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© 2010 Gebrüder Borntraeger, 70176 Stuttgart, Germany www.borntraeger-cramer.deDOI: 10.1127/0340-269X/2010/0040-0425 0340-269X/10/0040-0425 $ 16.65

Phytocoenologia, 40 (2–3), 205–241Stuttgart, November 15, 2010

Vegetation classifi cation and synecology of Bohemian Robinia pseudacaciastands in a Central European context

by Michaela VÍTKOVÁ and Ji�í KOLBEK, Pr�honice

with 15 fi gures and 5 tables

Abstract. Synanthropic forest and shrub vegetation dominated by Robinia pseudacacia spontaneously growing inthe open landscape and suburban areas in Bohemia (Czech Republic) was analysed. In total, 374 of 502 knownphytosociological relevés were synthesized – 302 relevés were sampled in the fi eld and 72 published relevés were obtained from the literature. Within the class Robinietea (including two orders and three alliances) one association is validated and two new subassociations are described: Arrhenathero elat ioris-Robinietum (including var. Cal-amagrostis epigejos), Poo nemoral is-Robinietum cardaminopsietosum arenosae and Poo nemoral is-Robinietum typicum. Currently, four associations of these Black Locust stands from Bohemia are recognised: Chel idonio-Robinietum, Poo nemoral is-Robinietum, Arrhenathero elat ioris-Robinietum, and Melico transsi lvanicae-Robinietum. The description of several units is amended with synecological charac-teristics (altitude, aspect, slope, bedrock, soil type, and important physicochemical and biochemical characteristics)and distribution both in Bohemia and, according to hints in the literature, also in Central European countries. The position of Black Locust stands in the syntaxonomical system is discussed, and nomenclatural types and diagnosticspecies combinations of higher syntaxa are given. The current level of knowledge of Black Locust stand classification fiin Central Europe is compared.

Keywords: Black Locust, Central Europe, classification,fi Czech Republic, distribution, phytosociology, Robinie-tea, synecological characteristics.

Introduction

Black locust (Robinia pseudacacia L.) is a controver-sial tree species which has been the subject of discus-sions among groups of foresters and conservationists worldwide for a long time. Its area of original distri-bution is the southeast of the USA (Fowells 1965).Black Locust created continuous secondary stands later after North American colonization. Its behav-iour corresponds to an early succession species – itpenetrates expansively with its aggressive root and trunk coppice shoots especially into areas damaged by fi re, clear-cuts, and abandoned fi elds and pastures, and thus it subsequently colonised the whole area of North America (Vadas 1914, Beck 1988, Huntley 1990). When colonizing open stands, Black Locust’srapid growth and the ability to fi x the nitrogen fromthe air are advantages that increase its competitiveabilities, and Black Locust can soon become a domi-nating species. As the amount of nitrogen in soil in-creases, within 20 – 30 years it ultimately functions as a limitating factor and the ratio of its occurrence inthe stand decreases to below 4%, favouring shadow-tolerant species (e.g. Boring & Swank 1984, Mon-tagnini et al. 1991, Elliott et al. 1998). In recentexperience this natural mechanism does not functionin secondary ranges (e.g. Kowarik & Langer 1994,Platen & Kowarik 1995). Because of the high helio-phytism of Black Locust, the possibility of its inva-

sion into closed natural forest communities is low. Incontrast, it is able to spread aggressively on steppeand sandy localities and in light, unclosed forests onslopes using its root and trunk coppice shoots andsubsequently to change the species composition of the herb layer, which is sensitive to its nitrification fieffect.

Black Locust was imported to Europe at the be-ginning of seventeenth century for gardening as oneof the fi rst North American tree species (e.g. Kolbek et al. 2004). It was already recommended for plantingin the eighteenth century for its long-lasting quality timber which is resistant to insects and fungi, rapidgrowth, easy reproduction, and ability to stabilizesoil (e.g. Göhre 1952, Wendelberger 1955), whichresulted in wide spreading of Black Locust mono-cultures to almost the whole world (e.g. Keresztesi 1988). According to Weber (2003), Black Locust ex-panded to the whole of Europe, except the northernpart and Mediterranean islands, and to North andSouth Africa, temperate Asia, Australia and NewZealand, Chile, and Argentina.

Unlike in Hungary and Germany, where it was used especially for stabilizing sand soils, fluvial de-flpositions, and aeolian sands, in the Czech Republic,except the Elbe lowland and a part of South Moravia,Black Locust was introduced mainly to former pas-tures on steep eroded hillsides along rivers (especially

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206 M. Vítková & J. Kolbek

Vltava, Berounka, Sázava, and Dyje; Fig. 1) and sur-rounding railways (Kolbek et al. 2004). Where Black Locust penetrated, it has always adapted very well to local conditions and created specifi c stands on diverse habitats, highly different from the autochthonouscommunities (e.g. Svobodová 1952, Wendelberger1954, 1955, Větvička 1961, Frantík 1985, Montag-nini et al. 1991, Peloquin & Hiebert 1999, Vítková 2004, Von Holle et al. 2006, Taniguchi et al. 2007).

We addressed the following aims: (1) Phytocoeno-logical classification of fi Robinia pseudacacia stands in comparison with Central European dates; (2) Com-parison and distribution of described units in Central Europe; (3) Evaluation of new sampled data from Bo-hemia; (4) Comparison of basal synecological charac-teristics of different Robinia pseudacacia communi-ties.

Vegetation classification of Black Locust fistands and their historical development in different countries

Classifying Black Locust stands in the phytosocio-logical system has been an activity of Central Euro-pean botanists since the second half of the twentiethcentury. The historical development was summarised

by Kolbek et al. (2004). The oldest and most exten-sive studies were carried out in Hungary. Pócs (1954)described association Robinietum pseudo-aca-ciae, which he further split according to successionalage and moisture regime. He submited also a proposalof the successional trend from the natural plant com-munities to Black Locust forests with dominatingBromus sterilis. Soó (1957, 1958) considered BlackLocust stands in Hungary as only a cultural con-soci-ation of different forest phytocoenoses and describedthem as Robinio-Brometum steri l is Soó 1957. According to the Code of Phytosociological Nomen-clature (Weber et al. 2000), publication of this asso-ciation is not effective. Fekete (1965) published asso-ciation Bromo steri l is-Robinietum and placedinto the class Quercetea pubescenti-petraeae.However, it was defi ned in the same coverage as thefiassociation Robinietum pseudo-acaciae Pócs 1954, hence it is only later synonym. The classification fiof Black Locust stands in Hungary was further stud-ied in 50’s and 60’s of 20th century by Tallós (1960), Timár & Bodrogközy (1959), Tuskó (1956), andMajer (1962, 1968). In the overview of plant commu-nities of Hungary, Borhidi (1996) classifi ed Black fiLocust stands as a lone-standing class, Robinie-tea Jurko ex Hadač & Sofron 1980, with one order,Chelidonio-Robinietal ia Jurko ex Hadač & So-

Fig. 1. Typical habitat of Black Locust stands growing on steep slopes in the Vltava valley (Central Bohemia).

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 207a

fron 1980, and two alliances: (1) Chelidonio-Rob-inion Hadač & Sofron 1980 (with two associations: Anthrisco cerefol i i-Robinietum Majer 1963 (recte 1962) and Chelidonio-Robinietum Jurko1963) and (2) Bal loto nigrae-Robinion Hadač &Sofron 1980 (with Balloto nigrae-Robinietum Jurko 1963 and “Bromo steri l is-RobinietumPócs 1954”, recte Robinietum pseudo-acaciae).Borhidi (2006) proposed in the online version of his later book (Borhidi 2003) to establish a new order, Bal loto nigrae-Robinietal ia Balogh & Borhidi2006, for Black Locust stands on dry, warm soils poor in nutrients. According to the Code of Phy-tosociological Nomenclature (Weber et al. 2000), publication of this order and association Anthriscocerefol i i-Robinietum is not effective. Borhidi (2003) further presented the associations Anthrisco cerefol i i-Robinietum and Chelidonio-Rob-inietum as very similar, and they could probably bemerged together.

Scamoni (1954) and Wendelberger (1955) were among the fi rst who studied the sociology of Black Locust stands in Germany and Austria. Scamoni (1954) distinguished three types of stands: 1. moistshadow nitrophilous stands with Chelidonium, 2. lighter grassy stands with Agrostis vulgaris, and 3.thermophilous stands with Brachypodium sylvati-cum. Wendelberger (1955) disaffirmed association-fial character of Black Locust stands. He marked themonly as degradation phases of natural forests, whichcould build relatively stabilised secondary communi-ties called “Sambuceto-Robinietum” or in allu-vial plains “Solidageto-Robinietum”. Accordingto their sociological structure they could be assignedto ruderal communities.

The current phytosociological classification of fiBlack Locust stands in Germany differs accordingto different authors. None of them recognize the class Robinietea. Kowarik (1995) introduced argu-ments about the forest character of Black Locuststands and proposed to classify them into the classQuerco-Fagetea within the lone-standing orderRobinietal ia , based on his experience of long-term research of the ecology and successional behaviour of Black Locust stands (e.g. Kowarik 1986, 1992,Kowarik & Langer 1994). Oberdorfer (1992)used a slightly different approach and included theseforests from South Germany in the alliance Berber-idion Br.-Bl. 1950 (class Querco-Fagetea, order Prunetal ia spinosae). He placed them within the community Galium aparine-Robinia pseudacaciaMüller 1966 and assigned var. Robinia pseudacaciato various shrub communities. By contrast Klauck (1988), due to the ruderal character of the herb layer,inclined to the opinion to place Black Locust standsinto the class Artemisietea vulgaris as a Sam-bucus-Robinia community with three vicariances –Eastern, Western, and Central European. Diesing& Gödde (1989) classifi ed them as a communitywith Robinia pseudacacia within the class Epilobi-etea angustifol i i , alliance Sambuco-Sal ic ion

caprae. Pott (1992) included all the communitieswith Robinia pseudacacia in Germany (like Mucinaet al. 1993 in Austria) in the class Galio-Urticeteaand order Lamio albi-Chenopodietal ia boni-henrici .

Willner & Grabherr (2007a) disagreed with the approach of Mucina et al. (1993) due to the differ-ent habitats and species composition of Black Locust stands in Austria. They described two clearly charac-terised associations: (1) Chel idonio-Robinietum on the fresh habitats and (2) Bal loto nigrae-Robinietum on the slightly dry to slightly freshhabitats. They included these two associations in the class Rhamno-Prunetea, alliance Arctio-Sam-bucion nigrae, but they gave an option to classify them into the lone-standing class Robinietea. They considered this classifi cation to be provisional, re-fiquiring further work.

Gilicka (1989) classifi ed Black Locust standsin Poland within the class Artemisietea vulgaris , order Galio-Calystegietal ia . Świerkosz (1993) included them in a different order and alliance within this class, namely Chelidonio-Robinietal ia and Chelidonio-Robinion. Matuszkiewicz (2004, unpublished) and Pawlaczyk (2008) included theassociation Chelidonio-Robinietum in the alli-ances All iar ion or Galio-All iar ion respectively.Black Locust stands were not mentioned at all in thelast published overviews of plant communities in Po-land (Matuszkiewicz W. 2001, Matuszkiewicz J. M.2001). In a West Polish index of plant units (Brzeg &Wojterska 2001) only Chelidonio-Robinietum waslisted, but without a classification into a hierarchicalfisystem.

Solomakha (2008) classifi ed Black Locust forests fiin Ukraine within the separate class Robinietea,order Chelidonio-Robinietal ia , which includesboth plantations and spontaneous stands of alloch-thonous woody species, for example Robinia pseuda-cacia, Populus nigra, Acer negundo, Eleagnus argen-tea, and Rhus typhina.

None of the Italian authors (Arrigoni 1997, Cor-betta & Pirone 1995, Gentile 1995, Mondino &Scotta 1987, Parolo 2000) recognized the class Robinietea. At the community level they classi-fi ed Black Locust stands according to elements of the fioriginal units within the orders Prunetal ia , Fag-etal ia sylvat icae or Quercetal ia pubescenti-petraeae. By contrast Theurillat et al. (1994) andAeschimann et al. (2004) included planted and natu-ral secondary Black Locust stands from the Alps in the lone-standing class Robinietea, order Cheli-donio-Robinietal ia , alliance Bal loto nigrae-Robinion.

Mititelu et al. (1973) classifi ed Black Locustfistands growing in Romania (Hanu-Conachi Na-ture Reserve) as the association “Bromo steri l is-Robinietum Pócs 1954” with six subassociations. According to many thermophilous taxa from the class Quercetea pubescenti-petraeae, BlackLocust stands were placed into this class as the order

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208 M. Vítková & J. Kolbek

Robinion pseudacaciae Csürös-Káptalan 1968,pursuant to Oprea (2004) and Sanda et al. (2008).Publication of this alliance is probably not effective(see the chapter Discussion). Covali (2009) includ-ed Black Locust stands growing in Moldova (ValeaMare, Zberoaia-Lunca) in the association Balloto nigrae-Robinietum (Galio-Urticetea, Chel i-donio-Robinietal ia and Balloto nigrae-Rob-inion).

In the Netherlands, Stortelder et al. (1999) stud-ied Black Locust stands marginally, describing them as a derivate community within the class Quercetearobori-petraeae.

In the Korean Peninsula, Black Locust forestswere mentioned in a few papers, especially by Cho et al. (1999), Cho & Kim (2005) and Kolbek & Ja-rolímek (2008). Only one association, Commeli-no-Robinietum pseudoacaciae Cho & Kim 2005, with six subassociations was described and doc-umented by phytosociological relevés. In comparison with Central Europaean Robinia-stands, the associa-tion of Northeast Asia represents an analogous unit to Chelidonio-Robinietum. From North Korea(Kolbek & Sádlo 1996), Robinia pseudoacacia is known as an accompanying species of the alliances Cosmo-Humulion japonicae, Plantaginion asiat icae, Zoysion japonicae and, on the river-side and coast, of Yellow Sea vegetation (Kolbek &Jarolímek 2008).

Historical development of Black Locuststand classifi cation in the territory of formerfiCzechoslovakia

Jurko (1963) was the fi rst to deal with Black Lo-cust stand classifi cation in the territory of the former fiCzechoslovakia. Within the class Robinietea two or-ders were distinguished:a) Chel idonio-Robinietal ia on nutrient rich,

deep, fresh clayey soils with favourable waterconditions (Chelidonio-Robinietum andSolidagini-Robinietum),

b) Bromo-Robinietal ia on low to medium nu-trient rich soils, usually shallow, rather dry, withskeleton or sand (Bal loto-Robinietum andBromo-Robinietum).

Jurko (1963) forgot to describe the alliances and his publication of both orders and also the class wasconsequently invalid (cf. Weber et al. 2000, Art. 8).Other associations from Slovakia were mentioned byŠčepka (1982), but all of them were only provisional:Sambuco nigrae-Robinietum, Urtico-Rob-inietum, Fumario off ic inal is-Robinietum , Aristolochio clematit is-Robinietum, andGalio aparines-Robinietum. Šimonovič et al.(2002) proposed next provisional association Ar-rhenathero elat ius-Robinietum based on fi vefiphytosociological relevés. The classifi cation of BlackLocust stands in Slovakia was further studied byBenčaťová & Benčať (2005, 2008).

Hadač & Sofron (1980) revised the classifi cationfipresented by Jurko (l.c.) and proposed preserving the class Robinietea. They merged both orders intoone: Chel idonio-Robinietal ia , and described two new alliances (Chelidonio-Robinion andBalloto nigrae-Robinion) without typifica-fition. Basic data were collected mostly in Slovakia,and only six phytosociological relevés from Bohemia were taken into account (from the Berounka valley, Sofron 1967).

The fi nal classififi cation proposed by fi Hadač & So-fron (1980): class Robinietea Jurko ex Hadač & Sofron 1980, order Chelidonio-Robinietal ia Jurko ex Hadač & Sofron 1980 includes two alliances: Chel idonio-Robinion Hadač & Sofron 1980 (with associations Chelidonio- Robinietum Jurko 1963, Impatienti parvif lorae-Robinietum Sofron 1967, Sol idagini-Robinietum (Wendelberger 1955) Jurko 1963), Bal loto nigrae-Robinion Hadač & Sofron 1980 (with associations Balloto nigrae-Robinietum Jurko 1963, Bromo steri l is-Robinietum Jurko 1963 ass. prov.).

Jarolímek & Šibík (2008) reached the same classi-fi cation asfi Hadač & Sofron (1980) only on the basisof numerical analysis of 48 phytosociological relevésknown from Slovakia. They completed the list of as-sociations with provisional associations described byŠčepka (1982). Svobodová (1952), Větvička (1961),Blažková (1961), and Sofron (1964, 1967) carried out the fi rst phytosociological studies on Black Lo-cust stands in Bohemia. The only validly describedassociation was characterised by Sofron (1967) asImpatienti parvif lorae-Robinietum from schist debris in the Berounka valley. He also provi-sionally classifi ed the association Poo nemoral is-Robinietum. Němec (1981) described three asso-ciations of the alliance Chelidonio-Robinion: Urtico dioicae-Robinietum, Poo nemora-l is-Robinietum and Rubo fruticosae-Rob-inietum without publishing nomenclaturaltypes. Vítková et a l . (2003) c lassi f ied the Black Locust stands in the Kř ivoklátsko PLA (Central Bohemia) . A new order Eu-phorbio cypariss iae-Robinietal ia with only one alliance Euphorbio cypariss iae-Robinionand association Melico transsi lvanicae-Rob-inietum were described for small-scale stands on ex-tremely dry rocky slopes, including low to shrubbyxerotherm Black Locust stands. The association Poonemoral is-Robinietum of the alliance Chelido-nio-Robinion was validated.

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 209a

Table 1. List of published names of units of Black Locust communities with phytosociological relevés or synoptic tables from Central Europe.

Unit Country Author Nr. of relsAristolochio clemati t is-Robinietum Slovakia Š�EPKA (1982) 1Arrhenathero-Robinietum Bohemia VÍTKOVÁ (2004) 21

Slovakia ŠIMONOVI� ET AL. (2001) 1Slovakia ŠIMONOVI� ET AL. (2002) 4

Balloto nigrae-Robinietum Austria WILLNER & GR RABHERR (2007) ST(42)R

Slovakia JURKO (1963) ST(13)Slovakia Š�EPKA (1982) 1Slovakia BEN�A�OVÁ & BEN�A� (2005) 1

Chel idonio-Robinietum Austria WILLNER & GR RABHERR (2007) ST(75)R

Bohemia KUBÍKOVÁ (1982) 1Bohemia KUBÍKOVÁKK (1997) 1Bohemia KOLBEK ET AL. (2003) 23Bohemia VÍTKOVÁ (2004) 140Germany KOWARIK (1986)K 1Poland SWIERKOSZ (1993) 9Poland PAWLACZYK (K PERS.COM.) 13

Slovakia Š�EPKA (1982) 1Slovakia JURKO & KONTRIŠ (1982) 8Slovakia ŠIMONOVI� ET AL. (2001) 1Slovakia ŠIMONOVI� ET AL. (2002) 11Slovakia BEN�A�OVÁ & BEN�A� (2005) 1Slovakia BEN�A�OVÁ & BEN�A� (2008) 1Slovakia KONTRIŠ ET AL. (2009) 7

Slovakia and Hungary JURKO (1963) 15Cardaminopsio arenosae-Robinietum Bohemia VÍTKOVÁ (2004) 28Fumario off ic inal is-Robinietum Slovakia Š�EPKA (1982) 1Galio aparines-Robinietum Slovakia Š�EPKA (1982) 1

Slovakia BEN�A�OVÁ & BEN�A� (2005) 1Slovakia BEN�A�OVÁ & BEN�A� (2008) 1

Impat ient i parvif lorae-Robinietum Bohemia SOFRON (1967) 7Bohemia KOLBEK ET AL. (2003) 5Bohemia VÍTKOVÁ (2004) 70

Melico t ranssi lvanicae-Robinietum Bohemia KOLBEK ET AL. (2003) 27Bohemia VÍTKOVÁ (2004) 32

Poo nemoral is-Robinietum Bohemia N�MEC (1981) 8Bohemia SOFRON (1964) 3Bohemia KOLBEK ET AL. (2003) 15Bohemia VÍTKOVÁ (2004) 123

Robinietum pseudo-acaciae Hungary PÓCS (1954) 17Hungary FEKETE (1965) 7Poland SWIERKOSZ (1993) 6

Romania OPREA (2004) 5Slovakia Š�EPKA (1982) 1Slovakia ŠIMONOVI� ET AL. (2001) 1Slovakia ŠIMONOVI� ET AL. (2002) 12Slovakia BEN�A�OVÁ & BEN�A� (2005) 1Slovakia BEN�A�OVÁ & BEN�A� (2008) 1

Rubo frut icosae-Robinietum Bohemia N�MEC (1981) 14Sambuco nigrae-Robinietum Slovakia Š�EPKA (1982) 1Solidagino-Robinietum Slovakia JURKO (1963) 1Urt ico dioicae-Robinietum Bohemia N�MEC (1981) 10

Slovakia Š�EPKA (1982) 1Slovakia BEN�A�OVÁ & BEN�A� (2005) 1

comm. Galium aparine-Robinia pseudacacia Germany OBERDORFER (1992) ST(92)R

comm. Sambucus nigra-Alliaria petiolata-Robinia Germany KLAUCK (1986)K 13

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210 M. Vítková & J. Kolbek

Material and methods

For classifi cation only those phytosociological rele-vés from successionally mature Black Locust standsgrowing in open landscape and suburban areas inBohemia were used. They originated from self-re-newing old plantations unimpacted by recent forest management from the end of the nineteenth to the middle of the twentieth century or from stands cre-ated by Black Locust invasion into natural communi-ties. Relevés from small-area stands in the urban ar-eas, from mixed forests with other tree species where Black Locust is not dominant, successionally young stands, plantations on dumpsites, and so on were not used. Further, phytosociological relevés from other Central European countries were not incorporatedbecause they do not match the above-listed criteria, or were available only as drafts or in incomplete pub-lications (published without accessory species or in asynoptic table with disagreement on a conception of association differentiation; see Table 1). An obtained data-set of phytosociological relevés from Central European Black Locust stands suitable for classifica-fition would be too small and thus unrepresentative (Table 1). Published data and unpublished relevés from Central Europe (Pawlaczyk, Poland; Böcker & Dirk, Germany) were used for a study of distribution and in comparison with individual syntaxa of Black Locust stands.

From 502 available phytosociological relevés from Bohemia 374 were used for syntaxonomical revision, of which 302 originated in our own fi eld research and another 72 from other authors (authors of relevés in Table 4B: MV = M. Vítková, VV = V. Větvička). Rel-evé material was collected in 1997 – 1999 using the Braun-Blanquet methods: seven-grade scale of abun-dance and dominance by Braun-Blanquet (1964). The composition of the moss layer was not deter-mined. Most of the relevés were recorded on plots of 250 m2; in some small-scale steppe stands the area was

reduced to 100 or 150 m2. Altitude, aspect, slope andbedrock were recorded for each relevé. The potential natural vegetation and names of the associations of natural communities were indicated for each unit ac-cording to the study by Neuhäuslová et al. (1998).The names of higher syntaxa follow Moravec etal. (1995) except Quercion petraeae and Caricihumil is-Quercetum (Kolbek et al. 2003). Thelocalities of all sampled and processed relevés were published by Vítková (2004, appendix 1).

A TURBO(VEG) database (programme version9.39, Hennekens 1996) was created from all thephytosociological relevés and tables. The nomencla-ture of taxa was used according to the programme(Ehrendorfer 1973). The only exceptions are Cra-taegus curvisepala and Rosa elliptica, which are used according to Neuhäuslová & Kolbek (1982). In Ta-ble 2 the diagnostic species are ordered according to the percentage in each unit and other species accord-ing to the total percentage in all relevés. The basicsplitting of the very diverse dataset was done by theagglomerative numerical classification in the SYN-fiTAX 2000 programme using the Ružička similaritycoeffi cient and thefi �-fl exible (fl � = - 0.25) method (Po-dani 2000). Indirect gradient analysis was comput-ed for the created groups using the CANOCO for Windows 4.0 programme (Braak & Šmilauer 1998). The more neutral term “diagnostic species combina-tion” (cf. Moravec et al. 1994, Jarolímek & Šibík2008) was used to characterise syntaxonomical units of Black Locust stands (association and higher syn-taxa) instead of dividing taxa into characteristic and differential. The term “differential species” was usedfor differentiation of subassociations (Koch 1926,Moravec et al. 1994).

Operating sequences for soil sample analyses (33localities) were chosen with respect to methods com-monly used for research in natural forest ecosystems (e.g. Moravec 2000, Kolbek et al. 2003). Values of ph (H2O), pH(KCl) and carbonate concentration

comm. Sambucus nigra-Robinia pseudacacia W and E Europe KLAUCK (1988)K ST(53)comm. Robinia-Calamagrostis epigejos Poland PAWLACZYK (K PERS.COM.) 9Robinia pseudacacia stands with Impatiens parvi�i ora�� Bohemia V�TVI�KA (1961) 8Robinia pseudacacia stands with Melica transsilvanica Bohemia V�TVI�KA (1961) 5Robinia pseudacacia stands with dominating grasses Bohemia V�TVI�KA (1961) 17shrubby Robinia pseudacacia stands on sunny rocks Bohemia V�TVI�KA (1961) 8moist Robinia pseudacacia stands on slope bases Bohemia V�TVI�KA (1961) 15Robinia pseudacacia stands Bohemia KLIKA (1949) 1

Bohemia BLAŽKOVÁ (1961) 7Bohemia KOLBEK ET AL. (1980) 2Bohemia FRANTÍK (1985)K 12

Bohemia and Slovakia SVOBODOVÁ (1952) 46Croatia HRUŠKA (1991) 10

Germany KOWARIK & LK ANGER (1994) 24R

Germany BÖCKER & DR IRK (K PERS.COM.) 12ST (x) … synoptic table consist of x relevés

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 211a

Column 1 2 2A 2B 3 4 C(%)Number of relevés 174 146 117 29 21 33Number of species 205 165 201 290 355 230

Chelidonio-RobinietumE2 - shrub layerSambucus nigra 97 49 56 24 38 3 100Ribes uva-crispa 54 30 30 28 38 18 100

E1 - herb layerGalium aparine 93 53 56 38 52 33 100Geum urbanum 91 38 34 52 29 15 100Urtica dioica 90 22 26 7 52 3 100Anthriscus sylvestris 68 14 17 . 5 3 100Poa trivialis 58 0.7 . 3 43 . 75Chaerophyllum temulum 50 12 15 3 14 3 100Taraxacum sect. Ruderalia 45 23 25 17 19 6 100Arctium minus agg. 25 0.7 0.9 . . . 50Lamium album 17 3 4 . . . 50

Poo nemoralis-RobinietumE2 - shrub layerPrunus spinosa 11 30 28 35 . 19 75

E1 - herb layerPoa nemoralis 22 95 97 90 43 58 100Viola arvensis 6 28 23 48 5 19 100Dryopteris filix-mas 2 23 22 24 . . 50Galeopsis pubescens 7 21 23 10 . 6 75Avenella flexuosa 2 18 15 31 5 7 100

Poo nemoralis-Robinietum cardaminopsietosum arenariaeE3 - tree layerQuercus robur 0.8 8 5 21 . . 50

E2 - shrub layerCrataegus monogyna 14 19 15 38 10 9 100

E1 - herb layerCardaminopsis arenosa 2 24 16 55 . 21 75Brachypodium pinnatum 6 19 12 45 . 9 75Hylotelephium maximum 2 12 6 38 5 6 100Festuca ovina . 8 0.9 38 10 . 50Viola collina 18 14 9 35 . 3 75Stellaria holostea 2 16 11 35 5 . 75Polygonatum odoratum 0.8 11 6 31 5 9 100Pimpinella saxifraga 2 7 0.9 31 . 6 75Lychnis viscaria . 6 3 17 . . 25

Arrhenathero elatioris-RobinietumE1 - herb layerArrhenatherum elatius 21 21 22 14 81 18 100Calamagrostis epigejos 2 14 13 17 57 6 100

Table 2. Synoptic table of the class Robinietea in Bohemia. The full data matrix computed contained 374 relevés and 475 species. Allspecies with frequencies lower than 10% in columns 1, 2, 3, 4 were excluded. Constancy was calculated only for associations, i.e. onlyfor columns 1, 2, 3, and 4 (without 2A and 2B). The subassociations 2A and 2B are displayed in italics.1 - Chel idonio-Robinietum2 - Poo nemoral is-Robinietum2A - Poo nemoral is-Robinietum typicum2B - Poo nemoral is-Robinietum cardaminopsietosum arenosae3 - Arrhenathero elat ioris-Robinietum4 - Melico transsi lvanicae-Robinietum

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212 M. Vítková & J. Kolbek

Elymus repens 15 3 4 . 38 3 100Poa angustifolia 10 4 5 . 38 15 100Agrostis capillaris 0.8 3 3 3 38 3 100Dactylis glomerata 8 3 3 3 33 . 75Festuca rubra agg. 2 3 3 7 33 3 100Dryopteris dilatata 5 3 3 3 29 . 75Anthoxanthum odoratum . 3 4 . 29 3 75

Melico transsilvanicae-RobinietumE2 - shrub layerRosa canina agg. 11 33 29 48 5 52 100

E1 - herb layerEuphorbia cyparissias 2 19 6 72 . 76 75Melica transsilvanica . 7 6 10 . 76 50Vincetoxicum hirundinaria 3 19 17 28 . 49 75Verbascum lychnitis 2 8 7 14 . 49 75Festuca pallens . 3 3 3 . 46 50Centaure stoebe . . . . . 46 25Thymus pulegioides . 1 . 7 . 42 50Artemisia campestris 0.8 . . . . 36 50Dianthus carthusianorum . 1 0.9 3 . 27 50Asperula cynanchica . 0.7 0.9 . . 27 50Aurinia saxatilis . . . . . 27 25Seseli osseum . . . . . 27 25

Chelidonio-RobinionE2 - shrub layerFraxinus excelsior 24 19 19 17 . 3 75

E1 - herb layerImpatiens parviflora 58 55 60 35 29 18 100Geranium robertianum 52 41 37 59 19 18 100Stellaria media 37 19 21 10 10 6 100Alliaria petiolata 29 24 24 24 5 . 75

Balloto-RobinionE1 - herb layerBallota nigra 43 12 12 10 . 6 75Bromus sterilis 28 6 8 . 29 . 75

Chelidonio-RobinietaliaE1 - herb layerChelidonium majus 50 41 46 21 52 15 100Rubus fruticosus agg. 43 56 58 48 86 33 100Galeopsis tetrahit agg. 39 51 52 48 48 15 100Moehringia trinervia 33 18 20 10 48 3 100

RobinieteaE3 - tree layerRobinia pseudacacia 100 99 99 100 100 55 100

E2 - shrub layerRobinia pseudacacia 73 88 92 76 100 91 100

E1 - herb layerRobinia pseudacacia 42 51 56 35 24 39 100Veronica sublobata 24 10 10 10 5 9 100Fallopia dumetorum 21 36 39 21 33 18 100Lamium maculatum 9 9 9 7 19 3 100Fallopia convolvulus 4 17 14 31 . 18 75

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 213a

OthersE3 - tree layerFraxinus excelsior 12 10 11 7 . 3 75Pinus sylvestris 2 6 4 14 . 15 75

E2 - shrub layerQuercus petraea 4 12 13 10 14 6 100Crataegus laevigata 2 6 7 . 5 18 100Crataegus sp. 16 19 19 21 . 12 75Cornus sanguinea 13 3 4 . . 3 75Humulus lupulus 13 0.7 0.9 . 10 . 75Ribes rubrum 7 2 3 . 15 . 75Cotoneaster integerrimus 2 8 6 17 . 18 75Pyrus pyraster 2 4 3 7 . 12 75Quercus robur 2 2 3 . . 15 75Euonymus europaeus 14 14 15 10 . . 50Juniperus communis . 3 0.9 10 . 12 50

E1 - herb layerRosa canina agg. 25 45 49 28 38 36 100Crataegus sp. 20 34 36 24 14 15 100Quercus petraea 14 47 51 31 57 24 100Thlaspi perfoliatum 12 4 5 . 5 9 100Allium vineale 10 10 9 10 10 9 100Allium oleraceum 8 6 6 3 5 12 100Prunus avium 4 10 11 3 10 3 100Fragaria viridis 4 8 7 14 10 18 100Fragaria vesca 4 8 5 17 14 18 100Sorbus aucuparia 2 4 5 . 10 3 100Sambucus nigra 57 31 34 17 14 . 75Fraxinus excelsior 29 23 24 17 . 3 75Euonymus europaeus 19 14 15 10 10 . 75Acer campestre 16 10 10 7 . 3 75Brachypodium sylvaticum 17 6 7 . 19 . 75Humulus lupulus 13 2 2 3 14 . 75Lapsana communis 9 10 11 3 . 3 75Prunus spinosa 8 23 21 28 . 12 75Rubus idaeus 4 5 4 7 10 . 75Silene latifolia subsp. alba 3 . . . 14 3 75Ornithogalum kochii 2 4 5 . 10 . 75Achillea millefolium 2 3 0.9 14 . 12 75Dactylis polygama 2 1 0.9 3 10 . 75Cotoneaster integerrimus 0.8 10 7 24 . 12 75Galeopsis ladanum 0.8 3 3 . . 12 75Teucrium chamaedrys 0.8 0.7 . 3 . 12 75Hypericum perforatum . 12 9 24 14 33 75Festuca rupicola . 3 2 10 5 12 75Viola odorata 17 12 13 10 . . 50Lactuca serriola 7 10 11 3 . . 50Holcus lanatus 0.8 . . . 10 . 50Eryngium campestre 0.8 . . . . 12 50Anthericum liliago . 8 5 21 . 21 50Asplenium septentrionale . 8 5 17 . 21 50Carex humilis . 3 . 17 . 24 50Hieracium pilosella . 2 0.9 7 . 15 50Potentilla incana . 0.7 0.9 . . 24 50Echium vulgare . 0.7 0.9 . . 21 50Centaurea triumfettii . 0.7 . 3 . 12 50Phleum phleoides . 0.7 . 3 . 21 50Sedum album . 0.7 . 3 . 18 50Aegopodium podagraria 12 . . . . . 25Sedum rupestre . . . . . 24 25

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214 M. Vítková & J. Kolbek

were measured according to Kubíková (1970), ex-changeable cation content according to Moravec(1965), and available phosphorus according to Olsenet al. (1954) and Olsen & Sommers (1982). Determi-nation of net N-mineralisation and nitrification ratesfiin soils were analysed according to standard ISO14238, where net N-mineralisation was found from concentrations of NH4 + and NO3- loss during miner-alisation from organic soil matter. The nomenclatureof the soils follows Němeček et al. (2001).

The RDA method was used to detect correlationsbetween site characteristics, physical-chemical soil properties, bedrock type, and individual syntaxa. In CANOCO, environmental variables (37) were used as species data, scaling, and standardizing were per-formed due to usage of different units. Thirty-threerelevés from localities of soil samples divided intofour groups according to syntaxa and coded as nomi-nal variables were used as environmental variables. Such design was applied in accordance to a high num-ber of environmental charakteristics in comparison with number of studied localities and recorded phy-tosociological relevés. The Monte Carlo permutationtest with 499 permutations (unrestricted permuta-tions) was used to test the correlation significance.fi

Results

Classification of fi Robinia pseudacacia standsa

In the studied stands (Fig. 2 – 3) Black Locust is an expressive dominant of the tree or shrub layers, often forming clear monocultures. Among the original spe-cies, Quercus petraea, Pinus sylvestris and Fraxinusexcelsior reach higher constancy. The latter species competes well with Black Locust mainly on mesic sites in Bohemia. Cover of the tree layer is usually higher than 50%. In xerotherm Black Locust standson rocky slopes (ass. Melico transsi lvanicae-Robinietum) the tree layer may not be developed. The shrub layer is formed partly by high shrubs (up

to approx. 6 m), which could almost have a tree habit (e.g. Sambucus nigra, Crataegus monogyna, Euony-mus europaeus), partly juvenile trees (mostly Fraxi-nus excelsior and Robinia pseudacacia), further low shrubs (Ribes uva-crispa, R. rubrum agg., Rubusfruticosus agg., R. idaeus) with a height up to 1.5 m.Lianas – Humulus lupulus, Clematis vitalba or Par-thenocissus quinquefolia – are also frequent in the shrub layer. The herb layer can reach a height of upto 1.5 m due to the presence of robust broad-leavedhemicryptophytes such as Urtica dioica, Anthriscus sylvestris and Arctium minus. Perennial grasses (e.g.Poa nemoralis, P. trivialis, Arrhenatherum elatius, Avenella flexuosafl ), low perennial hemicryptophytes (e.g. Geum urbanum, Lamium maculatum, L. album, Viola odorata), geophytes (e.g. Adoxa moschatellina,Allium sp. div., Gagea sp. div., Ranunculus ficariafisubsp. bulbifera, Ornithogalum kochii, Polygonatumodoratum), annual and biennial species with a longer vegetation period (e.g. Impatiens parviflorafl , Galeop-sis tetrahit, Bromus sterilis, Alliaria petiolata), some of them behaving like lianas (e.g. Galium aparine, Fallopia convolvulus, F. dumetorum), and ephemeralterophytes (e.g. Valerianella locustaVV , V. dentata, Stel-laria media, Cerastium glomeratum, Thlaspi perfolia-tum, Holosteum umbellatum) are also present in the herb layer. The moss layer is of minor importance inmost of the Black Locust stands. It provides highercover in moister localities with the stands of ass.Chel idonio-Robinietum; on drier sites (e.g. inPoo nemoral is-Robinietum or Melico trans-si lvanicae-Robinietum) Hypnum cupressiformecan occur on boulders or on rock outcrops.

Black Locust stands of Bohemia are characterisedby two strong aspects in the fi rst half of the vegeta-fition period. In the early spring, before the leaves of Black Locust appear (March to April), the standsare characterised by Thlaspi perfoliatum, Lamium purpureum, Viola odorata, Holosteum umbellatum, Valerianella locusta, Veronica sublobata, Aurinia saxatilis, Carex humilis, Gagea sp. div., Pulsatilla pra-tensis etc. Late spring (May to June) is characterised

Potentilla tabernaemontana . . . . . 24 25Convolvulus arvensis . . . . . 21 25Geranium sanguineum . . . . . 18 25Pulsatilla pratensis . . . . . 18 25Sanguisorba minor . . . . . 18 25Acinos arvensis . . . . . 15 25Stipa pennata . . . . . 15 25Stipa pulcherrima . . . . . 15 25Allium senescens subsp. montanum . . . . . 12 25Arenaria serpyllifolia . . . . . 12 25Bupleurum falcatum . . . . . 12 25Galium glaucum . . . . . 12 25Sedum acre . . . . . 12 25Sedum sexangulare . . . . . 12 25Next accesoric species 187 223 190 94 38 132

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 215a

by Robinia pseudacacia, Anthriscus sylvestris, Torilis japonica, Chelidonium majus, Chaerophyllum temu-lum, Alliaria petiolata, Ballota nigra, Anthericumliliago, Polygonatum odoratum, Verbascum lychnitis,and Vincetoxicum hirundinaria. The second half of the vegetation period lacks aspect-forming species;annual species are absent from the herb layer. Some species (e.g. Poa nemoralis, Stellaria holostea, Polygo-natum odoratum, Brachypodium pinnatum, Carexhumilis, Melica transsilvanica) spread locally in manystands and have higher sociability than in native natu-ral stands.

Black Locust stands have been found in Bohemia on sites of reconstructed units of Aceri-Carpi-netum, Melampyro nemorosi-Carpinetum,Sorbo torminal is-Quercetum, in warm areasof Potenti l lo albae-Quercetum, as well as incommunities on nutrient poorer substrates (Ti l io-Betuletum, Luzulo albidae-Quercetum,Viscario-Quercetum, Hieracio pal l idi-Pine-tum, and only in the Elbe lowland of Festuco ovi-

nae-Quercetum roboris). They also cover steep rocky slopes with vegetation of the alliance Alysso-Festucion pal lentis . Black Locust stands usually have fewer species than the native natural communi-ties on equivalent sites.

Classifi cation of Black Locust stands in Bohemia: Robinietea Jurko ex Hadač & Sofron 1980 Chel idonio-Robinietal ia Jurko ex Hadač & Sofron 1980 Chel idonio-Robinion Hadač & Sofron 1980 Chel idonio-Robinietum Jurko 1963 Poo nemoral is-Robinietum Němec ex Vítková & Kolbek in Kolbek et al. 2003 cardaminopsietosum arenosae subass. nova typicum subass. nova Bal loto nigrae-Robinion Hadač & Sofron 1980

Fig. 2. Distribution of Black Locust communities in Bohemia: Chelidonio-Robinietum and Melico transsi lvanicae-Rob-inietum.

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216 M. Vítková & J. Kolbek

Arrhenathero elat ioris-Robinietum Šimonovič, Šomšák & Nikodemová ex Vítková & Kolbek ass. nova Euphorbio cypariss iae-Robinietal ia Vítková in Kolbek et al. 2003 Euphorbio cypariss iae-Robinion Vítková in Kolbek et al. 2003 Melico transsi lvanicae-Robinietum Kolbek & Vítková in Kolbek et al. 2003.

Black Locust stands of the class Robinietea inBohemia

Diagnostic species of the class Robinietea Jurko ex Hadač & Sofron 1980 in the territory of Bohemia in-clude Robinia pseudacacia in the tree layer, Robinia pseudacacia, Sambucus nigra and Ribes uva-crispa inthe shrub layer and Poa nemoralis, Galium aparine, Impatiens parviflorafl , Geum urbanum, Urtica dioica,

Chelidonium majus, Rubus fruticosus agg., Geraniumrobertianum, Anthriscus sylvestris, and Galeopsis te-trahit agg. in the herb layer (ordered according toconstancy). Some accessoric species of the classesQuerco-Fagetea and Rhamno-Prunetea areusual in the tree and shrub layers. From the allianceBerberidion Rosa canina agg., Prunus spinosa, Cra-taegus monogyna, Euonymus europaeus, and Corylusavellana occur most frequently. They form densestands in some localities. Species of the order Lamio albi-Chenopodietal ia boni-henrici frequently occur in the herb layer. High constancy is character-istic except above mentioned also for Chaerophyllumtemulum, Ballota nigra and Poa trivialis, which can even dominate locally, and also Taraxacum sect. Ru-deralia, Alliaria petiolata, and Moehringia trinervia.Another group is formed by species remaining fromformer communities from the class Querco-Fage-tea (e.g. Dryopteris filix-masfi , Viola collina, Brachy-podium sylvaticum, and Stellaria holostea) or on

Fig. 3. Distribution of Black Locust communities in Bohemia: Arrhenathero elat ioris-Robinietum and Poo nemoral is-Robinietum.

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 217a

rocky sites from the class Festuco-Brometea (e.g. Aurinia saxatilis, Brachypodium pinnatum, Euphor-bia cyparissias, Festuca pallens, Melica transsilvanica,and Vincetoxicum hirundinaria).

Black Locust stands occur in Bohemia on various ecotope types characterised by diverse site condi-tions: 1. Mesophilous stands with nitrophilous shrubs

and herbs on alkaline rocks and soils with very favourable physical-chemical properties and good organic biomass humifi cation (Fig. 4).

2. Moderately high mesophilous stands with onegrass species dominating in the herb layer (most often Poa nemoralis), which prefer shallow, drysoils rich in skeleton on acid bedrocks, locally with protruding rocks in river valleys or in sur-roundings of water reservoirs (Fig. 1, 5).

3. Species-poor stands on strongly acid quaternary deposits with unfavourable physical-chemical soil conditions (Fig. 6).

4. Usually shrubby stands, light, rich in species on extremely dry rocky slopes of different mineralstrength (Fig. 7).

In Bohemia, the class Robinietea is divided into two basic groups (Fig. 8), which occur on differentecotopes and differ from each other in physiognomyand species composition:a) Large forest mesophilous stands classified within fi

the order Chelidonio-Robinietal ia withtwo alliances (Chelidonio-Robinion: eco-tope types 1 and 2; and Bal loto nigrae-Rob-inion: ecotope type 3); and

b) Small shrubby xerophilous stands, described by the order Euphorbio cypariss iae-Rob-inietal ia with only one alliance (Euphorbio cypariss iae-Robinion: ecotope type 4). Be-tween those two groups, open Black Locust stands of subass. Poo nemoral is-Robinietum car-daminopsietosum arenosae (ecotope type2) with some features of both groups are placed.Due to the occurrence of species of the classFestuco-Brometea, they were assigned to the xerophilous group (B) in Fig. 8. However, these xerothermophytes are present only a little andoccur strictly on protruding rocks. The describedsubassociation was placed into the mesophilousbranch (A) and was classified within the alliancefiChelidonio-Robinion according to the welldistinguished tree layer and higher constancy of nitrophilous species growing in suitable micro-site conditions with organic matter deposition.

Chelidonio-Robinietal ia Jurko ex Hada� &Sofron 1980

Mesophilous secondary Black Locust forests with ni-trophytes (Table 2, Fig. 2, 3).Nomenclatural typus of the order: Chelidonio-Robinion alliance, lectotypus hoc loco (Art. 20).

Diagnostic species combination

E3 (tree layer): Robinia pseudacacia,E2 (shrub layer): Ribes uva-crispa, Robinia pseudaca-cia, Sambucus nigra,E1 (herb layer): Chelidonium majus, Galeopsis tetra-hit agg., Galium aparine, Geum urbanum, Moehrin-gia trinervia, Rubus fruticosus agg., Taraxacum sect.Ruderalia, Urtica dioica.

This order includes mesophilous Black Locust for-ests, which grow up to 30 m high on favourable sites.The shrub layer is mostly well developed and is rich in species. The herb layer is characterised by a spe-cies-rich group of mesophilous perennial or annual ruderal taxa, spreading widely in some localities. Thestands of the order occur almost all over Bohemia ataltitudes up to 500 m a.s.l.

In Bohemia the order includes two alliances withthree associations.

Chelidonio-Robinion Hada� & Sofron 1980

Mesophilous Black Locust stands with occurrence of nitrophilous species on soils with various nutrients, soil texture, and moisture regimes.Nomenclatural typus of the alliance: Chel idonio-Robinietum Jurko 1963, lectotypus hoc loco (Art. 20).

Diagnostic species combination

E3 (tree layer): Robinia pseudacacia,E2 (shrub layer): Robinia pseudacacia,E1(herb layer): Alliaria petiolata, Anthriscus sylvestris,Fraxinus excelsior, Geranium robertianum, Geum ur-banum, Impatiens parviflorafl , Stellaria media.

This unit occurs almost all over Bohemia (Fig. 2, 3).It is common both on slopes in river valleys and ina wide variety of habitats on fl at terrain. The most distributed association in Central Europe is Che-l idonio-Robinietum Jurko 1963. Except forprovisional associations (e.g. Rubo fruticosae-Robinietum Němec 1981, Urtico-Robinietum Němec 1981 and Sambuco nigrae-RobinietumŠčepka 1982), invalid ass. Anthrisco cerefol i i-Robinietum Majer 1963 (recte 1962), ass. Impa-tienti parvif lorae-Robinietum Sofron 1967 (seebelow), ass. Poo nemoral is-Robinietum Němecex Vítková & Kolbek in Kolbek et al. 2003 (see alsobelow), and ass. Sol idagini-Robinietum (Wen-delberger 1955) Jurko 1963 are also included in this alliance. The last mentioned association is described only from eutrophic wet soils in habitats of alluvial forests in Slovakia (Jurko 1963). Jurko (1963) speci-fi ed this community as young and unsteady, requir-fiing subsequent research. Currently, no such stands are known in Slovakia (Kanka & Jarolímek, per-sonal comm.) or in Bohemia.

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218 M. Vítková & J. Kolbek

Fig. 4. Mesophilous stand of the Chelidonio-Robinietum with dominating Anthriscus sylvestris (North Bohemia).

Fig. 5. Light Black Locust stand of the Poo nemoral is-Robinietum in spring dominated by Poa nemoralis (Central Bohemia).

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 219a

Fig. 6. Species-poor stand of the Arrhenathero elat ioris-Robinietum in late summer dominating by Calamagrostis epigejos onaeolian siliceous sand (North Bohemia).

Fig. 7. Spring aspect with Aurinia saxatilis in shrubby xerotherm Black Locust stand of the Melico transsi lvanicae-Robinietum(Central Bohemia).

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220 M. Vítková & J. Kolbek

Chelidonio-Robinietum Jurko 1963

Mesophilous Black Locust forests dominated by ni-trophytes on alkaline or intermediate bedrocks; on acid bedrock mostly on lower parts of slopes (Table 2, 3; Fig. 4).Nomenclatural typus: Jurko (1963, relevé 14 in Table 1), lectotypus hoc loco.Syn.: Impatienti parvif lorae-Robinietum So-fron 1967.

Diagnostic species combination

E3 (tree layer): Robinia pseudacacia, E2 (shrub layer): Ribes uva-crispa, Robinia pseudaca-cia, Sambucus nigra,E1 (herb layer): Anthriscus sylvestris, Arctium mi-nus agg., Chaerophyllum temulum, Galium aparine, Geum urbanum, Lamium album, Poa trivialis, Taraxacum sect. Ruderalia, Urtica dioica.

Structure and species composition

This unit represents the mesophilous Black Locustforests, which have been treated like monoculture sprout forest in the past, and grow up to 25 m high on favourable sites. The shrub layer is very rich in species; 51 species were found. The upper shrub layer is usually characterised by Sambucus nigra,and the lower shrub layer is often accompanied by

Ribes uva-crispa. The herb layer cover depends onthe shrub layer cover, and is usually higher than 60%. A species-rich group of mesophilous ruderal taxa, which grow up to 1.5 m high, is characterised by highconstancy in the community. The moss layer is notdeveloped or only slightly present.

The species-rich units Aceri-Carpinetum or Melampyro nemorosi-Carpinetum representthe potential natural vegetation of these Black Locust forests; in Northwest Bohemia Potenti l lo albae-Quercetum may also occur.

Variability

The association is highly variable. The species com-position of the community differs in accordance to moisture regime, nutrient content in bedrock, soilprofi le thickness, the former community, and thefiseed bank in the surroundings. Mesophilous nitro-phytes always provide high cover, but other taxa cansometimes play the dominant role. The horizontalstructure is mosaic-like, with a facial alternation of populations of Urtica dioica, Stellaria media, An-thriscus sylvestris, Galium aparine, Bromus sterilis, Poa trivialis, and so forth. In other stands, only one distinctive dominant of high cover occurs (e.g. Urticadioica, Ribes uva-crispa, Ballota nigra and Anthriscus sylvestris – Fig. 4); in the warmest areas Anthriscus trichosperma is also found. Spreading Chelidonium majus dominates on acid bedrock on moister sites of

Fig. 8. Numerical classification of 374 relevés of the classfi Robinietea from Bohemia. Ružička’s coeffi cient andfi �-fl exible clusteringflmethod were used (�=-0.25).

Explanations: A – mesophilous stands; B – xerophilous stands; 1. Chel idonio-Robinietum; 2. Mixed group with Poo nemora-l is-Robinietum and less nitrophilous stands of the Chelidonio-Robinietum; 3. Arrhenathero-Robinietum; 4. Xerophilous stands (mostly Melico transsi lvanicae-Robinietum and Poo nemoral is-Robinietum cardaminopsietosum arenariae).

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Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 221a

the lower parts of slopes, along streams, or on river terraces. These stands differ from the abovemen-tioned ones both in habitat and low cover of diagnos-tic species, and could be described as var. Chelido-nium majus (Fig. 9). In contrast with Sofron (1967),Black Locust stands dominated by invasive alien Im-patiens parviflorafl were also classifi ed into this unit.Impatiens parviflorafl spreads invasively mostly in places with bare soil (e.g. where there is game animalimpact, fallen trees, or forestry management) or with an open herb layer. At these localities other herbalspecies are highly suppressed; the soil chemistry is changed as well (cf. Hofmeister 2002). Accordingto the high constancy of diagnostic species of the association Chelidonio-Robinietum, the lone-standing Impatienti parvif lorae-Robinietum Sofron 1967 was included into the unit described asvar. Impatiens parviflorafl (Fig. 9).

The stands are well distinguished from the associa-tion Poo nemoral is-Robinietum by high con-stancy, and usually also by cover of mesophilous an-nual or biennial ruderal species (Fig. 10).

Synecology (Table 3)

The association occurs on a wide variety of sites inBohemia. It is found on all aspects, on fl at terrain andon slopes up to 60º (in dingles). The stands of thisunit prefer alkaline or intermediate bedrocks withsoils well saturated with alkalis and low values of to-tal carbon and nitrogen (Fig. 11). The highest nitrifi-fication rate and the lowest ammonifi cation of all units of Black Locust stands were found in these habitats.On alkaline or intermediate bedrocks, they can occuron all parts of slopes, but on acid bedrock they oc-cur strictly on the lower parts. A favourable moistureregime allows good humifi cation of organic matter deposited here from the upper slope parts, and thecreation of a deeper A-horizon. Many of the localitiesare enriched by nutrients from adjacent fields. Thefilist of detected soil types is mentioned in Table 3. OnArenosols, only stands of var. Chelidonium majuswere found, mostly at the base of sandstone rocks.They can also grow in the fl at terrain on strongly acidquaternary deposits with clayey silt in the soil profile,fiwhich is the cause of a more favourable soil-moistureregime and better accessability of nutrients.

Distribution in Central Europe

The stands of the association occur almost all overBohemia at altitudes up to 500 m a.s.l. (Fig. 2). Jurko(1963) mentioned the Chelidonio-Robinietum aswidely distributed in the area of Slovakia. Accordingto Ščepka (1982) this association is the most typical and frequent in the southern part of the Eastern Slo-vak Lowland. His provisional associations Sambuco nigrae-Robinietum and Urtico-Robinietum also correspond through their species compositionto the association Chelidonio-Robinietum. The association Chelidonio-Robinietum wasmentioned further in Slovakia by Jurko & Kontriš

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eschweizerbartxxx_author

222 M. Vítková & J. Kolbek

(1982) in the Little Carpathians, by Šimonovič et al. (2001, 2002) from Záhorie PLA, by Benčaťová & Benčať (2005) in Pohronská pahorkatina hills, byBenčaťová & Benčať (2008) from Slovak Gate to the Danube, and by Kontriš et al. (2009) in Slov-enské Stredohorie Mts. The semi-moist Black Locust stands in Hungary (Majer 1968) could be also includ-ed in this association. These are the most productive Black Locust stands with a characteristic spring as-pect built with geophytes, which are replaced by tallnitrophytes (Galium aparine, Chelidonium majus,Anthriscus trichosperma, Urtica dioica, Clematis vi-talba, Galeopsis sp. div., etc.) in May. Borhidi (2003, 2006) classifi ed them as two similar associations: fiChelidonio-Robinietum and invalid Anthrisco cerefol i i-Robinietum Majer 1963 (recte 1962), a differentiation which will be studied at present. In Poland, Chelidonio-Robinietum was describedby Gilicka (1989) from Poznań, by Świerkosz (1993) from Wrocław, by Święs & Łuczycka-Popiel(1999) from the reservation Szwajcaria Podlaska, byPawlaczyk (2008) from Drawa National Park, and by Pawlaczyk (personal comm.) from Odra river

valley in Cedynia Landscape Park. This association is also known from Austria (Burgenland, Vorarlberg, Niederösterreich), where it invades abandoned fields fiand pastures or clear-cuts after oak forests in the col-line vegetation belt (Willner & Grabherr 2007a).It occurs also in Germany: Kowarik (1986) andKowarik & Langer (1994) recorded this associationfrom West Berlin. Some of the relevés could be clas-sifi ed as this unit from the deep, fresh soils in Saartal fi(Klauck 1986) and in the surroundings of Stuttgart(Böcker & Dirk, personal comm.).

Poo nemoralis-Robinietum N�mec exVítková & Kolbek in Kolbek et al. 2003

Light open Black Locust stands dominated mostly by Poa nemoralis on slopes with acid bedrock. (Tables 2, 3; Fig. 5).

Diagnostic species combination

E3 (tree layer): Robinia pseudacacia,E2 (shrub layer): Prunus spinosa, Robinia pseudacacia,

Fig. 9. PCA-ordination diagram for 174 phytosociological relevés of the Chel idonio-Robinietum. � var. Impatiens parviflorafl , var. Chelidonium majus.

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 223a

E1 (herb layer): Avenella flexuosafl , Dryopteris filix-fimas, Galeopsis pubescens, Poa nemoralis (dom.), Vi-ola arvensis.

Structure and species composition

Mesophilous Black Locust stands of forest character,with physiognomy determined by grasses (mostly Poa nemoralis; Fig. 5). Nitrophytes represent a spe-cies-rich group, but their cover is not high.

The tree layer usually consists only of Black Lo-cust with height of 12 – 16 m. The shrub layer is richin species (56 taxa), but its cover mostly does not exceed 20%. Poa nemoralis is a significant dominantfispecies of the herb layer, with its optimum in natu-ral forest communities, but is well adapted to con-ditions in Black Locust stands. In some localitiesthis species is replaced by Avenella flexuosafl , Elymus

caninus, Brachypodium pinnatum or B. sylvaticum.Some components of natural communities are pre-served, namely of Sorbo torminal is-Quercetumor Luzulo albidae-Quercetum; Melampyro nemorosi-Carpinetum mostly in stands with thehigher number of mesophytes. Seedlings of woodyspecies (incl. Robinia pseudacacia) are common inplaces with bare soil. Cover of the herb layer is usu-ally higher than 50%. Hypnum cupressiforme is themost frequent species of the moss layer; its cover isonly exceptionally higher than 10%.

Described association is differentiated from thestands of the association Chelidonio-Robinietumby low cover of mesophilous synanthropic species on rocky soils with a shallow A-horizon (Fig. 10). Dif-ferences in fl oristic composition with other associa-tions are given in the Fig. 12 and 13.

Fig. 10. PCA-ordination diagram for 320 phytosociological relevés of the Poo nemoral is-Robinietum and Chel idonio-Robinietum.

eschweizerbartxxx_author

224 M. Vítková & J. Kolbek

Fig. 11. RDA-ordination diagram for 33 soil sampling localities of the Black Locust stands.

Variability

1. The above mentioned stands of this association are characterised by rather stabilized ecology andphysiognomy across Bohemia and are described assubassociation Poo nemoral is-Robinietum typicum subass. nova hoc loco (Table 2).Nomenclatural typus: identical with the typusof the association name in Kolbek et al. (2003).Differential species: distinguished negative-ly by the absence of self differential species.The subassociation is represented by 14 relevésin Table 6.14.3 (Vítková et al. 2003) from the Křivoklátsko PLA (Central Bohemia). Only one relevé (no. 10) belongs to the following subas-

sociation. The Poo nemoral is-Robinietum typicum differs from the second subassocia-tion mainly in the absence of protruding rockhabitats with thermophilous fl ora. It is alsomore frequently invaded by Impatiens parviflorafl(Fig. 14).

2. Stands growing in the localities with protrudingrocks are slightly different. Distinctive featuresare tortuous growth of woody species (approx.10 m tall), a denser shrub layer with dominatingspecies from the class Rhamno-Prunetea, and a small but diagnostically signifi cant number of xerothermophytes of the class Festuco-Brom-etea and thermophilous oak forests. However,

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 225a

Fig. 12. PCA-ordination diagram for 167 phytosociological relevés of the Poo nemoral is-Robinietum and Arrhenathero elat ioris-Robinietum.

diagnostic species of the class Robinietea (es-pecially Geranium robertianum and Geum ur-banum) have a more important role. They are favoured by suitable microsite conditions withorganic matter deposition. These open stands are described as the separate subassociation Poo nemoral is-Robinietum cardaminopsi-etosum arenosae subass. nova hoc loco (Ta-bles 2, 4A, 4B).

Nomenclatural typus: relevé 9, Table 4A). Differential species:

E3 (tree layer): Quercus robur,E2 (shrub layer): Crataegus monogyna,E1 (herb layer): Brachypodium pinnatum, Car-daminopsis arenosa, Festuca ovina, Hylotelephi-um maximum, Lychnis viscaria, Pimpinella saxi-fraga, Polygonatum odoratum, Stellaria holostea,Viola collina.

The stands are interesting from a nature conser-vation point of view. It consists of preserved rel-icts of natural forest phytocoenoses. These repre-sent available potential for restoration of originalstands after felling the Black Locust stand or ter-mination of its life.

Synecology (Table 3)

The stands of this association occur on the upper andmiddle parts of slopes in river valleys with fully de-veloped river phenomenon (cf. Ložek 1988; Fig. 1) and in the surroundings of water reservoirs (Fig. 3).Bedrock is usually represented by rocks poor in minerals, mostly proterozoic schists, greywackes orgranitoids, with silicites and quartzites in some lo-calities. Intermediate bedrocks (e.g. andesites, Ordo-vik schists, amphibolites) and alkaline rocks (spilites)

eschweizerbartxxx_author

226 M. Vítková & J. Kolbek

are sometimes present resulting in the occurrence of Black Locust stands that are richer in nitrophytes.The A-horizon is in most cases shallow, dry, with ahigh content of skeleton, and it forms soil pockets insome cases. The unit prefers extremely to strongly acidic soils, usually moderately saturated, where alu-minium and hydrogen ions are important (Fig. 11).

Distribution in Central Europe

The association commonly occurs in river valleys of Bohemia. On suitable sites it appears outside the rivernetwork (Fig. 3). According to similar species com-position, some relevés on the shallow skeleton soils in Saartal, Germany (see Klauck 1986), or in the sur-roundings of Stuttgart (Böcker & Dirk, personal comm.) could be classifi ed within this association.

Balloto nigrae-Robinion Hada� & Sofron1980

Black Locust stands on minerally poorer, sandy, drier soils, occurring mostly in lowlands.Nomenclatural typus of the alliance: Bal loto ni-grae-Robinietum Jurko 1963 (Hadač & Sofron 1980: 248, holotypus)Diagnostic species combination (compared withBorhidi 2003, Fekete 1965, Hadač & Sofron 1980, Jurko 1963, Oprea 2004, Pócs 1954, Šimonovi

5č et

al. 2001, 2002, and Vítková 2004)E1 (herb layer): Arrhenatherum elatius, Ballota nigra,Bromus sterilis, Calamagrostis epigejos, Cynoglossum officinalefi , Dactylis glomerata, Elymus repens, Holos-teum umbellatum, Lactuca serriola, Poa angustifolia, Veronica arvensis, V. triphyllos.

Fig.13. DCA-ordination diagram for 179 phytosociological relevés of theFig 13 DCA ordination diagram for 179 phytosociological relevés of the Poo nemoral is-Robinietum (Poo nemoral is Robinietum ( subass.subass typicum, typicumsubass. cardaminopsietosum arenosae) and Melico transsi lvanicae-Robinietum.

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 227a

This alliance occurs in Bohemia only sparsely andis represented by one marginal association (see be-low). The stands of the typical association Bal loto nigrae-Robinietum Jurko 1963 are frequent in the lowlands of Slovakia (Jurko 1963, Ščepka 1982,Benčaťová & Benčať 2005), Hungary (Fekete 1965, Majer 1968, Borhidi 1996, 2003, 2006), and Austria (Willner & Grabherr 2007a), but they were nei-ther found nor documented in Bohemia. Pócs (1954) described from Hungary association Robinietum pseudo-acaciae, and later published association Bromo steri l is-Robinietum Fekete 1965 (auct.div.) is synonym. This association was also localised by Felföldy (1947) in the Nyírség area, by Fekete (1965) in the Gödöllö area (Hungary), by Świerkosz

(1993) in Wrocław (Poland), and by Šimonovič et al. (2001, 2002) in Záhorie PLA (Slovakia). Ščepka(1982) described provisionally the following asso-ciations: Fumario off ic inal is-Robinietum ,Aristolochio clematit is-Robinietum, and Galio aparines-Robinietum, which Jarolímek & Šibík (2008)

p included together with the associa-

tion Bromo steri l is-Robinietum in the alliance Bal loto nigrae-Robinion. Next, relevés data-sets from different communities of this alliance in Slovakia were published by Benčaťová & Benčať(2005, 2008) . Theurillat et al. (1994) and Aeschi-mann et al. (2004) mentioned the described alliancefrom the Alps. Except for the semi-moist habitats (cf. Majer 1968a ), most of the Black Locust stands in

FFig. 14. PCA-ordination diagram for 146 phytosociological relevés of the Poo nemoral is-Robinietum typicum and cardaminopsietosum arenosae.

eschweizerbartxxx_author

228 M. Vítková & J. Kolbek

Number of relevé 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 C(%)E3 - tree layerRobinieteaRobinia pseudacacia 5 4 4 4 4 4 3 4 3 3 4 3 4 4 4 4 3 3 3 4 100Querco-FageteaQuercus robur r . . . . . . . 2 1 . . . 2 . . . 2 . . 25Carpinus betulus r . . . . . . . . . . . . . . . . 2 . . 10Fraxinus excelsior . + . . . . . . . . . . . . . . . . . 1 10E2 - shrub layerPoo nemoralis-Robinietum cardaminopsietosum arenosaeRosa canina agg. . + . . 1 . . . + + + . + . + + . . + 1 50Crataegus monogyna . . . . . . + . + . + + . . . . + r . . 30Prunus spinosa . . 3 . + . . . . . 1 . . . . . . . . 1 20Cotoneaster integerrimus . + . . 1 . . . . . . . . . . . . . . 1 15RobinieteaRobinia pseudacacia 1 r 1 2 2 1 1 3 . . 3 . 2 . 1 . + 3 1 2 75Sambucus nigra . + 1 . . . . . . . . . . . . . . . . + 15Rubus fruticosus agg. . . + 1 . . . . . . . . . . . . . . . . 10Ribes uva-crispa . . . . . . . . + . . . . . . . + . . . 10Querco-FageteaFraxinus excelsior . 2 . . 1 2 . . . . . . . . . . . . . 1 20Sorbus torminalis + . . . . + . . . . . . . . . . . . . . 10Rhamno-PruneteaCrataegus sp. 1 . . . r . . . . . . . . . . . . . . 1 15Corylus avellana . + . . . . . . . . + + . . . . . . . . 15Juniperus communis . . . + . . . . . 1 . . . . . . 1 . . . 15Euonymus europaeus . + 1 . . . . . . . . . . . . . . . . . 10E1 - herb layerPoo nemoralis-Robinietum cardaminopsietosum arenosaePoa nemoralis 3 3 4 4 3 3 3 2 2 1 2 4 1 2 2 2 2 2 2 3 100Geranium robertianum + 1 1 + 2 1 . + 2 2 2 1 r + 2 . 1 . . + 80Cardaminopsis arenosa . + + + 2 1 + r + r + + . 1 . . . r + 1 75Geum urbanum . . + . . r . r + . 1 + . + 1 1 r + 1 . 60Hylotelephium maximum . + . . 1 . 1 . r r . . r . . + . . + 1 45Festuca ovina . . . . . . . . 2 . + . 3 3 2 2 2 1 + . 45Brachypodium pinnatum + . 2 . . . 3 . + . 1 1 . + . . . . + . 40Stellaria holostea r . r . . . 3 1 . + . + . . . + . . + . 40Viola collina . . + . . . . . . + . + . 1 r . + . + . 35Pimpinella saxifraga . . . . . . r r + . . . + r . . 1 . . . 30Polygonatum odoratum + 1 . . + . . 1 . . . . . . . . . 1 . . 25Poo nemoralis-Robinietum Viola arvensis r . + + . . r . 1 . 1 . 1 . 1 . + . 1 . 50Avenella flexuosa . 2 . . 1 2 . . . . . . r . 1 4 . 1 1 . 40Dryopteris filix-mas . 1 1 r 1 2 . . . . . . . . . . . . . . 25Galeopsis pubescens . . . + . . . . . + . . . . . . . . . + 15Chelidonio-Robinion, Chelidonio-Robinietalia Galeopsis tetrahit agg. 1 1 + . 1 + . + + . . . . r + . . + r . 55Galium aparine 2 + 1 . + + . 2 + . 2 . . . . . . . 1 . 45Rubus fruticosus agg. r 1 . . + 1 . . . . . . . . . . 1 + . 1 35Impatiens parviflora 1 1 . . 1 1 . . . . . . . . . . . . . 2 25Alliaria petiolata . . r r . . . . . . . . . . 1 . . 1 . r 25Chelidonium majus + . + . . . . . . . . . + . . . 1 . . . 20Moehringia trinervia r . . . 1 1 . . . . . . . . . . . . . . 15

Table 4A. Poo nemoral is-Robinietum cardaminopsietosum arenosae subass. nova (selected 20 relevés)

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 229a

Taraxacum sect. Ruderalia . . r r . . . . . . . . . . . . . . . 1 15Stellaria media . . + 2 . . . . . . . . . . . . . . . . 10Urtica dioica . . + + . . . . . . . . . . . . . . . . 10RobinieteaFallopia convolvulus . . . . 1 . 1 1 . + . . . . + . . . . 1 30Robinia pseudacacia . r + 1 1 . . . . . . . . . . . . . . 1 25Querco-FageteaCampanula persicifolia . . . . . . r . + . . r . + + . + . . . 30Fraxinus excelsior . 1 . . + + . . . . . . . . . . . . . + 20Quercus petraea . r . r 1 r . . . . . . . . . . . . . . 20Acer pseudoplatanus r . . r r . . . . . . . . . . . . . . . 15Tanacetum corymbosum . . r . + . . r . . . . . . . . . . . . 15Lychnis viscaria + + . . . . . . . . . . . . . . . . . . 10Carex muricata . . . 2 . . . . . . . . . . 2 . . . . . 10Melica nutans . . . + . . . . . . + . . . . . . . . . 10Festuco-BrometeaEuphorbia cyparissias . . + + . r 2 + + 1 . + 2 + 1 + + 1 . + 75Vincetoxicum hirundinaria . . . r . r + + . . . . . . . . . + . . 25Carex humilis . . . . . . . . 1 2 . + . . . . . . 2 . 20Melica transsilvanica . . + 1 . . . . . 3 . . . . . . . . . . 15Asplenium septentrionale . . + . r . . . . . . . . . . . . . . 1 15Fragaria viridis . . 1 1 . . . . . . . . . . . . . . . . 10Ajuga genevensis . . r 1 . . . . . . . . . . . . . . . . 10Thymus pulegioides . . + . . . . . . . . . . . . + . . . . 10Festuca rupicola . . . . . . . . . 2 . . . . . 3 . . . . 10Rhamno-PruneteaCrataegus sp. r . + . . r . . . . . . . . . . . . . + 20Rosa canina agg. . + . + . + . . . . . . . . . . . . . + 20Prunus spinosa . . 2 r . . . . . . . . . . . . . . . 1 15Cotoneaster integerrimus . . . . 1 + . . . . . . . . . . . . . + 15Euonymus europaeus . . + . r . . . . . . . . . . . . . . . 10Allium vineale . . r + . . . . . . . . . . . . . . . . 10Galio-UrticeteaFallopia dumetorum 1 + 1 1 . + . . . . . . . . . . . . . . 25Veronica sublobata . . 1 2 . . . . . . . . . . . . . . . . 10Lamium maculatum . . . . . . + . . . . . . . 1 . . . . . 10Other speciesE3 - tree layerPinus sylvestris . . . . . . . . . + . . . + . . . + . . 15E2 - shrub layerSambucus racemosa . r . . . . . . . 1 . . . . . . . . . . 10E1 - herb layerHypericum perforatum . . . 1 r . . r + . . . . . . + . . . + 30Asplenium trichomanes . . + . 1 + . . . . . . . . . . . . . + 20Calamagrostis epigejos . . . 1 . . . 1 . . . + + . . . . . . . 20Fragaria vesca . . . . + . . . . . . . . . 1 3 1 . . . 20Verbascum densiflorum . . . . . . . + + . . . . . . . . r 2 . 20Polypodium vulgare . 2 . . + 1 . . . . . . . . . . . . . . 15Arrhenatherum elatius . 1 . . 2 1 . . . . . . . . . . . . . . 15Viola canina . . + + . 1 . . . . . . . . . . . . . . 15Achillea millefolium . . . . . . . r . r . . 1 . . . . . . . 15Agrostis gigantea . . . . . . . . . . + . . . 1 . 2 . . . 15Sisymbrium loeselii . 1 . . . 1 . . . . . . . . . . . . . . 10

eschweizerbartxxx_author

230 M. Vítková & J. Kolbek

Hieracium pilosella . r . . . . . . . . . . . . . . . . 1 . 10Cerastium glomeratum . . 1 + . . . . . . . . . . . . . . . . 10Myosotis ramosissima . . + 1 . . . . . . . . . . . . . . . . 10Valerianella locusta . . + + . . . . . . . . . . . . . . . . 10Agrostis stolonifera . . + + . . . . . . . . . . . . . . . . 10Verbascum sp. . . r 1 . . . . . . . . . . . . . . . . 10Chenopodium album agg. . . . . . . . r . . . . + . . . . . . . 10Bromus tectorum . . . . . . . . 1 . . . . . . 2 . . . . 10

Fig. 15. DCA ordination diagram for 99 relevés of the allianceFig. 15. DCA-ordination diagram for 99 relevés of the alliance Balloto nigrae RobinionBalloto nigrae-Robinion according toaccording to BenBenččaaťťováová && BenBenččaaťť(2005, 2008), Fekete (1965), Svobodová (1952), Oprea (2004), Pócs (1954), Swierkosz (1993), Ščepka (1982), Šimonovič et al.(2001, 2002), and Vítková (2004): � Robinietum pseudo-acaciae, Bal loto nigrae-Robinietum, Arrhenathero elat ioris-Robinietum.

In one relevé only:E3: Betula pendula 18:2, Pinus nigra 17:2, Pyrus pyraster 1:r, Quercus petraea 19:2, Sorbus aria 18:+,E2: Prunus padus 5:2, Pyrus pyraster 4:2, Quercus petraea 19:+, Ribes alpinum 2:+, Ulmus minor 20:2,E1: Adoxa moschatellina 3:+, Agrostis capillaris 9:3, An-thericum liliago 20:+, Arabidopsis thaliana 3:1, Artemisia absinthium 13:r, Athyrium � lix-femina 2:r, Ballota nigra 20:r, Barbarea vulgaris 20:+, Bromus hordeaceus 13:3, Campanula rotundifolia 5:+, C. trachelium 11:r, Carduus nutans 17:r, Carpinus betulus 1:+, Centaurea triumfettii 16:+, Chaerophyllum temulum 10:1, Clinopodium vulgare 6:+, Conyza canadensis 13:r, Corylus avellana 6:r, Dactylis polygama 3:+, Dianthus carthusianorum 16:+, Festuca pallens 20:+, Fragaria moschata 14:2, Galium mollugo 6:r, Helichrysum arenarium 16:+, Hieracium murorum 2:r, Humulus lupulus 7:r, Impatiens glandulifera 20:+, Lapsana communis 20:+, Mahonia aquifolium 20:r, Oxalis stricta 17:+, Parthenocissus quinquefolia 20:r, Phleum phleoides 16:1, Potentilla argentea 3:+, Prunus padus 5:1, Pyrus pyraster 5:r, Ranunculus � caria subsp. bulbifera 3:+, Ribes uva-crispa 2:r, Rumex acetosa 5:+, Sambucus nigra 3:1, Securigera varia 20:+, Senecio viscosus 8:+, Silene nutans 6:+, Tanacetum parthenium 20: 1, Torilis japonica 1:r, Ver-bascum lychnitis 14:r, Veronica serpyllifolia 13:r, Vicia hirsuta 4:+, Viola odorata 6:1, V. reichenbachiana 5:1.

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 231a

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100

300

SW40

605

80.

amph

ibol

iteZd

ucho

vice

(Cen

tral B

ohem

ia)

10V

V23

0860

100

360

W40

5015

70.

rhyo

lite

and

daci

teD

ubov

ý vr

ch h

ill (C

entra

l Boh

emia

)11

VV

2308

6010

035

0S

3070

4060

.ba

salt

Hor

ní L

uh (C

entra

l Boh

emia

)12

VV

2408

6010

035

0S

3040

590

.ba

salt

Hor

ní L

uh (C

entra

l Boh

emia

)13

VV

0609

6010

025

0S

3060

1070

.gr

eyw

acke

Bru

nšov

(Cen

tral B

ohem

ia)

14V

V02

0760

100

230

NN

W45

600

50.

prot

eroz

oic

schi

stD

avle

(Cen

tral B

ohem

ia)

15V

V07

0960

100

340

W20

605

80.

prot

eroz

oic

schi

stC

hlum

ík h

ill n

ear S

koch

ovic

e (C

entra

l Boh

emia

)16

VV

2807

5910

032

0W

2560

510

0.

rhyo

lite

and

daci

teŽu

pano

vice

(Cen

tral B

ohem

ia)

17V

V21

0860

100

330

SW40

7015

70.

amph

ibol

iteZd

ucho

vice

(Cen

tral B

ohem

ia)

18V

V06

0960

100

250

WSW

3070

3050

.si

licite

Dav

le (C

entra

l Boh

emia

)19

VV

0207

6010

025

0S

2050

1550

.pr

oter

ozoi

c sc

hist

Sáza

va (C

entra

l Boh

emia

)20

MV

0107

9925

025

0SS

W45

6525

5520

prot

eroz

oic

schi

stLi

bic

e (C

entra

l Boh

emia

)

Cov

er [%

]

Tabl

e 4B

. Rel

evé

data

(Tab

. 4A

). E

xpla

nati

ons:

dot

s in

E0 =

cov

er is

not

det

erm

inat

ed.

eschweizerbartxxx_author

232 M. Vítková & J. Kolbek

Number of relevé 1 2 3 4 5 6 7 8 9 1 1 1 1 1 1 1 C 1 1 1 2 2 C0 1 2 3 4 5 6 (%) 7 8 9 0 1 (%)

E3 - tree layerRobinieteaRobinia pseudacacia 2 3 4 3 3 3 3 3 3 4 3 4 3 3 4 4 ## 3 3 3 3 3 ##E2 - shrub layerRobinietea, Chelidonio-RobinietaliaRobinia pseudacacia 1 3 1 + 1 2 2 2 2 2 2 2 2 1 1 2 ## 2 2 1 1 1 ##Sambucus nigra 3 . 2 2 2 4 . . 2 . . . . . 2 1 50 . . . . . 0Ribes uva-crispa . . . . 1 + . . . . . . . . 1 . 19 . . . . . 0E1 - herb layerArrhenathero elatioris-RobinietumArrhenatherum elatius 1 2 + + + + 1 + + 3 2 3 3 1 2 1 ## 1 . . . . 20Elymus repens + 2 + . + + . + . + . 2 1 . . . 50 . . . . . 0Dactylis glomerata + + . + + r . . 1 . . . . . 1 . 44 . . . . . 0Poa angustifolia . 3 . + + . . . . + 1 1 . + . . 44 . . . + . 20Calamagrostis epigejos . . . . . + 1 + . 3 4 + . . . 1 44 5 4 5 5 5 100Festuca rubra agg. + . . + 2 1 . 2 . . . . . . . + 38 + . . . . 20Agrostis capillaris . . + 2 + . + . . 2 1 . . . . . 38 1 . . + . 40Anthoxanthum odoratum . . . 1 . + 3 + . . . . . . . 2 31 1 . . . . 20Dryopteris dilatata . . . . . . . r . + . r . r . 1 31 . . . r + 40Balloto nigrae-Robinion, Chelidonio-Robinietalia, Robinietea Rubus fruticosus agg. 1 1 1 + 1 + 3 2 + + . + . . 1 3 81 + 1 + . + 80Galium aparine 2 1 1 . + 1 + 1 1 . + . + . 2 . 69 . . . . . 0Chelidonium majus 2 + 3 4 . + 1 . 3 . r . 1 2 + . 69 . . . . . 0Urtica dioica r r + . . r r r 1 1 1 . 1 . + . 69 . . . . . 0Poa trivialis 2 . 1 1 . . 1 + 1 + . . 2 . 2 . 56 . . . . . 0Poa nemoralis 1 . 2 . . 1 . 3 3 + . . 3 3 . 1 56 + . . . . 20Moehringia trinervia 1 . + 1 . . . 1 + 1 . 1 + + . . 56 . . . . r 20Galeopsis tetrahit agg. . . . . . . . + + 1 . + . r + 1 44 r + . r . 60Geum urbanum + . . + r . . 1 . . . . + . 1 . 38 . . . . . 0Bromus sterilis 1 . . . 2 r . . 1 . . 1 . . 2 . 38 . . . . . 0Impatiens parviflora 1 . . . . 1 . + 1 1 + . . . . . 38 . . . . . 0Ribes uva-crispa + . r + . . . . + r . . . . . . 31 . . . . . 0Taraxacum sect. Ruderalia . . . + r . . r . . . . . . r . 25 . . . . . 0Geranium robertianum . . . r . r . r + . . . . . . . 25 . . . . . 0Robinia pseudacacia . . . . r + . . . . . . . . + 1 25 . . . r . 20Chaerophyllum temulum . . . r r . . . . . . . . . 1 . 19 . . . . . 0Sambucus nigra . . . . . 2 . r 1 . . . . . . . 19 . . . . . 0Stellaria media . . . + . . . . . . . . . . 1 . 13 . . . . . 0Querco-Fagetea, Festuco ovinae-Quercetum roborisQuercus petraea r . . r . . . r + r . 1 r . . 1 50 . 1 1 + r 80Brachypodium sylvaticum . . . . . . . . 1 . . 2 2 . + . 25 . . . . . 0Hypericum perforatum . . . + + . . r . . . . . . . . 19 . . . . . 0Prunus avium . . . . r . . . . . . . . . r . 13 . . . . . 0Dactylis polygama . . . . . . . + . . . . 1 . . . 13 . . . . . 0Festuca ovina . . . . . . . . . . . 2 . 2 . . 13 . . . . . 0Rhamno-PruneteaRosa canina agg. r . . + + . r r r . . r . . r . 50 . . . . . 0Crataegus sp. + . . . + . . r . . . . . . . . 19 . . . . . 0Allium vineale . . . . 1 . . . . . . . . . 1 . 13 . . . . . 0Euonymus europaeus . . . . . . . . r . . . . . + . 13 . . . . . 0Galio-UrticeteaFallopia dumetorum . . . . r . . + . r 1 + 1 . 1 . 44 . . . . . 0Lamium maculatum . r . r 1 . . . . . . . + . . . 25 . . . . . 0Silene latifolia subsp. alba r r . . + . . . . . . . . . . . 19 . . . . . 0Humulus lupulus . . . . . . . . . . . . . . . + 6 r . . . 1 40

100 100

100 100

100

Table 5A. Arrhenathero elat ioris-Robinietum (rels 1 – 16), var. Calamagrostis epigejos (rels 17 – 21)

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 233a

Other speciesE2 - shrub layerQuercus petraea . . . . . . r r . . . . . . . 1 19 . . . . . 0Crataegus monogyna . . . . 2 . . r . . . . . . . . 13 . . . . . 0Ribes rubrum agg. . . . . . . . . . . . r . . + . 13 . . . . . 0Humulus lupulus . . . . . . . . . . . . . . + 1 13 . . . . . 0E1 - herb layerFragaria vesca . . . . . . . + . . . 1 . . + . 19 . . . . . 0Fragaria viridis . . . + 1 . . . . . . . . . . . 13 . . . . . 0Ornithogalum kochii . . . . + . . . . . . . . . + . 13 . . . . . 0Holcus lanatus . . . . . . . . . . 2 . . . 2 . 13 . . . . . 0Sorbus aucuparia . . . . . + . . . . . . . . . . 6 . r . . . 20Rubus idaeus . . . . . . . . . . . . . . . 2 6 . . . . 1 20

Nr. of Date Altitude Aspect Slope Geology Localityrelevé [m a.s.l.] [º] E3 E2 E1 E0

1 020697 230 - 0 20 35 55 5 fluvial sand and gravel Led ice (Central Bohemia)2 250697 230 - 0 50 25 80 1 fluvial sand and gravel Led ice (Central Bohemia)3 250697 230 S 5 55 15 60 5 fluvial sand and gravel Led ice (Central Bohemia)4 250697 210 W 15 30 40 70 1 marly sand Klene (Northern Bohemia)5 250697 210 S 15 40 25 30 1 marly sand Klene (Northern Bohemia)6 250697 210 - 0 50 80 10 1 fluvial sand and gravel Straškov (Northern Bohemia)7 250697 210 S 5 50 20 60 1 fluvial sand and gravel Straškov (Northern Bohemia)8 260697 210 - 0 50 15 60 40 fluvial sand and gravel P estavlky (Northern Bohemia)9 100797 180 ESE 10 50 30 75 1 fluvial sand and gravel Dolní Be kovice (Central Bohemia)

10 100797 175 - 0 60 10 85 5 fluvial sand and gravel Podvl í (Central Bohemia)11 100797 190 - 0 45 15 95 0 aeolian sands Bechlín (Northern Bohemia)12 100797 170 - 0 60 15 70 0 aeolian sands P edonín (Northern Bohemia)13 100797 155 - 0 50 10 90 0 fluvial sedimentary rock P edonín (Northern Bohemia)14 100797 150 - 0 45 5 70 0 aeolian sands Kozlovice (Northern Bohemia)15 030698 230 S 5 70 15 65 0 marlstone Brodce (Central Bohemia)16 040699 220 S 5 70 10 60 5 sandstone Jenichov (Central Bohemia)17 250697 210 - 0 40 20 90 0 fluvial sand and gravel Straškov (Northern Bohemia)18 100797 160 - 0 45 10 65 0 fluvial sand and gravel P edonín (Northern Bohemia)19 100797 165 - 0 30 5 80 0 fluvial sand and gravel P edonín (Northern Bohemia)20 100797 150 - 0 45 5 90 0 aeolian sands Kozlovice (Northern Bohemia)21 100797 170 - 0 40 5 90 0 aeolian sands Libotenice (Northern Bohemia)

Author of relevés: M. Vítková; Sampled area: 250 m2

Cover [%]

Table 5B. Relevé data (Table 5A)

In one relevé only:E3: Quercus petraea 14:+, E2: Crataegus laevigata 4:2, Frangula alnus 16:+, Ligustrum vulgare 8:+, Prunus avium 7:r, P. cerasusr4:r, Rosa canina agg.r 5:+, Rubus fruticosus agg. 11:+, Syringa vulgaris 8:r, r E1: Alliaria petiolata 13:+, Allium oleraceum 2:+, A. scorodoprasum 8:1, Alopecurus pratensis 2:+, Anthriscus sylvestris 13:r, Avenella r �exuosa�� 12:+, Campanula patula 15:+, Carex leporina 10:r, C. spicata r 11:1, Chenopodium album agg. 5:r, Cornus sanguinear 13:r, Corylus avellana r 1:r, Descurainia sophiar 15:r, rFalcaria vulgaris 5:1, Festuca pratensis 7:+, F. rupicola 2:+, Galium verum agg. 2:r, Glechoma hederacear 13:r, Hylotelephium rmaximum 4:+, Luzula multi�i ora�� 8:+, Pinus sylvestris 12:+, Polygonatum odoratum 4:+, Potentilla heptaphylla 4:r, P. reptans r 15:1, Pyrus pyraster 4:r, Ranunculusr � caria subsp. bulbifera� 1:+, R. polyanthemos 4:r, Rhamnus catharticar 5:r, Ribes rubrum agg. r 4:r, rRumex acetosa 3:r, R. obtusifoliusr 3:r, Scrophularia nodosar 14:r, Stellaria holostear 16:+, Thlaspi perfoliatum 15:1, Triticum aesti-TTvum 6:r, Ulmus laevisr 13:r, Veronica chamaedrys agg.r 8:+, V. of�ff cinalis� 8:r, V. sublobatar 1:1, Verbascum phoeniceum 15:+, Viola arvensis 6:r, V. hirtar 4:r, V. riviniana r 6:+.

eschweizerbartxxx_author

234 M. Vítková & J. Kolbek

Hungary could be classifi ed as the alliance Balloto nigrae-Robinion. New order Bal loto nigrae-Robinietal ia Balogh & Borhidi 2006 proposed by Borhidi (2006) is not effective published (see above).

The ordination diagram of all available Central European phytosociological relevés from associa-tions mentioned in the previous paragraph shown in Fig. 15 evidences subjective and often incorrect inclu-sion of separate relevés in the associations. The maxi-mum difference was determined by relevés placed into the association Robinietum pseudo-acaciae Pócs 1954 (see Discussion).

Arrhenathero elatioris-Robinietum Šimonovi�, Šomšák & Nikodemová ex Vítková& Kolbek ass. nova hoc loco

Black Locust stands, poorer in species, dominated bygrasses, on strongly acid quaternary deposits (Table2, 3, 5A and 5B).Nomenclatural typus: relevé 10, Table 5ADiagnostic species combination

E3 (tree layer): Robinia pseudacacia, E2 (shrub layer): Robinia pseudacacia,E1 (herb layer): Agrostis capillaris, Anthoxanthum od-oratum, Arrhenatherum elatius, Calamagrostis epige-jos, Dactylis glomerata, Dryopteris dilatata, Elymus repens, Festuca rubra agg., Poa angustifolia.

Structure and species composition

This community represents productive Black Locustmonocultures, with the tree layer rising up to the height of 30 m. The shrub layer is composed mostly of juvenile forms of Black Locust (cover usually be-low 25%). The herb layer is often dense (cover 60 to95%) and its physiognomy is determined by peren-nial grasses. The moss layer is usually missing.

The potential natural vegetation on these sites is represented by pine-oak forests of the associationFestuco ovinae-Quercetum roboris , and onnutrient richer substrates by linden-oak forests of theTil io-Betu letum association.

Variability

The stands occurring on linden-oak forest sites arericher in species than in pine-oak forest sites. They are characterised by a higher ratio of nitrophilous taxaand higher cover of the shrub layer; Sambucus nigra dominated at some places. The stands on pine-oakforest sites are poor in species. Calamagrostis epige-jos, with cover between 70 and 95%, spreads at some localities of both types, causing extreme depletion of the herb layer, while mesophilous taxa completely disappear (Fig. 6). These stands are documented inTable 5A, relevés 17 – 21, as var. Calamagrostis epige-jos. The moss layer is completely missing.

Species-rich stands of this community on siteswith Til io-Betuletum differ from the stands of

Poo nemoral is-Robinietum in ecological con-ditions (landform, bedrock, soil type etc.) as well asspecies composition (see Fig. 12). The stands of as-sociation Robinietum pseudo-acaciae are richerin spring geophytes, terophytes, and thermophilousspecies (Fig. 15).

Synecology (Table 3)

The community occurs at low altitudes, where BlackLocust is not harmed by frostbite and has a straighttrunk and a shapely crown. The stands usually occur on fl at terrain on strongly acid quaternary deposits,on aeolian siliceous sands, or on sandy gravel ter-races (see Fig. 11), and they are mostly moderatelysaturated (Table 3). Some of the lowest values of the nitrifi cation rate were detected in soils of these Black fiLocust stands. In the case of occurrence of clayey siltin the soil profile, soil characteristics become morefifavourable and the number of nitrophytes increases.

Distribution in Central Europe

The stands of the association occur in Bohemiamostly in the Elbe lowland (Fig. 3). Šimonovič etal. (2001, 2002) provisionally classifiedfi the younger open Black Locust stands in the aeolian minerallypoor siliceous sands (pH(H2O) 3.99) in the ZáhoriePLA (Slovakia) as the association Arrhenatheroelat ius-Robinietum. Passarge (1967) describedstands with identical physiognomy on nutrient poorsand habitats in Germany (Scheeren) with dominat-ing grasses Calamagrostis epigejos, Festuca ovina,Agrostis capillaris and Poa pratensis. According to eight unpublished relevés, Pawlaczyk (personal comm.) detected a grass community with dominating Calamagrostis epigejos without nitrophilous plantson sandy soils in the Odra river valley in Cedynia Landscape Park (NW Poland). Similar stands withCalamagrostis epigejos were also described by Majer(1968) in Hungary, where they grow over the clear-ings after clear-cuts on dry sites.

Euphorbio cyparissiae-Robinietal ia Vítková in Kolbek et al. 2003

Low, tortuous and xerophilous Black Locust stands of thermophilous habitats (Table 2, Fig. 7).Nomenclatural typus of the order: Euphorbio cy-pariss iae-Robinion Vítková in Kolbek et al. 2003

Diagnostic species combination

E3 (tree layer): Robinia pseudacacia (may not be de-veloped),E2 (shrub layer): Robinia pseudacacia, Rosa canina agg.,E1 (herb layer): Artemisia campestris, Asperula cyn-anchica, Aurinia saxatilis, Centaurea stoebe, Dian-

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 235a

thus carthusianorum, Euphorbia cyparissias, Festuca pallens, Melica transsilvanica, Poa nemoralis, Seseli osseum, Thymus pulegioides, Verbascum lychnitis,Vincetoxicum hirundinaria.

The physiognomy of xerophilous Black Locust stands is characterised by species of natural phytocoenoses of similar sites, especially xerotherm herb vegetationof the class Festuco-Brometea, thermophilous shrubs of the class Rhamno-Prunetea and ther-mophilous oak forests of the alliance Quercion pubescenti-petraeae.

The mesophilous stands of the order Chelido-nio-Robinietal ia differ from the order Euphor-bio cypariss iae-Robinietal ia in the high con-stancy (and often also cover) of diagnostic speciesof the class or order, and also of species of the class Galio-Urticetea. These taxa are mostly mesophi-lous annual or biennial ruderal species and include the invasive alien Impatiens parviflorafl .

The stands of order Euphorbio cypariss iae-Robinietal ia were found mainly in the valleys of rivers. On suitable habitats with rocky terrain theyalso appear outside the river network (Fig. 2).

These xerophilous Black Locust stands were not yet mentioned in Central European literature. How-ever, it may be possible to classify some stands de-scribed by Oberdorfer (1992) from South Germany and from the surroundings of Stuttgart (Böcker &Dirk, personal comm.). According to the floristic flcomposition in the synoptic table, xerophilous BlackLocust stands probably also occur in the territory of Austria (Willner & Grabherr 2007b).

The order includes only one alliance Euphorbio cypariss iae-Robinion Vítková in Kolbek et al. 2003 and association Melico transsi lvanicae-Robinietum with the same diagnostic species as theorder.

Melico transsilvanicae-Robinietum Kolbek & Vítková in Kolbek et al. 2003

Low to shrubby xerotherm Black Locust stands on thermophilous rocky slopes (Tables 2, 3; Fig. 7).

Diagnostic species combination

Identical with diagnostic species combination of theorder.

Structure and species composition

Black Locust is the dominant woody species in theshrub layer, and usually occurs here at the border of its physiological abilities. The crowns are un-shaped, the trunks are deformed and the trees (butmostly shrubs) are of dwarf stature (3 – 5 m), rarely higher. Xerophilous woody species (e.g. Prunus spi-nosa, Crataegus laevigata, Cotoneaster integerrimusand more thermophilous species of the Rosa genus) compete well with Black Locust in some localities.

The protected shrub Juniperus communis also occurs it some places.

At most of the localities the herb layer is ratherdense (cover more than 60%) and very rich in spe-cies. Elements of the class Festuco-Brometeadominate here, accompanied by species of light andxerophilous forests.

VariabilityVV

This unit includes various Black Locust stands on sites of thermophilous shrubby vegetation or of xe-rotherm herb vegetation. At the core of the associa-tion are mature stands with long-term stabilisationby vegetative renewal of Black Locust. DistinctiveBlack Locust stands grow on extreme substrates – proterozoic chert or olivine nephelinite. Some standsfunction as important refuges for rarer species (e.g.Allium strictum, Anthericum liliago, Stipa joannis,Carex humilis, C. caryophyllea, Hieracium bauhini) which occur with less vitality or not at all in contact stands. Differences in comparison with associationPoo nemoral is-Robinietum are given in theFig. 13.

Synecology (Table 3)

Stands prefer slopes with partially bare rocks in riv-er canyons. Parent material with different mineralstrength and method of formation comes close to the surface (amphibole-biotitic granodiorite, andesite,diabase, spilite, olivine nephelinite, proterozoic chert,proterozoic schist). The potential ability of nitrifica-fition rises to a high value, while the distribution of nitrophilous taxa in those xerophilous stands is prob-ably suppressed by low soil moisture. Carbonates have been detected only in spilite and olivine neph-elinite. The concentration of available phosphorus islow at all localities.

The correlation of Melico transsi lvanicae-Robinietum with environmental characteristics in Fig. 11 is only a result of “fl attening” of the multi-dimensional space into a 2D picture. The centroid of the association reaches a high value (0.9444) on the3rd ordination axis, while site parameters are foundon the fi rst two axes. Only two localities lying on op-posite gradient poles were used for construction of Fig. 11.

Distribution in Central Europe

Distribution of described association is identical todistribution of the order Euphorbio cypariss iae-Robinietal ia (see above).

eschweizerbartxxx_author

236 M. Vítková & J. Kolbek

Discussion

Position of the class Robinietea in hierarchysystem

Some authors (Horvat et al. 1974, Ellenberg 1996, Matuszkiewicz W. 2001, Matuszkiewicz J. M. 2001, Rivas-Martínez et al. 2001) do not clas-sify Black Locust stands into the classifi cation sys-fitem of plant communities at all. In vegetation over-views from Central Europe (Pott 1992, Mucina et al. 1993, Willner & Grabherr 2007a, Jarolímek & Šibík 2008

993,), the classifi cation as well as rank in

the hierarchical classifi cation system is done only fiprovisionally, based on a restricted relevés data-set.Authors from various countries reaching similar con-clusions, usually only at the association level, classi-fied Black Locust stands totally differently at higher filevels. The fi rst group of authors disclaims their for-fiest character and classifi es them within the classes Galio-Urticetea (Mucina et al. 1993, Pott 1992, Covali 2009) or Artemisietea vulgaris (Gilicka 1989, Świerkosz 1993, Pawlaczyk 2008) because of the high constancy of nitrophytes in the herb layer. Protagonists of the separate status of Black Locust stands classify them within the class Robinietea(Jurko 1963, Hada(( č & Sofron 1980, Theurillat et al. 1994, Borhidi 1996, 2003, 2006, Aeschimann etal. 2004, Jarolímek & Šibík 2008, Solomakha 2008

9 , 3, ,,)

or the lone-standing alliance Robinion pseuda-caciae Csürös-Káptalan 1968 of the class Querce-tea pubescenti-petraeae (Mititel u et al. 1973, Oprea 2004, Sanda et al. 2008). According to theCode of Phytosociological Nomenclature (Weber et al. 2000), publication of this alliance is probably not effective. A search for the paper with the original di-agnosis of the alliance (“Robinion pseudacaciae Csürös-Káptalan 1968” cited in later articles) was un-successful even when done by Romanian colleagues(Oprea, personal comm.). Kowarik (1995) suggest-ed the establishment of the new order Robinietal ia within the class Querco-Fagetea.

Jurko (1963) defends a forest character of Black Locust stands. Kowarik (1995) argues for the pos-sibility of their unambiguous assignment to forestformations using qualitative and quantitative aspects.The same opinion is shared by Klauck (1986), who detected a steady microclimate (temperature, relativeair moisture, and evaporation in herb and shrub lay-ers) inside the Black Locust stand in comparison to the neighbouring grass community. He suggests that this is due to the similar spatial structure of Black Lo-cust forests and natural forests, although he disagreeswith their classifi cation within forest communitiesdue to the occurrence of ruderal species. Platen &Kowarik (1995) confi rm the forest character of Black Locust stands on the basis of comparison of speciescomposition of vegetation, spider and Carabidae populations between dry sandy grasslands, Black Lo-cust stands of various ages and stands dominated by native pioneer broad-leaved woody species.

Another key objection of opponents of the class Robinietea deals with the absence of character spe-cies (Mucina et al. 1993). In typical Black Locust stands of the order Chelidonio-Robinietal ia ,dominated nitrophilous taxa play an important role, differentiating the stands from other forests, while occurring in other communities as well. Such speciescannot thus be used as character species, but only asdifferential species (cf. Moravec et al. 1994). Kowar-ik (1995) proposed using Robinia pseudacacia as the only character species of the Black Locust stands,but only when the spontaneously developed BlackLocust stands are separated carefully from plantedand forestry treated cultures. He recommended theirfurther specifi cation to create a group of differentialspecies that can indirectly confirm fifi xation of nitro-figen by Black Locust. There could be, according toKowarik (1995), sharp differentiation of Black Lo-cust stands from other forest communities not onlydue to fl oristic composition, but ecologically as well.He compares this situation to units that have onlyone character species or are distinguished only by acombination of differential species. Moravec et al.(1994) stated that it is possible to distinguish certain vegetation units from others by a combination of dif-ferential species.

Jarolímek & Šibík (2008) compared the classifica-fition sharpness of vegetation classes of Slovakia. Theclass Robinietea reached a relatively high value of the Sharpness Index (S = 40.17), in comparison withthe class Galio-Urticetea, which is the secondleast sharp (S = 4.73) due to the occurrence of manytaxa with a wide ecological range. Indeed, they foundthe highest similarity of the Robinietea class toGalio-Urticetea, but no xerophilous Black Locuststands appeared in the analysed set. Different sizes of data-sets in the compared classes could cause furtherdistortion.

No strong evidence has been given so far to make a defi nite classififi cation of Black Locust stands into the fihierarchical system possible. The relatively low num-ber of phytosociological relevés is a problem for themajority of Central European countries when someexisting types of Black Locust stands are excluded.The number of available phytosociological relevés isuneven in different Central European countries, andmany of them are recorded only in synoptical tables without the possibility of checking their creation ac-cording to original data. The character of the datathus does not empower the overall synthesis fromall Central European countries. Based on analysis of the Bohemian data-set, the proposed position of theseparate class Robinietea seems to be the best solu-tion. Integration of Black Locust stands presented byKowarik (1995) into the class Querco-Fagetea, although a unique order, is not suitable with respectto the species-rich shrubby Black Locust stands onrocky slopes that are characterised by the high fre-quency, and often cover as well, of species of the classFestuco-Brometea (association Melico trans-si lvanicae-Robinietum).

eschweizerbartxxx_author

Vegetation classi� cation and synecology of Bohemian � Robinia pseudacacia 237a

Data validation

The existing description of the class Robinieteawas incorrect. Validation of the order and descrip-tion of new alliances was carried out by Hadač &Sofron (1980), but according to the Code of Phy-tosociological Nomenclature (Weber et al. 2000),since 1979 a publication is valid only when the no-menclatural type of syntaxa is given (Art. 5). Within the class Robinietea, that qualification is not ful-fifi lled for the orderfi Chelidonio-Robinietal ia , nor for alliances Chelidonio-Robinion or Bal-loto nigrae-Robinion. The alliance Bal loto nigrae-Robinion so far has the only validly pub-lished association Balloto nigrae-Robinietum,which can therefore be marked as a holotypus (Art.18). In agreement with Article 20, we propose assum-ing the association Chelidonio-Robinietum as a nomenclatural typus, namely a lectotypus, for the alliance Chelidonio-Robinion, which contains three validly published associations so far. Similarly,the order Chelidonio-Robinietal ia consists of two alliances, from which we recommend assumingthe alliance Chelidonio-Robinion (Art. 20) as a lectotypus again. Since January 1st 1980, marking of acharacter and/or differential species is an obligatory component of the original diagnosis of a syntaxa of higher rank than an association (Weber et al. 2000,Art. 8). However, Hadač & Sofron (1980) give onlydiagnostic species, and Jurko (1963) did not mark the diagnostic species of the order Chelidonio-Rob-inietal ia . Therefore we propose diagnostic spe-cies combination of the order Chelidonio-Rob-inietal ia as follows: Chelidonium majus, Galeopsistetrahit agg., Galium aparine, Moehringia trinervia, Ribes uva-crispa, Robinia pseudacacia, Rubus frutico-sus agg., Sambucus nigra, Stellaria media, Taraxacum sect. Ruderalia, and Urtica dioica.

Proposal of the classification of Black Locustfistands in Central Europe

Robinietea Jurko ex Hadač & Sofron 1980 Chelidonio-Robinietalia Jurko ex Hadač & Sofron 1980 (mesophilous secondary Black Locust forests with nitrophytes) Chelidonio-Robinion Hadač & Sofron 1980 (mesophilous Black Locust stands with occurrence of nitrophilous species on soils with various nutrients, soil texture, and moisture regimes) Chelidonio-Robinietum Jurko 1963 (Syn.: Impatienti parviflorae-Robinietumfl Sofron 1967) Poo nemoralis-Robinietum Němec ex Vítková & Kolbek in Kolbek et al. 2003 cardaminopsietosum arenosae subass. nova typicum subass. nova Solidagini-Robinietum (Wendelberger 1955) Jurko 1963

Classification of Black Locust stands at thefilower hierarchical level

Jarolímek & Šibík (2008) mentioned a list of pairsof alliances with the highest similarity for the Slovakregion. The alliances Chelidonio-Robinion andBalloto nigrae-Robinion were the most simi-lar of all, but only 46 phytosociological relevés were used for the classifi cation. Despite this, a question ap-pears of whether it would not be better to join bothalliances of the order Chelidonio-Robinietal ia into only one. On the other hand, Borhidi (2006)proposed establishing a new order Bal loto nigrae-Robinietal ia with one alliance, Bal loto nigrae-Robinion. In Bohemia, only a few stands of thisalliance occur. However, the results of preliminary classification of available Central European phyto-sociological relevés, which could be included into the alliance Bal loto nigrae-Robinion and asso-ciation Chelidonio-Robinietum, acknowledged the possibility to merge both alliances. For definitefidecision, it would be needed to have more phytoso-ciological relevés from more countries.

At the lowest hierarchical level, the authors clas-sify Black Locust stands differently as well. While a proportion of authors include all stands in one com-munity (e.g. Diesing & Gödde 1989, Oberdorfer1992, Pott 1992, Mucina et al. 1993) or associa-tion (e.g. Fekete 1965, Brzeg & Wojterska 2001,Sanda et al. 2008), other authors (e.g. Ščepka 1982, Jarolímek & Šibík 2008, Solomakha 2008)

g choose

a very narrow association concept according to the dominant species. Alternation of dominant speciesis very common in mesophilous Black Locust stands where fl oristic composition is usually the same, and taxa covers vary in accordance with the soil moisture regime, soil profi le thickness, the former community,the seed bank in the surroundings, and so on. Due to the high constancy of diagnostic species of the as-sociation Chelidonio-Robinietum, the second approach with separate narrow units can hardly be accepted.

The questionable conception has also associa-tion Robinietum pseudo-acaciae (“Bromo steri l is-Robinietum”) according to different authors. Only the phytosociological relevés from Hungary (Fekete 1965) and Romania (Oprea 2004)are conformable with the original diagnosis in agree-ment with Pocs (1954) (Fig. 15: squares inside thecircle). Next authors (e.g. Ščepka 1982, Benčaťová & Benčať 2005, 2008) included in this associationthe relevés with dominating Bromus sterilis and oth-ers nitrophilous taxa, but with absence of spring geophytes, terophytes, and thermophilous species (Fig. 15: squares outside the circle). These relevés tend to be classifi ed as the association Chelidonio-Robinietum.

eschweizerbartxxx_author

238 M. Vítková & J. Kolbek

Balloto nigrae-Robinion Hadač & Sofron 1980 (Black Locust stands on minerally poorer, sandy, drier soils, occurring mostly in lowlands) Balloto nigrae-Robinietum Jurko 1963 Arrhenathero elatioris-Robinietum Šimonovič, Šomšák & Nikodemová ex Vítková & Kolbek ass. nova Robinietum pseudo-acaciae Pocs 1954 (Syn.: Bromo sterilis-Robinietum Fekete 1965) Euphorbio cyparissiae-Robinietalia Vítková in Kolbek et al. 2003 (low, tortuous and xerophilous Black Locust stands of thermophilous habitats) Euphorbio cyparissiae-Robinion Vítková in Kolbek et al. 2003 (low, tortuous and xerophilous Black Locust stands of thermophilous habitats) Melico transsilvanicae-Robinietum Kolbek & Vítková in Kolbek et al. 2003.

The separate status of the associations Solidagini-Robinietum and Balloto nigrae-Robinietum needs tobe verifi ed by systematic research in the future. Simi-larly, the merger of the alliances Chelidonio-Rob-inion and Balloto nigrae-Robinion would requireevaluation of more extensive data set, which is notavailable so far.

Acknowledgements. Participation of the authors was possibledue to the Czech research project AVOZ 60050516. The publish-ing of the article was supported by Grant No. IAAX00050801of the Grant Agency of the Academy of Sciences of the Czech Republic. We are indebted to L. Balogh, A. Borhidi (Hungary), I. Jarolímek, R. Kanka, M. Valachovič (Slovakia), P. Pawlaczyk, F. Święs (Poland), A. Oprea with his colleagues (Romania), andtwo anonymous reviewers for consultations concerning theclassification of Black Locust stands and procuration of neededfipapers, respectively. We are much obliged also to P. Pawlaczyk(Poland), R. Böcker, and M. Dirk (Germany) for their disclo-sure of unpublished relevés. Further, we appreciate the help of J. Tonika and L. Borůvka with the correction of soil and geo-logical terms; J. Sofron, Z. Svobodová, and V. Větvička with the location of the published relevés; A. Keményová with transla-tion of the Hungarian papers; O. Vítek with partial translationinto English and creation of Fig. 2 and 3; and I. Rajznoverová, I. Jarošincová, H. Zbuzková and O. Nováková with technical assistance. Language revision was carried out by the Proof-Reading-Service.com, United Kingdom.

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Address of the authors:Michaela Vítková and Jiří Kolbek, Institute of Botany, v. v. i., Academy of Sciences of the Czech Republic, 252 43 Průhonice, Czech Republic.E-mails: vitkovi@seznam.cz, jiri.kolbek@ibot.cas.cz