The course of resin canals in the shoots of conifers

The course of resin canals in stem cortex and the continuity between resin canals in leaves and those in stem cortex were investigated. The present paper is the first of three parts of the investigation. In this paper, fundamental features of resin canals and actual resin canal patterns in the Taxaceae, Cephalotaxaceae and Podocarpaceae are reported. From the observation of serial transections of shoots, composite diagrams and three-dimensional models of resin canal patterns are drawn. Central canals, if present, run vertically in stem cortex and sometimes divide, end blindly or unite each other. The distance between two adjacent central canals fluctuates rhythmically in connection with the vascular supply from the stem to leaves. The resin canal patterns of the families are classified into four types. Those ofTaxus, Nothotaxus and three species ofDacrydium belong to the Taxus type, those ofTorreya andCephalotaxus to the Torreya type, those ofDacrydium elatum, Podocarpus alpinus, P. elatus, P. elongatus andP. neriifolius to the Dacrydium type, those ofP. macrophyllus, P. nagi andP. koordersii to the Podocarpus type.     

松科植物属间C值和种子千粒重与进化关系初探

在检索C值数据库和种子数据库的基础上,对松科具有完整C值和种子千粒重等数据信息的5个属108种植物进行统计分析,并利用线性回归模型分析了二者之间相关性。结果显示,松科植物C值从9.5~36 pg呈正态分布,种子千粒重从1.2~1 269 g呈偏态分布,种间千粒重相差悬殊,高达1058倍。ANOVA分析表明,松属C值平均为25.79 pg,显著高于其它4个属;落叶松属C值最低（12.53 pg),显著低于云杉属（18.44 pg）,而云杉属、冷杉属和雪松属C值间却无显著性差异。结合前人研究结果,从C值的角度分析,松科5属间的演化顺序为：松属→云杉属→雪松属→冷杉属→落叶松属。松科5属间种子千粒重均值差异显著,松属种子均值最大（123.7 g）,其次为雪松属（84.5 g）和冷杉属（26.4 g）,最后为落叶松属（5.5 g)和云杉属(3.9 g)。除松属外,其它4个属ANOVA分析,雪松属种子千粒重显著高于云杉属、冷杉属和落叶松属（P<0.01),而云杉属和落叶松属种子千粒重间无显著差异(P>0.05)。对数回归分析和线性相关性分析表明,松科108种植物的C值和种子千粒重回归方程为：y=3.145x-7.248,决定系数（R²）为0.346,存在显著正相关性（P<0.000 1）,线性相关系数(γ)高达58.8%,这说明松科植物随着C值增大,其种子千粒重呈明显增加趋势。综上所述,松科属间种子千粒重差异显著,且随着属间进化程度的提高,C值和种子千粒重均呈明显下降趋势,其具体机理尚有待于结合其它功能性状做深入分析。     

Comparative anatomy of resin ducts of the Pinaceae

 Resin ducts are common in the Pinaceae. The comparative anatomy of stems and leaves of 50 species and two varieties from ten genera has been investigated. The structure and distribution of resin ducts differ among genera. Resin ducts occur in foliage leaves of ten genera of Pinaceae. Cortical resin ducts are absent in the stems of Pseudolarix and Larix. Resin ducts only occur in the secondary xylem of stems of Pinus, Picea, Cathaya, Larix, Pseudotsuga and some Keteleeria species. All of the epithelial and sheath cells are alive and thin-walled in the resin ducts of stem cortex and mesophyll. Except for Pinus the epithelial cells of resin ducts in the secondary xylem of stems have thick, lignified walls. Comparative study shows there are obvious differences in the resin ducts of different genera; apparent differences do not exist, however, in the resin ducts of different species of the same genus. According to the structure and distribution of the resin ducts in ten genera of Pinaceae, a synoptical arrangement of the genera is given and generic relationships of the Pinaceae are discussed. Received: 12 September 1995 / Accepted: 14 March 1996     

Characteristics of Wood Structure in Gymnosperms and Their systematic Significance

The systematic positions and taxonomic ranks of orders and families in Gymnosperms, especially those in Coniferopsida, are analysed and discussed in this paper based on the evolutionary trends in the wood structure. The opinions of the present authors are as follow: 1. The separation of the Araucariaceae from the Coniferae and establishment of the Araucariales are reasonable,because the intertracheid pitting in the wood is the Araucarioid type and there are no pits on both horizontal and end walls of ray parenchyma cells in the family. 2. The position of the genus Ginkgo in Cheng’s system is acceptable. Ginkgo is more similar to Coniferae than to Cycadaceae in the wood structure. 3. According to the characteristics of wood structure, arrangement of the Podocarpaceae, Cephalotaxaceae and Taxaceae between the Araucariaceae and the Pinaceae is reasonable. Among these families, the Cephalotaxaceae and Taxaceae are more closely related to each other in the view of the spiral thickenings which often appear on the inner wall of wood tracheids. 4. Further evidence for the establishment of the Sciadopitysaceae is provided. For example, most of cross-field pits in the wood of the Sciadopitysaceae are window like, while some of them are of the Lemon type or the Subtaxodioid type; bordered pits are of the Araucaria B type. 5. The characteristics of wood structure in the genus Platycladus differ greatly from Thuja. The former has cross field pits of the Cupressoid type, bordered pits of Araucaria B type and warty layer on the inner surface of tracheids. All of these characteristics have added further evidence for the separation of Platycladus from Thuja. 6. Based on the structural characteristics of woody rays in the Pinaceae, the most primitive genera are Abies, Keteleeria and Pseudolarix, while more advanced ones are Cedrus and Tsuga, and even more advanced ones are Pseudotsuga, Cathaya, Picea and Larix, all of which share normal resin canals. The most advanced genus is Pinus which is also of normal resin canals. Pinus can be divided into three subgenera, Haploxylon, Parry and Diploxylon, according to the presence or absence of dentation and warty layer in wood tracheids. 7. It is reasonable to place the genus Amentotaxus in the Taxaceae, because membrane of bordered pits in the genus is similar to that in the other four genera of the Taxaceae, both of the Araucaria type. 8. The present authors agree with Cheng’s (1978) treatment of Sect. Heopeuce in Tsuga, based on the fact that Tsuga longibracteata has traumatic resin canals and warty layer. Reducing Pinus hwangshanensis into P. taiwanensis, made by Cheng, is reasonable because of the similarities between P. hwangshanensis and P. taiwanensis in the wood structure. The establishment of a new subgenus, Patty, for Pinus bungeana is suitable based on chemotaxonomy, morphology and the distinct warty layer on the innersurface of wood tracheids.     

松科4属植物茎初生结构比较研究

研究比较观察了松科云杉属的云杉（Ｐｉｃｅａ ａｓｐｅｒａｔａ）、油杉属的油杉（Ｋｅｔｅｌｅｅｒｉａ ｆｏｒｔｕｎｅｉ）、雪松属的雪松（Ｃｅｄｒｕｓ ｄｅｏｄｅｒａ）、松属的海南五针松（Ｐｉｎｕｓ ｆｅｎｚｅｌｉａｎａ）和大明松（Ｐ．ｔａｉｗａｎｅｎｓｉｓ Ｈａｙａｔａ ｖａｒ．ｄａｍｉｎｇｓｈａｎｅｎｓｉｓ）５个种的幼茎初生结构。结果表明,皮下层的细胞层数、皮层细胞的组成,树脂道的分布,鞘细胞的排     

Cytogenetic and molecular characterization of the Abies alba genome and its relationship with other members of the Pinaceae

Genome size, karyotype structure, heterochromatin distribution, position and number of ribosomal genes, as well as the ITS2 sequence of the internal transcribed spacer (ITS) were analysed in silver fir (Abies alba Mill.). The analysis also included characterization of the Arabidopsis-type of telomeric repeats in silver fir and in related species. The results were compared with results from other species of the Pinaceae, to evaluate phylogeny and chromosomal and molecular evolution in the Pinaceae. Integrated chromosomal data provided insights into chromosome and karyotype evolution in the Pinaceae. The evolutionary trend for GC-rich heterochromatic blocks seems to involve loss of blocks that are not associated with rDNA. Similarly, numerous large blocks of interstitial plant telomeric repeats that are typical for all analysed species of the genus Pinus were not observed in the evolutionarily younger genera, such as Abies, Picea and Larix. On the contrary, the majority of telomeric sequences in these three genera appeared confined to the chromosome ends. We confirmed the current position of Abies and Tsuga in subfamily Abietoideae and the position of Pinus in the subfamily Pinoideae based on ITS2 sequences. Pseudotsuga is placed together with Larix into the subfamily Laricoideae. We conclude that the current position of the genus Picea in the subfamily Abietoideae should be reconsidered and, possibly, the genus Picea should be reclassified as a separate subfamily, Piceoideae, as recently proposed.     

Tricholoma matsutake Y1 strain associated with Pinus densiflora shows a gradient of in vitro ectomycorrhizal specificity with Pinaceae and oak hosts

Tricholoma matsutake produces commercially valuable yet uncultivable matsutake mushrooms during an ectomycorrhizal association with coniferous trees. In the Far East, most matsutake are harvested in managed Pinus densiflora forests. To determine whether T. matsutake has host plant specificity, we synthesized mycorrhiza in vitro between T. matsutake Y1 that originated from a P. densiflora forest and various Pinaceae and oak hosts. The strain Y1 formed a continuous Hartig net, a sign of ectomycorrhization, in the lateral roots of Pinus sylvestris, Pinus koraiensis, Pinus parviflora var. pentaphylla, Picea glehnii, Picea abies, and Tsuga diversifolia seedlings in vitro, which resembled those formed with the natural host Pinus densiflora. The strain conferred a discontinuous Hartig net with Pinus thunbergii, Picea yezoensis, Abies veitchii, and Larix kaempferi. However, no such development by this strain was observed on the roots of Quercus serrata, unlike T. bakamatsutake B1, a false matsutake that is symbiotic with oak trees. The data suggest that T. matsutake can be associated with diverse conifers but may establish ectomycorrhizal relationships only with specific host plant species.     

长白山云冷杉林幼苗幼树空间分布格局及其更新特征

长白山云冷杉针阔混交林是我国东北主要的森林类型之一,其乔木树种幼苗幼树的结构和动态决定着未来林分的结构和生长动态。在长白山地区设置一块具有代表性的云冷杉针阔混交林幼苗幼树更新样地,统计分析幼苗幼树更新特征,绘制地径结构图、树高结构图及其空间分布图。运用点格局分析中的单变量O-ring统计方法,分析更新树种的空间分布格局;用双变量O-ring统计方法,分析更新树种种间的空间关联性。研究结果表明:(1)更新树种组成有冷杉(Abies nephrolepis)、色木槭(Acer mono)、紫椴(Tilia amurensis)、红皮云杉(Picea koraiensis)、红松(Pinus koraiensis)、蒙古栎(Quercus mongolica)、春榆(Ulmus japonica)7种,其中以冷杉、色木槭为主,更新幼苗幼树的地径近似呈倒J型分布,树高结构近似呈双峰分布;(2)所有更新树种、冷杉、色木槭在小尺度1—10 m的范围内呈聚集分布,随着尺度增加,聚集程度减弱,逐渐趋于均匀分布和随机分布,紫椴、云杉和红松在空间所有尺度上以随机分布为主;(3)更新树种之间的空间关联性在小尺度范围上正关联性比较多,较大尺度范围上负关联性比较多,随着尺度增加,空间关联性减弱。     

Alterations of the resin canal system of <Emphasis Type="Italic">Pinus pinaster</Emphasis> seedlings after fertilization of a healthy and of a <Emphasis Type="Italic">Hylobius abietis</Emphasis> attacked stand

Changes in resource availability and biotic and abiotic stress may alter the defensive mechanisms of pine trees. The effect of fertilisation on the resin canal structure of Pinus pinaster seedlings established in two trials in NW Spain, one attacked by Hylobius abietis and the other non-attacked, was studied. The leaders of 50 plants were destructively sampled and the resin canal density, the canal area and its relative conductive area in the phloem and xylem were assessed. Experimentally increased nutrient availability significantly decreased resin canal density in the phloem of the seedlings in the two analysed trials, where unfertilised seedlings presented up to 30% more resin canal density than the fertilised seedlings (mean value ± SEM = 0.32 ± 0.02 resin canals mm⁻² in the fertilised plants versus 0.45 ± 0.04 resin canals mm⁻² in the control plants). Fertilisation had no effect on the resin canal system in the xylem, but significantly increased tracheid size. Significant differences of resin canals among sites were observed mainly in the xylem; the resin canal density was 1.7-fold greater in the attacked site than in the non-attacked site. The similar structure of phloem resin canals in both sites supports that phloem resin canals are constitutive mechanisms of defence in P. pinaster, whereas xylem resin canals would be constitutive mechanisms but also inducible mechanisms of resistance following the attack of pine weevils or bark beetles. XM and LS equally contributed to this paper.     

Mycorrhiza-like interaction by Morchella with species of the Pinaceae in pure culture synthesis

 Isolates from two species of Morchella were tested for ability to form mycorrhizae in pure culture synthesis with Arbutus menziesii, Larix occidentalis, Pinus contorta, Pinus ponderosa, andPseudotsuga menziesii. Ectomycorrhizal structures (mantle and Hartig net) formed with the four species of the Pinaceae but not with A. menziesii. Results are compared to previous studies on morel mycorrhizae and discussed in an ecological context. Accepted: 23 October 1999     

The significance of lenticels for successful<Emphasis Type="Italic"> Pityogenes chalcographus</Emphasis> (Coleoptera: Scolytidae) invasion of Norway spruce trees [<Emphasis Type="Italic">Picea abies</Emphasis> (Pinaceae)]

In this study we tested whether Pityogenes chalcographus L. beetles preferably infest their host tree, Picea abies (L.) Karst., through lenticels or not. In a second step the resin canal system, which is important for defence against bark beetles, was investigated under lenticels and under lenticel-free periderm. Beetle colonisation tests on breeding logs resulted in 93.2% successful invasions through lenticels, 4.2% through lenticel-free periderm, 1.2% through wounds, and 1.4% near branches (n=1,606). Three main reasons why beetles invade preferable sites are discussed, taking the function of lenticels to supply gas exchange of the interior tissues into consideration. First, attractive volatiles are more easily detectable over the lenticels, and this could be a stimulus for the bark beetles to invade their hosts through these structures. Secondly, the very loosely arranged tissues of the lenticel and bigger intercellular spaces in the cortex and secondary phloem under the lenticel permit the beetles easier movement towards the vascular cambium. Thirdly, the total resin canal area of both primary and secondary canals and the number of secondary resin canals were reduced under lenticels. A higher resin canal density beneath lenticels would constitute a barrier to gas movement.     

庞泉沟自然保护区寒温性针叶林演替优势种格局动态分析

以华北落叶松和青杆为主的寒温性针叶林是庞泉沟自然保护区的重要林型,也是主要保护对象之一。通过空间代时间构建庞泉沟自然保护区寒温性针叶林演替的时间序列,序列1是华北落叶松单优群落稳定发展的过程,序列2是从华北落叶松单优群落演替至华北落叶松-青杆共优群落再到青杆单优群落的过程,采用点格局分析法和与Monte-Carlo拟合检验对演替过程中华北落叶松和青杆的分布格局及其相互关系进行了研究。结果表明:(1)序列1在华北落叶松单优群落稳定发展过程中,华北落叶松的分布格局由集群分布趋向于随机分布,甚至在0—2.5 m上表现为均匀分布,驱动力是种内竞争引起的自疏现象。序列2从华北落叶松单优群落演替至青杆单优群落过程中,华北落叶松的分布格局同样是由集群分布趋向于随机分布,驱动力主要来自于由于青杄侵入扩散而形成的种间竞争;青杄集群的尺度逐步增大,驱动力主要是种群拓殖和种内竞争。(2)二者种间关系,在华北落叶松单优群落阶段无明显相关,共优阶段由于竞争在0—6.5 m尺度上呈显著正相关,在青杄单优群落,青杄竞争获胜,在0—2.5 m尺度上无明显相关,二者种间关系的变化主要来自于群落剩余资源驱动下的种内种间竞争。     

Fossil records of subsection Pinus (genus Pinus,Pinaceae) from the Cenozoic in Japan

Extant pines of subsection Pinus (section Pinus, genus Pinus, Pinaceae) are predominantly distributed in Eastern Asia. However, the extent of diversification in the section has yet to be fully clarified. We reviewed fossil records of subsection Pinus from Japan and collected permineralized materials, in which anatomical details are preserved for better understanding of the diversification. Our results suggest that this subsection appeared in Japan no earlier than the Middle Eocene, with extant species (i.e., Pinus densiflora and Pinus thunbergii) appearing around the beginning of the Pleistocene. Pinus fujiii (Early Miocene to Early Pleistocene) is inferred to have a close affinity to P. thunbergii based on the medial arrangement of its leaf resin canals. Additionally, P. fujiii has a similar cone morphology to those of extant species living in China, bridging the morphological gap between P. thunbergii and Chinese relatives of P. thunbergii as inferred by molecular phylogenetic analyses. Our results also suggest that taxonomic revisions of Pinus miocenica and Pinus oligolepis are required among the Japanese fossil species reported to date.     

Comparative Anatomy of Resin Canals of the Needles of Picea Grown in China

The comparative anatomy of the leaves of 21 species and 5 varieties of the genus Picea has been investigated with special reference to their resin canals. Resin canals are found in the needles of all Picea species under study. They are always arranged in longitudinal series, mostly being interrupted by transverse partitions of mesophyll except in a few species where they exhibit two disinterrupted canals. The number of the canal segments in a needle is quite variable in different species, and even in different needles of the same species. Their arrangement, according to the frequency of the occurrence of two-canals, one-canal and no canal in the middle transection of the needles and from the observation of the clearing specimens, can be classified into four groups (fig. 1). It would seem preferable that the characteristic difference of the resin canals, which might be of texonomic importance, lie in their discrepancy in the relative distribution of the resin canals rather than in the number or the relative length of the canals in each needle. This study also suggests that those with two canals lying along the lateral angles of the needles might be considered as the primitive types, from which other forms have been derived. Long canals may be broken into short cystic segments or subsequently degenerated into residue canals located only on one side of the needles.     

Areography of the Gymnosperms of China (1) Distribution of the Pinaceae of China

Nine of 10 genera and 119 of approximately 240 species of the Pinaceae occur in China, including 67 endemic species and two endemic genera. In this paper, the distributional maps of all the genera of the Pinaceae are presented (fig. 1-8). The horizontal and vertical distributions of species in each genus are discussed. The analysis of the distribution patterns of the genera indicates that some genera, such as Keteleeria, Tsuga, Pseudotsuga, Cathaya and Pseudolarix, are restricted to the area south of the Qinling Mountains and the Huaihe River, and the others, i. e. Picea, Abies, Larix and Pinus, extend northward to northeastern China. However, all of the genera except Keteleeria and Pinus are not found in very dry areas and tropical mountainous regions of China. The monotypic genera, Cathaya and Pseudolarix, are distributed in eastern and central China. The genus Keteleeria consists of 10 species, 7 of which are concentrated in southern Guizhou, northern Guangxi, southwestern Hunan and easternmost Yunnan. The distribution of the remaining 6 genera shows the maximum concentration in western Sichuan and northwestern Yunnan. (Figs. 2-8). Furthermore, more than third of species of the Pinaceae (37.8%) are also concentrated in western Sichuan and northwestern Yunnan. where a great variety of habitats and different topographic features occur. It is apparent that to conduct our systematic and evolutionary studies on this family in these region is especially needed. The relations between the areal size and the tolerance of species are discussed. The distributions of macrofossils and microfossils of the genera of the Pinaceae ia China are given, and it has been proved that areas of most genera of the family were considerably larger in the past. than at present.     

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.     

The relationships between the surface arboreal pollen and the plants of the vegetation in China

This paper attempts to test the relationships between the surface arboreal pollen and the plants across the entire vegetation of China. Twelve genera from major arboreal plants are selected from 680 pollen taxa in a total of 803 surface pollen assemblages for this excise. The pollen data are quantitatively compared with the gridded vegetation map on spatial similarity, direction and distance of pollen dispersal, and pollen accuracy for indicating the plant distribution density.The results show that the distributions of the pollen taxa can reflect the major patterns of the plants. Abies, Larix, Picea, Pinus, Quercus and Betula exhibit high spatial similarities between pollen and source plants. The accuracy for Tsuga, Platycarya, Fagus and Cyclobalanopsis is relatively low, probably due to the low pollen productivities and the absence of samples from southern China. In the disagreement part of the pollen-plant accuracy percentage, the pollen taxa Larix, Platycarya, Fagus and Cyclobalanopsis have high proportions in underestimating plants, likely a “convergent-type” in the pollen deposition; while pollen Tsuga, Quercus, Pinus, Betula and Ulmus have high proportions in overestimating plants, likely a “divergent-type” in the pollen dispersal.Pollen dispersal estimates suggest that southward transportation of the 12 pollen taxa is dominant, and the transport distance can be up to ca. 1000 km. This is mainly caused by several months of persistent north winds from the winter monsoon, particularly in spring seasons in eastern China. The second highest pollen dispersal direction is eastward, which occurs mainly in the boreal or sub-boreal plants of Larix, Abies, Picea, Betula and Acer and is likely influenced by the Westerlies in the mid-latitudes of China. Summer-green and evergreen Platycarya, Fagus and Cyclobalanopsis are mainly dispersed to the north, reflecting the influence of the Asian summer monsoon in eastern and southern China. Results of spatial differentiation analysis using accuracy weighting suggest that the low percentage portion (1-10%) in pollen of Abies, Picea, Betula and Quercus and high percentage portion (>40%) in pollen Pinus are acceptable for the plant representatives.These results, based upon the analyses of surface pollen and modern vegetation types of China, can provide important information for testing the various methods for reconstructing and modelling the environmental or climatic parameters during the recent geological past from palaeovegetation data.     

Late Embryogeny of Pinus bungeana

Except for Pseudolarix, which is endemic to China, the late embryogeny of ten genera of Pinaceae has been reported before in the diffferent degree. Among them the mature embryos of Keteleeria evelyniana are different from those of the others in having well-developed cotyledons and a very short hypocotyl. As far as information we have is concerned there are three types in structure of the mature embryos of Pinaceae. The first type occurs in Keteleeria and Cedrus, which have very well-developed cotyledons; the second one appears in Picea and Larix, in which the hypocotyl and the root cap are equal in length; the third one, to which Pinus bungeana belongs, includes all the other genera of Pinaceae. The last type is of a prominent hypocotyl in the mature embryos. It is interesting to note that the mature embryos of Torreya grandis and Keteleeria evelyniana are very similar in having specially developed cotyledons, while the proportions of the various tissues in the mature embryo in Fokienia of Cupressaceae as well as Taxus and Amentotaxus of Taxaceae are similar to those of Pinus bungeana. The pith and secretory cells are usually present in the mature embryos of Pinaceae. Although no pith is present in those of Metasequoia and Taiwania of Taxodiaceae, the secretory cells generally occur in their embryos in Fokienia of Cupressaceae and Taxus and Pseudotaxus of Taxaceae, neither pith nor secretory cells are present in their hypocotyl. From above, the structures of mature embryos among Pinaceae, Taxodiaceae and Cupressaceae are different from one another to some degree. The most outstanding feature of the matur embryos in Pinus bungeana is that the shoot apex is very well developed, with a high H/D ratio, about 0.83 on an average, even up to 0.96 in some case. Above-mentioned H/D ratio of Pinus bungeana is rare in the mature embryos of conifers. Gifford (1943) reports that the average H/D ratio of shoot apex of Ephedra altissima is from 0.44 to 0.68, while in 5-year-old branch apex of Pinus ponderosa, the average ratio is about 0.25, and that in l5-year-old branch apex is about 0.35. For the apex of the dormant short shoots of Pinus densiflora the average ratio is about 0.35, but that of shoot apex is about 0.52 when new buds have just formed. According to the present data about shoot apices of both branches and mature embryos, the average ratio of shoot apex of mature embryos in Pinus bungeana is the largest one. From the present investigation the shoot apex of mature embryos of Pinus bungeana exhibits four distinct tissue zones, i.e. the apical initials, the central mother cell zone, the peripheral tissue zone and the rib meristem. It is worthy of note that the shoot apex of Pinus strobus may be divided into five zones, including transition zone between central mother cells and rib meristem (Owston, 1968). Four zones are recognized in the shoot apex of Pinus lambertiana and P. ponderosa, without transition one (Sacher, 1954). From cytological zonation, the shoot apex of mature embryos in Pinus bungeana is rather similar to that of Pinuslambertiana and P. ponderosa.     

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.