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.     

Turonian Pinaceae of the Raritan Formation, New Jersey

 Fossil vegetative and reproductive structures from deposits of the Raritan Formation in New Jersey (Turonian, Upper Cretaceous, ∼90 MYBP) include ferns, gymnosperms, and angiosperms. Gymnosperms collected from this formation have been known since the beginning of the 20th century. Fossil leaves, wood and seed cones have been are identified as belonging to the Cupressaceae, Pinaceae, and Taxodiaceae. In the present contribution, we describe a series of fossil dwarf shoots, leaves and a pollen cone belonging to the family Pinaceae. Fossils are charcoalified with preserved three-dimensional structure and excellent cellular detail. The dwarf shoots are assigned to a new species Prepinus crossmanensis sp. nov. and to the previously described Pinus quinquefolia Jeffrey. The new species Prepinus crossmanensis differs in size, shape, presence of hypodermis, sclerenchyma and stomata in the cataphylls, and number and shape of needle leaves from previously known species. Also, isolated leaves were found that were assigned to the new species, Prepinus raritanensis sp. nov. The new species is differentiated by the size and shape of stomata, the presence of number of layers of the hypodermis; and the cell-shape and number of layers of the mesophyll and transfusion tissue. A previously undescribed male cone, Amboystrobus cretacicum gen. and sp. nov., has an axis with spirally attached microsporophylls, each bearing two abaxial ovoid microsporangia. The pollen grains are monosulcate and bisaccate (eusaccate), with an elliptical corpus, granulate exine sculpture, and honeycomb-like wall structure. Received March 21, 2000 Accepted November 13, 2000     

SEASONAL SECONDARY GROWTH IN NEEDLE LEAVES OF PINUS STROBUS AND PINUS BRUTIA

Mature needles and elongating current year's needles of Pinus strobus growing in Massachusetts and P. brutia growing in Israel were collected monthly or bimonthly for seasonal analysis of leaf cambial activity. Mature needles produced secondary phloem but no xylem, and, regardless of the season, had a cambial zone from 2 to 3 cell layers wide. In the current year's needles maturation was basipetal and the procambium differentiated into primary xylem, primary phloem, and the phloem-producing vascular cambium before needle maturity. One- and 2-year-old needles of Pinus strobus produced slightly over 4 cell layers of phloem between April 15 and September 1 of 1983, with a peak production rate of about 2 cell layers per month in May and early June. One-year-old needles of P. brutia produced about 6 phloem cell layers in 1983, with phloem being produced throughout the year except in midsummer. This was contrasted by fall and winter dormancy in needles of P. strobus.     

THE FATE OF THE DWARF SHOOT APEX IN BRISTLECONE PINE (PINUS LONGAEVA)

The fate of the pine dwarf shoot (DS) apex after needle initiation has been controversial. Dwarf shoot primordia of Pinus longaeva were examined to determine the developmental basis for DS with and without interfoliar buds. Interfoliar buds are microscopic buds derived from the original terminal apex of the DS. In October, all the DS primordia are similar in size and appearance. However, as the needles elongate in the following June the apices of more proximal DS decrease in size, such that by July there is a clear diminishing size gradient of apical domes in going from the most distal to the most proximal positions. The distal DSs start to form bud scales in July and have fully formed interfoliar buds by mid-August. In contrast, those DS apices lacking protective bud scales at needle maturity become suberized and can never proliferate into long shoots. The distal placement of interfoliar buds may be due to a group effect, where each developing DS inhibits the more proximal DSs in the long shoot terminal bud.     

MOBILIZATION BY PINUS RESINOSA CONES AND SHOOTS OF C14-PHOTOSYNTHATE FROM NEEDLES OF DIFFERENT AGES

Current-year, 1-year-old, 2-year-old, and 3-year-old needles of 25- to 30-year-old Pinus resinosa trees were separately exposed to C¹⁴O₂ at various times during the growing season. Currently produced C¹⁴-photosynthate was preferentially mobilized in the following order: second-year cones > current needles > current internodes > first-year conelets. A changing seasonal pattern was shown in sources of current photosynthate for growth of cones and shoots. One-year-old needles were the major source of current photosynthate for growth of both cones and shoot internodes. During June the 2- and 3-year-old needles contributed appreciable amounts of current photosynthate to both cones and developing shoots, but after late June their contribution was slight. The supply of carbohydrates to all tissues, except 2nd-year cones, from the three age classes of old needles declined late in the season as tissues mobilized increasingly more carbohydrates from current-year needles. Nevertheless, the bulk of the C¹⁴-photosynthate produced by current-year needles was retained by them. The preferential mobilization of carbohydrates by reproductive tissues over vegetative tissues is emphasized as is the importance of both reserve and currently produced carbohydrate for growth of various tissues.     

The morphology,taxonomy, and occurrence of the genus <Emphasis Type="Italic">Pseudoprotophyllum</Emphasis> Hollick (Platanaceae) in Late Cretaceous floras of Northern Asia

An extended generic diagnosis of Pseudoprotophyllum is given on the basis of new data from Late Cretaceous floras of Northern Asia. Both peltate and apeltate morphological types are included in the genus. The morphological diversity is described in the type species P. boreale (Dawson) Hollick from the Cenomanian-Turonian of North America and four North Asiatic species: P. minimum I. Lebed. , P. sibiricum I. Lebed., P. hatangaense Abramova, and P. giganteum Sveshn. et Budants. The geographical and stratigraphic analyses show that Pseudoprotophyllum evolved in Northern Asia since the Cenomanian until Campanian and was restricted to the Siberian-Canadian floristic region with a warm-temperate humid climate.     

云南松上散斑壳属一新种

根据子囊果的形态及其在基质中的位置和松针上线纹的特征,鉴定出四川二郎山云南松针上散斑壳属的一个新种,即四川散斑壳(Lophodermium sichuanense D.X.Qiu et Liu)。文中对该新种的形态特征作了汉文和拉丁文描述。     

PERMINERALIZED PINACEOUS LEAVES FROM THE UPPER CRETACEOUS OF HOKKAIDO

Three fragments of fascicles of a possible five-needle pine are described from the Upper Cretaceous Yezo Group (Santonian/Senonian) of Hokkaido. Specimens from the Omakizawa, Oyubari, Yubari City, the Koyanozawa, Ikushumbetsu, Mikasa City, and the Sankebetsugawa, Haboro are preserved in calcium carbonate nodules containing abundant ammonites. Leaves borne in apparent fascicles of five measure 0.7–0.8 mm in radial and 0.8–1.0 mm in tangential diam and are represented by short fragments of isolated needles. Thick-walled epidermal cells on these amphistomatic leaves resemble the underlying uniform hypodermis. Two external resin canals are situated near the lateral corners toward the adaxial surface. Only slightly plicate mesophyll cells in a layer one to two cells thick border on an elliptical endodermis with thickened outer cell walls. Two to three layers of transfusion tissue surround the double vascular strand. Two vascular bundles are separated by one cell layer of sclerenchyma fibers. Small patches of abaxial and adaxial sclerenchyma fibers have also been observed. Leaves most closely resemble those of Pinus leiophylla Schl. et Cham. Subgenus Pinus, Section Pinea, Subsection Leiophyllae and P. montezumae Lamb. Subgenus Pinus, Section Pinus, Subsection Ponderosae and are described as a new species P. hokkaidoensis sp. nov. Stockey and Ueda. Close anatomical comparisons are made with this leaf and previously described permineralized Upper Cretaceous pine needles from Hokkaido and eastern North America.     

Anatomical changes with needle length are correlated with leaf structural and physiological traits across five Pinus species

The genus Pinus has wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade‐offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species: Pinus echinata, Pinus elliottii, Pinus palustris, Pinus taeda, and Pinus virginiana. We found that increasing needle length contributed to a trade‐off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity.     

Effects of previous defoliation on pine looper larval performance

1 During outbreaks of the pine looper, Bupalus piniarius, its host, Pinus sylvestris, is severely defoliated. The larvae of this geometrid normally feed on mature needles. However, because trees are totally defoliated during outbreaks, the next generation is forced to feed on current-year needles. 2 Bupalus piniarius larvae feeding on previously defoliated trees may show lower performance either because of induced resistance or because larvae have to feed on needles not normally fed upon (current instead of mature). 3 These hypotheses were tested in an experiment where larvae were reared on (i) shoots naturally defoliated the previous year, and thus, bearing only current-year needles, (ii) non-defoliated shoots where larvae had access only to current-year needles, and (iii) control shoots with access to both current and mature needles. 4 There was no support for the induction hypothesis. Survival was lower on naturally defoliated shoots than on control shoots (81.3 vs. 90.9%), but survival was lower also on non-defoliated shoots where larvae had access only to current-year needles (78.8%). Data on larval feeding distribution showed a strong preference for mature needles. 5 Needle nitrogen concentration of current-year needles was 38% higher on defoliated trees than on non-defoliated trees. 6 It is concluded that defoliation affected larval performance primarily through the removal of the preferred type of needles and not because of an induced resistance. Effects of increased concentrations of allelochemicals in defoliated shoots, if present, were probably cancelled out by increased nitrogen concentrations.     

SECONDARY GROWTH IN NEEDLE LEAVES OF PINUS LONGAEVA (BRISTLECONE PINE) AND OTHER CONIFERS: QUANTITATIVE DATA

Needle leaves of Pinus longaeva can be accurately dated and remain alive on branches for 30 or more yrs, making this species ideal to study secondary growth in leaves. Field observations and regression analysis of needle age versus mean needle length both indicate primary (elongation) growth of needles is completed in the first year. Statistical analysis of cell counts for one- to 33-yr-old needles indicate production along the entire length of needles of 1.0-1.7 cell layers per year of secondary phloem, but no secondary xylem. Microscopic measurements and cell counts reveal that with advancing needle age phloem increases radially and transfusion tissue buckles, but the number of endodermal cells and xylem width do not change. Living phloem remains constant in amount (ca. 9 layers) with advancing needle age, indicating yearly replacement of old by new phloem. For comparison to P. longaeva, needle leaves were also studied for ten other conifer taxa with maximum needle longevities ranging from 3 to 19 yrs. In needles of each taxon no secondary xylem is produced, but secondary phloem production occurs throughout the post-elongation lifespan of the needles regardless of maximum needle longevity.     

Cretaceous and Eocene fossils of the rare extant genus Synneuron Lundstrom (Diptera: Canthyloscledidae): evidence of a true Pangean clade

The first two fossil species of the canthyloscelid genus Synneuron are described based on compression wings. Synneuron eomontana sp. nov. is described from the Middle Eocene Coal Creek Member of the Kishenehn Formation, in the USA, and Synneuron jelli sp. nov. is described from the Lower Cretaceous Koonwarra Fossil Bed of the Korumburra Group, in Australia. The wings are illustrated and compared to the extant species of the genus, to species of the three other recent genera of Canthyloscelidae and to an anisopodid. A phylogenetic analysis of the relationships between the species of Synneuron was performed. The Eocene fossil S. eomontana appears as sister of the pair of recent Holarctic species of the genus, while the Australian Cretaceous species S. jelli is sister of the clade with the species of Synneuron of the northern hemisphere. The sister group of Synneuron is the canthyloscelid clade (Hyperoscelis + Canthyloscelis), for which a middle Jurassic fossil is known. At the early Cretaceous, Gondwana was already separated from Laurasia and the disjunction between the species of Synneuron in Australia and the northern hemisphere clade of the genus suggest a true pangeic origin for the genus. The biology of the canthyloscelid larvae is shaped by its trophic specialization—xylosaprophagous. This suggests that the transition from the Pangean Jurassic gymnosperm-dominated forests to the late Cretaceous angiosperm-dominated forests may be related to the low recent diversity of Synneuron or of the canthyloscelids in the world—and maybe to the extinction of the genus in the southern hemisphere. This major turnover of the vegetation type along the Cretaceous may be also somehow related to the complete extinction of other groups of flies strictly associated with gymnosperms, as may be the case of the lower brachyceran family Zhangsolvidae. This speculation needs additional corroboration from other groups, that will become available with the combination of systematics, paleontology and biogeographical information of different early Cretaceous clades.     

Longevity of needle fascicles of Pinus longaeva (Bristlecone pine) and other North American pines

Summary Variation in leaf longevity of gymnosperms has received surprisingly little attention despite its likely adaptive significance. Pinus longaeva, a pine of arid, subalpine environments in the western United States, has the record among conifers for needle longevity, with a maximum dwarf shoot (needle fascicle) retention time of up to about 45 years. Most low elevation pines have dwarf shoot retention times (DSRs) of two to four years. Literature data for the 37 species of Pinus native to the United States and Canada and field data for eight taxa (21 populations) of pines growing at various elevations in California each show a strong positive correlation between elevation and DSR, respectively, r=+0.65, df=35, p<0.001 and r=+0.82; df=19, p<0.001. We propose that extended needle fascicle longevity represents an adaptation to arid and especially high elevation environments. Field data from native stands and common gardens indicate that differences between taxa in DSR relate to both genetic and environmental factors. When grown at the same sites certain species (eg, P. longaeva, P. monophylla) had much longer DSRs than others, indicating a genetic basis for differences in needle fascicle longevity. For six of seven taxa that were each studied at more than one elevation there was a statistically significant increase in DSR in going from the lowest to the highest elevation site, indicating strong environmental control of needle fascicle longevity.The physiological control of dwarf shoot senescence and abscission is poorly understood. For P. longaeva dwarf shoots of a particular age class are not shed simultaneously; rather there is a more or less gradual attrition of dwarf shoots from the long shoot. Although different types of long shoots of pines are known to differ physiologically, for P. longaeva there was no consistent difference in DSR on various types of lateral long shoots (eg, vegetative, pollen cone-bearing, seed cone-bearing), nor was there a statistically significant difference in DSR on trunks versus on their lateral long shoots. In addition, for P. contorta ssp. bolanderi and P. muricata needle fascicle longevity was not affected by the degree of edaphically induced dwarfing (ie, stunting) of the trees.     

Reappraisal of two of Witham's species of fossil wood with taxonomical and nomenclatural notes on Planoxylon Stopes, Protocedroxylon Gothan and Xenoxylon Gothan

Fossil wood collections at the Natural History Museum, London, were searched for the type material of two important species, described as early as 1831 by Witham, on the basis of material at least partly furnished by Nicol and collected in the Liassic of Whitby (Yorkshire, UK). The names given to these two species by Witham are the basionyms for more than ten other species names, while these species names also provide syntypes for several generic names. Despite this, the original material has been only rarely and partly revised, leading to much confusion.We managed to locate some of the original material. Several topotypes were also studied. On the basis of this review we propose here a neotypification for Peuce huttoniana Witham, a lectotypification for Cupressinoxylon barberi Seward and a lectotypification for Tiloxylon Hartig. We evidence several taxonomical and nomenclatural synonymies, assign Peuce lindleyana Witham to Protocedroxylon and P. huttoniana to Xenoxylon Gothan, and introduce the new combination Xenoxylon huttonianum (Witham) nov. comb. as the correct name for Xenoxylon ellipticum Gottwald & Holleis ex Schultze-Motel.A topotype of Araucariopitys americana Hollick & Jeffrey was also reviewed, which, together with previous results, leads us to suggest that Araucariopitys Hollick & Jeffrey, Protocedroxylon Gothan and Planoxylon Stopes should not be considered as taxonomical synonyms. This point is crucial to the interpretation of the fossil record of Early Abietinae.     

A new species of <Emphasis Type="Italic">Nilssoniocladus</Emphasis> Kimura et Sekido from the Lower Cretaceous of the Markovsky peninsula (Southern Primorye)

An imprint of shoot Nilssoniocladus Kimura et Sekido bearing leaves Nilssonia Brongniart was found for the first time in the Lower Cretaceous deposits of the Markovsky peninsula near the city of Vladivostok. A new species, Nilssoniocladus anatolii, is identified. The finds of such shoots in organic connection with leaves are rare, and new data supplement our knowledge on the diversity, stratigraphic and geographic distribution of the genus Nilssoniocladus.     

THE DEVELOPMENTAL ANATOMY OF LONG-BRANCH TERMINAL BUDS OF PINUS BANKSIANA

A study of the composition of long-branch terminal buds (LBTB) of Pinus banksiana Lamb. and the yearly periodicity associated with their formation, development, and elongation was undertaken. Each LBTB has lateral bud zones and zones of cataphylls lacking axillary buds. When present, staminate cone primordia differentiate from the lowest lateral buds in the lowest lateral bud zone of the LBTB. Ovulate cone primordia and lateral long-branch buds can differentiate from the upper lateral buds in any lateral bud zone. When both types of buds are present, lateral long-branch buds are uppermost. Dwarf-branch buds occur in all lateral bud zones. During spring LBTB internodes elongate, new cataphylls are initiated, dwarf branches elongate, needles form and elongate, pollen forms and is released, and ovulate cones are pollinated. During summer buds form in the axils of the newly formed cataphylls. By early fall the new LBTB are in overwintering condition and the four types of lateral buds are discernable. The cytohistological zonation of the LBTB shoot apex is similar to that of more than 20 other conifer species. Cells in shoot apices of pine are usually arranged in distinct zones: apical initials, subapical initials, central meristem, and peripheral meristem. Periclinal divisions occur in the surface cells of the apex; therefore no tunica is present. At any given time, shoot apex volume and shape vary among LBTB in various positions on a tree. In any one LBTB on a tree, shoot apex shape changes from a low dome during spring to a high dome during summer to an intermediate shape through fall and winter.     

Chemical Composition of Pinus sibirica (Pinaceae)

Siberian pine (Pinus sibirica), also known as Siberian cedar pine and Siberian cedar, is an important plant that has been long used as a source of natural compounds and materials (wood, needles, soft resin, turpentine, colophony). Its chemical composition has been studied well enough; however, to our surprise, no articles that compile the phytochemical data have been published so far. Presumably, this is due to the fact that most of the studies were published in journals difficult to access and not indexed by search systems. This review, for the first time, presents a systematic compilation of available data of secondary metabolites occurring in the needles, shoots, bark, wood, seeds, and oleoresin of Pinus sibirica.     

ANATOMICALLY PRESERVED SEED CONES OF THE PINACEAE FROM THE EARLY CRETACEOUS OF VIRGINIA

A collection of anatomically preserved conifer cones from the Early Cretaceous of Virginia contains two new species of the extinct pinaceous genus Pityostrobus. Superficially, the fossil cones resemble those of modern species of Picea. However, the fossils reveal such a peculiar mixture of anatomical features that they cannot be assigned to any extant genus of the Pinaceae. One of the new species, Pityostrobus hueberi, is most comparable with Pityostrobus corneti Alvin from the Early Cretaceous of Belgium. Pityostrobus virginiana, the other new species, differs from all other members of the genus in only slight—but nonetheless significant—attributes. Pityostrobus hueberi and P. virginiana are the first species of this genus to be reported from Early Cretaceous sediments of the North American Atlantic Coastal Plain. As such, they increase our knowledge of the structural variation exhibited by ancient members of the Pinaceae.     

STABILITY OF NUCLEAR DNA CONTENT DURING ADVENTITIOUS SHOOT FORMATION IN PINUS TAEDA L. TISSUE CULTURE

Attempts to preserve selected genotypes through clonal propagation may be hampered by genetic instability produced during the tissue culture phase of the propagation procedure. To investigate the genetic condition of loblolly pine (Pinus taeda L.) tissue cultures, nuclear DNA content in embryos grown in vitro and in regenerated buds were examined using two-wavelength microspectrophotometry. Embryos were grown on chemically defined media containing cytokinin as the sole growth regulator. Nuclear DNA content remained at or between 2 C and 4 C with no evidence of polyploidy during nearly 3 months of continuous culture. Benzylaminopurine, even at concentrations which were supraoptimal for shoot regeneration, had no effect on nuclear DNA content. Regenerated shoots also had stable nuclear DNA contents which correspond to a diploid condition. These results are of significance for the development of mass propagation techniques for this commercially valuable species.