Journal of v egctation Sciel/cc 4: 279-282; 288-291.1993 © lA VS; Oputus Press Uppsala. Printed in SIVeden
279
Vegetation science in Hungary Fekete, Gábor!
*
& Tóthmérész, Béla2
+lnstitute of Ecology and Botany. Hungarian Academy of Sciences, Vácrától, H-2163, Hungary; 2Department of Ecology, L. KOSSlIt/1 University of Sciences, Debrecen, H-40JO, Hungary: *Tel. + 362760147;
Introduction This paper presents a concise account of the traditions, main results and perspecti ves of vegetation science in Hungary. It includes the history of plant sociology which, as in other countries in Central Europe, was dominated for a long time by the Braun-Blanquet (ZürichMontpellier) approach. Local studies and some specialised fields, e.g. cryptogamic and weed communities, will not be included. Aspects of community ecology will be treated in the companion paper by Kovács-Láng (1993). Only a few publications in Hungarian have been included (see Soó 1978 for a bibliography). References are combined with those in the next paper; App. 1, listing the main centres of research, follows after the References.
Fax +36 2760110
Important contemporaries of Soó were Pál Magyar, Imre Máthé, Zoltán Hargitai and Lajos Felföldy. Magyar, a forester, was a pioneer in the classification of forest vegetation. Máthé emphasised the phytogeographical aspect of phytosociology, introduced experiments in plant communities, and started production studies in the 1950's. Felföldy was the first to describe epiphytic cryptogam communities. He wrote a compendium on plant sociology, which was both theoretical and a good practical guide to field work as weil. One of the most prominent representatives of classicai plant sociology was Bálint Zólyomi. He published much in the 1950's and 1960's, mostly in German and treated special problems such as post-glacial vegetation history in relation to zonality and relic communities (Zólyomi 1953, 1957, 1987).
The early period The 'flourishing' time of classicai phytosociology Vegetation science in Hungary originated from the 19th-century plant formation approach (i.a. Kerner, Borbás, Bernátsky). Plant sociology as an independent discipline was introduced in Hungary in 1925 by Rajmund Rapaics in a book meant for the general public. Both European-based static phytosociology and Americaninspired dynamical ecology were included. This view was quite unusual at that time in Central Europe. Rezső Soó recognized the perspecti ves of the ZürichMontpellier School quite early; he founded the 'Phytogeographical School' in Debrecen and became a leading personality in the field. Most of his field work was carried out in the 1930's and 1940's. He had a vast lexical knowledge and natural ingenuity for synthesizing floristic, geographical and sociological facts. His magnum opus is a handbook in six volumes (Soó 19641980), including a history of Hungarian vegetation, a detailed syntaxonomic description and chorology of Hungarian plant communities, and a fairly complete 'biological flora'. Soó belonged to the c1assical phytosociologists who worked mainly qualitatively.
In the early 1950' s a new generation of geobotanists emerged. They were trained in phytosociology and phytogeographical mapping at Vácrátót by Soó & Zólyomi; new textbooks appeared, i.a. by Soó and Jávorka. In subsequent years many detailed phytosociological case studies were carried out, following a uniform approach, where the use of constant, characteristic and differential species was combined with a precise physiognornical description. Also, textural analysis was developed and spectra of life-fo rms and distribution types used. Representative examples are Tibor Simon (1957) and Attila Borhidi (1963-1965). Thus, the 1950's were a golden age for classical phytosociology. Phytosociology was of ten combined with vegetation mapping, although few maps were completed. ExampIes from the series 'The Vegetation ofHungarian Landscapes' include Simon (1957), Pócs et al. (1958), and Kovács (1962). Other map types include the elimatezone map of Borhidi (1961), and the widely used map (1 : 1 500000) of Hungary's potential natural vegetation
Current research & teaching
280
by Zólyomi,
which appeared
in the National
Atlas. For a
sociology
developed
during the 1950' s and 1960' s, par-
review of vegetation mapping. see Fekete (1980). A range of other aspects was included in the phytosociology of this period. Borhidi (e.g. 1963) paid attention to spatial and geographical relationships. He also applied a comprehensi ve phytosociological methodology in subtropical and tropieal regions as weIl. His reeent book on Cuba (Borhidi 1991) is perhaps the most complete and cornprehensive work concerning the phytogeography and vegetation ecology of a Latin American country.
ticularly in relation to forestry (Jakucs, Zoltán Baráth, Horánszky, Fekete, Pócs). Based on Zólyomi's work, it was concerned with the syntaxonomical description and mapping of forest types, using topographical and geological maps. Examples are the study of the Bükk Mountains (Zólyomi et al. 1955) and the monograph on Castanea satlva forests (Csapody 1969). These studies led to practical advice on sylviculture such as producti on estimates, natural renewal of stands and reforesta-
Gábor Vida (e.g. 1963) described altitudinal differences in montane beech forests. Tamás Pócs (1960) followed a phytogeographical approach and related the mixed forests of south western Hungary to the Baltic mixed forest zone. He also included quantitative aspects
tion. The effarts were appreciated by Hungariari forestry authorities. Unfortunately, this ecological foundation of forestry was abandoned later on. During the entire period of development of Hungarian plant sociology one therne has received special
in his analyses, sueh as rninirnal area. Fekete (1965) studied steppe-woodlands, unique for Central Europe and Fekete & Zólyomi (1966) studi ed mountain zonation in a sirnilar comparalive way. Adolf Horvát (1972) described the species-rich comrnunities of the Mecsek Mountains. An excellent description of the mire region in the Duna-Tiszaköze was made by Magda Járai-
attention and caused much debate, i.e. the original vegetation of the Alföld (the Hungarian Plain). Argu ments from the fields of floristics, vegetation geography and culture history have been used in various ways in order to achieve a convincing reconstruction. The Austrian plant geographer Anton Kerner von Marilaun, in his (1863) book on the plant life of the Danube countries, reg ard ed the plain as the western most extension of the climatic, treeless steppes of southern Russia. Early Hungarian geographers such as Rapaics argued that the
Kornlódi (1958). Her palynological analyses were summarised in a book on the Holocene environment of Hungary (Járai-Kornlódi 1987). Margit Kovács (e.g. 1962) described the dynamic nature ofwet meadows and related forest vegetation to soil and rnicroclirnate (Kovács 1975). György Bodrogközy (1960, 1965, 1970) studied relations between plant community and soil type in saltaffected areas and floodplains. Szujkó-Lacza (1962, 1964) described the species-pcor plant communities of andesitic bedrocks in northern Hungary. Pál Jakucs (1959,1961, 1968 and especially 1972) studied spatial and ternporal patterns in relation to geomorphology and microclirnate and developed ideas on cyclic succession and transitional environments with fringe communiries between forest and open vegetation. István and Vera Kárpáti (1969, 1975a) paid attention to dynarnics and related the succession of gallery forests to the geomorphological evolution of floodplains. Simon (1977) was the first to apply the degree of polyploidy as a criterion in Hungarian phytosociology. Pócs, Simon and András Horánszky fonned a group of 'reformers', who introduccd statistical methods for the eoruparisori and regrouping of sociological tables. Horánszky (1964) posed the interesting question, to what extent we can accept the sociological con ven ti on of fixed 'boxes' fitting certain piant-community types. The next generation of phytosociologists worked mainly with the traditional approach. It inc\udes Debreczy (1968) and Isépy (1970). Seregélyes (1974) used quantitative rnethods and was one of the first to use a graph-theoretical approach to cluster reIevés. An important
and successful
applied branch of plant
original landscape was forested and turned into steppe through human activities. As we gradually discovered, the situation is more complicated. Soó clairned in 1945 that the Alföld belonged to the forest-steppe region, the semi-arid elimate of which could facilitate the development offorests, steppes, and even mires, entirely depending on the local groundwater conditions. However, this forrnation-based view was valid mainly for the sanddune area in the Alföld, It does not apply to the extensive loess areas, which remained largely unknown untillittle of the original vegetation was left. Its reconstruction was accomplished by Zólyomi (e.g. 1957) and led to a more complex interpretation including edaphic dependence, history of human influence and dynamical status of the various cornrnunities, The lesson learnt is that the reconstruction of the potential natural vegetation requires both syndynamical and causal ecological approaches as weil as detailed floristic information. In summary. Hungarian plant sociology up to the 1960' s developed into a local variant of the ZürichMontpellier School, with equal ernphasis on syntaxonomical
and nomenclatural
aspects and a physiognomic
description. On the whole, it rernained a static approach; successional schernes were created only as by-products of descriptive studies. The bibliography of Hungarian synbiology between 1900 and 1972 by Soó (1978) gives an idea of the popularity of the different subdisciplines (number of publications, caregories taken from Soó):
.,
/
" Current~esearch History of phytosociology Terminology, methodology Case studies, vegetation rnonographs Phytosociology with geographical aspects Phytosociology with reference to pedology Sociology of lichens and bryophytes Segetal communities Syntaxonomy Syndynamics, experimental phytosociology Quantitative phytosociology
Broadening of the approach, theoretical development
14 23 395 17 65 96 6S 108 48 27
methodological
and
From the mid-1960's onwards many researchers gave up classical plant sociology. One important reason for this shift was the big influence Angio-American quantitative plantecology had on Hungarian ecologists.lt made many colleagues reali se that traditional sociology was too qualitative and therefore too subjective. Another reason was the impact of, and governmental support for the International Biological Programme. The shift was so drastic that it resulted in a certain discontinuity in Hungarian vegetation description. Quantitative plant sociology started with István Précsényi (e.g. 1964), who introduced statistical considerations in sampling and the study of distributional patterns. A central figure in this development is Pál Juhász-Nagy (1976, 1984; Juhász-Nagy & Podani 1983, etc.) who contributed and still contributes to the theoretical and methodological foundation of ecology at large. He also developed a family of information theory models with 'characteristic functions' for, i.a. 'florula diversity', 'local distinctiveness' and 'associatum'. These models are based on species Iists of sampling units of increasing size. Characteristic areas can be defined in terms of the maxima and minima of these functions. A new generation of quantitative plant ecologists arose under the influence of Précsényi and JuhászNagy. Amongst the first were Edit Molnár and János Nosek, who worked on spatial processes in a Festucetuni vaginatae grassland (Molnár & Nosek 1979, 1980; Nosek 1976, 1986). Zol tan Szöcs (1979) developed computer programmes to study spatial patterns by computer simulation. This, and other work on spatial analysis was reviewed by János Podani (1984a). Podani (1984b, 1987, 1989a,b; Podani & Dickinson 1984; Kenkel, JuhászNagy & Podani 1989) developed some new methods on sampling and c1uster analysis. His program package SYN-TAX (Podani 1988) is much used both in Hungary and abroad. Podani (1985) also used non-floristic descriptors in classification; he showed that the subsequent evaluation of character states may recover aspects of vegetation structure that are not apparent otherwise.
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281
Results of multivariate analysis can be compared and integrated with the traditio nal syntaxonomical approach, as is exemplified in the revision of the Fagion illyricum alliance (Török, Podani & Borhidi 1989). The concepts and models of Juhász-Nagy were applied by Sándor Bartha (1990, 1991, Bartha & Horváth 1987; Szollát & Bartha 1991). He devised a simplified, computerized sampling procedure for belt transects. The characteristic information functions prov ed to be useful for detecting the development of topographic patterns during primary succession. Stabilized and abiotically controlled cornmunities can becharacterized as weil, e.g. by the invariance of characteristic-ordering relationships. Since the application of this model for the deterrnination of the characteristic areas requires the analysis of many sam pl ing units, a three-way multivariate technique has been developed (Camiz& Gergely 1990) for describing the behaviour of this model. Another development is a cornpu ter simulation for the study of the effects of local constraints on plant population dynamics by Tamás Czárán (1984, 1989). Then, Czárán & Bartha (1989) cornpared simulation results with real field data. They generated community-point patterns using life-table data, dispersion parametérs and radii of competitive effects. They concluded that the spatiallimits of dispersal and competition may result in the increased persistence of weak competitors, may moderate the realized competitive effects of strong species, and may shape the spatial coalition structure of the community. A newapplication of numerical methods is that of Péter Csontos (1986), who studied the 'autocoenology' of lmpatiens parviflora. Autocenology, a concept parallel to autecology, provides a possibility to bridge phytosociology and population biology. Demographical and dispersal attributes of the local population were recognized as driving forces in succession. In a study of sandy grassland, Fekete, Tuba & Melkó (1988) distinguished elementary 'cenosystems' and developed causal schemes to indicate the effect of accompanying species on the reproductive allocation of the dominant population. Another new development is concerned with sociological pattern analysis. László Körrnöczi & Andrea Balogh (1990) used dominance shifts along belt transects to detect vegetation boundaries. Tibor Standovár (1985, 1986) studied both vegetation and soil patterns in mountain meadows and arrived at direct causal-analyticaI questions to be investigated. In the field of vegetation dynamics new developments started after a symposium on cenological suecession in 1982. A number of permanent plots was established, which serve as a framework for observing long-term spatio-temporal changes and results of sophisticated experiments. A wide range of approaches
Current research & teaching
282
.,
was developed. Précsényi (1981) used a comparisen of traditional sociological tables to show a non-Iinear change in diversity during succession, Diversity changes were also followed dllring regeneration succession on permanent plots by students of
OutIook
Précsényi andJakucs, and Béla Tóthrnérész,
ence' is characterized by some distinctive features. One is the emphasis on theoretical foundations, another is the arial ysis of spatial processes, in relation to the anal y-
a high disorder
notably Mária Papp, Ilona Mészáros After deforestation of an oakwood,
was observed,
which decreased
as early
At present, vegetation science is a diverse field of science, both regarding methods and prograrnmes. It may be stated that the 'new Hungariari Végetarion Sci-
as in the third year (Papp 1984, 1987). Mészáros (1990) described spatial and temporal changes in the herbaceous
sis of vegetation structure. As to the near future, some new lines of research
vegetation of an ecotone between beech forest and a clear-cut area using permanent transects. Tóthmérész ( 1987, 1991) studyi ng recovery patterns of the vegetation after clear-cutting, found that ecological features of species were the most sensitive indicators of successional changes, rnore sensitive than species abundance. István Bagi (1985, 1987a,b; Bagi & Körrnöczi 1986) used the short-lived species of riverbed communities (Nanocyperion) to demonstratc that the successive ap-
obvious and will certainly be further developed in Hungary. First, research of plant populations will be extended and linked to studies on the community level, with a special emphasis on the occurrence of mosaics of various communiries. Integration of population biology into vegetation science, as seen above, is expected to take place along various lines. Of special iruportanec here, as documented by new, unpublished results, is the usefulness of the concept of 'cenopopulations' in ana-
pearance of communiries in these zones is different, while the succession in itself is divergent. Degradation processes were studied as weil. The
Iysing the 'in situ' demography and dynamics of natural grassland populations, The development of vegetation science will also be
effect of grazing by cattle and geese on grassland was analysed by Précsényi and others (Précsényi 1990). The critical time of regeneration of a Cynodonti-Festucetum pseudovinae grassland is at least 7 yr (Matus & Tóthmérész 1990). The latter authors also developed a new method to reveal the coalition structure of communities based on the combination of association analysis, plexus
stimulated by practical needs. Nature conservation is an exarnple here. Plant sociology and vegetation dynamics may be regarded as indispensable for documentation, community diagnostics and landscape reconstruction. As an example, conservationists recognized the suitability of Juhász-Nagy's information theoretical func-
graph methcd and Principal Coordinate Analysis; both positive and negative associations were less frequent in the grazed situation th an in the non-grazed one. Török (1989) assessed the rate of disturbance caused by tourists and over-populatiori of game on grassland. Klára Virágh (1982, 1986, 1987) followed the effects of the selective removal of the dominant species by leafherbicides in an old perennial grassland. The degree of community recovery was assessed by comparing
community.
sirnilarity
gress in Hungary,
relations,
many textural
and structural
com-
tions as tools for detecting Another
fine structural
promising
changes
are
in the
field is the application
of island biogeography to the ecology of isolated patches of natural végetation. Recently, the study of sociological 'islands' produced some interesting results relevant to conservancy as weil. There is a great dernand for accurate vegetation maps which serve as a basis for recording the status quo of the vegetation and to document and evaluate changes in it. The application of remote sens ing is making proas weil as some methods
for evaluat-
munity attributes and population recovery patterns in control and herbicide-disturbed plots (Virágh 1989a,b). In an intact stand she demonstrated between-year structural changes due to fluctuating weather conditions, as László Körmöczi (1989) did in another grass land. Some new experimental approaches are promising for the study of population processes. Artificial two-
ing rnaps. Inevitably, the fast spread of computers will further promote the development of numerical vegetation science. Efforts are made to establish computerized data
species communities offer possibilities for describing interactions within and between populations ('noise' is negligible here as cornpared with natural communities). Edit Molnár described intra- and interspecific competition and their relative importance in pure and mixed stands of Amaranthus chlorostachys and Chenopodium albuin.
tion: the ideas of László ürlóci and his associates development are influential in Hungary.
banks for both vegetation and flora. Computer-assisted methods tend to be more and lllore important as to sampling procedures, data process ing and data evalua-
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