EMu. HAx)A~

Institute of Landscape Ecology, Czechoslovak Aeadem F of Sciences, Bo~ivojova $5, IS0 00 Praha 3, Czechoslovakia

S e c o n d a r y V e g e t a t i o n Success ion in t h e Be l i ansk~ T a r r y Mts . T h r e e D e c a d e s a f t e r Graz ing

K e y w o r d s

Mountain vegetation, Changes after grazing, Polygono-Tri.setion, Epilobion angusti]olii, Card/uo- Urticion

A b s t r a c t

I:[AD~.~ E. (1990): Secondary vegetation succession in the Beliansk6 Tatry Mrs. three decades after grazing. -- Folia Geobot. Phytotax. , Praha, 25: 349--356. -- Vegetation changes in seminatural plant communities in the Beliansk6 Tarry Mrs. have been studied three decades after grazing or mowing. The changes are slow, they have reached the association level only after about 20 years.

I N T R O D U C T I O N

The establishment of the Tatra National Park (TANAP) has eliminated the impact of two important antropogeneous activities, influencing mountain vegeta- tion: grazing and mowing on the one hand and wood cutting on the other hand, whereas the trampling of vegetatin by tourists has increased in the last years. As we know very little about the reactions of our high mountain vegetation on limestone in formerly grazed and mown areas after several years of protection we have tried to bring some material on this problem from a well studied valley in the Tatra of Bela, the "dolina Siedmieh pramefiov" (the Seven Springs Valley).

Secondary vegetation succession was studied in the same valley by J. ~ARDA et al. (1963). They made i.a. observations on vegetation changes during 5 years in permanent 1 m z plots in the surroundings of the Prote~ chalet in "Urticetum dioicae" and "Alehemilletum Tastoralis". Changes in the intact Urticetum during 5 years showed small, "mainly quantitative changes without any clear trend in succession with the exception of Rubus idaeus, whose abundancy increased during the experiment from 0 to 4 %. In the plot where the original community of Urtica

350 FOLIA GEOBOTA~ICA ET PHYTOTAX01~OMICA, 25, 1990

dioica was total ly destroyed, Carduu~ per~onata dominated after the first year o f observation and disappeared in the following years, whereas in the 4th year Rubu8 idaeus already cevered 25 % of the area. Urtica did not restore its dominance during 4 years of the experiment.

Changes in "Alchemilletum paztoralis" in the intaot permanent plot during 4 years were mainly quantitative with the exception of some synanthropie species, which disappeared during the observation.

Although the observatins of ~ R I ) A et al. were interesting, they did not give clear results concerning vegetation succession in the after-grazing l~eri~d in the limestone area.

METHODS

~MA~DA et al. used the usual method for studying plant succession, i.e. permanent 1 m z plot~. This is a relatively accurate method, but in mountains and especially in terrain where avalanches fall nearly every year, the establishment of permanent plots for a long observation period is difficult or even illusory. Besides, the small area observed by such a method gives only fragmentary information about the whole community. We have therefore applied another method. In the years 1956--58 plant communities in the valley were mapped (Bi~wznvA et I~AI~A& Ms.} and in the subsequent years changes in the communities were observed by taking 4--5 releves in the limits of the previously mapped communities. Thus we were able to observe qualitative and semiqantitative ch~ges, using the ll-grade scale of DO~m~-HADA~ (HADA~ et aL 1969}.

The following colleagues took part in this investigation: in the year 1977 Dr. D. FI~EROV~, Dr. P. C~x~w, CSc., Dr. I. POXOR~/L and Dr. A. pwr~r, in the year 1988 Dr. J. LEP~, CSe., Dr. O. RAUCH, CSc. and Dr. P. K o v ~ CSc. We would like to thank to the Director of the TANAP for enabling us to use the Prete~ chalet in both cases.

RESULTS AND DISCUSSION

Anthropogenous a n d anthropo~oogenous impact on vegetation in the valley of Seven Springs was and partly sti!l is of a varied character. In the year 1920, the whole slope exposed t o the south was almost totaUy deforested. Artificial afforestation had but little success in the first 3 - -4 decades. In 1955 we found on the "Rubanisko" 0.e. the clearing) a typical clearing vegetation with dominating Digitalo-Calamagrostierum arundinaceae SILL. 1933 on ridges and Rubo-Chama- enerietum angustifolii HADA5 et al. 1969 in depressions. Reforestation was difficult part ly because of the oompetition of Chamaenerium resp. Calamagrostis, but mainly because of the destruct'ion of fresh buds of spruoe b y a heavy snowdrift in winter. Not until reforestatin concentrated on areas well protected from wind in the neighbourhood of the remaining woods, did planted trees start to cover the clearing gradually.

In 1977 the Rubo-Chamaenerietum had already disappeared from the depressions. Instead we found either Geo-Dactylidetum H A D ~ 1981 (see Tab. 1. GD) or Sene- cior~i-Rubetum idaei H~DA5 et al. 1969, whereas the Digitalo-Calamagrostietum had hardly changed. Even in 1988 the Digitalo-Calamagrostietum did not show radical changes. In the relevd below the Jelenia skala rock the following species were listed by J. I~pw Calamagrostis arundinacea, Digitalis grandiflora, Chamerion angustifoliura, Galium 8chultesii, Rubus idaeus, Fragaria vesca, Heracleum sphodyi-

HADA~: SECONDARY SUCCESI01~" IN BEZIAI~S]~:]| TJkTB~Y 351

T a b l e I . Success ion c h a n g e s f rom Alchem~llo-F~stucetum ~ra~ensiv in 1956 ( A F 56) to Geo.Dac- tylidetum slovenicae in 1977 (OD 77) a n d Oeo-Dactylidetum in 1988 (GD 88).

A F 65 G D 77 GD 88

Alchemilla monticola OPIz Agrostis tenuis SIBT~ Geum rivale L. Achillea mille/olium L. Veronica chamaedrys L. Taraxacum o//icinale agg. Alopecurus pratensis L. Hyperioum maougatum CR~.TZ Acetosa alpestris (JAcq.) A. L S v ~ Dactylis glomerata L. q- D. slovenica DoMlu~ Banunvulus re,pens L. Urtica dioica L. Myosotis sylvatica HOFYM. Deschampsia cespitosa (Z.) P . BEAuv. 1Festuca rubra L. Cardaminopsi8 halleri (L.) ]~_~YEK Galium schultesii VEs~ Gerani/um phaeum L. Tri fol ium pratense L. Fcst~ca pratensis HUDS. Phleum pratense L. Glechoma hirsuta W ~ D S T . e t Kxw. Poa pratensis L. Vicia sepium L. AegoTodium podagrarla L. Ran~nculus aoris L. Poa trivialis L. Rumex obtusi]olius L. Cerastium holosteoides F ~ s ,~tellaria graminea L. Carum carvi L. Eu,phrasia montana JO~D. Thymus 8 p. Leontodon hispidus L. Plantago media L. Elytrigia repens (L.) Dv, sv . Potentilla aurea (CI~A~TZ) B~CK Prunella vulgaris L. Leontodon autumnalis L. Origanum vulgarr L. Chaerophyllum hirvu~um L. Epilobium alpestre (JACQ.) K~OCKER Carduus personata (L.) J,,,cQ. Lathyrus praten~is L. Luzula luzuloides (LA~.) DANDY et W r ~ M o ~ Pinvpinella major (L.) ttUDS. qeran~um sylvaticum L. Melandrium dioicum (L . )Coss . e t GERm. Cruciata glabra (L.) EH:aEND Senecio /uchsii G ~ L . Senvcio jacquinianus Rc~B.

5.6 4.4 1.6 1.2 4.4 2.6 2.0 0.8 1.4 1.8 1.6 0.8 0.6 0.6 1.0 0.4 0.2 3.6 2.2 7.2 1.2 1.6 1.0 0.4 0.4 2.2 1-8 0.~ 0.4 1.6 1.2 1.6 0.6 0.8 0.6 1.4 0.2 0.4 0.4

2.2 2.6 4.4 1.8 3.2 1.2 1.0 3.4 0.8 6.0 0.2 2.0 0.4 0.2 1.4 1.4 2.6 3.0 0.6 1.0 0.4 0.4 0.8 0.2 0.6

3.0 4.6 2.0 1.4 2.0 LO 1.6 0.8 0.4 0.2 0.6 0.4

0.5 0.5 3.5 0.9 1.4 0.5 0.7 2.2 3.0 6.2 0.5 1.7 2.5 0.5 1.0 0.5 1.5 2.2

0 .5 0.5 0.7 0.5

0.2 2.5 2.2 2.0 2.0 0.5 1.0 1.2 0.7 0.7 0.5 0.5

352 FOLIA GEOBOT&NICA ET PHMTOT&XONOMIC&, 25, 1000

Table 1 (Cont~.)

AF 65 GD77 GD 88

Heracleum sphandyliavm L. A~lelica eylvesrria L. Fil ipendula ulrnaria (L.) MAXIM. ,~cro'phularia se, o,polii H o r n Chaerophyllum aromatieum L. I~mium cupreu~ SCHO~ Alchemilla crinita BusEB Clinopodium vulgare L. Oentiana asclepiadea L. Trisetmm flavescena (L.} P. B ~ u v . B68torta maior S. F. GI~kY Carwpa,~ula Srae.b.e2ium L. Aiuga reptana L. Gallium album M--z.~a Frc~wria veaca L. Calamagrosti, ar~ndinaeea (L.) ROTH Carez m,uricata L. Galeopais tarahit L. ~ h u s idaeus L. Alca~emilla aeutiloba O~iz S~aria nemorum L. Senecio subal1~inua Koc~ Hylotelephimm~ argutum (Haw.} HOlXm Erysimum hun~avioum ZA.~A~. Aconitum firmum (RCl:m.) N:~r;~. Cardamine impa$iens L. Valeriana saanbuei]olia ~ i~ ic . Chamaerion anyusti/olimm (L.) ~[OLU~ Carex aylvatica 1:[UPS. Milium e]/usum L. Poa alpina L. Galeopsis *peciosa MILL.

0.4 0.4 0.4 0.2 1.0 0.2 2.0 2.0 1,4 0.4 0.2 0.4 0.4 0.2 0.2 0.2 0.2 0.4

0.5 0.2 0.2 0.7 0.2 0.2 1.0

1.0 3.0 1.2 1.7 1.0 0.2 0.5 0.5 0.2 0.5 0.2 0.2 0.7 0.2

i um, H ylotelephium argutum, Dactyl is dovenica, Acetosa alpestris, Senecio ]acquinianus La thyrus pratenais, Stavhy8 alpina a n d some o ther species, typ ica l for this asso- ciation, besides e.g. Ghaerophyllum h i r su tum, Ch. aromaticum, Polygonatum verticil- la tum and Cimic i fuga foe$ida, showing t h a t the soil had become sl ight ly moister af ter refores ta t ion o f t h e clearing.

I n 1988 prac t ica l ly the whole clearing (with the except ion o f the uppe rmos t par~) was reforested main ly b y spruce. The relev4 17/88 t aken b y P. K o v ~ 22. 6. 88 below the horizontal pa th on the slope no t far be low the chalet , shows a Calamagrostio-cult i-Piceetum, a depaupe ra t ed forest c o m m u n i t y , filling a gap left b y the fo rmer Galamcuyrostio arundinaceae.Abie te tum HOaVAT 1950:

10 • 10m, E340 ~o, E215 ~o,E*60 % , E 0 5 ~o Ea: Picea abies 6, Sorbua aueuparia 2, Larix europea 1, Sal/~ silesiaca 1

Et:/~ubus ideaeua 4, ~orbus auev4paria +

H/~D~k~: SECOI~DARY SUCOESION IN BELIKNSK]~ TATRY 353

E1: Calamagroztis arundinacea 6, Senecio jac~inianus 4, Myosotis syiva~ca 4, Dentar~a glandulosa 3, Arhyrium flli, z-fernina 3, Geum rivale 3, Hylotelephium arffutum 2, Urtica d/ioica 2, Dactyl~s doven~oa 3, Pulmonaria obsc, ura 2, Galium schultesii 3, ]Prim~da datior ~ubsp. carpat~ca 2, Epilobi/um cdpestre 1, Frazjaria yeses 2, Geranium robertianum 2, Asarum europaeum 2, Phy- te~ma ~oicaturn 1

E,j: Hypn~rn oupressi]orme 4

I t is not without interest that in more windprotected ecotopes natural forest regeneration was not so slow as in the area deforested artificially. In the year 1956 an avalanche destroyed totally the edge of the spruce forest near the path from the chalet to the Oval ~lab. In the following years there was a rapid succession from Rubo-Chamaenerietum HAVA~ et al. 1969 to Senecioni-Rubetum idaei HAD&~ et al. 1969 and to the forest, so that in 1988 we found a well developed wood community belonging to the all. Chrysanthemo-Piceion (KRA~X 1933) Bi~.z~A et HAvA~ 1969 with both tree and herb vegetation well developed.

The valley bottom along the ttlbokff petok brook as well as the Ov~i tlab was grazed by 700--1 300 sheep for 2--5 weeks up to 1954 and mown once a year up to 1956--7. In 1956--60 we found there a semicultural meadow community, described as Alchemillo.Festucetum pratensis HADA~ vt M. 1969, belonging to the all. Polygono-Trisetion Bm-BL. et Tx. ex M~RSCmLLL 1947. After anthropozooge- nous impact had ceased (with the exception of tourist trampling) this community gradually disappeared and a new association, the Geo-Dactylidetum 81ovenicae HADA~ 1981 or even Senecioni.Rubetum idaei developed in its place after two decades. In the third decade this community did not change much, the changes being mainly the disappearance of the most synanthropic elements, as shown in Tab. 1. Introduced species i.e. species occurring in the investigated ares only in secondary of seminatural communities (Festuca pratensis, Ranunculus avris, Achilles millefolium, Taraxacum officinale agg., Stellaria grcminea, Alopecurus praSer~iz, Phleum praten~e, Carum carvi, Ranunculu~ repent, Pea prat~n~is, Leon. todon hispidus, L. autumnalis, Rumex obtusifolius, Elytrigia repens and Prunella vulgarly) disappeared quickly in the observed communities. In 1956 15 introduced species grew in the Alchemillo.Festucetum pratensis, the Geo-Dactglidetum in 1977 contained only 7 of them, and the same association in 1988 had 6 introduced species only. The decrease of introduced species is more conspicuous if their abundanoy is ooml~red. The sum of their abundaney figures was in 1956:25.2 in 1977:6.4 and in 1988: 3.8. The JACCARD index of similarity of the observed communities changed in the following way : I j between Alchemillo.Festu~dum pratensis (1956) and Geo-Dactylidetum (1977 resp. 1988) was 25.4 and 18.7 respecti- vely, corresponding to differences on the association level, whereas I j between Geo-Dactylidetum 1977 and the same community in 1988 was 53.2, indicating that both communities belong to the same association.

The vegetation in the Ov~i ~lab above the chalet is covered by herbaceous and grass vegetation, as frequent avalanches maintain this depression treeless. A ty- pical easily identifiable community was Alchemillo.De~cham~sietum ce~pitosae H ~ D ~ et al. 1969 on a small ridge not far from the chalet, whose floristie composi- tion was influenced by sheep grazing and b:r avalanches. In 1977 the Hyperivo- Deschampsietum HAD~ 1981 was found in its place (see Tab. 2): The changes in the following 11 years (34 years after the grazing had c~sed) were insignificant,

3 5 4 I~0LIA GEOBOTANICA ET PHYTOTAX0t~ 'OMICA, 25, 1990

Tab le 2. Success ion changes f rom Alchemillo-Deschampsietum in 1956 (AD 56) to Hyperico- -Deschampsietum in 1977 (HD 77) a n d Hyperico-DesvhamTsietum in 1988 (HD 88)

A D 56 H D 77 H D 88

D~chamlzsia cespitosa (L.) P. BEAUV. Alchamilla monticola 0PIz ttypericura maaulatum Cm~Tz Agrostis tsnuis Sr~TH. Luzula luzuloide8 (LAM.) D ~ v Y e t W ~ L ~ . Veronica chamaedrys L. Cardaminop#is halleri (L.) HAY~X Urtica dioica L. Ac~tosa a2pestris (JAcO.) ~ . LSVE Aconitum firmum ( R c ~ . ) N~.rr.~. Epilobi~m alp~tre (JAcQ.) K . R O C ~ Ckrysosplenium alt~rnifolium L. Myosotis sylvaticu HOI~FM. qeum rivalr L. Pimpindla major (L.) Hubs. Carduus laersonat~ (L.) JACob. Alopseurus pra~asis L. Fcs$uca pratensis H U b s . Aehillea mille/olia~ra L, PMeatm pratenae L. 8ener 8ubalpinus K o c h Campanula glomerata L. ssp. ]atrae (BoRB.) DOST. Biswrta major S. F. G~AY Poa trivialis L. Primula elatior (L.) HILL Festuca rubra L. Geranium phaeum L. Viola bi/lora L. Calaznagrostis villosa ( C H ~ x ) G ~ r,. Banunwd~ acris L. Phle'um rhaeticum ( H u ~ . ) I~USCm~.ST Potentilla aurea (CR.) BECX Gampanv2a tatrc~ BORB~S Thymus sp, Galium aniaolahyllum VrrL. Trifolium pratemse L. Soldanr carpatiea Vr~aH. Carum carvi L. ~en~vio iacquinianus RCHB. Angelica eylvestris L. Heracleum sphondylium L. Galium sehultesii VEST Gahw'psis speciosa Nfrr,~, Chamaerion an~sti/ol~um (L.) HoI~lys Eryslmum hungaricum Za.~AL. Gc~ranium sylvatioum L. Aconitum vari~atum L. Thali~rum aquilegii/olium L. Leontodon hislai&us L. Alchemilla erinita B u s ~ , ~haero.ph/yllum hirsubum L.

8.8 7.0 4.4 1.2 1.4 4.0 4.4 2.0 3.2 1.0 4.2 0.5 1.6 1.7 § 3 . 2

1.6 3.5 0.6 2.0 0.4 0.2 + 0.5

0.6 1.7 0.4 2.2 + 0.5

0.5 1.5

6.8 1.2 5.2 2.6 2.0 1.0 2.8 4.2 2.0 1.0 1.0 0.6 0.2 0.8 0.2

+ 2.8 2.4 2.8 3.4 0.2 2.2 0.6 0.4 0.6 3.4 0.2 0.8 0.2 0.4 0.4 0.4 0.2 0.6 0.2 0,4 0.1 0.4 0.4 + 0.1 + 0.2

2.8 1.6 1.0 1,0 0.8 1.0 1.0 0.8 0.8

4.2 0.8 0.8 1.2 0.4 0.6 0.2 0.2 1.2 0.4 0.1 1.0 0.4

2.7 1.2 0.7 2.5 0.5 1.5 0.2 0.5

HADA~: SECONDARY SUCCESlON IN BELIAI~ISKI~ TATRY 355

Table 2. (Cont.)

AD 56 HD 77 HD 88

Cirsium arvense (L.) SceP. ~enecio ]uchsil GmEL. Lathyrus pratensis L. l~anunculus reports L. ~tellaria graminea L. Phyteuma s.picatum L. Gentians aselepiadea Scox,. Rubus idaeu. L. Cr~viata glabra (L.) EHI~END. Ae~jopodium podagraria L. Taraxacum o]]icinale agg. Valerians ~ambuci/olia ~ELAX. Alchemilla acutiloba OPIz

0.1 0.8 0.4 1.0 1.0

o:7 0.2 2.5 0.5 0.5 0.2 0.5 1.2

the introduced element was however less numerous. The Alvhemillo-Deschampsie- turn had 6 introduced species, the ttyperico-Deschampsietum only 2 of them. The JACOARD index of similari ty between the AlchemiUo-Deschampsietum and Hyperico. Deachamp, ietum in 1977 and 1988 is 29.2 and 22.9 respectively, whereas between the ttyperico-De~champsietum in 1977 and 1988 it is twice a~ high.

The Geranio-Urticetum dioicae HXDA5 1969, SO common below the chalet, chang- ed in 1988 to Carduetum personatae MoB~IU 1967 or to 8enecioni-Bubetum idaei HAl)AS 1969, with the exception of the enclosure, where sheep were kept before the 1954 (the so called kogiar). In 1956 a well developed Geranio-Urticetum was there; the same association was found there in 1977 (HADA~): Urtica dioica 9, Carduus personata 4, Zamium cupreum 2, Myosotis sylvatica 3, Scrophularia sco,polii 3, ~ubus idaeus 2, Chaerophyllum aromaticum 1, Galeopsis speciosa 1, Senecio jacquinianu8 2, Cardamine impatiens 1, Epilobium montanum 2, Galeopsis bifida 1, Fragaria ve,ca + , Acer pseudoplatanus juven. 5, in all 14 species. In 1988 P. K o v ~ found there the following community: Urtica dioica 9, Carduus personata 4, Rubu$ idaeus 2, Lamium cuTreum 3, Ohaerophyllum aromaticum 1, Stachys alpina 2, Stellaria nemorum 1, Geranium phaeum l, t typnum cupressiforme 1, in all 9 species. During 32 years this community has n o t changed quMitatively.

SUMMARY

Changes in anthropc-zoogenous plant communities in the valley of S e v e n Springs (Tatra of Bela) have been studied over a period of 30--32 years. Succession in the windexposed "Rubanisko" (clearing) were slow. The t~ubo-Chamaenerietum changed into Geo-Dactylidetum or Senecioni-l~ubetum idaei, whereas the Digitalo- Calamagrostietum arundinaceae changed very little even after 3 decades.

The Alchemillo.Deschampsietum in the Oval ~lab was succeeded by Hyperico- Deschampsietum after 2 decades. I ts changes after the subsequent 11 years were insignificant.

356 F O L I A G E O B O T A I ~ I C A ET P H Y T O T A X O N O M I C A , 25, 1990

In the valley bot tom the Alchemillo-Festucetum pratensia was succeeded by Oeo-Dactylidetura after 20 years or was invaded by Senecioni-Rubetum. In all cases the introduced flora-element became less numerous.

L I T E R A T U R E C I T E D

HADA5 E. (1981a): Z m 6 n y ve vege tac i k o t l i n y S iedmich p ramef iov (Belansk6 T a t r v ) 4v6 desl t i lotf po skonSeni p a s t r y . - - In : ~lov~tk a horskA pi~iroda vo 20. s to le tL - - Sborn . l~ef. Konf . ~p in - d l e r h v Ml:~n 1977, H r a d o c KrAlov6 1981: 7 3 - - 7 6 .

HADA~ E. (1981b): Z m ~ n y vogetaco Dol iny S iedmich p r a m e f o v v Bel iansk~,ch T a t r h c h za m i n u - l #eh 20 let. - - Zborn . ReL z k o n s k 39. v~,roSiu uzAkononia T A h l A P u a k 25. v( ' r . Ta t r zahsk io - go P a r k u Na rodowego , T a t r anskA L o m n i c a 1981 : 256- - 261.

HADA5 E. e t al. (1969): Die P f l anzengese l l s cha f t en des Tales Dol ina S i edmich p r a m e f o v in de r Be l ae r Ta t ra . - - Vogethc ia 0.SSR, B2 - - SAV Bra t i s l ava .

~MARDA J . a kol. (1963): D r u h o t n 6 spo le~ ,ns tvA ra s t l fn v T a t r a n s k o m n~4rodnom p a r k u . -- ,~port, B ra t i s l ava .

R e c e i v e d 3 J a n u a r y 1989