Broad-leaved forests in the North-Western Caucasus. I. Order Carpinetalia betuli P. Fukarek 1968

N. Ye. Shevchenko, T. Yu. Braslavskaya



Braun-Blanquet classification of the piedmont and mountain broad-leaved forests (formed by Carpinus betulus, Quercus petraea/hartwissiana/robur, Fraxinus excelsior) was done basing on 224 relevés collected in 20142018 in the North-Western Caucasus (N 43,544,8, E 38,541,5; Fig. 2). DCA-ordination of the data corresponds to their correlation with environment variables (absolute elevation, geographical coordinates, tree canopy density) in Landolts ecological scales was carried out.

Suballiance Tamo communisCarpinenion betuli suball. nov. prov. and new lower syntaxa are proposed (Table 1) within the alliance CrataegoCarpinioncaucasicaePassarge 1981. Nomenclature type of the suballiance is the ass. Tamo communisCarpinetum betuli ass. nov. (Table 3, holotypus is relevé 4: authors number 83, author N. E. Shevchenko, 19.07.2016, N 44.257, E 39.760, 352 m above sea level, slope 5NE) with three variants: typica (Table 3: 115), Staphylea colchica (Table 3: 1627) and Festuca drymeja (Table 3: 2838). Also, there are ass. Aro maculatiCarpinetum betuli (Table 4: 115; holotypus is relevé 5: authors number 427, author N. E. Shevchenko, 12.05.2017, N 44.471, E 40.516, 455 m above sea level, slope 6NE) as well as Abies nordmannianaCarpinus betulus community (Table 4: 1627) in new suballiance. Not too common in the North-Western Caucasus are ash (Fraxinus excelsior) forests which have no suffiicient floristic peculiarity and are considered as facies in associations Tamo communisCarpinetum betuli var. typica and Aro maculatiCarpinetum betuli.

The specificity of the studied forests in comparison with hornbeam and oak forests in the Central Caucasus (Georgia), North Turkey, the Balkans and the Crimea (Passarge, 1981a; Korzhenevskiy, 1982; Didukh, 1996; Korkmaz et al., 2008; Košir et al. 2013; Çoban, Willner, 2019; Novak et al., 2019) is that the North-Western Caucasus forest flora includes (Table 2), besides European species of temperate broad-leaved forests (Acer campestre, Euonymus europaea, Carex sylvatica, Convallaria majalis, Rubus caesius), also southern European species of thermophilous broad-leaved forests (Acer tataricum, Cornus mas, Ligustrumvulgare, Lonicera caprifolium, Tamus communis, Vincetoxicum scandens,Hedera helix, Festuca drymeja), and reduced set of species which are character for Euxinian and Caucasian forests(Quercus hartwissiana, Tilia begoniifolia, Rhododendron luteum, Daphne caucasica, Staphylea colchica, Smilax excelsa, Paris incompleta, Polygonatum orientale, Lathyrus roseus, Campanula alliariifolia, but without Daphne pontica, Epimedium pubigerum, Erica arborea, Ostrya carpinifolia, Salvia forskahlei, Vaccinium arctostaphylos).

DCA-ordination (Fig. 8) showed that the differences in species composition of the broad-leaved forest syntaxa are due to both absolute elevation (vector Elev in Fig. 7) and geographic longitude (vector E) of the relevés. So, forests of ass. Aro maculatiCarpinetum betuli and community Abies nordmannianaCarpinus betulusare situated, in general, at higher positions than forests of ass. Tamo communisCarpinetum betuli, and the first syntaxon is situated east of the two last ones. Floristic difference between these syntaxa corresponds with parameters assessed by values of Landolts scales: soil aeration (vector D), climate continentality (vector K) and light regime (vector L). The lowest α-diversity is in ass. Tamo communisCarpinetum betuli, and the highest is in the ass. Aro maculatiCarpinetum betuli and community Abies nordmannianaCarpinus betulus it (Fig. 9).

Associations Carpino betuliQuercetum petraeae Grebenshchikov et al. 1990 and Rhododendro luteiQuercetum petraeae Grebenshchikov et al. 1990, earlier described on small sets of relevés in the North-Western Caucasus (Grebenshchikov et al. 1990) within the alliance Carpino betuliQuercion petraeae Grebenshchikov et al. 1990 (now invalid due to absence of stated holotypus), are very close to the new ass. Tamo communisCarpinetum betuli. Recently described (also on small sets of data) five associations (Akatova, Ermakov, 2020), within the alliance CrataegoCarpinion caucasicae Passarge 1981, are valid. Therefore, further clarification and optimization of the North-Western Caucasus oak-hornbeam forest classification is required, having in mind the final decision on the alliance/suballiance names and diagnosis. Their belonging to the order (Carpinetalia betuli or Rhododendro ponticiFagetalia orientalis) is also a debatable question, because researchers working in such forests on Balkans, in the Crimea and the North Turkey have come to different conclusions. The analysis of species with constancy 6080 % in 224 relevés from the studied area reveals 8 diagnostic ones of the order Carpinetalia betuli vs. only 3 diagnostic ones of the order Rhododendro ponticiFagetalia orientalis, that allows to assign these oak-hornbeam forests to the first order. The Abies nordmannianaCarpinus betulus community is intermediate between these two orders but after trees from shade-tolerant fir undergrowth become, the canopy mature will be mixed that is character for forests of the order Rhododendro ponticiFagetalia orientalis.

Key words: mountain and piedmpont broad-leaved forests, North-Western Caucasus, Braun-Blanquet classification approach, alliance CrataegoCarpinion caucasicae

Section: Articles

How to cite

Shevchenko N. Ye., Braslavskaya T. Yu. 2021. Broad-leaved forests in the North-Western Caucasus. I. Order Carpinetalia betuli P. Fukarek 1968 // Rastitelnost Rossii. 42: 118145.

Received July 13 2020. Signed for printing December 30 2021


Agroklimaticheskiy spravochnik po Krasnodarskomu krayu [Agricultural-climatic handbook for the Krasnodarsiy Region] 1961. Krasnodar. 466 p. (In Russian).

Akatova T. V., Bibin A. R., Grabenko E. A., Zagurnaya Yu. S. 2016. Klyuchevye biotopy ekspluatiruemykh lesov Krasnodarskogo kraya i Respubliki Adygeya (Severo-Kavkazskiy gornyy region) [Key biotopes of exploited forests in the Krasnodarskiy Region and Adygeja Republik (the Northern Caucasus mountain ecoregion)]. Ustoychivoe lesopolzovanie. 3: 2935. (In Russian).

Akatova Yu. S., Ermakov N. B. 2020. Soobshchestva shirokolistvennykh lesov nizhney chasti lesnogo poyasa basseyna r. Belaya (Severo-Zapadniy Kavkaz) [Communities of broadleaved woodlands in the lower part of the forest belt of the Belaya river basin (North-Western Caucasus). Plant biology and Horticulture: theory, innovation. 156: 6578. (In Russian).

Altukhov M. D., Litvinskaya S. A. 1989. Okhrana rastitelnogo mira na Severo-Zapadnom Kavkaze [The conservation of plants and vegetation in the North-West Caucasus]. Krasnodar. 189 p. (In Russian).

Biota ecosystem Bolshogo Kavkaza [Biota of the ecosystems in the Bolshoy Caucasus]. 1990. oscow. 221 p. (In Russian).

Bitsin L. V., Ilin L. I., Maltsev M. P. 1966. Forests of the Northern Caucasus. Lesa SSSR. T. 3 [Forests of the USSR. Vol. 3]. oscow. P. 261313. (In Russian).

Bondarenko S. V. 2010. The characteristic of forests of the Aphyps river basin (the North-Western Caucasus). Botanicheskiy zhurnal. 95(2): 215225. (In Russian).

Braun-Blanquet J. 1964. Pflanzensoziologie. Grundzüge der Vegetationskunde. 3 Aufl. Wien; New York. 865 s.

Cherepanov S. K. 1995. Sosudistye rasteniya Rossii i sopredelnykh gosudarstv (v predelakh byvshego SSSR) [Vascular plants of Russia and adjacent states (within the former USSR)]. St. Petersburg. 992 p. (In Russian).

Chytrý M., Tichy L., Holt J., Botta-Dukát Z. 2002. Determination of diagnostic species with statistical fidelity measures. Journal of Vegetation Science. 13(1): 7990.

Chytrý M., Otypková Z. 2003. Plot sizes used for phytosociological sampling of European vegetation. Journal of Vegetation Science. 14(4): 563570.

Çoban S., Willner W. 2019. Numerical classification of the forest vegetation in the Western Euxine Region of Turkey. Phytocoenologia. 49(1): 71106.

Didukh Ya. P. 1996. Nemoral forests in the Mountain Crimea of the class Querco-Fagetea Br.-Bl. et Vlieg. 1937. Ukrainian Phytosociological Collection. Ser. A. Iss. 3. Kiev. P. 3451. (In Ukrainian).

Elagin I. N. 1953. Oak forests on the west extreme of north slope of the Caucasian Ridge. Shirokolistvennye lesa Severo-Zapadnogo Kavkaza [Broad-leaved forests in the North-Western Caucasus]. oscow. P. 187243. (In Russian).

Ermakov N. B. 2012. Prodromus of higher syntaxa of Russian vegetation. Sovremennoye sostoayniye osnovnykh kontseptsiy nauki o rastitelnosti (Mirkin B. M., Naumova L. G.) [Contemporary state of main concepts in vegetation science (by Mirkin B. M., Naumova L. G.)]. Ufa. P. 377483. (In Russian).

Euro+Med Plantbase. The information resource for Euro-Mediterranean plant diversity. (date of access: 20.03.2021).

Gigauri G. N. 1986. Shade tolerance of Abies nordmanniana, Picea orientalis and Fagus orientalis. Lesovedenie. 3: 5056. (In Russian).

Grebenshchikov O. S. 1974. O nekotorykh vidakh graba, ikh soobshchestvakh i geograficheskom rasprostranenii [On some hornbeam species, their communities and geographical range]. Botanicheskiy zhurnal. 59(7): 10131019. (In Russian).

Grebenshchikov O. S., Shanina A. A., Belonovskaya E. A. 1990. Forests on the west extreme of the Bolshoy Caucasus. Biota ecosystem Bolshogo Kavkaza [Biota of the ecosystems in the Bolshoy Caucasus]. oscow. P. 6384. (In Russian).

Grossgeym A. A. 1948. Rastitelnyy pokrov Kavkaza [Vegetation blanket of the Caucasus]. oscow. 267 p. (In Russian).

Grudzinskaya I. A. 1953. Broad-leaved forests in the piedmont of the North-West Caucasus. Shirokolistvennye lesa Severo-Zapadnogo Kavkaza [Broad-leaved forests in the North-West Caucasus]. oscow. P. 5186. (In Russian).

Gulisashvili V. Z. 1979. Structure peculiarities of mountain forests. Gornye lesa [Mountain forests]. Moscow. P. 2940. (In Russian).

Gulisashvili V. Z., Makhatadze L. B., Prilipko L. I. 1975. Rastitelnost Kavkaza [The Caucasus vegetation]. Moscow. 234 p. (In Russian).

Gvozdetskiy N. A. 1963. Kavkaz: ocherk prirody [The Caucasus: an essay on nature]. Moscow. 264 p. (In Russian).

Hammer Ø., Harper D. A. T., Ryan P. D. 2001. PAST: Paleontological statistics software package for education and data analysis.Palaeontologia Electronica. 4(1): 19. (date of access: 20.06.2020).

Ignatov M. S., Afonina O. V., Ignatova E. A., Abolina A., Akatova T. V., Baisheva E. Z., Bardunov L. V., Baryakina E. A., Belkina O. A., Bezgodov A. G., Boychuk M. A., Cherdantseva V. Ya., Czernyadjeva I. V., Doroshina G. Ya., Dyachenko A. P., Fedosov V. E., Goldberg I. L., Ivanova E. I., Jukoniene I., Kannukene L., Kazanovsky S. G., Kharzinov Z. Kh., Kurbatova L. E., aksimov . I., Mamatkulov U. K., Manakyan V. A., Maslovsky O. M., Napreenko M. G., Otnyukova T. N., Partyka L. Ya., Pisarenko O. Yu., Popova N. N., Rykovsky G. F., Tubanova D. Ya., Zheleznova G. V., Zolotov V. I. 2006. Check-list of mosses of East Europe and North Asia. Arctoa. 15: 1130.

Korkmaz H., Yalçin E., Kutbay H. G., Berk E., Bilgin A. 2008. Contribution to the knowledge of the syntaxonomy and ecology of macchie and forest vegetation in Paphlagonia, North Anatolia, Turkey. Acta Botanica Gallica. 155(4): 495512.

Korotkov K. O., Morozova O. V., Belonovskaya E. A. 1991. The USSR Vegetation Syntaxa Prodromus. Moskow. 346 p.

Korzhenevskiy V. V. 1982. Sessile-oak forests on southern macro-slope of the Main Ridge of the Crimea Mountains. Trudy Nikitskogo botanicheskogo sada [Proceedings of the Nikitskiy botanical garden]. 86: 3646. (In Russian).

Košir P., Casavecchia S., Čarni A., Škvorc Ž., Zivkovic L., Biondi E. 2013. Ecological and phytogeographical differentiation of oak-hornbeam forests in southeastern Europe. Plant Biosystems. 147(1): 8498.

Landolt E., Baumler B., Erhardt A., Hegg O., Klotzli F., Lammler W., Nobis M., Rudmann-Maurer K., Schweingruber F. H., Theurillat J.-P., Urmi E., Vust M., Wohlgemuth T. 2010. Flora indicativa Okologische Zeigerwerte und biologische Kennzeichen zur Flora der Schweiz und der Alpen. 2nd edition. Haupt. 378 p.

Makunina A. A. 1986. Fizicheskaya geografia gornykh regionov SSSR [Physical geography of mountain areas in the USSR]. Moscow. 166 p. (In Russian).

McCune B., Mefford M. J. 2006. PC-ORD. Multivariate Analysis of Ecological Data. Version 5.10. MjM Software, Gleneden Beach, Oregon, U. S. A.

Metodicheskie podkhody k ekologicheskoy otsenke lesnogo pokrova v basseyne maloy reki. [Methodical approaches to ecological evaluation of forest blanket in a small-river basin]. 2010. Moscow. 383 p. (In Russian).

Mucina L., Bültmann H., Dierßen K., Theurillat J.-P., Raus T., Čarni A., Šumberová K., Willner W., Dengler J., Gavilán García R., Chytrý M., Hájek M., Di Pietro R., Iakushenko D., Pallas J., Daniëls F. J. A., Bergmeier E., Santos Guerra A., Ermakov N., Valachovič M., Schaminée J. H. J., Lysenko T., Didukh Y. P., Pignatti S., Rodwell J. S., Capelo J., Weber H. E., Solomeshch A., Dimopoulos P., Aguiar C., Hennekens S. M., Tichý L. 2016. Vegetation of Europe: hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Applied Vegetation Science. 19 (Suppl. 1): 3264.

Novák P., Zukal D., Kalníková V., Chytrý K., Kavgacı A. 2019. Ecology and syntaxonomy of Colchic forests in south-western Georgia (Caucasus region). Phytocoenologia. 49(3): 231248.

Ostapenko B. F. 1968a. On forest typology of northern slope of the Bolshoy Caucasus. Lesotipologicheskie issledovaniya (Trudy Kharkovskogo selskokhozyaystvennogo instituta. T. 72(109)) [Studies on forest typology: Proceedings of Kharkov Agricultural Institute. Vol. 72(109)]. Kharkov. P. 345. (In Russian).

Ostapenko B. F. 1968b. Classification of forest types and typological zoning of northern slope of the Bolshoy Caucasus. Lesotipologicheskie issledovaniya (Trudy Kharkovskogo selskokhozyaystvennogo instituta. T. 72(109)) [Studies on forest typology: Proceedings of Kharkov Agricultural Institute. Vol. 72(109)]. Kharkov. P. 45110. (In Russian).

Ostapenko B. F. 1968c. Diagnostics and economic arranging of forest types on northern slope of the Bolshoy Caucasus. Lesotipologicheskie issledovaniya (Trudy Kharkovskogo selskokhozyaystvennogo instituta. T. 72(109)) [Studies on forest typology: Proceedings of Kharkov Agricultural Institute. Vol. 72(109)]. Kharkov. P. 111257. (In Russian).

Otypková Z., Chytrý M. 2006. Effects of plot size on the ordination of vegetation samples. Journal of Vegetation Science. 17(4): 465472.

Passarge H. 1981a. Carpineta im kartalinischen Kaukasus. Phytocoenologia. 9(4): 533545.

Passarge H. 1981b. Über Fagetea im kartalinischen Kaukasus. Feddes Repertorium. 92(56): 413431.

Peet R. K., Roberts D. W. 2013. Classification of natural and semi-natural vegetation. Vegetation ecology. 2ndd. E. van der Maarel, J. Franklin (eds.). Chichester. P. 2870.

Roleček J., Tichý L., Zelený D., Chytrý M. 2009. Modified TWINSPAN classification in which the hierarchy respects cluster heterogeneity. Journal of Vegetation Science. 20(4): 596602.

Safarov I. S., Olisaev V. A. 1991. Lesa Kavkaza: sotsialno-ekologicheskiye funktsii [Caucasian forests: social and ecological functions]. Vladikavkaz. 271 p. (In Russian).

Shirokolistvennye lesa Severo-Zapadnogo Kavkaza [Broad-leaved forests in the North-Western Caucasus]. 1953. oscow. 384 p. (In Russian).

Sosnin L. I. 1939. Forest types in the Caucasus State Reserve. Trudy Kavkazskogo gosudarstvennogo zapovednika. Vyp. 2 [Proceedings of the Caucasus State Reserve. Iss. 2]. oscow. P. 582. (In Russian).

Spravochnik lesotakcatsionnykh normativov dlya Severnogo Kavkaza (utverzhden prikazom Rosleskhoza. 10 ot 19.01.1995) [Handbook of forestry-taxation norms for the North-Western Caucasus (approved by order of Russian Forestry Service, N 10 on 19.01.1995)]. 1995. Moscow. 90 p. (In Russian).

Tichy L. 2002. JUICE, software for vegetation classification. Journal of Vegetation Science. 13(3): 451-453.

Theurillat J.-P., Willner W., Fernández-González F., Bültmann H., Čarni A., Gigante D., Mucina L., Weber H. 2021. International Code of Phytosociological Nomenclature. 4th edition. Applied Vegetation Science. 24(1): 162.

Ugurlu E., Roleček J., Bergmeier E. 2012. Oak woodland vegetation in Turkey a first overview based on multivariate statistics. Applied Vegetation Science. 15(4): 590608. 2012.01192.x.

Zernov A. S., Alekseev Yu. Ye., Onipchenko V. G. 2015. Opredelitel sosudistykh rasteniy Karachaevo-Cherkesskoy Respubliki [Key for identifying of vascular plants in the Karachaevo-Cherkessk Republik]. Moscow. 459 p. (In Russian).

Zonn S. V. 1950. Gorno-lesnye pochvy Severo-Zapadnogo Kavkaza [Mountain forest soils in the North-Western Caucasus]. MoscowLeningrad. 333 p. (In Russian).