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1 INVASION OF OPUNTIA HUMIFUSA AND O. PHAEACANTHA (CACTACEAE) INTO PLANT COMMUNITIES OF THE KARADAG NATURE RESERVE Valentina V. Fateyga 1, Nataliya A. Bagikova 2 ============== RESEARCH ARTICLES ============= 1 T.I. Vyazemsky Kaadag Scientific Station Natue Reseve of RAS, Russia valentina_vt@mail.u 2 The Ode of the Red Banne of Labou Nikitsky Botanical Gadens National Scientific Cente of RAS, Russia nbagik@mail.u Received: The esults of a study of Opuntia humifusa and O. phaeacantha natualised in the Kaadag Natue Reseve (southeasten pat of the Cimean Peninsula) ae pesented. Thee, the lagest coenopopulations of Opuntia plants ae confined to the «biostation» teitoy (bodeing with the pak, administative buildings and housing estate). Twelve localities wee descibed in the Kaadag Reseve. These diffe by phytocoenotic chaacteistics, aea and floistic composition. Seven localities include only O. humifusa plants; fou ones include only O. phaeacantha individuals; and both the species ae pesent on the twelfth locality. The total numbe of individuals of each species and ontogenetic stuctue of the population wee studied in each locality. The total numbe of O. humifusa individuals in the Kaadag Reseve is moe than 600 plants within the «biostation» teitoy, while the total numbe of O. phaeacantha plants is about 400 individuals. Studying of the plant communities has been caied out accoding to the Baun-Blanquet method. Opuntia plants fom deivate communities within degaded steppes, phyganoid-steppes, and semi-deset badland phytocoenoses almost at all studied localities. A significant numbe of synanthopic species (including alien plants) was found within these communities. Opuntia plants ae able to self-epoduce pedominantly vegetatively. Self-seeding epoduction occus less fequently. Both species can be consideed as invasive plants because they have a high adaptive capacity. Key wods: alien plants, Cimea, Kaadag Natue Reseve, Opuntia, phytoinvasion, plant communities Intoduction Recently, the invasion of alien species has become one of the most significant poblems and is consideed as a theat to biodivesity of natual ecosystems (Vinogadova et al., 2010). Such invasions ae also known in biodivesity hotspots Potected Aeas. One of them is the Kaadag Natue Reseve. It is located on the Black Sea coast on the southeasten pat of the Cimean Peninsula. The Reseve was founded in Besides natual landscapes it houses the aea of the Kaadag biological station (founded in 1914). It includes administativeeconomic buildings, a pak and housing stock. The Kaadag Natue Reseve was ascibed to the highest pioity categoy on Cimean biodivesity conseving (Biodivesity Suppot Pogam, 1999). The change of climatic epochs, mountainbuilding pocesses and physical-geogaphical featues of the Kaadag Natue Reseve have contibuted to the foming of both its phytocoenotical and floistic divesity. The vegetation of the Kaadag Reseve is divese. It foms two indistinctly expessed belts: 1) the lowe belt pesented by pubescent oak (Quecus pubescens Willd.) foests and steppes; and 2) the uppe belt pesented by dumast 26 oak (Quecus petaea (Matt.) Liebl.) foests, honbeam (Capinus oientalis Mill.) foests and ash (Faxinus excelsio L.) foests. In geneal, foests cove about 50% of the Kaadag Reseve s aea, while steppes cove about 25% of it. Phyganas (7.5% of the total aea) and savannoid communities (about 2% of the total aea) occu on sepaate spots amongst the massifs of foests and steppes. In addition, halophytic communities ae distibuted along the sea coast (Didukh & Shelyag-Sosonko, 1982). Within the aea of the Kaadag Reseve, 1165 vascula plant species, including 31 endemics, ae known (Mionova & Fateyga, 2015) fom the in total 2536 species of the Cimean floa (Yena, 2012), including 106 endemics. Despite the obvious consevation value of the Kaadag Natue Reseve, intensive economic activity has been caied out at almost all its aea until This fact, in tun, has contibuted to the penetation of alien plant species thee, as well as to thei spead along the natual landscapes of othe pats of the Kaadag Reseve. Accoding to ecent data, about 50 alien plant species ae egisteed within the Kaadag Natue Reseve (Kamenskikh & Potapenko, 2012; Mionova & Fateyga, 2015), o

2 moe than 90 alien plant species including acheophytes (Bagikova, 2013a). Many of the aliens ae capable to self-epoduction unde the conditions of the Kaadag Natue Reseve. Some of these plants have accidentally been penetated into the Reseve. Othes have been cultued puposefully. Species of the genus Opuntia Mill. ae classified within the second goup. They fom the object of ou pesent study. Opuntia Mill. is one of the ichest genea in the family Cactaceae, with 200 to 250 species in consistent with diffeent authos (Bitton & Rose, 1919; Andeson, 2001; Rebman & Pinkava, 2001). Unde natual conditions epesentatives of the genus ae known fom the Ameican continent (Mah, 2001; Nyffele, 2002; Bulot, 2007; Majue & Evin, 2008). Due to numeous xeomophic adaptations most species ae confined to aid habitats of subtopical and topical natual belts. Mexico is a cente of the genus divesity (Mah, 2001; Rebman & Pinkava, 2001; Casas & Babea, 2002; Majue & Evin, 2008). Some Opuntia species ae able to withstand tempeatues below feezing duing the winte peiod (Loik & Nobel, 1993; Nobel & Bobich, 2002). It allows them to penetate into the colde egions. Plants may be found thee mainly unde the tees canopy o on the slopes with a southen exposue o fom thei leewad side (Loik & Nobel, 1993). Thus, Opuntia plants ae able to exist unde a wide ange of tempeatue conditions and in diffeent plant communities: fom desets to topical foests and alpine meadows (Mauseth, 2006). Opuntia species have long been cultivated in some counties due to thei taste and decoative qualities (e.g., Mah, 2001; Casas & Babea, 2002; Majue, 2007; Ee et al., 2009). They wee fistly intoduced fom Mexico to Spain, and then to othe Euopean counties (Ee et al., 2009). The modeate Mediteanean climate poved to be vey favouable fo thei cultivation, distibution and natualisation (Babea et al., 1992; Casas & Babea, 2002; Ee et al., 2009; Tashev, 2012). Then Opuntia species penetated into the counties of Asia and Afica, and into Austalia (Mah, 2001). To date, of the 27 invasive Opuntia species, the lagest numbe of natualised ones has been ecoded in Spain, South Afica and Austalia (Novoa et al., 2014). Thee ae cases whee some Opuntia species caused significant economic damage to entie continents. The classic example is an invasion of Opuntia sticta (Haw.) Haw. and O. vulgais Mill. into Austalia (Fawley, 2007). Fo the fist time Opuntia penetated thee in 1787, togethe with a Bazilian settle. And it gadually spead ove a wide teitoy 27 of pastues and famlands. By the ealy 20th centuy the aea occupied by Opuntia was seveal million hectaes (Fawley, 2007). An ecological catastophe was pevented only in 1925, when Cactoblastis cactoum (Beg, 1885) was bought in Austalia in ode to contol the Opuntia invasion (Zimmeman et al., 2000; Fawley, 2007; Pembeton & Liu, 2007). This moth has become a biological baie fo the futhe spead of Opuntia plants. The invasion of Opuntia ficus-indica (L.) Mill. in South Afica in the 1990s has led to a significant agicultual cisis in this egion (Dean & Milton, 2000). Most likely, the histoy of Opuntia s intoduction in the Cimea began in the 19th centuy. In most cases, it was associated with the Nikitsky Botanical Gaden. Although, thee is an opinion that the intoduction of Opuntia plants was induced by Geman colonists in the 18th centuy (Byalt et al., 2009). To date, Opuntia species have distibuted ove a significant teitoy of the southen coast of Cimea fom Sevastopol to Koktebel and in some aeas of the Cimea steppe (Bagikova & Ryff, 2014b). Accoding to the latest published data (Bagikova & Ryff, 2014b), eight Opuntia species ae consideed as self-enewing plants on the Cimean Peninsula. Fom the published memois of Ye.A. Sludsky (son of the fist diecto of the Kaadag biological station) it has been found out that Opuntia appeaed in Kaadag at the beginning of the 19th centuy. They wee planted by A.I. Bachinsky, who «walked though Kaadag in foests, valleys and mountains, and in some places he planted Opuntia cacti to acclimatise them in the wild» (Sludsky, , p. 69). Late V.N. Wuczeticz planted them also nea the biological station. Accoding to some data (Voinov, 1930, 1968; Anisimova, 1939), thee wee attempts to intoduce thee Opuntia species. Of them, to date, we identified plants of two species, Opuntia humifusa (Raf.) Raf. (Fig. 1B) and the ed-floweed fom of O. phaeacantha Engelm. (Fig. 1D). Both species gow in the wild in the Kaadag Natue Reseve, including the biostation teitoy. It should be noted that many Opuntia species belong to taxonomically difficult goups. Even within the native ange, eseaches (e.g., Majue, 2007; Majue, Evin, 2008; Majue et al., 2012) often distinguish many vaieties within a single species. Many specialists indicate on objective poblems associated with accuate identification of alien species within thei seconday ange. Among them is Yu.K. Vinogadova with co-authos (2010) who emphasise that most infomation about oigin, type of ange, ecological popeties, etc. is associated with the taxon name.

3 And alien species ae often epesented by atypical foms within thei seconday ange. In addition, infomation about the hebaium specimens collected in natual habitats is hadly available. Fig. 1. Opuntia humifusa (A, B, E, G) and O. phaeacantha (C, D, F, H) in the Kaadag Natue Reseve. A, C communities with paticipation of each Opuntia species (A locality 9, C locality 2); B, D floweing plants; E, F fuits; G, H juvenile plants. 28

4 Pobably, difficulties in identification of Opuntia species have led to a situation whee intoduced and natualised species in the Cimea wee listed unde diffeent names in numeous publications. Fo example, the intaspecific taxonomic position has not been finally established fo the ed-floweed fom of O. phaeacantha, which is cultivated in the vicinity of Koktebel and in Nikitsky Botanical Gaden. But it is known in the wild in the Kaadag Reseve. This is caused by the fact that it is most likely a cultiva, diffeent fom the ancestal taxon (o taxa) by one o moe taits. These plants could be witten out accoding to Westen Euopean catalogues of the ealy 20th centuy (whee they appeaed) as a vaiety of Opuntia camanchica Engelm. & J.M. Bigelow va. uba (see Bitton & Rose, 1919, p. 144). This Opuntia species has been noted unde the mentioned name in many publications of Cimean eseaches (Anisimova, 1939; Kamenskikh & Mionova, 2004; Emisaliyev & Skopintseva, 2008; Kamenskikh & Potapenko, 2012). Plants gowing in Kaadag and Koktebel have been classified as a «f. uba» within O. phaeacantha accoding to the monogaph «Tees and Shubs of USSR» (Zamyatnin, 1958). They have also been consideed within O. phaeacantha va. camanchica (Engelm. & J.M. Bigelow) L.D. Benson, accoding to some web-souces: «Plantaium» ( «All Cimean Plants» ( u/). Othe specialists have consideed this Opuntia species without the epithet «uba»: eithe as an independent species O. camanchica (Voinov, 1930; Mionova & Kamenskikh, 1995; Byalt et al., 2009), o as a vaiety, O. phaeacantha va. camanchica (Voinov, 1968; Vasilyeva & Seov, 2009; Bagikova, 2013a,b), o aely as O. phaeacantha without indicating its infaspecific status (Byalt, 2004, 2012; Yena, 2012). The oigin and authoship of the epithet «uba» ae not clea; it is absent in the moden taxonomic databases such ae «IPNI» ( and «Topicos» ( The diagnosis and location of the type fo this taxon ae not known. Theefoe, futhe nomenclatual, mophological and molecula-genetic studies ae needed in ode to detemine moe accuate the status of the ed-floweed fom of O. phaeacantha. This applies to some othe Opuntia species natualised in the Cimea as well. We have focused on the Opuntia invasion in the Kaadag Natue Reseve because detailed investigations of this poblem have not been caied out peviously. The aim of the pesent study is to establish the invasion degee of two Opuntia species in diffeent plant communities within the Kaadag Natue Reseve. 29 Mateial and Methods In ode to estimate the distibution extent of Opuntia species within the Kaadag Natue Reseve, we used ou own data fo 2014, as well as the esults of fagmentay eseaches conducted in pevious yeas. Population studies of two Opuntia species and geobotanical suveys of plant communities with thei paticipation wee caied out on 12 localities in Octobe Novembe The localities wee situated within the teitoy of the biological station and the aeas of the Kaadag Reseve adjacent to it (Fig. 2). In most cases, a studied plot had an aea of the whole Opuntia locality. Only a geobotanical suvey in locality 12 was caied out on five plots with an aea of 25 m 2. In ode to descibe the plant communities, we investigated the geneal coveage (visually), the floistic composition and the species abundance. We estimated the abundance of each species in accoding to the 7-point scale of Baun-Blanquet (Baun- Blanquet, 1964; Mikin et al., 1989). Due to a lack of classification fo the Kaadag vegetation accoding to the ecological-floistic appoach, we have caied out the identification of plant communities with Opuntia species in accodance with the existing syntaxonomy of the vegetation of Cimea (Kozhenevsky et al., 2003; Bagikova, 2016) and Euope (Mucina, 1997; Rodwell et al., 2002; Mucina et al., 2016). Syntaxa names ae given accoding to cuent equiements of phytosociological nomenclatue (Webe et al., 2000). In ode to estimate the ontogenetic stuctue of populations, we distinguished thee goups: a) geneative individuals; b) viginile individuals, appeaing fom ooting of vegetative pats of paent plants; c) individuals of seed oigin (juvenile and immatue plants). Each individual o goups was consideed as one counting unit. In ode to detemine the total numbe of O. humifusa and O. phaeacantha individuals, we used method of total counting. We noted the pesence / absence of Opuntia seedlings in each locality without taking into account thei numbe. The identification of Opuntia species has been caied out accoding to both classical and moden monogaphs on taxonomy of the family Cactaceae (Bitton & Rose, 1919; Andeson, 2001), as well as on the base of special web-souces ( Species names ae povided accoding to «The Plant List» ( theplantlist.og/). Names fo othe plant species of spontaneous Cimean floa ae povided accoding to Yena (2012).

5 Fig. 2. Situation of the localities with Opuntia humifusa and O. phaeacantha within the «biostation» teitoy in the Kaadag Reseve (explanations ae in Table 1; backgound maps wee taken fom Collected hebaium specimens wee deposited into the Hebaium of the Nikitsky Botanical Gaden (YALT). Live plants ae peseved in the Succulents collection of the Nikitsky Botanical Gaden. Photogaphs wee made using the digital cameas Sony DSC-H1, Sony DSC- HX200 and Canon PoweShot A570 IS. Results Only single individuals of both Opuntia species wee found in the least distubed habitats. These ae 1) along the ecological path of the Kaadag Reseve and 2) on the coastal 30 clayey-gavelly slopes of the Beegovoy idge adjacent to it. The lagest coenopopulations in the Kaadag Natue Reseve ae situated within the teitoy of the biological station whee 12 localities with Opuntia plants wee studied. These ae diffeent on the base of phytocoenotic chaacteistics, aeas (Table 1) and floistic composition (Table 2). Opuntia humifusa was found in seven localities, O. phaeacantha was found in fou. In one locality, both species wee egisteed. Opuntia phaeacantha was noted in the localities 1, 2, 4, 5, 12, O. humifusa in the localities 3, 6 12 (Table 1 and 2).

6 Table 1. Geneal chaacteistics of studied localities in the Kaadag Natue Reseve Locality Gowing place Co-odinates; altitude Steepness (degees) / exposue of the slope Aea, Individuals numbe m2 pe locality, units Species 1 Below the house of the T.I. Vyazemsky s sanatoium complex 2 Above hostel 3 Above hostel 4 Between the oad and the idge Beshtash 5 Unde the idge Beshtash 6 On the oadside nea the boile house 7 Nea the shop To south of the laboatoy building To south of the laboatoy building Between the laboatoy building and the meteoological station Below the meteoological station Seaside slopes above the beach 44º N 35º E; 50 m a.s.l. 44º N 35º E; 39 m a.s.l. 44º N 35º E; 37 m a.s.l. 44º N 35º E; 35 m a.s.l. 44º N 35º E; 32 m a.s.l. 44º N 35º E; 22 m a.s.l. 44º N 35º E; 18 m a.s.l. 44º N 35º E; 11 m a.s.l. 44º N 35º E; 12 m a.s.l. 44º N 35º E; 14 m a.s.l. 44º N 35º E; 16 m a.s.l. 44º N 35º E; m a.s.l / SSE O. phaeacantha / SE O. phaeacantha / SE O. humifusa 20 / W 10 2 O. phaeacantha / SE O. phaeacantha 5 / W 10 4 O. humifusa / SE O. humifusa 30 / SSW O. humifusa / SSW O. humifusa / SSW O. humifusa 5 10 / S O. humifusa / SSE ; 116 O. humifusa; O. phaeacantha Locality 1 is situated below the house of the T.I. Vyazemsky s sanatoium complex, on the gently sloping plot in tansfomed pistachio spase foest with todden heb laye. Within the phytocoenosis, we found species (Table 2), which diagnose xeothemic and semi-xeothemic hebaceous communities of the class Festuco-Bometea B.-Bl. & Tx. ex Soó Paticipation of synanthopic species and Atemisia tauica Willd. indicate the violation and tansfomation degee of the community in both this and othe localities. Within this locality, thee wee 59 geneative, 160 vegetative, and no less than 40 juvenile and immatue individuals of O. phaeacantha. Young plants wee found at a distance of m fom the 31 main goups. We egisteed an abundant fuiting. At the time of obsevation, a pat of the fuits has fallen off, but fom 2 to 5 fuits have still emained on most segments (Fig. 1F). Locality 2 is situated above the hostel of the biological station, on a slope with a degaded steppe vegetation neaby locality 1 and delimited fom it by a dit oad (Fig. 1C). Within this locality, thee ae eight geneative, 17 viginile (non-seeded) and at least six juvenile and immatue individuals of O. phaeacantha. Young (seed oigin) plants and single 2 4-segment vegetative individuals wee found among stones below the main goup (at least 10 m in diamete), as well as unde the canopy of tees and shubs on a faily steep slope.

7 Table 2. Floistic composition and coveage of species accoding to Baun-Blanquet s scale in plant communities with Opuntia species in the Kaadag Natue Reseve Designation of locality a 12b 12c 12d 12e Geneal coveage, % Total numbe of species pe locality Achillea nobilis L Agopyon cistatum (L.) Gaetn. subsp. pectinatum (M. Bieb.) Tzvelev *Ailanthus altissima (Mill.) Swingle Alcea tauica Iljin Alyssum totuosum Waldst. & Kit. ex Willd *Amaanthus etoflexus L. Anisantha tectoum (L.) Nevski + Atemisia lechiana Stechm Atemisia tauica Willd Aspaagus veticillatus L. + Aspeula tenella Degen Atiplex auchei Moq. Atiplex micantha C.A. Mey Atiplex oblongifolia Waldst. & Kit. 1 + Bassia postata (L.) Beck + Bassia sedoides (Pall.) Asch. + Beta tigyna Waldst. & Kit. + Bothiochloa ischaemum (L.) Keng + Bupleuum bachiatum K. Koch Bupleuum exaltatum M. Bieb. + Camphoosma monspeliaca L Cappais hebacea Willd Centauea capina Steven + *Centauea diffusa Lam. Centauea salonitana Vis. Centauea steilis Steven + + Chenopodium album L. + + Convolvulus cantabica L. + + Cocus angustifolius Weston + Cynanchum acutum L. 1 Dactylis glomeata L Dasypyum villosum (L.) P. Candagy + *Descuainia sophia (L.) Webb. ex Pantl + Dianthus maschallii Schischk. + Diplotaxis tenuifolia (L.) DC. Echium italicum L. subsp. biebesteinii (Lacaita) Geute & Budet Elytigia caespitosa (K. Koch) Nevski subsp. nodosa (Nevski) Tzvelev Elytigia obtusifloa (DC.) Tzvelev + 4 Elytigia epens (L.) Nevski + Epheda distachya L Eodium cicutaium (L.) L He. + Eysimum cuspidatum (M. Bieb.) DC. + Euphobia mysinites L Falcaia vulgais Benh. Festuca valesiaca Gaudin Galatella linosyis (L.) Rchb. f. + + Galatella villosa (L.) Rchb. f Galium humifusum M. Bieb. + Galium xeoticum (Klokov) Pobed. + Hedea helix L. + Heliotopium ellipticum Ledeb. Heniaia bessei Fisch. ex Honem. + Jasminum futicans L. + Lactuca viminea (L.) J. Pesl & C. Pesl + + Lappula sp. + Lepidium daba L. + Limonium platyphyllum Lincz. + Limonium scopaium (Willd.) Stank Linaia genistifolia (L.) Mill. + + Medicago cetacea M. Bieb. 1 + Medicago falcata L Medicago igidula (L.) All

8 Designation of locality a 12b 12c 12d 12e Geneal coveage, % Total numbe of species pe locality Melica tanssilvanica Schu *Opuntia humifusa (Raf.) Raf *Opuntia phaeacantha Engelm Onithogalum pyenaicum L. + Otites densifloa (d Uv.) Gossh. + Phlomis heba-venti L. subsp. pungens (Willd.) Maie ex De Filipps + *Pinus butia Ten. va. pityusa (Steven) Silba 1 Pistacia mutica Fisch. & C.A. Mey. 3 Plantago lanceolata L *Platycladus oientalis (L.) Fanco 4 *Potulaca oleacea L. Potentilla sp. *Punus ceasifea Ehh. Pyus elaeagifolia Pall. + Rapistum ugosum (L.) All. Rosa sp. Salvia nemoosa L. subsp. pseudosylvestis (Stapf) Bonm. + Scozonea mollis M. Bieb. Secuigea vaia (L.) Lassen + Seseli totuosum L ?Solanum alatum Moench Stachys cetica L. subsp. velata (Klokov) Geute & Budet Stipa capillata L. 1 Taaxacum eythospemum Besse Teucium chamaedys L. 1 Teucium polium L Vicia anatolica Tuill Zygophyllum fabago L. + + Note: * alien species of the Kaadag Natue Reseve;? oigin (native o alien) is uncetain; species is extemely ae, coveage is <1%; + species ae, coveage is 1 2%; 1 individuals ae numeous, coveage is 2 5%; 2 numbe of individuals is lage, coveage is 5 25%; 3 with any numbe of individuals, coveage is 25 50%; 4 with any numbe of individuals, coveage is 50 75%; 5 with any numbe of individuals, coveage is moe than 75%. 33 Locality 3 is situated below locality 2, on a steep slope. It is chaacteised by a stongly damaged vegetation cove epesented by plant species (Table 2), which diagnose synanthopic plant communities belonging to the classes Atemisietea vulgais Lohmeye et al. ex von Rochow 1951 and Stellaietea mediae Tx. et al. in Tx. ex von Rochow 1951 (accoding to Bagikova, 2016) o Chenopodietea B.-Bl. in B.-Bl. et al (accoding to Mucina et al., 2016). Opuntia humifusa is epesented hee by fou lage individuals, which each up to 5.5 m in diamete. In addition, we found six pe-geneative plants. Of them, fou individuals had a vegetative (fomed fom boken segments) oigin and two plants had a seed oigin. The Opuntia individuals ae chaacteised by an abundant fuiting (in aveage of 3 5 fuits pe segment) (Fig. 1E). Howeve, seedlings o juvenile plants wee not found. The pimay vegetation in localities 1 3 was pesented by feathe-fescue steppe communities belonging to the class Festuco-Bometea. They develop on heavy clays in conditions of the Kaadag Natue Reseve. Low-gowing annual and peennial hebaceous plants pedominate in these phytocoenoses. The pesence of Atiplex oblongifolia, Limonium scopaium, Epheda distachya and species of the genus Bassia All. indicates the slight soil salinity. The tansfomation degee of these phytocoenoses is indicated by synanthopic species and also by Atemisia tauica which was egisteed with high constancy and abundance. Locality 4 is situated on a wateshed slope, between the oad and the idge Beshtash. Opuntia phaeacantha occus hee in udeal-steppe communities typical fo badlands o semi-deset landscapes close to them. Pehaps, this phytocoenosis can be classified as the association Ataphaco-Cappaidetum Kozhenevsky & Klyukin 1988 of the alliance Ataphaco-Cappaidion Kozhenevsky 1988, descibed fom Cimea (Kozhenevsky et al., 2003) within the Mediteanean class Pegano hamalae- Salsoletea vemiculatae B.-Bl. & O. de Bolòs 1958 (Mucina, 1997; Rodwell et al., 2002; Mucina et al., 2016), although it has some similaities with plant communities of the Aal-Caspian class Atemisietea lechianae Golub 1994 (Golub, 1994) too. This locality diffes by both the smallest aea and abundance of Opuntia plants. Only two O. phaeacantha individuals wee found hee. One plant is geneative (with single fuits), anothe one is vegetative.

9 Locality 5 is situated neaby locality 4, on a deset-steppe wateshed flysch slope below the idge Beshtash. This phytocoenosis is less distubed than the fist fou habitats abovementioned. It epesents a basal goup, because it contains fagments of seveal classes in accodance with the classification of Cimean vegetation (Kozhenevsky et al., 2003). Thee ae petophytic-steppe (Festuco-Bometea) and badland (Pegano hamalae-salsoletea vemiculatae) classes with paticipation of sepaate elements of shale scees (Thlaspietea otundifolii B.-Bl. 1948). Nineteen geneative O. phaeacantha plants wee found hee. Abundant fuiting (2 5 fuits pe segment) was egisteed. Locality 6 is situated at the oadside. This phytocoenosis is an anthopogenic deivative of the steppe community on heavy saline clays. Two geneative and two vegetative O. humifusa plants wee found on this locality. Fuiting (1 3 fuits pe segment) was egisteed. Locality 7 is situated on a flowebed nea the shop. Theefoe, the pimay vegetation (steppe with elements of semi-deset vegetation of badlands) is highly anthopogenically tansfomed hee. Opuntia humifusa had peviously been planted hee. And in time it spead unde the cowns of Platycladus oientalis plantings. Fifteen geneative and ten vegetative Opuntia plants wee found on this locality. Individuals of seed oigin and seedlings wee not egisteed. Locality 8 is situated to south and east of the laboatoy building, on the seaside on a gavelly-clayey denudation slope. It is chaacteised by a significant paticipation of semi-shubs and gamineous plants in the heb laye composition. Apat fom O. humifusa, anothe alien species, Ailanthus altissima, was found hee. Locality 9 is situated to south of the laboatoy building, on a steep gavelly-stony denudation slope. It is chaacteised by a high degee of anthopogenic tansfomation. Ailanthus altissima thickets occupy the uppe pat of the slope. Its lowe pat is almost completely coveed by O. humifusa (Fig. 1A). The floistic compositions of the plant communities in the localities 8 9 ae simila to the ones in the phytocoenoses of the seaside cliffs belonging to the ode Onosmo polyphyllae-ptilostemonetalia Kozhenevsky 1990 (Mucina, 1997; Rodwell et al., 2002; Mucina et al., 2016), which was descibed in Cimea. It is typical fo the foothills and low-hill teains of Southeasten Cimea and some points in the Southen coast. It contains open communities of xeophytic dwaf semishubs 34 on slopes, composed by vaious ock fomations with weak and modeate anti-denudation esistance (Kozhenevsky et al., 2003). The plant community in locality 9 is moe anthopogenically tansfomed than the one in locality 8. Locality 10 is situated west of locality 9, in the zone of a seaside cliff. The phytocoenosis descibed hee is an ecotone o deivative plant community. It is close to badlands phytocoenoses. In accodance with classifications of the vegetation of Cimea (Kozhenevsky et al., 2003) and Euope (Mucina, 1997; Rodwell et al., 2002; Mucina et al., 2016), this phytocoenosis belongs to the Mediteanean halo-nitophilic semi-deset shub vegetation of the class Pegano hamalae-salsoletea vemiculatae, although it has elements of plant communities belonging to the ode Onosmo polyphyllae- Ptilostemonetalia, class Festuco-Bometea, etc. Locality 11 is situated on a elatively flat plot. Within its boundaies we distinguished the badland community with deivatives of steppe vegetation located on heavy saline clay soils. Its floistic composition is stongly diffeent to the ones in othe studied plant communities (Table 2). In total, we egisteed 198 geneative and 51 vegetative (13 plants have seed oigin, 38 ones ae a esult of vegetative epoduction) O. humifusa individuals within the localities The plants ae well-developed, with abundant fuiting. We egisteed seedlings and juvenile individuals. Locality 12 has the lagest aea (about 3200 m 2 ). It is situated on a denudation slope with a steppefied pistachio spase foest and plantations of Pinus butia va. pityusa. The vetical stuctue of the vegetation is pluistatal, the hoizontal one is mosaic. On the base of floistic composition, plant communities descibed hee with both Opuntia species may be classified as basal o deivative goups because these occupy ecotones and contain a complex of species fom diffeent Mediteanean types of vegetation. Some of them ae phytocoenoses close to plant communities of the alliance Atemision aboescentis Géhu & Biondi in Géhu et al (Rodwell et al., 2002) o Ataphaco-Cappaidion (Kozhenevsky et al., 2003) fom the class Pegano hamalae-salsoletea vemiculatae, which include semi-shub semi-deset halo-nitophilic goups on clay soils in the Westen Mediteanean (Rodwell et al., 2002) and in Cimea (Kozhenevsky et al., 2003). Futhemoe, we found hee species fom plant communities of the alliance Ptilostemonion Kozhenevsky 1990, descibed in South-Easten Cimea. These communities ae typical fo denu-

10 dation slopes on cabonate-fee ocks (included in ode Onosmo polyphyllae-ptilostemonetalia Kozhenevsky 1990). In addition, we egisteed species of Mediteanean ephemee vegetation types (class Theo-Bachypodietea B.-Bl. ex A. de Bolòs y Vayeda 1950), petophytic steppes (class Festuco- Bometea), as well as synanthopic and alien species (Table 2). We would like to list species which wee found outside the boundaies of the established plots. These plants wee egisteed with low abundance (, aely +), but they chaacteise the descibed biotopes. These ae Agopyon cistatum subsp. pectinatum, Ailanthus altissima, Aspaagus veticillatus, Cappais hebacea, Centauea salonitana, Dianthus maschallii, Lactuca viminea, Rapistum ugosum, Rosa sp., Salvia nemoosa subsp. pseudosylvestis, Scabiosa micantha, Teucium chamaedys, as well as Aegilops ovata L., Camelina sp., Caduus pycnocephalus L. subsp. albidus (M. Bieb.) Kazmi, Lens nigicans (M. Bieb.) God., Lycopsis sp., Papave sp., Scabiosa micantha Desf., Senecio leucanthemifolius Poi. subsp. venalis (Waldst. & Kit.) Geute and Valeianella coonata (L.) DC. In total, within locality 12 we egisteed 229 geneative and 70 vegetative O. humifusa plants, as well as 72 geneative and 44 vegetative O. phaeacantha individuals. The lagest numbe of seedlings and juvenile plants of both Opuntia species wee found hee (Fig. 1G, H). Thus, Opuntia humifusa was noted fom diffeent ecotopes. This species pefes plots with spase and low hebage in semi-natual and anthopogenically damaged communities. The plants occu in compact goups which makes it often difficult o even impossible to distinguish the individuals within such a goup. Theefoe, cutains of plants often have not clea boundaies. O. humifusa popagates pedominantly in the vegetative manne (by ooting of vegetative pats of plants). Seed epoduction occus elatively aely despite of abundant fuiting (in aveage, 4 7 fuits pe segment). The total abundance of O. humifusa in all studied localities is moe than 600 individuals. Opuntia phaeacantha also gows in diffeent ecotopes. In many of them, it becomes a dominant species. Opuntia phaeacantha popagates pedominantly in the vegetative manne (by ooting of vegetative pats of plants). The plants abundantly bloom and fuit (in aveage, 3 6 fuits pe segment ae obseved). Individuals of seed oigin occu moe common than in O. humifusa populations. The total abundance of this species in all studied localities is about 400 individuals. 35 Discussion Opuntia humifusa plants wee found in the Kaadag Natue Reseve in the belt of pistachio and junipe spase foests at an altitude of m above sea level, while O. phaeacantha plants wee found thee at an altitude of m above sea level. Thus, both studied Opuntia species inhabit the seacoast zone, pedominantly on slopes of middle steepness of the south-east and south-west exposue, occupying the lowe altitudinal belt of vegetation. This fact distinguishes the studied Opuntia populations of those known in moe southen aeas of the Mediteanean, whee these ae able to penetate to altitudes up to 800 m (Ee et al., 2009). Opuntia humifusa is the most common species on the Cimean Peninsula. It pefes well-aeated skeletal vaiants of dy clayey and loamy cabonate and cabonate-fee soils. Opuntia humifusa often foms deivative goups in diffeent vegetation classes, which include communities of stony outcops (Koeleio-Coynephoetea canescentis Klika in Klika & Novák, 1941), spase foests (Quecetea pubescentis Doing-Kaft ex Scamoni & Passage 1959) and steppe phytocoenoses (Festuco-Bometea and Theo-Bachypodietea), as well as phyganoid communities floistically ae close to phytocoenoses of the ode Onosmo polyphyllae-ptilostemonetalia (Bagikova & Ryff, 2014a). In almost all studied localities in the Kaadag Natue Reseve, Opuntia plants fom deivational goups in degaded steppe, fyganoid-steppe and semi-deset badlands communities on heavy, slightly saline, gavelly-clay bown soils. Apat fom Opuntia species, the main dominant species hee is Atemisia tauica, in combination with vaious synanthopic plants, belonging pedominantly to the family Chenopodiaceae (Chenopodium alba, Atiplex micantha, A. auchei, A. tataica). The studied plant communities have a high invasibility, i.e. susceptibility to invasion of alien species (Mikin & Naumova, 2014). These contain at least ten alien plant species (see Table 2). It is about 11% of the total numbe of alien plants in the Kaadag Natue Reseve. Amongst these, apat fom Opuntia species, Ailanthus altissima is consideed as an invasive species. Ou data ae in accodance with esults obtained fo O. humifusa populations in Southen Bulgaia (Tashev, 2012) whee this plant inhabits degaded oak spase foest with seconday xeophytic goups in the heb laye. In the Kaadag Natue Reseve, the lagest O. humifusa coenopopulations wee found on the teitoy of the biological station and steep denudation slopes adjacent to it. This species inhabits natually o anthopogenically

11 distubed phyganoid-steppe and badland communities, which ae floistically close to phytocoenoses of the ode Onosmo polyphyllae-ptilostemonetalia and class Pegano hamalae-salsoletea vemiculatae. Single O. humifusa individuals have aleady invaded in almost undamaged communities. They gow in junipe spase foests with slightly degaded hebage on the idge Kaagach (westen pat of the idge Beegovoy) (Fateyga & Fateyga, 2011). Also, O. humifusa plants ae known in pistachio spase foests on seaside gavelly-clay slopes. These phytocoenoses ae floistically close to communities of the class Theo-Bachypodietea and ode Onosmo polyphyllae-ptilostemonetalia. The invasion of the ed-flowe fom of O. phaeacantha in Cimea is known in the Kaadag Natue Reseve only. In this aea, plants fom deivative goups in phytocoenoses of diffeent vegetation classes. The highest vitality of individuals was egisteed on wateshed slopes of flysch low-hill teains in degaded steppe communities of the class Festuco-Bometea in combination with elements of phytocoenoses belonging to the classes Pegano hamalae-salsoletea vemiculatae and Thlaspietea otundifolii. Paticipation of Opuntia species in phytocoenoses of diffeent vegetation classes and tansitional natue of goups demonstate the divesity of ecological, including climatic, conditions of the Kaadag Natue Reseve. Most of the studied phytocoenoses with Opuntia species belong to xeothemic communities, attibuting to Mediteanean vegetation classes. In geneal, they chaacteise the climate of the Kaadag Natue Reseve. So, it is consideed as vey aid, hot, with vey mild wintes; it has the tansitional chaacte fom the sub-mediteanean climate typical fo the westen pat of the southen coast of Cimea to the modeately continental and modeately hot aid climate, typical fo the steppe pat of the Cimean Peninsula (Kostenko et al., 2011). The study of Opuntia species in the Kaadag Reseve shows some featues of thei distibution in diffeent biotopes. We obseved both single plants and goups fomed by individuals of vegetative and geneative oigin. Sometimes, the invasive plants fomed cutains up to seveal metes in diamete. The stuctues of communities and populations status indicate that both species being found in damaged habitats ae invasive plants because these have ovecome geogaphical and biological baies. Paticipation of single Opuntia plants in natual communities indicates that both invasive species ae able to ovecome ecological-coenotic baies. Theefoe, these species 36 will be classified as tansfomes if they will continue thei expansion in futue. To date, the Opuntia populations occupy insignificant aeas in the Kaadag Natue Reseve. That is why they do not cause any significant ecological damage to natual ecosystems. Howeve, we should not undeestimate the potential and oppotunities fo distibution of Opuntia species. It is enough only to look at the histoy of Opuntia dispesal in othe counties and continents. Fo example, Opuntia plants wee fistly bought in South Afica in In the late 18th centuy the plant was successfully enteed into cultue. In 1942, Opuntia plantations have coveed an aea of 9000 km 2 in the Kaoo and the savannas of South Afica. And only in the late 20th centuy, this plant has become a eal agicultual catastophe (Dean & Milton, 2000). In ode to spead ove the Austalian teitoy and to become a eal ecological catastophe thee, Opuntia species took time about 140 yeas (fom 1789 to 1925) (Fawley, 2007). At pesent, O. humifusa is the invasive species in Austalia, South Afica and nine Euopean counties. Opuntia phaeacantha is known fom Austalia and thee counties of Cental Euope (Tashev, 2012; Novoa et al., 2014). The tem «lagphase» means the time fom the appeaance of an alien species to its dispesal in conditions of a seconday ange (Vinogadova & Mayoov, 2015). We suggest that on the Cimean Peninsula the lag-phase fo O. humifusa is most likely 110 yeas, taking into account that the fist eliable data on its self-epoduction dates back to the fist half of the 20th centuy, while this species appeaed in the Cimea in the ealy 19th centuy (Anisimova, 1939). The lag-phase fo O. phaeacantha is most likely even shote, because less than 80 yeas passed fom its intoduction to natualisation in the Kaadag Natue Reseve. In ode to get a bette undestanding of the invasion pocesses of epesentatives of the genus Opuntia into natual communities of the Cimea, complex studies ae needed coveing taxonomic, mophological, floistic, geobotanical and othe appoaches. Due to the fact that collection and hebaisation of cacti is a time-consuming and sometimes painful pocess, many botanists neglect epesentatives of this family duing the pepaation of hebaium collections of the floa of cetain egions (Majue, 2007). Reseaches should pay moe attention to the collection of hebaium mateial of this genus since many Opuntia species belong to taxonomically difficult goups. Hebaium mateial is needed in ode to avoid eos in detemining and, subsequently, misintepetation of the ate and degee of the distibution of cetain taxa.

12 Conclusions Based on the fact that the two studied Opuntia species wee oiginally planted in Kaadag in the ealy 20th centuy and since then thei abundance inceased, they have ovecome the geogaphical, biological and patly ecological baies. It can be stated that they ae invasive species in the Kaadag Natue Reseve, which sometimes demonstates popeties of the plants-tansfomes. This is caused due to thei adaptation to the natual conditions of the seconday ange, the ability to self-epoduce vegetatively (basic) and by seeds. They have a high invasive potential. The invasion degee of vaious cacti into the natual communities of Cimea has not sufficiently been studied yet. To date, it seems elevant to monito the futhe expansion of both natualised and, possibly, new species of the genus Opuntia ove thee. Acknowledgements The authos ae gateful to L.P. Mionova fo assistance in conducting of the eseach, L.E. Ryff fo consultations duing the aticle pepaation, A.V. Fateyga, S.Yu. Kostin and K.L. 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