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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

NEW TAXODIACEOUS SEED CONES FROM THE UPPER CRETACEOUS OF NEW JERSEY

Nephrostrobus cliffwoodensis gen. et sp. nov., Nephrostrobus bifurcatus sp. nov., and Rhombostrobus cliffwoodensis gen. et sp. nov. are described based on anatomical studies of seed cone fragments from the Upper Cretaceous Magothy Formation of New Jersey. These species belong to the Taxodiaceae. As in Sequoia, Sequoiadendron. and Metasequoia, the vascular strands supplying the scale and bract in Nephrostrobus are about equal. These vascular strands are arranged in a reniform pattern resembling that found in Metasequoia, but the orientation differs by 180 degress. Nephrostrobus cliffwoodensis and Nephrostrobus bifurcatus differ from each other in the branching of the complex trace and associated resin canals. Rhombostrobus cliffwoodensis resembles Cunninghamia in the arrangement of vascular strands and associated resin canals in its bract-scale complexes. However, the relative amount of bract and ovuliferous scale making up the complex is more like that in Sequoia. This combination of cone features does not occur in any of the extant genera. Nephrostrobus and Rhombostrobus are not considered to be ancestral to any of the extant taxodiaceous genera, but are extinct members of an ancestral complex from which the extant genera were derived.     

CONE AND OVULE ONTOGENY IN TAXODIUM AND GLYPTOSTROBUS (TAXODIACEAE – CONIFERALES)

Seed cones in Taxodium distichum and Glyptostrobus pensilis occupy the position of permanent shoots and are initiated in the summer preceding spring pollination. Morphological features are similar in the two genera, reflecting their close taxonomic relationship. Ovule complexes originate as two (rarely more) ovule primordia in the axil of each fertile bract but without any indication of a preceding discrete ovuliferous scale. When the nucellus, integument, and micropyle are well developed, a series of up to ll abaxial lobes forms at the base of each ovule pair. They become fused by basal growth. After pollination the common basal meristem of lobes and bract extends by intercalary growth to form the conspicuous “ovuliferous scale” of the mature cone; the lobes enlarge and exceed the ovules. Despite the topographic similarity in the cones of both genera, there are differences in vasculature such that the vascular traces to the axillary complex originate directly from the axial cylinder in Glyptostrobus but from the bract trace in Taxodium. The complex vasculature of the mature cone develops late and primarily as an expression of intercalary growth.     

STRUCTURE AND AFFINITIES OF ATHROTAXITES BERRYI BELL,AN EARLY CRETACEOUS CONIFER

Compressions and impressions of leafy twigs, pollen cones, and seed cones of Athrotaxites berryi are abundant in certain layers of the Kootenai Formation (Aptian) in Montana and the Lower Blairmore Formation in adjacent Alberta. The twigs are densely covered by helically arranged leaves that are about 2 mm long and wide. Pollen cones are borne laterally on ultimate branch segments. Some are sessile, while others terminate a minute lateral branch. The cones are 3–4 mm in diam and about 10 mm long. Each sporophyll has a stalk that is about 0.7 mm long and an upturned laminar tip that is 1–1.5 mm long by 1 mm wide. At least two pollen sacs are attached to the abaxial side of each sporophyll. Seed cones are borne terminally on lateral branches that are often curved. These cones are about 10 mm at their widest diameter and about 15 mm long. Each bract and associated ovuliferous scale are fused to form a wedge-shaped complex that is 4–5 mm long. The complex is 0.7 mm wide at its base and expands to about 2.5 mm wide and thick near its apex. The tip of the complex narrows abruptly to a point and terminates in a spine that is about 0.5 mm long. At least one seed occurs on the adaxial side of each complex. Athrotaxites berryi belongs to the Taxodiaceae. It resembles modern Athrotaxis cupressoides but differs from it in too many aspects to be included in the modern genus.     

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.     

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.     

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.     

A taxodiaceous seed cone from the Triassic of Antarctica

A silicified seed cone is described from the lower Middle Triassic of A silicified seed cone is described from the lower Middle Triassic of Antarctica. The cone measures up to 3.4 cm long and 1.4 cm wide, and consists of helically arranged cone scales attached to a eustelic axis. Bract and ovuliferous scale are approximately of equal length and fused at the base. The bract is entire and vascularized by a single trace. The ovuliferous scale contains five distal lobes, each vascularized by a terete strand that divides to form a smaller trace to each of the five inverted ovules. Ovules are small and flattened with the three-parted integument attenuated into oppositely positioned lateral wings. The Triassic specimens are compared with both extant and fossil conifer seed cones and believed to have their closest affinities within the Taxodiaceae.