ri^ DE GRUYTER
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HACQUETIA 13/2 • 2014, 279-284
DOI: 10.2478/hacq-2014-0008
SYNTAXONOMIC AND NOMENCLATURAL NOTES ON THE SCREE VEGETATION OF
cAucASuS
Elena A. BELONOVSKAYAi, Ladislav MUCINA^* & Jean-Paul THEURILLAT^
Abstract
We present description of a new class for the high-altitude screes of the Caucasus (the Lamio tomentosi-Chaero-phylletea humilis) and two alliances such as the Scrophulario minimae-Symphyolomion graveolentis (Chaerophylle-talia humilis, Lamio tomentosi-Chaerophylletea humilis) and the Scrophulario variegatae-Epilobion dodonaei (Epilobi-etaliafleischeri, Thlaspietea rotundifolii).
Key words: Georgia, alpine vegetation, Russian federation, riverine terraces, syntaxonomy. Izvleček
V članku predstavljamo opis novega razreda na visoko ležečih meliščih na Kavkazu (Lamio tomentosi-Lamietea to-mentosi) in dveh zvez - Scrophulario minimae-Symphyolomion graveolentis (Chaerophylletalia humilis, Lamio tomentosi--Chaerophylletea humilis) in Scrophulario variegatae-Epilobion dodonaei (Epilobietaliafleischeri, Thlaspietea rotundifolii). Ključne besede: Gruzija, alpinska vegetacia, Ruska federacija, rečne terase, sintaksonomija.
1. introduction	give way to the Drypidetea spinosi which, in our
syntaxonomic concept, also includes the Scro-The communities of the Thlaspietea rotundifolii	phulario-Helichrysetea italici 8. Brullo et al. 1998 (Braun-Blanquet 1948, Zollitsch 1966, Englisch et	(Brullo et al. 1998) and the Onosmo polyphyllae-al. 1993, Valachovič et al. 1997 etc.) are a common	Ptilostemonetea Korzhenevsky 1990 (see Golub et appearance in scree habitats of temperate, boreal	al. 2011 for the latest syntaxonomic synthesis). and subarctic zones of Europe, where they could	In Caucasus, Onipchenko (2002a: Table 3.1) be found in all vegetation altitudinal belts, from	described the Chaerophylletalia humilis and placed the planare and colline belts as high as the alpine	this order within the Thlaspietea rotundifolii, aland subnival belts. In the mediterranean-climate	though on page 18 he also noted that practically regions of the Iberian Peninsula they are replaced	all diagnostic species of the class are missing in by the Phagnalo-Rumicetea indurati (Rivas Goday	the Caucasian communities. Onipchenko (l.c.) et Esteve 1972) Rivas-Mart^nez, Izco et Costa	also suggested that there were some diagnostic 1973 (e.g. Ortiz & Puglar 2000, Rivas-Mart^nez et	species of the European alliances and orders and al. 2011) and in the (sub)mediterranean regions	therefore he decided to classify his new order of the Central and Eastern Mediterranean, they	(Chaerophylletalia humilis) within the Thlaspietea
1	Institute of Geography, Russian Academy of Sciences, Staromonetny 2, 109017 Moskva, Russian Federation; E-mail: belena53@mail.ru
2	Iluka Chair in Vegetation Science & Biogeography, School of Plant Biology, The University of Western Australia, 35 Stirling Highway, Crawley WA 6009, Australia; E-mail: Laco.Mucina@uwa.edu.au
3	Centre alpien de Phytogeographie, Fondation J.-M. Aubert, Case postale 71, CH-1938 Champex-Lac, Switzerland; Laboratoire de Biogeographie, Section de Biologie, Universite de Geneve, 1, ch. de l'Imperatrice, Case postale 60, CH-1292 Chambesy, Switzerland; E-mail: jean-paul.theurillat@unige.ch
* corresponding author
rotundifolii. In fact only Oxyria digyna, Crypto-gramma crispa and Epilobium dodonaei would qualify at the latter group. Interestingly, these species occur in the Caucasian scree communities at lower altitudes (perhaps (sub)montane to subalpine zone). The high altitudes (alpine to subnivale zones) of the Caucasus range are home to endemic-rich scree communities that appear to be floristically very different from the Thlaspietea rotundifolii of Europe (see also Nakhutsrishvili & Gagnidze 1999; nakhutsrishvili 2003).
Belonovskaya (2012; based on earlier paper by Belonovskaya & Korotkov 2002) developed the onipchenko's (2002) remark on the poor floristic relationship of the Caucasus scree communities and those of Europe further and introduced a new syntaxonomic concept - the Veronico telephi-ifoliae-Lamietea tomentosi, including one new order (Veronico telephiifoliae-Lamietalia tomentosi), with two new alliances (Drabo scabrae-Eunomion rotundifoliae, Scrophulario minimae-Symphyolo-mion graveolentis). Belonovskaya's (2012) paper however have not engaged fully with the syntax-onomy proposed by onipchenko (2002) and all the suggested new syntaxonomic units (class, order and the alliances) are rendered either invalid according to the International Code Phytosocio-logical Nomenclature; Weber et al. 2000) or (in one case) should be considered synonymous with an earlier syntaxonomic concept of onipchenko (2002).
The prime aim of this paper is to validate the syntaxon names (or introduce a new one) and contribute to clarification of the syntaxonomic relationship of the onipchenkos's syntaxa and those suggested by Belonovskaya (l.c.) in the light of the new syntaxonomic-nomenclatural synthesis undertaken within so called 'EuroVeg-Checklist' (Mucina et al. in prep.).
2. NOMENCLATURE OF THE NAME 'VERONICO TELEPHIIFOLIAE-LAMIETEA TOMENTOSI'
We agree that the endemic-rich, high-altitude scree communities of the Caucasus should be considered as a syntaxonomic concept different from the Thlaspietea rotundifolii. However, name 'Veronico telephiifoliae-Lamietea tomentosi' (coined to accommodate these high-altitude Caucasian scree communities) has been invalidly (Art. 2b of the ICPN) published in Belonovskaya (2012:
967) because the name of the unique order - the 'Veronico telephiifoliae-Lamietalia tomentosi - include in the original diagnosis of the new class, was invalidly published at the time of the publication of the class name.
Indeed, the original diagnosis of the Veronico telephiifoliae-Lamietalia tomentosi contains two new alliances: the Drabo scabrae-Eunomion rotun-difoliae and Scrophulario minimae-Symphyolomion graveolentis. There are four new associations in the original diagnosis of the Drabo scabrae-Eu-nomion rotundifoliae and three new associations in the Scrophulario minimae-Symphyolomion graveolentis (Belonovskaya 2012). All these seven associations are validly published because their original diagnosis contains only the type releve (in Russian) presented in the Table 1 in Belonovskaya (2012) and each type releve containing the name-giving taxa of respective association. on the other hand, both names of the new alliances are invalidly published because their type, indicated in the bottom of the Table 1 is given in Russian not matching the demand of the Art. 5 of the IPCN (Weber et al. 2000) that requires (expressis verbis) using the Latin terms typus or holotypus. Because of this technicality, the original diagnosis of the 'Veronico telephiifoliae-Lamietalia tomentosi containing no validly published alliances, the name remains invalidly published (Arts. 2b and 8 of the IPCN).
Besides, even if the new alliance Drabo scabrae-Eunomion rotundifoliae (Belonovskaya 2012) was validly published, the latter alliance name would have become automatically a superfluous name for the validly published Chaerophyllion humilis Onipchenko 2002 which was listed by Belonovskaya (2012) in the synonymy of the Drabo scabrae-Eunomion rotundifoliae on p. 968. Similarly, if the new order Veronico telephiifoliae-Lamietalia tomen-tosi were validly published, it would have been a homotypical synonym for the validly published Chaerophylletalia humilis Onipchenko 2002. It appears that syntaxonomically this order matches the Belonovskaya's 'Veronico telephiifoliae-Lami-etalia tomentosi'. Onipchenko (2002a: 168) listed both name-giving taxa (Lamium tomentosum and Veronica telephiifolia; syn. V. minuta C.A. Meyer; see Fig. 1) in the diagnosis of Chaerophylletalia humilis.
In order to avoid confusion in future, we introduce a new name for the invalid syntaxonomic concept of the 'Veronico telephiifoliae-Lamietea tomentosi' and by recognising the role of Chae-
Figure 1: Selected character species of the Lamio tomentosi-Chaerophylletea humilis: A: Lamium tomentosum, B: Veronica minuta, C: Chaerophyllum humile. Broad valley of the Tergi (Terek) River near the town of Stepantsminda in Caucasus, Republic of Georgia with extensive terraces supporting communities of the Murbeckiello huetii-Epilobion dodonaei (D); Epilobium dodonaei photographed on the alluvial terraces of the Tergi (Terek) River, Georgia. Photo credits: A-C photos is Mikhail Dokukin (Nalchik, Kabardino-Balkaria, Russia), D-E: L. Mucina.
Slika 1: Izbrane značilnice razreda Lamio tomentosi-Chaerophylletea humilis: A: Lamium tomentosum, B: Veronica minuta, C: Chaerophyllum humile. Široka dolina reke Tergi (Terek) River v bližini mesta Stepantsminda na Kavkazu, Gruzija z ekstenzivnimi terasami, kjer se pojavljajo združbe zveze Murbeckiello huetii-Epilobion dodonaei (D); Epilobium dodonaei je fotografiran na aluvialni terasi reke Tergi (Terek), Gruzija. Foto: A-C Mikhail Dokukin (Nalchik, Kabardino-Balkaria, Russia), D-E: L. Mucina.
rophyllum humilis as one of the local dominating species, we coin the name and concept of the La-mio tomentosi-Chaerophylletea humilis (for the full protologue see below).
4. description of the
NEW syntaxa and THE NEW
syntaxonomic scheme for the Caucasian scree vegetation
3. THE case of MURBECKIELLIOn HUETII onipchenko 2002
The concept and name murbeckiellion huetii was introduced for the first time by onipchenko (2002a) who classified two associations within this alliance - the Hyalopoo ponticae-Oxyrietum digynae onipchenko 2002 from siliceous, snow-rich alpine moraine habitats and the Scrophulario variegatae-Epilobietum dodonaei which was designated as the typus of the alliance. The latter association, however, was not validly described because the typus of the association has not been assigned. The designation of the typus of the Scrophulario variegatae-Epilobietum dodonaei was attempted by onipchenko (2002b) who selected (releve) '26/95, Table 3.3' in onipchenko (2002a) as the typus. This attempt was, however, invalid since there are two releves designated as 26/95: 0 26/95 and 1 26/95 in Table 3.3. Because the choice of the typus was not unequivocal, this validation cannot be recognised, hence both the Scrophulario variegatae-Epilobietum dodonaei and the murbeckiellion huetii remain invalidly described.
The murbeckiellion huetii is a syntaxonomi-cally heterogeneous concept. The Hyalopoo pon-ticae-Oxyrietum digynae onipchenko 2002 is flo-ristically (and ecologically) closer to the Hya-lopoion ponticae Rabotnova et onipchenko in onipchenko 2002 than any syntaxonomic unit of the Lamio tomentosi-Chaerophylletea humilis. The Scrophulario variegatae-Epilobietum dodonaei, on the other hand should be classified with the Epi-lobietaliafleischeri; it is characteristic of gravel alluvial banks of mountain streams and Epilobium dodonei plays a major role in these community (see Fig. 1A-E). It is therefore not our intention to validate the Murbeckiellion huetii, but rather coin a new name that would better represent the vegetation of gravel terraces of Caucasus - the Murbeckiello huetii-Epilobion dodonaei (for the pro-tologue see below).
The low-altitude Murbeckiello huetii-Epilobion dodonaei is classified here into the Epilobietalia fleischeri Moor 1958 nom. cons. propos. (Raus et al. in prep.) of the Thlaspietea rotundifolii.
Lamio tomentosi-Chaerophylletea humilis Belonov-skaya, Mucina et Theurillat classis nova hoc loco
Holotypus: Chaerophylletalia humilis onipchenko 2002 (onipchenko 2002a: 18, 21)
synonym: Veronico telephiifoliae-Lamietea tomento-si Belonovskaya 2012 (Arts. 2b & 8 of the ICPN) Diagnostic taxa: Cerastium multiflorum, Corydalis alpestris, Jacobaea erucifolia subsp. arenaria, Jurinea humilis, Lamium tomentosum, Veronica telephiifolia
Chaerophylletalia humilis Onipchenko 2002
synonym: Veronico telephiifoliae-Lamietalia to-mentosi Belonovskaya 2012 (Arts. 2b & 8 of the ICPN)
Chaerophyllion humilis onipchenko 2002
synonym: Drabo scabrae-Eunomion rotundi-foliae Belonovskaya 2012 (Arts. 5 & 8 of the ICPN)
Diagnostic taxa: Cerastium alpinum, Chae-rophyllum humile, Delphinium caucasicum, Draba scabra, Eunomia rotundifolia, Poten-tilla gelida, Saxifraga moschata, Tephroseris integrifolia, Veronica minuta
Veronico minutae-Chaerophylletum humilis onipchenko 2002 Cruciato tauricae-Chaerophylletum humilis Belonovskaya 2012 Saxijrago sibirica-Alopecuretum sericei Be-
lonovskaya 2012 Myosotido alpestris-Potentilletum gelidae Belonovskaya 2012
Scrophulario minimae-Symphyolomion gra-veolentis Belonovskaya, mucina et theurillat all. nova hoc loco
Holotypus: Cruciato tauricae-Pseudovesica-rietum digitatae Belonovskaya 2012 Diagnostic taxa: Cruciata taurica, Scrophu-laria minima, Symphyoloma graveolens
Cruciato tauricae-Pseudovesicarietum digitatae Belonovskaya 2012 Ranunculetum arachnoidei Belonovskaya 2012
Anthemidetum creticae Belonovskaya 2012 Cerastietum alpini Belonovskaya 2012
Thlaspietea rotundifolii Br.-Bl. 1948
Epilobietaliafleischeri Moor 1958 nom. cons. propos.
Murbeckiello huetii-Epilobion dodonaei Be-lonovskaya, Mucina et Theurillat all. nova hoc loco
Holotypus: Scrophulario variegatae-Epilo-bietum dodonaei Onipchenko ex Belono-vskaya, Mucina et Theurillat ass. nova hoc loco; Holotypus:: onipchenko 2002a: Table 3.3, releve 1 10 94
Synonym: Murbeckiellion huetii Onipchenko 2002 nom. inval. (Arts. 5 & 8 of the ICPN)
Diagnostic species: Epilobium dodonaei, Murbeckiella huetii
5. ACKNOWLEDGEMENTS
We thank Thomas Raus (Berlin, Germany) for checking the nomenclature and taxonomy of the diagnostic species mentioned in our paper. Marie Lochman (Perth, Australia) kindly scanned slides and Mikhail Dokukin (Nalchik, Russia) for the photos used in Figure 1. Zaza Kikvidze identified the locality in the Tergi River valley. LM thanks Gia Nakhutrishvili (Tbilisi, Georgia) and Georg Grabherr (Vienna, Austria) for making the visit to Caucasus in 1990 possible.
6. REFERENCES
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Received 2.12.2013 Revison received: 6.2.2014 Accepted 6.2.2014