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Thermophilous oak forests of the steppe and forest-steppe zones of Ukraine and Western Russia

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We present a formal classification of thermophilous oak forests of the steppe and forest-steppe zones of Ukraine and Russia. Using 45 sources (synoptic tables; some from Central and Western Europe were also included for comparative purposes), we classified the data using cluster analyses, followed by post-classification tools aimed at formal identification of the optimal number of clusters and fidelity-based table sorting. Db-RDA ordination and a CART were used to identify the lead putative climatic drivers of the vegetation patterns. Of the six clusters identified by our classification procedures, two clusters are interpreted here as new alliances (Betonico-Quercion, Scutellario-Quercion). Some new associations classified into these alliances were also either validated or described as new. We further show that the Quercion petraeae is of heterogenous nature and the position of the units previously classified as the Potentillo albae-Quercion should be re-evaluated. NMDS was used to analyse the patterns of the phytocoenologic elements (diagnostic species of relevant syntaxonomic classes) in the six clusters. This analysis revealed that the classification of the Ukrainian and Russian thermophilous oak forests into the Quercetea pubescentis class is untenable and remains open to further scrutiny.

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Classification and Regression Tree


Distance-based Redundancy Analysis


Non-metric Multidimensional Scaling


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Appendix: Descriptions and validation of new syntaxa

Appendix: Descriptions and validation of new syntaxa

Betonico officinalis-Quercion roboris Goncharenko & Semenishchenkov all. nov. hoc loco

Dry-mesic floristic-rich oak forests in the Sarmatian region.

Misapplied names: Quercion petraeae sensu auct. ucrain. et ross., non Quercion petraeae Issler 1931; Potentillo albae-Quercion petraeae sensu auct. ucrain. et ross., non Potentillo albae-Quercion petraeae Jakucs in Zólyomi 1967

Holotypus: Galio tinctoriae-Quercetum roboris Goncharenko 2003

Diagnostic taxa: Quercus robur Clematis recta, Cytisus ruthenicus Ajuga genevensis, Allium oleraceum, Anthericum ramosum, Asperula tinctoria, Betonica officinalis, Campanula persicifolia, Digitalis grandiflora, Melampyrum nemorosum, Origanum vulgare, Potentilla alba, Securigera varia, Serratula tinctoria, Trifolium alpestre, Turritis glabra, Veronica chamaedrys, Vicia sepium, Vincetoxicum hirundinaria, Viola hirta

Scutellario altissimae-Quercion roboris Goncharenko all. nov. hoc loco

Steppic oak forests in the Pontic region.

Misapplied name: Aceri tatarici-Quercion sensu auct. ucrain. et ross., non Aceri tatarici-Quercion sensu Zólyomi 1957.

Holotypus: Fritillario ruthenici-Quercetum roboris Onyschenko, Dyakova & Karpenko ex Goncharenko ass. nov. (for the valid description of the association see below).

Diagnostic taxa: Quercus robur Acer tataricum, Cerasus fruticosa, Crataegus rhipidophylla agg., Prunus spinosa subsp. dasyphylla Aegonychon purpurocaeruleum, Carex michelii, Euphorbia semivillosa, Festuca stricta subsp. sulcata, Melica picta, Phlomis tuberosa, Scutellaria altissima, Vicia pisiformis, Vincetoxicum scandens

Daphno cneori-Quercetum roboris Goncharenko ass. nov. hoc loco

Xeromesophytic oak-pine forests on boreal terraces of the Dnieper River of the Central Ukraine.

Incl.: Convallario majalis-Quercetum roboris daphneosum cneori Gayova & Korotchenko 2013 (Gayova and Korotchenko 2013, p. 28)

Note: The association typifies pine-oak forests supported by the boreal terraces of the Dnieper River in Central Ukraine (near Cherkassy). (Gayova and Korotchenko 2013) classified this vegetation as the subassociation Convallario majalis-Quercetum roboris daphneosum cneori. We suggest that due to distinct floristic composition showing more xerophilous elements (e.g. Anemone patens, Daphne cneorum, Dracocephalum ruyschiana, Teucrium chamaedrys) when compared with other associations of the alliance Convallario majalis-Quercion roboris (see columns 18 – 20, Table S3); this syntaxon deserves the rank of an association.

Scorzonero ensifoliae-Quercetum roboris Sokolova ex Semenshchenkov ass. nov. hoc loco

Xeromesophytic oak forests in depressions on sandy terraces (arens) of the Don River in the steppe zone

Synonym: Scorzonero ensifoliae-Quercetum roboris Sokolova 2011 nom. inval. (arts. 3b & 5).

Holotypus: Russian Federation, Rostov Region, Tarasovsky District, near Mityakinskaya village; 48.5996° N lat., 39.7182° E long.; Habitat: sandy terrace of the Don River; Relevé area: 225 m2. Cover of tree layer: 70 %. Cover of shrub layer: 3 %. Cover of herb layer: 40 %. The species cover-abundance codes follow Braun-Blanquet (1964). Date: 2 August 2009. Author: T. Sokolova.

E1: Quercus robur 4, Malus sylvestris +, Pyrus pyraster +, Ulmus minor +. E2: Euonymus verrucosus 2, Acer tataricum +, Crataegus rhipidophylla +, Rosa sp. +, Rubus caesius +. E3: Aegopodium podagraria 3, Vincetoxicum hirundinaria 1, Agrimonia eupatoria +, Agrostis tenuis +, Arctium nemorosum +, Bromopsis inermis +, Calystegia sepium +, Campanula rapunculus +, C. trachelium +, Chelidonium majus +, Convallaria majalis +, Daucus carota +, Fragaria viridis +, Galeopsis tetrahit +, Galium aparine +, Geum urbanum +, Heracleum sibiricum +, Polygonatum odoratum +, Pulmonaria obscura +, Scorzonera ensifolia +, Stellaria media +, Trifolium medium +, Urtica dioica +, Viola suavis +, Vincetoxicum scandens +

Fritillario ruthenici-Quercetum roboris Onyschenko, Dyakova et Karpenko ex Goncharenko ass. nov. hoc loco

Syn.: Fritillario ruthenicae-Quercetum roboris Onyschenko, Dyakova & Karpenko 2007 nom. inval. (arts. 3b & 5)

Holotypus: Onyshchenko et al. (2007, p. 92, Tab. 1, relevé 1)

These are steppic oak forests on carbonate soils formed over chalk or limestone bedrocks, originally described from the National Landscape Park “Svyati Gory” in the Donetsk region in Ukraine. This association is possibly identical to the type Aceri tatarici-Quercetum pubescentis-roboris rossicum reported by Zólyomi (1957) for the Donetsk Ridge using the data from Ukrainian phytosociologists Gryn’ (1940) as well as from Kotov and Karnaukh (1940).

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Goncharenko, I., Semenishchenkov, Y., Tsakalos, J.L. et al. Thermophilous oak forests of the steppe and forest-steppe zones of Ukraine and Western Russia. Biologia (2020). https://doi.org/10.2478/s11756-019-00413-w

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  • Aceri tatarici-Quercion
  • Betonico officinalis-Quercion roboris
  • Eastern Europe
  • Quercetea pubescentis
  • Quercion petraeae
  • Scutellario altissimae-Quercion roboris
  • Thermophilous oak forests