes an actin depleted core which contains cytoplasm and organelles, including small amyloplasts, and extends back 36 m from the tip. Behind the core region, the distinct actin layer is absent as microfilaments are present throughout the pollen tube. Organelle zonation is not always maintained in these conifer pollen tubes. Large amyloplasts will fill the pollen tube up to the growing tip, while the distinct layer of microfilaments and cytoplasm beneath the plasma membrane is maintained. The distinctive microfilament arrangement in the pollen tube tips of this conifer is similar to that seen in tip growth in fungi, ferns and mosses, but has not been reported previously in seed plants.     

Western white pine (Pinus monticola Dougl.) reproduction: I. Gametophyte development

Male and female gametophyte development are described from light and transmission electron microscope preparations of ovules from first and second year Pinus monticola Dougl. seed cones. In the first year of development, pollen tubes penetrate about one-third the distance through the nucellus. The generative cell and tube nucleus move into the pollen tube. The megagametophyte undergoes early free nuclear division. First-year seed cones and pollen tubes become dormant in mid-July. In the second year, seed cones and pollen tubes resume development in April and the pollen tubes grow to the megagametophyte by mid-June. Early in June the generative cell undergoes mitosis, forming two equal-size sperm nuclei that remain within the generative cell cytoplasm. The generative cell has many extensions and abundant mitochondria and plastids. The megagametophyte resumes free nuclear division, then cell wall formation begins in early July. Cell wall formation and megagametophyte development follow the pattern found in other Pinaceae. Three to five archegonial initials form. The primary neck cell divides, forming one tier of neck cells. Jacket cells differentiate around each central cell. The central cell enlarges and becomes vacuolate; then vacuoles decrease in size and the cell divides, forming a small ventral canal cell and a large egg. Plastids in the central cell engulf large amounts of cytoplasm and enlarge. This process continues in the egg, and the peripheral cytoplasm of the egg becomes filled with transformed plastids. Mitochondria migrate around the nucleus, forming a perinuclear zone. The wide area of egg cytoplasm between these two zones has few organelles. A modified terminology for cells involved in microgametophyte development is recommended. Received: 9 December 1999 / Revision accepted: 30 April 2000     

PICEA WOLFEI,A NEW SPECIES OF PETRIFIED CONE FROM THE MIOCENE OF NORTHWESTERN NEVADA

Several silicified ovulate cones from the late middle Miocene (Barstovian) represent a new species, Picea wolfei Crabtree. This is the second species of Picea for which structurally preserved seed cones are known to be reported from the Tertiary. The cones are 5.0–8.0 cm long and 1.5–2.0 cm at their greatest diameter. Ovuliferous scales are inserted helically around the cone axis and are recurved at their point of divergence. Each scale is broadly obovate to spatulate with a rounded apex and bore two seeds adaxially. The bract subtending the scale is 4.5–7.3 mm long and is fused to the scale for 1.4–2.0 mm. Each bract has an inflated keel-like base which projects abaxially between the seeds of adjacent scales. The fossil cones superficially resemble those of the extant Picea breweriana, yet differ from them anatomically. The new species also resembles Picea lahontense, a fossil compression from the Miocene Trout Creek Flora of south-central Oregon, but the different modes of preservation preclude meaningful comparison. Picea diettertiana, the only structurally preserved fossil cone of this genus previously described, is quite dissimilar in that it lacks a sclerotic pith.     

Saussurea pseudograminea sp. nov. (Asteraceae) from the Qinghai–Tibetan plateau,China

Morphological characteristics, as well as ultrastructure of pollen grains, chromosome numbers and karyotype analysis have been used to establish a new species of Saussurea from the Qinghai–Tibetan plateau. The new species, Saussurea pseudograminea Y. F. Wang, G. Z. Du et Y. S. Lian is easily distinguished from the similar S. graminea Dunn by having 2–3 capitula, involucre 0.7–1.2 cm in diameter, smaller pollen grains, pollen surface with larger and denser spines, achenes 4.0–5.5 mm long, 32 chromosomes, and a karyotype formula 2n = 2x = 32 = 18m + 10sm + 4st, whereas S. graminea has solitary capitula, involucre 1.2–2.0 cm in diameter, larger pollen grains, pollen surface with smaller and sparser spines, achenes 3–4 mm long, 28 chromosomes and a karyotype formula 2n = 2x = 28 = 6m + 20sm + 2st. The new species is distributed in Dianzhangou, Awanchang, and Gamaliang mountain regions of Maqu county in Gansu province.