THE RELATIONSHIP AND ORIGIN OF TSUGA HETEROPHYLLA AND TSUGA MERTENSIANA BASED ON PHYTOCHEMICAL AND MORPHOLOGICAL INTERPRETATIONS

Morphological intermediacy between Tsuga heterophylla and Tsuga mertensiana has stimulated controversy concerning the relationship between these two hemlock species and has led to wide acceptance of hybridization as a causative factor of intermediate forms in areas of distributional overlap. In this study, comparative’ chemical analyses were used to test the theory of hybrid derivation of intermediates. Chromatographic analyses were further applied to the phylogenetic ranking of several North American and Asian species of Tsuga and to the examination of the hypothesis that Tsuga mertensiana is a derivative of intergeneric hybridization involving Tsuga heterophylla and Picea sitchensis. Chromatographic data derived from extensive sampling in the western Cascades of Washington led me to conclude that hybridization of Tsuga heterophylla and T. mertensiana is a possible but rare phenomenon and is not necessarily indicated by morphological intermediacy. Phylogenetically both of these species are typical representatives of Tsuga and are distinct from species of Picea. There is, however, some support for the derivation of Tsuga mertensiana by intergeneric hybridization.     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Development of nuclear microsatellite markers for the Japanese conifers Tsuga diversifolia and T. sieboldii (Pinaceae)

Nuclear microsatellite markers were developed for the two Tsuga species native to the Japanese Archipelago, Tsuga diversifolia and T. sieboldii, and a population with genetic affinities to T. diversifolia on Ulleung Island, Korea. Tsuga diversifolia and T. sieboldii are widespread dominant trees of temperate and subalpine forests in Japan but to date no genetic markers have been developed for these species. Fifteen polymorphic loci were developed and characterized, of which 14 are reliably amplified in each taxon. Across both species and the Ulleung Island population, the number of alleles per locus ranged from 3 to 26 (average = 13.93) and observed heterozygosity ranged from 0.005 to 0.935 (average = 0.535). In addition, all 15 loci were successfully amplified in a single accession of the Chinese species, T. chinensis. These markers will be useful for investigating the species’ biogeography, range‐wide genetic diversity, conservation genetic issues and potential for hybridisation.     

Effect of bryophytes on survival of conifer seedlings in subalpine forests of central Japan

Bryophytes form the major seedbed for coniferous trees in the subalpine forests of central Japan. Field experiments were conducted on the floor of a closed stand dominated byTsuga on Mt Fuji to examine the seedling survival ofAbies veitchii andTsuga diversifolia in seven substrate types in relation to the morphological characteristics of the seeds and seedlings. NeitherAbies norTsuga seedlings could survive on dwarf-bamboo litter beyond 2 years after the seed rain.Abies seedlings survived not only in all types of bryophyte communities but also in larch litter. In contrast, the survival ofTsuga seedlings was restricted to communities of smaller-statured bryophytes found on logs. The seeds and 1st year seedlings ofA. veitchii were larger than those ofT. diversifolia. Abies seedlings produced hypocotyls taller than any bryophyte community and radicles which were long enough to penetrate into the moist humus layer, whereasTsuga seedling radicles presumably penetrate into humus only in bryophyte communities forming a thin mat on logs. The effect of bryophytes on survival of tree seedlings therefore depends on the relationships between the morphology of seeds and seedlings and the structure of bryophyte communities.     

Leaf‐morphology and leaf‐anatomy in Ephedra altissima Desf. (Ephedraceae,Gnetales) and their evolutionary relevance

Leaves in most extant Ephedra ‐species represent only rudimentary scales without chlorophyll. Photosynthesis is completely restricted to green shoots. Only some species, e.g. Ephedra altissima, develop folious leaves even when being mature. Morphology and anatomy of cotyledons, primary leaves, and subsequent folious leaves of Ephedra altissima were examined particularly with special focus on their vasculature. The results show that the reduction of the leaves is achieved by an extreme reduction of the adaxial leaf surface. Most parts of the leaf surface are therefore developed by the abaxial side. Apart from a reduced lamina, a thick cuticle and sunken stomata are the only xerothermic adaptations in leaves of Ephedra altissima. A hypodermis and endodermis are also not developed. The anatomical results indicate that ancestors of modern Ephedra‐ species might have evolved in a more humid climate, contrasting to the arid habitats in which extant Ephedra‐ species are native today. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Foliar terpenoids in Tsuga species and the fecundity of scale insects

Summary The relationship between the reproductive success of two Japanese scale insects, Fiorinia externa Ferris and Nuculaspis tsugae (Marlatt) (Homoptera: Diaspididae) and the concentrations of 15 terpenoids in needles of Tsuga sieboldii, the Japanese host, and T. canadensis, the North American host, was investigated during 1981 and 1982 in a field plot of 8-year-old trees in New Haven, CT, USA. Both scales produced significantly more eggs per female on T. sieboldii than on T. canadensis. Stepwise multiple regression analyses indicated that the variation in fecundity within both scales was strongly associated with variation in the terpenoid profile between tree species.General patterns of phytochemical variation between the two Tsuga species based on differences in the concentration of terpenoids having similar chemical structures were revealed by the multivariate statistical technique, principal components analysis. The volatile leaf oil profile of T. sieboldii was relatively richer in terpene alcohols, while that of T. canadensis was relatively richer in terpene hydrocarbons and terpene acetates. The individual terpenoids were then assigned to one of five groups based on chemical structure and regression analyses were repeated; fecundity of both scales increased with increasing concentration of terpenoid alcohols. Fecundity of F. externa was negatively associated with the relative concentration of acyclic terpenes but the opposite was true for N. tsugae. Analysis of foliar terpenoids may provide a basis for predicting the relative susceptibility of Tsuga species to attack by F. externa and N. tsugae.     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

南溪楼梯草,云南荨麻科一新种

该文描述了自云南东南部发现的荨麻科楼梯草属一新种,南溪楼梯草(Elatostema nanxiense)。此新种与田林楼梯草(E. tianlinense)相似,但其叶具三出脉,边缘具浅圆齿或小钝齿,雄总苞苞片较少,8枚,排成一层,背面有1～3条纵肋,只1枚在顶端具角状突起而与后者相区别。此外,此新种与盘托楼梯草系的广布种盘托楼梯草(E. dissectum)的区别在于其茎被糙伏毛,叶呈椭圆形,具三出脉,雄头状花序的花序梗较短,雄总苞苞片较少,呈宽卵形或横长方形,背面有1～3条纵肋,雄小苞片有缘毛。     

Forest succession in the subalpine region of Mt. Fuji,Japan

Subalpine forest succession was studied on Mt. Fuji, Japan, where various types of forests in different successional phases occur owing to volcanic action. Ninety stands were subjected to ordination using an index (SI) defined by the relative basal area and the life span of component woody species, and the cover of canopy layer of the sample stands. Two different sequences of sample stands were found. One was from deciduous scrubs, through Larix kaempferi forests and Abies forests, to Tsuga diversifolia forests, and the other from Abies-Tsuga thickets to Abies forests. Through analyses of the forest structure and composition, soil survey and identification of fallen logs, the former sequence was recognized as the primary sere and the latter as a regeneration sere following gap formation. During forest succession, basal area reached a maximum in the seral phase with a multi-layered structure. The Tsuga forests, whose understory is restricted to a moss layer, were regarded as the climax. The death or fall of Tsuga stems resulted in gaps, which were subsequently occupied by Abies-Tsuga thickets. The second Abies forests were distinguished from the ones in the primary sere by the occurrence of Dryopteris and Cacalia and the lack of Rhododendron in the understorey. Both Abies forest types included Tsuga saplings. Thus, a cyclic relation is supposed between Abies and Tsuga.Nomenclature follows Ohwi (1975) and Nakaike (1982) for vascular plants, Iwatsuki & Noguchi (1973) for mosses, Inoue (1981) for hepaticae, Kashiwadani (1981) for lichens, respectively. Abies veitchii, A. mariesii were lumped as Abies spp.I wish to express my sincere gratitude to Prof. Toshio Hamaya, Tokyo, for the cordial guidance and encouragement. I also thank Prof. M. Numata and Dr. M. Ohsawa, Chiba, Prof. K. Okutomi, Tokyo, Dr. K. Suzuki, Tokyo, Dr. M. Suzuki, Kanazawa, and Mr. H. Taoda, Kumamoto, for their valuable advice and discussions.     

Dispersal versus climate: Expansion of Fagus and Tsuga into the Upper Great Lakes region

Pollen records for American beech (Fagus grandifolia) and eastern hemlock (Tsuga canadensis) compiled from 50 sites in Michigan and Wisconsin, USA, show that both species entered the Upper Great Lakes region about 7000 yr B.P., reaching their western and southwestern boundaries between 2000 and 1000 yr B.P. Fagus advanced northward into lower Michigan as a continuous front, except where Lake Michigan posed a geographic barrier. Colonies were established on the far side of the lake after a 1000 year lag, implying that longdistance dispersal across a 100-km wide barrier can occur. The Fagus range may not have been in equilibrium with climate for one or two thousand years before this time, when seeds were dispersed across the lake to Wisconsin. Tsuga seeds may have been dispersed 150 km or more from Ontario to reach Upper Michigan. Scattered colonies were established 6000–7000 yr B.P. on either side of Lake Michigan, which did not pose a significant barrier to this wind-dispersed species, Tsuga spread rapidly over a large region prior to 5000 yr B.P. Subsequent expansion to the west occurred more slowly, and may reflect gradual climatic changes in northern Wisconsin during the second half of the Holocene. Tsuga's range may have been limited by dispersal, rather than climate, for an unknown length of time prior to 5000 yr B.P. During this period Tsuga was expanding its range rapidly. The study shows, however, that it is difficult to devise rigorous tests to distinguish between dispersal limitations and climate as factors limiting range limits in the past.This work has been supported by the U.S. National Science Foundation.This work has been supported by the U.S. National Science Foundation.     

Hepatics from Rovno amber (Ukraine), 6. Frullania rovnoi,sp. nov.

A fossil species of the extant liverwort genus Frullania Raddi is described and illustrated, based on a single inclusion in a piece of Rovno amber (Ukraine) that shares its age with Late Eocene Baltic amber, its northern contemporary. Frullania rovnoi is characterised by leaves with a rounded dorsal lobe and the absence of ocelli. The ventral lobe is inflated and forms a saclike lobule, which is bell-shaped and somewhat constricted above the mouth. The bifid underleaves have several blunt teeth or angulations along the shoulder. The Rovno fossil differs sufficiently from morphologically similar species preserved in Baltic and Bitterfeld amber as to be described as new to science. The shape of the lobules and underleaves, as well as the absence of ocelli, indicate an affiliation to F. sect. Australes, hitherto represented in Eocene amber inclusions solely by F. schumannii (Casp.) Grolle. The Rovno fossil is distinguished from extant species of F. subg. Australes and from F. schumannii by having roughly and irregularly dentate-angulate underleaf margins.     

Aluminum,Fe, Ca,Mg, K,Mn, Cu,Zn and P in above- and belowground biomass. I.Abies amabilis andTsuga mertensiana

In a mature mixed subalpine stand ofTsuga mertensiana andAbies amabilis, significantly higher Al levels were found in foliage, branch and root tissues ofT. mertensiana.Tsuga mertensiana had significant increases in Al, Ca and Mn levels with increasing foliage age. In current foliage,T. mertensiana had lower levels of Ca, similar levels of Mg and P, and higher levels of Mn thanA. amabilis. Both tree species had Cu and Fe present at higher levels in branch than foliage tissues. Fine roots had the highest concentrations of Al, Fe and Cu but the lowest Ca and Mn concentrations of all tissues analyzed. In the roots of both species, phloem tissues always had significantly higher Al levels than xylem. Fine roots (< 1 and 1–2 mm) ofT. mertensiana had higher Al levels than were found inA. amabilis. Roots greater than 2 mm in diameter exhibited no significant differences in Al levels in phloem or xylem tissue betweenA. amabilis andT. mertensiana. The two species show a clear difference in their ability to accumulate specific elements from the soil.     

New species of Hypolytrum Rich. (Cyperaceae) from the Neotropics

Five new species ofHypolytrum Rich. from South America are described and illustrated.Hypolytrum amorimii andH. jardimii, both of sect.Bullata T. Koyama, are species with pseudopetioles and colored leaves, and endemic to the rain forest of southeastern Brazil.Hypolytrum bahiense (sect.Hypolytrum), a species with a lax synflorescence and two, free and lightly scabridulous floral bracts, is endemic to the rain forest of southeastern Bahia, Brazil.Hypolytrum leptocalamum (sect.Hypolytrum), a species with lax synflorescence and spike ellipsoid to cylindrical, is restricted to area of tepuis in the Guayana Highland (Brazil, Guyana, and Venezuela).Hypolytrum paraense (sect.Hypolytrum), a species with three, partly connate and densely scabridulous floral bracts, is restricted to the rain forest of the Amazon Basin, in the state of Pará, Brazil.     

The ultrastructure of Cerebropollenites from the Jurassic and Cretaceous of Asia

The dispersed pollen taxon Cerebropollenites recovered from the Jurassic of Afghanistan and Cretaceous of China was investigated ultrastructurally. Cerebropollenites lacks the differentiation of the proximal and distal hemispheres, a faint proximal triradiate mark, and an equatorial fringe, all features found in extant Tsuga, however, the gross pollen morphology, and the wall ultrastructure of Cerebropollenites suggests an affinity with the extant genus Tsuga (Section Micropeuce). The differences observed between Cerebropollenites and Tsuga are no greater than the differences observed between the pollen of the two Sections of Tsuga, Hesperopeuce and Micropeuce. The occurrence of Cerebropollenites in the Jurassic and Cretaceous is, thus far, the only fossil evidence that the genus Tsuga may have been present in the Mesozoic, and suggests that the Pinaceae, which is unequivocally recognized in the Cretaceous, may have had an earlier origin.     

A molecular systematic study ofCathaya, a relic genus of thePinaceae in China

The systematic position ofCathaya, a relic genus of thePinaceae, was discussed based on therbcL gene sequence. The sequence data were analysed with PAUP and MEGA programs. The great genetic distance value betweenCathaya and any other genus of thePinaceae showed thatCathaya was a distinct and isolated genus. The most parsimonious Fitch tree and neighbor-joining tree showed thatCathaya was distantly related to the clade comprisingAbies, Keteleeria, Pseudolarix andTsuga, and a sister group relationship betweenCathaya andPinus was weakly supported.Pseudotsuga is closely related toLarix. In theAbies-Keteleeria-Pseudolarix-Tsuga clade,Abies has a close relationship toKeteleeria whilePseudolarix is relatively closely related toTsuga.     

Coexistence mechanisms of evergreen,deciduous and coniferous trees in a mid-montane mixed forest on Mt. Emei,Sichuan, China

Structure and regeneration of a mid-montane (2200 m a.s.l.)mixed forest codominated by evergreen (Lithocarpuscleistocarpus), deciduous (Acer flabellatum)andconiferous (Tsuga chinensis, Abies fabri, andTaxus chinensis) trees were analyzed in a 40m× 60 m plot on Mt. Emei, Sichuan, China. Plant communitystructure and composition varied depending on topographic micro-habitat withinthe plot. Four topographic communities (topo-communities) were distinguishedwith dominant species corresponding to topography: (1)Abies – valley bank, (2) Acer– lower steep slope, (3) Lithocarpus – uppergentle slope, and (4) Tsuga – ridge. The coexistencemechanisms of the evergreen, deciduous, and coniferous trees were determined byidentifying the regeneration process characteristic of each dominant species,asrepresented by their seedling dispersion patterns and seedling establishmentalong the topographic gradients. The saplings and seedlings of the dominantswere distributed differently according to the topography:Lithocarpus under the canopy of parent trees and in gapsofthe upper gentle slope and ridge, Acer mainly in bothwell-lit and shady sites on the lower steep slope and on fallen logs,Abies on the valley bank but only in well-lit sites,Tsuga on the ridge, Taxus mostly onrock and well-lit sites on the slopes. Distribution of surviving saplings andseedlings was also related to the species of nearby canopy trees. We suggestthat Lithocarpus, Tsuga and Acer tendto be self-replacing in their own topographic habitats, andAbies survives as a fugitive by occupying occasionalsuitable gaps. The variation in soil conditions, particularly nutrients, withtopography affected seedling establishment and the growth of trees. Thesuccessional change of quantitative species composition, as predicted by theMarkovian model, shows the mixed forest to be in a sustained climax stage.     

Anatomy, palynology and nutlet micromorphology of Turkish endemic Teucrium sandrasicum (Lamiaceae)

In this study, the anatomical features of the leaf and stem, besides the pollen and nutlet characteristics of Teucrium sandrasicum are investigated. T. sandrasicum, belonging to sect. Teucrium, is an endemic perennial herb growing on serpentine around Muğla province. The anatomical studies on T. sandrasicum revealed that the stem shares the general characteristics of the Labiatae family. The leaves clearly exhibit xeromorphy due to features such as the distribution of stomata on the lower surface (hipostomatic), the occurrence of guard cells below the epidermis (xeromorphic type), inrolled margins, thick cuticle layer, thick outer epidermal cell wall, a high density of trichomes and thick palisade layer of the mesophyll. The anatomical studies showed that the upper epidermal cells of the leaf include many spherocrystals. The pollen grains are prolate, medium in size, 3-colpate with verrucate ornamentation. The nutlets are ellipsoid with a reticulate-verrucate surface. The results have proven that T. sandrasicum is different from the other species of the sect. Teucrium because of the branched trichomes on the stem and the lack of eglandular trichomes on the nutlets.     

庆元香科科,浙江唇形科一新种

该文描述了采自浙江南部庆元县的唇形科(Lamiaceae)香科科属(Teucrium L.)一新种:庆元香科科(T.qingyuanense),并附有线描图.庆元香科科与峨眉香科科(T.omeiense)和香科科(T.simplex)接近,与峨眉香科科的区别在于其茎、花序轴和花梗均密被倒向短柔毛,花萼外面被短柔毛和腺点...     

Liaoxia Cao et S.Q. Wu (Gnetales): ephedroids from the Early Cretaceous Yixian Formation in Liaoning, northeastern China

Gnetalean compression-impression fossils are described from the Early Cretaceous Yixian Formation, Liaoning Province, north-eastern China, and assigned to six species of Liaoxia Cao et S.Q. Wu. The fossils have opposite-decussate phyllotaxis and cones comprising 2–12 pairs of bracts. Ovulate cones have seeds typically in a distal position. The species differ from each other and from previously described fossils in the absence or presence of leaves, shape of cones and seeds, and shape and position of cone bracts. The species of Liaoxia are probably close relatives of extant species of Ephedra L., but diagnostic reproductive details that could confirm this hypothesis are not preserved. The restricted information in the fossils and the poorly understood morphological diversity of extant Ephedra, prevent assignment of the fossils to any particular subgroup of Ephedra, as well as an explicit exclusion of them from the extant genus.