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Dry grasslands are highly diverse vegetation types of great importance for livestock production in rural Balkan areas. We analysed a large data set of phytosociological relevés of dry grasslands (Festuco-Brometea and Festucetea vaginatae classes) in Serbia to produce the first overview of its classification, distribution, environmental conditions and biodiversity patterns. Phytocoenological relevés from relevant literature sources and our own investigations were stored in the Vegetation Database of Serbia (EU-RS-002) and the Balkan Dry Grassland Database (EU-00-013). After heterogeneity-constrained random resampling, the final dataset contained 1,897 relevés and 1,323 species. Species composition was classified hierarchically by the beta flexible method. We used species ecological indicator values for the estimation of the ecological conditions. Floristic and vegetation diversity and the conservation relevance of various dry grassland types, based on an assessment of endemic and protected species, were analysed. We identified 11 clusters, which were well characterised by their species composition and ecology. The first three clusters included loess and sand steppe grasslands mostly found in the Pannonian part of Serbia (Festucion rupicolae and Festucion vaginatae). The next three clusters consisted of Balkan ultramafic rocky grasslands of the order Halacsyetalia sendtneri, Balkan submediterranean mountain steppe grasslands on calcareous substrates, belonging to the order Astragalo-Potentilletalia and grasslands of the Balkan alliance Saturejion montanae on limestone. The third group of relevés comprised Balkan alliances of dry grasslands on deep soils, the Chrysopogono-Danthonion and sub-continental steppes of the alliance Festucion valesiacae in hilly areas of Serbia, mostly in the thermophilous oak zone. According to ordination analysis (DCA), the main floristic gradient was largely determined by temperature and moisture. The Festuco-Brometea class exhibited high floristic diversity (1,323 plant species) and very high conservation relevance in view of the large number of Balkan endemic species (204). A total of 233 species and subspecies protected by national legislation within the studied vegetation were recorded.
Cryophytic steppes in the Minusinskaya intermountain basin containing plant species that are predominantly distributed in the alpine zone such as Androsace dasyphylla, Dryas oxyodonta, Festuca sphagnicola, Kobresia myosuroides, K. filifolia, Minuartia verna, Oxytropis bracteata, Sagina saginoides, Papaver nudicaule, Patrinia sibirica, Pedicularis lasiostachys, Pulsatilla ambigua, Saussurea schanginiana, which are considered remnants of the Pleistocene vegetation. Based on 89 relevés, we classified cryophytic steppes using the Braun-Blanquet method within two phytosociological classes: Central Asian steppes of the Cleistogenetea squarrosae and West Palearctic steppes of the Festuco-Brometea. Three associations (Androsaco dasyphyllae-Caricetum pediformis, Pulsatillo patentis-Caricetum pediformis and Bupleuro multinervi-Helictotrichetum desertori) with three subassociations and three variants were described with respect to their phytosociological affinities and ecology. DCA ordination showed floristic differences between syntaxa, while correlations of DCA axes and floristic and environmental variables detected substrate type and temperature regime as presumably main drivers for vegetation differentiation. Another driver for vegetation differentiation seems to be continentality of the climate. Small scale distribution of cryophytic steppes were mapped using satellite images with resolution of 1.8 m. Cryophytic steppes always occupy only small areas in landscapes, on convex parts of undulated microrelief of mountain slopes and summits characterised by drought in summer and deep soil freezing in winter. These special micro-ecological conditions play an essential role for the existence of alpine flora in the Minusinskaya intermountain basin.
We studied the dry grasslands of shallow, skeletal soils (Sedo-Scleranthenea, Koelerio-Corynephoretea) in northern Europe, based on a combination of new relevés from southern Oland (Sweden, n = 182) and Saaremaa (Estonia, n = 73) as well as a comprehensive evaluation of literature data, of which 65 suitable relevés were directly included in our analyses. Apart from a few vague indications of acidophytic Sedo-Scleranthenea communities (order Sedo-Scleranthetalia), all data refer to basiphytic communities (Alysso-Sedetalia); our analyses are thus focussed on the latter. The Nordic Alysso-Sedetalia communities proved to be quite different from their temperate counterparts and thus are included in a separate alliance, Tortello tortuosae-Sedion albi, which forms the northern counterpart to the central European Alysso-Sedion. Within the northern alliance, we distinguish two suballiances. The more widespread central suballiance Tortello tortuosae-Sedenion albi inhabits different types of base-rich substrata in both natural and anthropogenic sites, and is comprised of the Cladonio symphicarpiae-Sedetum albi and the Ditricho flexicaulis-Sedetum acris. The second suballiance Tortello rigentis-Helianthemenion oelandici is restricted to the alvar sites (= treeless limestone plateaus) in Oland, Gotland, Västergötland and Estonia. It is characterised by several endemic taxa and a large number of cryptogams typical of alvar. It is comprised of four associations, Crepido pumilae-Allietum alvarensis, Fulgensio bracteatae-Poetum alpinae, Helianthemo oelandici-Galietum oelandici and Gypsophilo fastigiatae-Globularietum vulgaris. All six Nordic associations are described in detail with respect to their floristic composition, ecology, distribution and lower-ranked units, and each is represented by a vegetation table. The floristic differences within the Nordic communities are worked out in a synoptic table. Whereas several vegetation scientists have pointed out that vegetation types occurring at the limits of their distribution ranges in northern Europe are generally difficult to classify, our application of the Braun-Blanquet approach, which is based on a priori separated structural types and the general application of the central syntaxon concept, has enabled us to characterise and adequately define all Nordic communities. The Tortello-Sedion associations are two to three times as species-rich as those of the Alysso-Sedion and are among the most diverse small-scale plant communities ever described. We discuss the reasons for this exceptionally high plant diversity and the peculiar species mixture in the Tortello-Sedion and compare the relationship between Alysso-Sedion and Tortello-Sedion to the situation of other Nordic syntaxa of predominantly temperate vegetation types. Our results further underline the uniqueness of Baltic alvars and their paramount importance for conservation at the European level.
Thermophilous deciduous forests of Quercetalia pubescentis are widespread in Bosnia and Herze-govina (B&H), where they occupy about 11% of the national territory and account for about 20% of the total forest area. This paper provides their formalized classification and nomenclatural revision, based on 274 relevés from the literature and 399 relevés collected during intensive field research. The Cocktail method was used for supervised classification, which resulted in 17 associations recognized for B&H. Additionally, one new association emerged after semi-supervised classification of relevés not classified by Cocktail. The following associations were recognized, and characterized by species composition, ecology and distribution: 1. Querco pubescenti-Carpinetum orientalis; 2. Rusco aculeati-Carpinetum orientalis; 3. Carici hallerianae-Quercetum pubescentis; 4. Cruciato glabrae-Carpinetum orientalis; 5. Seslerio autumnalis-Quercetum pubescentis; 6. Aristolochio luteae-Quercetum pubescentis; 7. Asparago tenuifolii-Quercetum pubescentis; 8. Seslerio autumnalis-Ostryetum carpinifoliae; 9. Rusco aculeati-Ostryetum carpinifoliae; 10. Querco pubescenti-Ostryetum carpinifoliae; 11. Quercetum frainetto-cerridis; 12. Fraxino orni-Quercetum cerridis; 13. Lathyro nigri-Quercetum petraeae; 14. Aceri obtusati-Quercetum petraeae; 15. Cytiso hirsuti-Quercetum petraeae; 16. Festuco drymejae-Quercetum petraeae; 17. Potentillo micranthae-Quercetum petraeae; 18. Seslerio autumnalis-Quercetum petraeae. The validity and legitimacy of associations were checked and they were validated and corrected as needed, strictly following the rules of the International Code of Phytosociological Nomenclature. A complete list of synonyms has been given for every association, with an indication of the article of the Code according to which the name must be rejected. The associations were assembled into four groups, following the criterion of dominant species in a tree layer, in order to present differences and similarities in floristic composition between associations of the same type, i.e., dominated by the same tree species. An ecogram was drawn displaying the relative ecological range of each association along soil pH and moisture gradients. The largest number of associations (13) occurs in the Mediterranean region; the Dinaric and Pre-Pannonian regions each harbour ten associations, while there are only five associations in the Transitional Illyrian-Moesian region. Seven syntaxa previously reported for thermophilous deciduous forests of B&H were not recognized during the analysis. A list is given of all nomina nuda that could not be resolved and ascribed to synonymy with accepted associations.
The grey dunes along the present-day depositional coasts of Latvia are among the most diverse dune ecosystems in the Baltic Sea region. This paper focuses on the grey dune plant communities, their classification based on the Braun-Blanquet approach and their ecology and distribution in Latvia. A total of 3,430 phytosociological relevés were sampled in all coastal sections in the period from 1994 to 2008. The vegetation was classified using TWINSPAN analysis. Diagnostic species were determined for each syn taxon using statistical measures of fidelity (phi coefficient and Fishers exact test). The phytogeography of associations and variants is described on the basis of a distribution range analysis of plant species. In total, three associations of the class Koelerio-Corynephoretea occur in the grey dunes. The Corniculario aculeatae-Corynephoretum canescentis and the Caricetum arenariae belong to the alliance Corynephorion canescentis and the order Corynephoretalia canescentis. The Festucetum polesicae belongs to the alliance Koelerion glaucae in the order Sedo acris-Festucetalia. With six variants, it is the phytosociologically most diverse association of the grey dunes: typical variant, variant of Gypsophila paniculata, variant of Koeleria glauca, variant of Thymus serpyllum, variant of Epipactis atrorubens and variant of Corynephorus canescens.
Our results show phytosociological differences among five coastal sections due to geomorphology and coastal processes. The most diverse grey dune plant communities occur on the open Baltic Sea coast where all three associations are represented. The most common vegetation type on all coastal sections in Latvia is the typical variant of the Festucetum polesicae. The Epipactis atrorubens variant of this association is widely distributed, too. It has its major occurrences along the Gulf of Riga and the Irbe Strait. The Corniculario aculeatae-Corynephoretum canescentis was found in only two locations. Stands of the Festucetum polesicae var. Koeleria glauca occupy wider areas on old dunes, while the Festucetum polesicae var. Gypsophila paniculata is more typical on young dynamic dunes with intensive sand drift. The results of the phytogeographical analysis of vegetation data show that under the influence of climate, especially with dominance of the oceanic and transformed oceanic air masses, grey dune plant communities on the Latvian coast are mainly sub-oceanic to sub-continental. The most continental community is the Festucetum polesicae var. Koeleria glauca, while the Caricetum arenariae and the Corniculario aculeatae- Corynephoretum canescentis have the strongest oceanic character.
New vegetation data of dry grasslands in the Western Carpathians and the northern Pannonian Basin
(2010)
The paper presents new vegetation data from dry grassland sites in the biogeographical regions of the Western Carpathians and the northern Pannonian Basin, mainly belonging to the alliances Bromo pannonici-Festucion pallentis and Festucion valesiacae (Festuco-Brometea). The 124 phytosociological relevés were sampled between 2005 and 2009 in Slovakia, the SE Czech Republic, NE Austria, and N Hungary. They are classified into 16 associations and four transitional vegetation types. The paper also brings new information on the distribution of grassland associations in the study area. A new locality of the rare association Teucrio botryos-Andropogonetum ischaemi was confirmed. During our recent investigation of historical sites of the Alopecuro pratensis-Festucetum pseudovinae (Cynosurion cristati, Molinio-Arrhenatheretea) in the Slaná river floodplain, only one occurrence could be confirmed. Geographical principles in distribution of dry grassland associations and classification of the Stipa pulcherrima-dominated stands are also discussed.
In the lower siliceous uplands of Central Europe, various types of nutrient-poor grasslands are widespread and grow intermingled. These species-rich grasslands, often dominated by taxa of the Festuca ovina aggregate, comprise various phytosociological classes. They are remnants of a historic rural lands - cape and are of conservation importance. Few studies on such grasslands are available and there has been disagreement in assigning them to appropriate habitat types or syntaxa. We investigated such nutrient-poor grasslands in the lower Aar valley (Middle Hesse, Rhenish Massif). We surveyed 104 vegetation plots distributed throughout the valley and identified syntaxa to (sub)association level. We carried out supervised classification combining cluster analysis, a priori assignment to classes based on prevailing diagnostic species, and regional refinement based on phi-value maximisation of the units. As a result, we classified five associations within four classes: Polytricho piliferi-Festucetum tenuifoliae/Galio harcynici-Deschampsietum flexuosae and Festuco rubrae-Genistelletum sagittalis (Calluno- Ulicetea), Jasiono montanae-Festucetum ovinae (Koelerio-Corynephoretea), Gentiano-Koelerietum pyramidatae (Festuco-Brometea) and Arrhenatheretum elatioris (Molinio-Arrhenatheretea). Ecologically, soil acidity (resulting from Ca concentrations of the bedrock) was the main cause of floristic dissimilarity of the grasslands and thus community differentiation. Many stands grew on soils with intermediate pH and showed a peculiar mixture of basiphilous and acidophilous species. We conclude that (i) our approach of supervised classification yields convincing reproducible results when a syntaxonomic system is adapted top-down to a geographically restricted area, (ii) nutrient-poor siliceous grasslands dominated by taxa of the Festuca ovina aggregate can be well assigned to ecologically meaningful syntaxa, and (iii) the nutrient-poor siliceous grasslands of the Lahn-Dill Highlands are of high conservation relevance and in urgent need of protection.
A phytosociological survey of weed (segetal) vegetation in Slovakia was performed. A total of 508 relevés were sampled in 2002–2008. The aims of this study were to determine the actual distribution of the segetal communities, to analyze their floristic structure, and to evaluate their relationships to selected environmental factors. Thirteen plant communities of the class Stellarietea mediae were distinguished by cluster analysis; 11 communities were included in the subclass Violenea arvensis (Lathyro tuberosi-Adonidetum aestivalis, Consolido-Anthemidetum austriacae, Euphorbio exiguae-Melandrietum noctiflori, Veronicetum trilobae-triphyllidi, Lamio amplexicauli-Thlaspietum arvensis, Taraxacum sect. Ruderalia community, Spergulo arvensis-Scleranthetum annui, Myosotido-Sonchetum arvensis, Echinochloo-Setarietum pumilae, Galinsogo-Setarietum, and Stachyo annui-Setarietum pumilae) and two in the subclass Sisymbrienea (Portulacetum oleraceae and Setario viridis-Erigeronetum canadensis). Communities were characterized by diagnostic, constant, and dominant species and their structure, ecology, and distribution were estimated. The species composition of these communities was documented in synoptic and association tables. DCA ordination and analysis of variance was used to determine the main environmental factors of floristic differentiation and to determine ecological and structural differences among the communities. The analyses showed that the most important factors affecting floristic composition and classification of the weed communities are their time of development (agroecophase), the type of crops and altitude.
Marginal associations, i.e. floristically impoverished associations at the margin of the distribution area of a higher syntaxon, form a problem in vegetation classification, because true character species are lacking. We propose a new approach for the classification of such marginal associations, making use of the notion of 'chorological tension zones'. In the absence of true character species, the species from other syntaxa of the same formation can be used as such. Our proposal is to use the species group from every formation-true class only once within every marginal alliance, to limit the number of possible marginal associations. This approach is illustrated in a classification of the retamoid thickets in the Netherlands. On the basis of a numerical-subjective classification of the relevant species in the scrub layer and the evaluation of relevant literature, we conclude that the broom and gorse thickets in the Netherlands can be assigned to the Cytisetea scopario-striati Rivas-Mart. 1974, which is represented by four associations, each of which is characterised by the species of other scrub classes. The Ulici europaei-Sarothamnion scoparii Doing ex Weber 1997 is represented by the Rubo plicati-Sarothamnetum scoparii Weber 1987 and the Crataego monogynae-Cytisetum scoparii R. Haveman, I. de Ronde & J.H.J. Schaminée ass. nov., the Ulici europaei-Cytision striati Rivas-Mart., Báscones, Díaz, Fern. Gonz. & Loidi 1991 by the Frangulo alni-Ulicetum europaei De Foucault 1988 and the Rubo ulmifolii-Ulicetum europaei J.-M. Géhu ex R. Haveman, I. de Ronde & J.H.J. Schaminée ass. nov. This classification is based on a restricted dataset though, and a revision, based on a larger dataset from a wider region has to prove the tenability of the classification.
Dry open rocky grassland vegetation on shallow ultramafic soils in the Central Balkans represents typical secondary grasslands, which have developed mainly in the zone of thermophilous mixed deciduous broadleaved and pine forests. Although all relevant national and regional syntaxonomic reviews classify these rocky grasslands within the distinct order Halacsyetalia sendtneri, the syntaxonomic position of the order in different systems of classification has varied in the past. Considering this as well as the fact that there have been no synoptic works on this specific vegetation type, we gathered all available data on the order Halacsyetalia sendtnerii from the serpentinites of the Western and Central Balkan Peninsula for its critical evaluation. The results obtained in our analyses allowed us to propose a new syntaxonomic concept, which is partly in accordance with previously published syntaxonomic schemes. Two alliances can be distinguished: Centaureo kosaninii-Bromion fibrosi and Potentillion visianii, for which the diagnoses, diagnostic and constant taxa are given. Furthermore, we discussed the syntaxonomic position of the order Halacsyetalia sendtneri with respect to the classes Festuco-Brometea and Koelerio-Corynephoretea, as within the analysed associations, many taxa diagnostic for the class Koelerio-Corynephoretea were registered. The thermophytic pioneer grasslands and therophyte sward communities included in the alliance Thymion jankae nomen. inval., characterised by the absence of typical species of the order Halascyetalia sendtneri and presence of taxa diagnostic for the class Koelerio-Corynephoretea, are temporarily left within the order Halacsyetalia sendtneri. Finally, we provided nomenclatural adjustments for the analysed associations when necessary, although a conclusive judgement regarding all the associations currently included within the analysed order can only be made after more detailed field surveys including data on cryptogams as well as joint analyses including all floristically and ecologically similar syntaxa (e.g. Astragalo-Potentilletalia, Festucetalia valesiacae).
During the last decade, three new acidophilous forests associations were detected in the Mecsek Mts (SW Hungary), and described as acidophilous beech wood (Sorbo torminalis-Fagetum (A. O. Horvat 1963a) Borhidi et Kevey in Kevey 2001), acido-mesophilous oak wood (Luzulo forsteri-Quercetum petraeae (A. O. Horvat 1963a) Borhidi et Kevey 1996) and acido-xerophilous oak shrubland (Genisto pilosae-Quercetum polycarpae (A. O. Horvat 1967) Borhidi et Kevey 1996). In this article two further new associations are described: the acidophilous oakwood of the Mecsek (Viscario-Quercetum polycarpae Kevey, ass. nova) and the acido-mesophilous oakwood of western Hungary (Campanulo rotundifoliae-Quercetum petraeae (Csapody 1964) Kevey, ass. nova). These associations are related to the acidophilous forests of the Balkan Peninsula based on the infrequent presence of sub-Mediterranean species. A detailed comparative study of these new associations with the earlier known ones permitted to develop a reshaped classification of the syntaxonomy of these units, creating four new suballiances: within the frame of Quercion farnetto I. Horvat 1938 the suballiances Luzulo forsteri-Quercenion polycarpae Kevey, suball. nova and the typical Quercenion farnetto Kevey, suball. nova, in the frame of Quercion petraeae Zolyomi et Jakucs 1957 the suballiances Luzulo multiflorae-Quercenion petraeae Kevey, suball. nova and the Quercenion petraeae Kevey, suball. nova.
This paper presents the results of a study on psammophilous grasslands, conducted during the years 1994-1997 and in 2008 in the Masurian Lake District (NE Poland). The study aimed at the determination of floristic composition, phytosociological diversity and habitat characteristics. Field work resulted in 511 phytosociological releves. Soil samples were taken to determine pH, hydrolytic acidity, sum of basic cations, cation exchange capacity, content of CaC03 and total carbon. In 2008, the presence of communities dominated by grey-hair grass (Corynephorus canescens) in the previously established sites was checked. TWINSPAN was used for hierarchic classification of the association Comiculario aculeatae-Corynephoretum canescentis (syn. Spergulo morisonii-Corynephoretum canescentis). Accordingly, the association is divided into four subtypes (typical subtype, subtype with Cladonia mitis, subtype with Thymus serpyllum and subtype with Festuca ovina) with nine variants. Field observations showed that the communities analyzed have been under strong anthropogenic pressure, causing their degeneration and decrease in area. The most common human uses of psammophilous grassland communities are housing development, road construction, afforestation, sand extraction, wood storage and unauthorized dumping. In contrast to the above types of land use, seasonal use of grasslands for recreation purposes (as sunbathing spots or as sports fields) does not lead to their degeneration and may have a beneficial influence on their maintenance as a component of the local landscape.
Bramble scrubs are among the least known and understood vegetation types in Europe. In the Dutch National Vegetation Overview, three associations belonging to the Lonicero-Rubion silvatici were distinguished, viz. the Rubetum grati, Rubetum silvatici, and Rubetum pedemontani. During several vegetation mapping projects and Rubus excursions, a distinct type of bramble scrub was recorded repeatedly in the Campine in the province Noord-Brabant in the southern part of the Netherlands. In this paper, this scrub is described as a new association, the Rubetum taxandriae Haveman, de Ronde & Weeda, with R. taxandriae, R. campaniensis, and R. baronicus as character species, and R. insectifolius as regional character species. Variation, ecology and distribution of this new association are given and discussed, and two subassociations are distinguished. The differences with the Rubetum silvatici, to which this community was believed to belong, are discussed. Based on an earlier analysis of the centres of diversity of the genus Rubus in the Netherlands, it is supposed that the Rubetum silvatici in its circumscription in the Dutch National Vegetation Overview can be divided in more regionally distributed communities, partly as subassociations of the Rubetum silvatici, partly as independent associations.
Many current approaches to the formal definition of vegetation units have the disadvantage of leaving a large amount of relevés unclassified. In this paper I propose a new method for the unambiguous assignment of relevés, which is based on the summarised cover value of diagnostic species. In the first step, a relevé is assigned to the class with the highest cover score. For this purpose, the character and differential species of the class as well as the character species of all subordinated syntaxa are considered diagnostic. Once the class has been determined, the assignment proceeds successively to the lower ranks. This method, which may be called “summarised percentage cover approach”, uses solely the diagnostic species of syntaxa for the assignment of relevés to vegetation units, making additional formal definitions unnecessary. As a test data set, I used 487 relevés of fringe vegetation and grasslands of nutrient-poor sites sampled in the Vienna Woods (Wienerwald), Austria. All relevés were provisorily classified at alliances level, mostly following the assignment of the original authors. TWINSPAN and DCA were applied to evaluate this preliminary classification. Diagnostic species for both the alliance and the class level were identified using the total cover value ratio as fidelity measure. The subjective classification was largely confirmed by the numerical methods. On basis of the summarised cover of Trifolio-Geranietea and Festuco-Brometea species, between 64% (Geranion sanguinei) and 99% (Seslerio-Festucion pallentis) of the relevés were reassigned to the same class as in the original classification. The fact that a considerable amount of relevés originally classified as Geranion sanguinei was reassigned to grasslands reflects the problems in delimiting fringe communities rather than a poor performance of the assignment method. The “summarised percentage cover approach” allows not only for the unequivocal assignment of relevés to an existing classification system, but may also help to improve classifications by clarifying the delimitation of higher syntaxa.
Transitional between the Pannonian Basin and the Eastern Carpathians the Transcarpathian Ukraine (Transcarpathia) has a diverse geology including Quaternary sediments, volcanites, limestones and flysch, and its climate at low altitudes is relatively warm and humid. We conducted a field survey in June 2016 focusing on mesophilous forest vegetation along a transect covering almost the whole lowaltitudinal part of Transcarpathia. We recorded 54 relevés in the field and further digitized 22 relevés from literature. Using classification analysis, we distinguished three clusters of oak-hornbeam forests (alliance Carpinion betuli; 1–3) and three clusters of scree and ravine forests (alliance Tilio platyphylli-Acerion; 4–6): (1) Slightly wet Pannonian oak-hornbeam forests occurring in the lowland part of the region (Circaeo-Carpinetum); (2) Mesophilous oak-hornbeam forests (Carici pilosae-Carpinetum); (3) Xero-mesophilous oak-hornbeam forests (a drier subtype of the Carici pilosae-Carpinetum and the association Primulo veris-Carpinetum); (4) Mesophilous scree forests (Phyllitido-Aceretum); (5) Forests of steep slopes (Aceri-Tilietum) and transitions to mesophilous oak-hornbeam forests (Carici pilosae-Carpinetum); (6) Cool and wet scree and ravine forests (Arunco dioici-Aceretum pseudoplatani). Using indirect ordination analysis, three environmental variables (altitude, heat load index and slope) were identified as factors of significant influence on the species composition. These factors well distinguish oak-hornbeam forests from scree and ravine forests.