Classification and Phytogeography of Larch Forests of Northeast Asia

This study focuses on a phytosociological system of the extensive Larix gmelinii and Larix cajanderi dominated forests and woodlands that span the nemoral, boreal and subarctic zones in northeast Asia in the Russian Far East, Eastern Siberia and northern China, an area that covers more than 5 million km². A total of 299 relevés representing different parts of the larch forest range were used to classify communities in this area into the boreal Rhytidio rugosi-Laricetea sibiricae and Vaccinio-Piceetea and to the nemoral Querco mongolicae-Betuletea davuricae. The hierarchical system of lower syntaxa of larch forests and woodlands of northeast Asia includes 24 associations and nine subassociations. Nineteen associations and two alliances – Rhododendro aurei-Laricion cajanderi and Rhododendro daurici-Laricion gmelinii – are described for the first time. Two associations were classified into Dictamno dasycarpi-Quercion mongolicae of Lespedezo bicoloris-Quercetalia mongolicae representing the nemoral class Querco mongolicae-Betuletea davuricae. All larch communities in the nemoral zone represent different post-fire-event stages. Eleven associations occurring in humid conditions of the boreal zone and in the areas influenced by the Pacific Ocean are classified into Abieti nephrolepidis-Piceion jezoensis, Piceion jezoensis and Pino pumilae-Piceion jezoensis representing boreal forests of humid maritime regions belonging to the order Abieti veitchii-Piceetalia jezoensis. In this area seral larch communities occupy zonal sites, and primeval larch forests may occur only in edaphic conditions restricting the distribution of zonal dominants, Picea jezoensis and the species of Abies. The area of continuous permafrost within the boreal zone having continental and ultracontinental climate conditions is occupied by primeval larch forests united in Ledo palustris-Laricion cajanderi and Rhododendro aurei-Laricion cajanderi of Ledo palustris-Laricetalia cajanderi, and on sites with dry soils with a deeper frozen horizon in Lathyro humilis-Laricion cajanderi of Lathyro humilis-Laricetalia cajanderi.     

Range-habitat relationships of vascular plant species at the taiga forest-steppe borderline in the western Khentey Mountains,northern Mongolia

World distribution of 488 out of 619 vascular plant species known from an area of 500 km² within the western Khentey Mountains, northern Mongolia is analyzed. Most species belong to Eastern Asian (29%) or Asian, Eurasian, or circumpolar temperate species (24%) supporting the classification of the Khentey Mountains as part of the temperate zone of Eurasia. Seventeen percent of species are boreal plants. Circumpolar temperate-boreal (9%), Central (-Eastern) Asian (9%), Continental mountain species (5%), Middle-Central Asian (3%), arctic-alpine (3%) as well as Western Eurasian and western Siberian species are of lower significance for the flora of the western Khentey Mountains. Eastern Asian species occur in all types of habitats, whereas plants of other distribution types are focused on certain habitats. Boreal species preferably grow in the dark taiga, which prevails in the upper montane belt and on northern and eastern slopes of the most humid parts of the lower montane belt. Temperate and temperate-boreal species prefer subtaiga forests, which are found on northern and eastern slopes in drier parts of the lower montane belt as well as in upper parts of sun-exposed, southern and western slopes of the lower montane belt. Central (-Eastern) Asian and Middle-Central Asian species primarily inhabit forest steppe habitats, such as meadow and mountain steppes, Ulmus pumila open woodlands and dry Pinus sylvestris forests on steep, southern slopes.     

Phytosociological affiliation of Annex II species Tephroseris longifolia subsp. moravica in comparison with two related Tephroseris species with overlapping distribution

The phytosociological affiliation of Tephroseris longifolia subsp. moravica, species of European importance, was studied in relation to two closely related species of the genus Tephroseris which have overlapping distribution within the Western Carpathian Mts: T. intergrifolia and T. crispa. The main aim was to compare plant communities inhabited by the three taxa, to assess the major environmental gradients responsible for variation in their distribution and to estimate ecological indicator values for Tephroseris longifolia subsp. moravica. T. longifolia subsp. moravica was recorded in nine localities in the Slovakia and Czech Republic where it occurs in very specific site conditions of ecotone habitats. Its phytosociological affiliation is restricted to grasslands of the alliances Bromion erecti and Arrhenatherion elatioris and to the ecotone vegetation between these grasslands and beech forests. T. integrifolia occurs most frequently in the Diantho lumnitzeri-Seslerion, Bromion erecti and Quercion pubescenti-petraeae alliances. T. crispa occurs predominantly in communities of the Calthion palustris alliance and Scheuchzerio-Caricetea fuscae, Mulgedio-Aconitetea and Montio-Cardaminetea classes. The major gradient responsible for variation in species composition of communities inhabited by the studied taxa was associated with moisture and nutrient content. The vascular plant-based ecological indicator values for Tephroseris longifolia subsp. moravica calculated from phytosociological relevés with its occurrence were set for light — 6, temperature — 5, continentality — 4, moisture — 5, soil reaction — 6 and nutrients — 5. We conclude that the studied taxon has intermediate relationship to the most of the studied factors in comparison with two related species, T. crispa and T. integrifolia.     

The mountain forests of british columbia and the american northwest: Floristic patterns and syntaxonomy

An overview of the montane and subalpine forests ofAbies lasiocarpa, Picea engelmannii, Abies amabilis andTsuga mertensiana of northwestern North America is provided based on 541 relevés of forest vegetation from coastal and inland British Columbia, Washington and northwestern Montana. A new classification is proposed based on an examination of the floristic patterns across these communities. Owing to their broadscale floristic similarities, coastal subalpineAbies amabilis-Tsuga mertensiana forests and inland forests ofAbies lasiocarpa, Picea engelmannii, P. glauca, Thuja plicata andTsuga heterophylla are combined into a single order,Thujetalia plicatae. The floristic patterns within this order in turn support the recognition of four alliances. These are (1)Tsugion mertensianae, including coastal and inland subalpine forests ofTsuga mertensiana, Abies amabilis, A. lasiocarpa andPicea engelmannii, with deep winter snowpacks and an often luxuriously developed shrub layer; (2)Abieti-Tsugion heterophyllae all. nov., including mesic inland montaneThuja plicata-Tsuga heterophylla andAbies lasiocarpa-Picea engelmannii forests and related southern borealAbies lasiocarpa-Picea glauca communities; (3)Gymnocarpio-Abietion lasiocarpae, including moist coastal and inland forests dominated by a fern-rich understory andOplopanax horridus; and (4)Lysichito-Chamaecyparidion nootkatensis, which includes wet coastal subalpine forests withLysichiton americanum. A total of five new associations are described, bringing the number of recognized associations within the order to twenty-five. An extensive synonymy and crosswalk to other classification systems is provided.     

Classification of ultracontinental boreal forests in central Yakutia

Using the Braun-Blanquet approach, five associations of boreal forests were distinguished in central Yakutia, the most continental part of eastern Siberia. Ecological features of the syntaxa were explained with the use of the DCA ordination of 50 relevés. All available data from eastern Siberia were involved in the study for syntaxonomic analysis. Central Yakutian boreal forests were classified into two classes:Rhytidio-Laricetea sibiricae Korotkov etErmakov 1999 — ultracontinental light coniferous hemiboreal forests, andVaccinio-Piceetea Br.-Bl. inBr.-Bl. et al. 1939 — typical coniferous taiga forests of northern Eurasia. A new concept of higher syntaxonomic units of the classVaccinio-Piceetea in eastern Siberia has been developed. Three orders represent the diversity of taiga forests: (1)Cladonio-Vaccinietalia Kielland-Lund 1967 (with alliancesHieracio umbellati-Pinion sylvestris Anenkhonov etChytry 1998 andSaxifrago bronchialis-Pinion sylvestris all. nov.) — light coniferous boreal forests occurring in dry and moderately dry oligotrophic sites in various climatic sectors of Northern Eurasia; (2)Lathyro humilis-Laricetalia cajanderi ord. nov. (with alliancesAulacomnio acuminati-Laricion cajanderi all. nov. andRhododendro daurici-Laricion gmelinii all. nov.) — zonal boreal forests with xeric elements, which are typical of regions of northern Asia with cold, dry ultracontinental climate; (3)Ledo-Laricetalia cajanderi ord. prov. (with allianceLedo-Laricion cajanderi prov.) — North Eurasian boreal forests occurring in cold sites with excessive soil moisture, sometimes water-logged. Phytogeography and ecology of these orders are discussed in comparison with other regions of northern Asia.     

Thermophilous oak forests in the Czech Republic: Syntaxonomical revision of theQuercetalia pubescenti-petraeae

Using the Braun-Blanquet approach, a syntaxonomical revision of the order ofQuercetalia pubescenti-petraeae, based on the synthesis of 634 relevés, distinguished three alliances of the thermophilous oak forests in the Czech Republic: (1)Quercion pubescenti-petraeae (3 communities:Pruno mahaleb-Quercetum pubescentis, Lathyro versicoloris-Quercetum pubescentis, Corno-Quercetum), which includes peri-alpine and peri-Carpathian oak forests on calcareous bedrock, (2)Aceri tatarici-Quercion (2 communities:Quercetum pubescenti-roboris, Carici fritschii-Quercetum roboris), which comprises Pontic-Pannonian oak forests on loess or sand that occur in southern Moravia only, (3)Quercion petraeae (5 communities:Potentillo albae-Quercetum, Brachypodium pinnatum-Quercus robur community,Sorbo torminalis-Quercetum, Genisto pilosae-Quercetum petraeae, Asplenio cuneifolii-Quercetum petraeae), which includes Central European thermophilous oak forests distributed outside the range ofQuercus pubescens and some other submediterranean species.     

Plant communities of mires in the western part of the Krkonoše mountains,Czechoslovakia

Mire plant communities of the W. Krkono?e mountains may be grouped in 11 associations and 3 alliances:Caricion canescentis-fuscae, Rhynchosporion albae andOxycocco-Empetrion hermaphroditi. The ecology, structure and geographical distribution of the communities, as well as the ecology of the species occurring in them are discussed.     

Notes on syntaxonomy of cultural forest communities

It is suggested that communities of cultivated forests can be incorporated into the system of “natural” wood communities either as facies, variants or subassociations, if they differ below the association level; if they differ on the association level, a new association is suggested, with the prefix “culti-” before the planted dominant tree species. Most of our planted forest communities differ on the level of association or less; the communities ofRobinia are the exception. The authors agree withJurko (1963) in classifying them as individual associations, alliances, order and class.     

The high-rank syntaxa of the rock-cliff and scree vegetation of the mainland Greece and Crete

A revision was undertaken of the high-rank syntaxa of the vegetation occurring in rock fissures, ledges and screes of mainland Greece and Crete. All published phytosociological relevés available were collected and subjected to numerical classification and ordination. Four orders (Androsacetalia vandelii, Onosmetalia frutescentis, Potentilletalia speciosae andPetromaruletalia pinnatae) comprising 8 alliances (one of them new) were distinguished within theAsplenietea trichomanis (rock fissures, clefts, and ledges). The scree vegetation was classified partly within theThlaspietea rotundifolii (Drypidetalia spinosae with 2 alliances) and partly within theDaphno-Festucetea (Saturejo-Scutellarietalia—a new order, with 2 alliances). Ordination revealed clear differences in floristic composition of the alliances distinguished. The studied chasmophytic vegetation of Greece contains a very high proportion of endemics which serve as diagnostic species of the syntaxa. Further, an analysis of chorological species spectra suggested that the chorological homogeneity of a syntaxon should be considered an important diagnostic feature at high-syntaxon levels.     

Endemische Phytoassoziationen aus den Rumänischen Karpaten

The importance of endemic species for plant communities in the Roumanian Carpathians Mts. has been studied and discussed. Associations of the classAsplenietea trichomanis endemic in the South Carpathians were surveyed; they were classified into some endemic alliances (Gypsophilion petraeae, Micromerion pulegii, andSilenion lerchenfeldianae). Their locus classicus, nomenclatoric type and present distribution were also given.     

Classification of meadows of the South Siberian uplands and mountains

The classification of South Siberian meadows using the Braun-Blanquet approach was carried out on the basis of a geographically wide-ranging data set from the West Siberian Plain and the Altai-Sayanian mountain systems. Two main phytosociological types of meadows were distinguished within the classMolinio-Arrhenatheretea. The natural Siberian forest meadows were included in the orderCarici macrourae-Crepidetalia sibiricae comprising two alliances (Crepidion sibiricae, Aconito barbati-Vicion unijugae) and 7 associations. The anthropogenic types of dry meadows were included in the European-West Siberian orderArrhenatheretalia, comprising one alliance (Festucion pratensis) and 5 associations. DCA ordination revealed clear differences in the floristic composition of the higher units and in the diagnostic importance of the main ecological and phytosociological species groups.     

Diversity of forest vegetation across a strong gradient of climatic continentality: Western Sayan Mountains, southern Siberia

Southern Siberian mountain ranges encompass strong climatic contrasts from the relatively oceanic northern foothills to strongly continental intermountain basins in the south. Landscape-scale climatic differences create vegetation patterns, which are analogous to the broad-scale vegetation zonation over large areas of northern Eurasia. In their southern, continental areas, these mountains harbour forest types which potentially resemble the full-glacial forests recently reconstructed for Central Europe. To identify forest vegetation–environment relationships in the southern Siberian mountain ranges, forest vegetation of the Western Sayan Mountains was sampled on a 280 km transect running from the northern foothills with oceanic climatic features to the continental Central Tuvinian Basin in the south. Based on the species composition, vegetation was classified into hemiboreal forests, occurring at drier and summer-warm sites with high-pH soil, and taiga, occurring at wetter, summer-cool sites with acidic soil. Hemiboreal forests included Betula pendula-Pinus sylvestris mesic forest, Larix sibirica dry forest and Pinus sylvestris dry forest. Taiga included Abies sibirica-Betula pendula wet forest, Abies sibirica-Pinus sibirica mesic forest and Pinus sibirica-Picea obovata continental forest. Hemiboreal forests were richer in vascular plant species, while taiga was richer in ground-dwelling cryptogams. Vegetation–environment relationships were analysed by indirect and direct ordination. Winter and summer temperatures and precipitation exerted a dominant influence on species composition. Soil pH was also an important correlate of species composition, but this factor itself was probably controlled by precipitation. At a more local scale, the main source of variation in species composition was topography, producing landscape patterns of contrasting plant communities on slopes of different aspects and valley bottoms. The response of tree species to major environmental factors was expressed with Huisman–Olff–Fresco models. Larix sibirica appeared to be most resistant to drought and winter frosts, Pinus sibirica was adapted to low temperatures both in winter and summer, and Picea obovata had an intermediate response to climate. Betula pendula, Pinus sylvestris and Populus tremula were associated with the warmest sites with intermediate precipitation, while Abies sibirica was the most moisture-demanding species, sensitive to deep winter frosts.     

Chemical and 13C-n.m.r. studies of an arabinogalactan from Larix sibirica L.

An arabinogalactan isolated from the wood of Larix sibirica L. was investigated by methylation analysis, partial hydrolysis, enzymic oxidation, and ¹³C-n.m.r. spectroscopy. The structural conclusions arrived at by ¹³C-n.m.r. spectroscopy were consistent with the data from methylation analysis. The polysaccharide is highly branched and similar in structure to those of arabino-3,6-galactans isolated from other Larix species.     

Soil-moisture conditions indicated by field-layer plants help identify vulnerable forests in the forest-steppe of semi-arid Southern Siberia

High variability in soil-moisture conditions is typical for semi-arid forest-steppe ecosystems where precipitation varies greatly over time. Plant species that inhabit these environments integrate responses to broadly fluctuating wetness conditions. Indirect assessment of contrasting habitat wetness based on plant indicator values, species frequency, and species coverage was carried out in two sites representing the larch (Larix sibirica) and pine (Pinus sylvestris) forest-steppe communities. For the larch forest-steppe, we found that plant community composition and spatial structure depended strongly on wetness. In addition, we found that the vegetation was clearly differentiated into forest stands and steppe communities, depending on the slope aspect. There was also a strong correlation between dissimilarities of species composition and differences in habitat wetness revealed in the larch forest-steppe. In contrast, soil properties, such as gravel and stone content were found to be a key factor in the spatial distribution of plant species composition in the pine-forest-steppe communities. Indirect assessment of moisture conditions in the forest-steppe habitats, based on the field-layer plant species, was found to be preferable for indicating soil water deficits in the forest. Furthermore, as long-term observational data is often lacking, indirect assessment of the forest-steppe vegetation provides an opportunity to identify vulnerable forests at the marginal distribution. Based on indirect assessments of soil-moisture conditions, and taking into account differences in potential drought resistance between larch and pine forests, we concluded that increasing aridity will cause the replacement of Siberian larch by Scots pine in the South Siberian forest-steppe landscape. Consequently, in the future it is likely that forest-steppe typological diversity will decrease, and the semi-arid landscape may become more monotonous.     

Malaxis chica,a new species of Orchidaceae from Nuevo León,Mexico

A new species from northern Mexico,Malaxis chica, is described and illustrated. It is distinguished by its small, unifoliate habit, subumbellate inflorescences, and sagittate lip with long basal lobes.Malaxis chica is compared to three species—M. corymbosa, M. salazarii, andM. rosei—with which it shows some morphological similarity.     

Refugium of the Boreal Forests of the Circumpolar Urals

The results of studies of the vegetative cover and ontogenetic composition of the cenopopulations of Picea obovata Ledeb. and Larix sibirica Ledeb. in the Gorno-Khadytinskii Forest Reserve, which is the refugium of the northern taiga forests of Western Siberia, have been summarized. According to the Brown–Blanke system, two associations of forests were identified and described, as well as two tundra associations, for comparison. The unique association of the Aconito septentrionalis–Piceetum obovatae taiga forests Zaugolnova et Morozova 2009 were analyzed in detail, and a comparison of these associations was carried out in the Gorno-Khadytinskii and Pechoro-Ilychskii reserves, where these forests are the reference standards of the zonal-type forests. An estimate of the ontogenetic composition of the cenopopulations of P. obovata Ledeb. and L. sibirica Ledeb. is given, as well as the absolute age of the trees of these species.     

How heterogeneous are the cloud forest communities in the mountains of central Veracruz, Mexico?

The montane forest in central Veracruz, Mexico is a hotspot of biodiversity. We asked whether lower and upper montane forests could be distinguished in this ecoregion. Variables of vegetation and seasonality in precipitation were tested across 14 sites between 1,250- and 2,550-m elevations. A total of 1,639 individuals and 128 tree species was recorded. There was a unimodal pattern in the richness of species, genera, and families; their richness was positively correlated with precipitation in the wettest quarter of the year, though there were no differences in the basal area and density. Rarefaction, species turnover, nonmetric multidimensional scaling, and a cluster histogram suggest two major groups: lower elevation forests that are less diverse, have low beta diversity and are more similar in composition, with Clethra macrophylla, Liquidambar styraciflua, and Quercus lancifolia as indicator species; and higher elevation forests that are more diverse, have high species turnover, and include forests with Quercus corrugata and Prunus rhamnoides, and forests with Fagus grandifolia, Persea americana, and Ternstroemia sylvatica as indicator species. However, other communities (an Oreomunnea mexicana at the upper site, and a limestone site in the lower forests), exemplify the high regional heterogeneity. We conclude that elevation and seasonality in precipitation produce a directional change in richness and indicator species, but not in vegetation structure. Lower montane forests differed from cloud forests at upper elevations. However, other factors should be included—mainly biogeographic affinities, historic and recent anthropogenic disturbance—to conclusively distinguish them. Montane forest can still be considered very heterogeneous and very high in beta diversity.     

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。     

INFLUENCE OF SITE AND STAND FACTORS ON THE DISTRIBUTION OF CRUSTOSE LICHENS OF THE CALICIALES IN A SUBOCEANIC SPRUCE FOREST AREA IN CENTRAL NORWAY

The distribution of crustose Caliciales has been surveyed in 100 spruce forest patches in Sør-Trøndelag, central Norway. Relationships between occurrence of the species and a number of site and stand variables were analysed by detrended correspondence analysis (DCA) and direct gradient analysis. Species diversity was significantly higher in old forests and in forests at higher altitudes compared to young forests and forests at lower altitudes. Old trees and snags are considered to be the most important structural components in old forests promoting species diversity of the Caliciales. Threatened or vulnerable species, such asChaenotheca gracillima, Cybebe gracilenta, Sclerophora coniophaeaandS. peronellawere confined to forest on rich soils showing no correlation with forest stand age.Chaenotheca brachypodaandC. trichialiswere found to be the most typical old forest species among the Caliciales. Humidiphilous species are considered to be less affected by forestry in a humid climate. A change in forestry practice towards methods imitating the natural dynamic processes is considered necessary to maintain species diversity of the Caliciales in boreal forests.