Montañas Hercínicas

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PIRINEOS 151-152

K ey wo rds: Her cy n ian mo u n ta in s , sp ec i e s d iv e r s i ty , an emo - o r o g r ap h ic

system

1. Introduction

Biodiversity is manifested at all levels of the biological hierarchy: from

g en es to sp ec ie s , co mmu n i t i e s an d eco sy s t ems ( SOLB R I G, 1 9 9 1 ;

GRO OM BRIDG E, 1992). W hilst shidie s of diversity at the gene level require

exper imenta l p rocedures which can on ly be appl ied to a l imi ted number o f

model o rgan isms, a t the species and communi ty levels d ivers i ty i s s tud ied

using observat ional -co mpara t ive me thod s (MA YR, 1982). The la t te r enab les a

sa t isfac tory evaluat ion of reg ional and g lobal d ivers i ty , and the management

and co nser vatio n of biotic resourc es. A n assessm ent of biological diversity in

practice is therefore often reduced to observations and comparisons of species

presence or absence. Identif ication of the centres of biodiversity (WWF &

lUC N, 1994) and var ious ho tspo ts o f species b iod ivers i ty have rece ived m uch

at ten t ion la te ly . Most mounta in reg ions, owing to the i r physica l and b io t ic

diversity, belong to the species-r ich centres and hotspots.

The priorit ies identif ied at the EST Workshop at Monte Bondone in 1995

for Eu ropea n m oun ta in b iod ive rs i ty research were : (1) a syn thesis o f cur ren t

data wi th regard to the b iod ivers i ty in mounta in hab i ta ts in Europe, (2 ) an

identif ication of gaps in scientif ic knowledge and (3) the init iation of co

opera t ive research programmes.

The availabil i ty of data for synthesis is a result of centuries long natural

h is to ry research in Europe. A cursory look a t the da ta would reveal tha t bo th

p lan t s an d an ima l s sh o wed a co n t r a s t b e tween lo wlan d s an d u p lan d s wi th

regard to the numb er o f species and var ie ty of comm uni ty types per un i t a rea

and ho w impac ts of p resen t-day imm igra t ions an d h is to r ica l fac tors affected

the specia t ion , immigra t ion and surv ival o f v iab le popula t ions. In a wider

contex t , the impor tance of mounta in d ivers i ty fo r human socie ty and in

genera l , i t s in f luence on the development o f na tura l h is to ry can be shown

(JENÍK, 1996).

In Europe, the highest biological diversity is associated with the Pyrenees,

A lps and C aucasus ; i .e . the h igh -mo unta in systems s i tua ted a t the t ransi t ion

zones be tween the tempera te and submedi ter ranean zones (MEUSEL

et al.,

1965). The A lps hav e a partic ularly long histor y of scientific exp loratio n in

we stern Eu rope (OZ END A , 1988) wi th an ac t ive na tu ra l h is to ry research

predat ing the Linnaeus per iod . The geomorpholog ica l and p lan t d ivers i ty o f

the A lps wi th the ir c . 350 endem ic species of vascu lar p lan ts (PAW LOWSKI,

1970) has m uch a t t rac ted bo tan is ts and p lan t socio logy and the syn ta xono my

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B O D IV E R SITY O F T HE H E R C Y N IA N M O U N T A IN S O F C E N T R A L E U R O P E

of communi t ies o f the Zur ich-Montpel l ie r School were born in these

mo u n ta in s ( B R AUN- B LA NQUET, 1 95 1) .

In the scientif ic sh ad ow of the A lps rema in the so called m idd le-

mo unta ins o f C en tra l Europe . These lower ranges ( see m ap on the inner s ide

of the cover in OZEN DA , 1994) belo ng geologically to a m uc h o lder

He rcyn ian (Variscan) system tha n the A lps. The y stretch from the Massif

C en tra l in France to the wes ternm ost ou t l ie rs of the Western C arpath ia ns in

Poland and Czech ia . They r ise on ly marg inal ly above the a l t i tude of na tura l

tree growth today and provide excellent examples of refuges of arctic-alpine

po pul atio ns of the presen t Post-glacial era. They offer, in the absence of stro ng

hu m an im pacts , a system to s tudy the in terac t ion betw een physica l and b io t ic

fac tors . These middle-mounta ins have a lso p layed an impor tan t ro le in the

development o f un ivers i t ies and o ther learned inst i tu t ions of Germany,

France , A ust r ia , C zech ia and Poland .

2.

Brief historical notes

The explora t ion of species d ivers i ty in the C en tra l Europe an mid dle-

mo unta ins b egan by the ac tiv i ties of herbal is ts L . C am erar ius and P. Jesen ius

in the O re M oun tain s at the end of the ló^'̂ cen tury (HEYNERT , 1964). C .

SCHWENCKFELDT f i r s t publ ished enumerat ions of b io ta occur r ing in the

Sud etes in 1600. A fter the 30-year war, in the second half of the 18* cen tury

florist ic researc h becam e active again (e.g.: MA TTU SC HK A , 1776-1777). A

unique scientif ic expedition of the Royal Bohemian Society into the Giant

Mountains in 1786 (JIRASEK et al, 1791) returned with long l ists of plants

from sp ecies-r ich sites. I t also discovered new species for science, e.g. Poa laxa,

a species later recognised as a typical component of the distant North

A mer ic an mou nta in s ( JENÍK, 1986). Nu m erou s f lo ra wo rks covered the

northern (Silesian) side of the Sudetes (for a list see JENÍK, 1961).

C o m p a r a t i v e s t u d i e s i n t h e 19 ^ c e n t u r y h i g h l i g h t e d e s s e n t i a l

d i f f e r en ces in sp ec ie s co mp o s i t i o n b e tween lo wlan d s an d u p lan d s an d

descr ibed the d is t r ibu t ion of var ious fo rmat ions ( today ' s b iomes) in cer ta in

mo u n ta in r an g es . B o tan ica l r e sea r ch h ad a lway s b een mo r e ad v an ced th an

zo o lo g y , ma in ly o win g to th e e f f o r t s o f p h y s i c i an s , h e r b a l i s t s an d

apothecar ies search ing for medic inal herbs . Ear ly exp lora t ion of wi ld l i fe

was mad e b y h u n te r s an d n a tu r a l i s t s , wh o d esc r ib ed g ame an d p r ed a to r y

mammals , b i r d s an d r ep t i l e s , b u t mo d e r n zo o lo g ica l r e sea r ch can o n ly b e

da ted b ack to the be g in n in g of the 19'^ cen tury . The s t ud y of the

in v e r t eb r a t e f au n a —co n cen t r a t in g o n so me g r o u p s o f i n sec t s— s t a r t ed

much la ter (KIESENWETTER, 1847) and s lowly , the d i f ference in

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PIRINEOS 151-152

b i o d i v e r s i t y b e t w e e n h i g h - m o u n t a i n s a n d p a r t i c u l a r m i d d l e - m o u n t a i n

r an g es b ecame mo r e ap p a r en t .

Du r ing the 20* cen tury , a num ber of taxonomic summ ar ies reconf i rmed

the h igh species- r ichness in some of the Hercynian ranges, par t icu lar ly in

those which r i se above the t imber l ine . A summary of the i r charac ter is t ics i s

g iven in the A ppen dix . Progress in an ima l and p lan t taxon om y and the

availabil i ty of floristic data w ith a goo d coverag e over the who le of C entra l

Eu r o p e en ab led co mp ar a t iv e s tu d ie s an d th e co mp i l a t io n o f p l an t

d is t r ibu t ion maps (MEUSEL et al., 1965 ; 1978). Other deve lopm ents inc luded

sma l l - sca l e v eg e ta t io n map p in g ( MI KYSKA, 1 9 6 8 ) , d e sc r ip t io n o f

communities (by using phytosociological relevés) and a classif ication of

syntaxonomic units (OBERDORFER, 1977-1992; MORAVEC et al., 1983), and

compara t ive eco log ica l evaluat ions of mounta in ecosystems, based on the

latter (JENÍK, 1961).

Presen t ing an overv iew of the b iod ivers i ty i s f raught wi th d i f f icu l ty

a r i s in g fr o m C en t r a l Eu r o p e ' s cen tu r i e s l o n g p o l i t ic a l f r ag men ta t io n .

A d min i s t r a t iv e b o u n d a r i e s d iv id in g r id g es in to so v e r e ig n t e rr i t o ri e s h av e

const ra ined f ie ld s tud ies , separa ted co l lec t ions of specimens and forced

u n n a tu r a l b o u n d a r i e s i n map p in g . Fo r ex amp le , i n d iv id u a l g eo g r ap h ic

( landscape) un i ts were of ten d iv ided and each s ide ' s b io t ic communi t ies

were descr ibed in local publ ica t ions a t opposi te s ides o f borders , somet imes

in d i f feren t languages wi th the in format ion se ldom crossing the border .

Research has been f raught v^ i th semant ic inconsis tencies , and topographic

n ames , n o men c la tu r e o f p l an t s an d an ima l s , an d map p in g u n i t s o f

vegeta t ion have var ied much . In add i t ion , about ha l f the 20* cen tury has

been af fec ted b y mi l i ta r isa t ion , the tw o Wor ld Wars and la ter by the erec t ion

of the I ron C ur ta in .

Biogeographical research suffered from the absence of unif ied toponyms

an d r e l ev an t t o p o g r ap h ica l map s . Th e Cen t r a l Eu r o p ean mid d le - mo u n ta in s

sti l l lack a consensus on oronyms (JENIK, 1998). For example, the prominent

range s i tua ted a long the borde r d iv id ing A ust r ia , Ge rma ny and C zech ia is

ca l led Bohmerw ald in Ge rma n speak ing A ust r ia , bu t Hin terer Bayer ischer

Wald in Germa ny, and Su mav a in C zech ia . The usage of these oronym s of ten

over laps and much confusion is caused in the re levan t da ta banks, reg ional

li terature and when an English equivalent is required (a suitable one for the

above ma y be Bohem ian Forest) . A nother low er range to the NW along the

same nat ional boundary is ca l led Oberpfa lzerwald on the German s ide , ye t

the C zech s ide bears the nam e C esky les, a hom ony mo us eq uivalen t o f the

A ust r ian nam e Bohm erwald . It i s wo r th no t ing tha t in the m aps of C laudiu s

Ptolemaios the same area was called Silva Gabreta, a forgotten unifying name

only recen t ly re in t roduc ed for a new n atura l h is to ry book ser ies .





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Owing to the above, in format ion about the d ivers i ty o f species ,

communities and ecosystems, is scattered in the scientif ic l i terature of the

four ne igh bour in g countr ies : A ust r ia , C zech ia , Ge rma ny and Pola nd . Only a

few large-scale synthetic works, such as those of HEGI et al. (1966 et seq),

TUTI N et al. (1964-1983), FIRBA S (1949; 1952) and ELLEN BERG (1996)

provide integrated information over the entire area. Two recent publications

by UNE SC O and lU C N (PRICE, 1995 ; lUC N, 1995) have a lso g iven a

t r an sb o u n d a r y t r ea tmen t of th e Her cy n ian mid d le - mo u n ta in s .

3.

Species diversity in Central Europe

There are ab out 80000 kno w n species of all organ ism grou ps (Table 1) in the

C zech p a r t o f C en t r a l Eu r o p e ( e .g . VA VR OUSEK & MO LD A N, 1 9 8 9;

PLESNÍK, 1999). This is only about 5 % of the world total number of species

described so far (GROOMBRIDGE, 1992). In temperate Europe, the most

species r ich groups (fungi and invertebrates) are associated with old growth

forests a nd can hav e a hig h r ichness within a single comm unity, i . e . hig h a lpha

diversity (sens u WHITTA KER, 1970). Ho wev er, old gro wt h forest stands are

rare and the majori ty of the remain ing wo odlan ds hav e been t ransformed in to

mo nod om inan t s tands wi th few t racheophyte species . A zonal non- forest

eco sy s t ems , wh o se sp ec ie s co mp o s i t i o n su b s t an t i a l ly v a r i e s a lo n g

environmental gradients and over patchy soils (beta diversity

sensu

Whittaker ,

1970),

a lso have a h igh b iod ivers i ty . The d ivers i ty o f au to t rophic

tracheophytes may be used as an estimator of overall biodiversity and the

non-forest fo rbs an d gram inoids wi th the i r nume rous associa tes can prov id e a

proxy in es t imat ing species- r ichness in the m ounta ins .

C losed wo odl and s covered > 90 % of the landsca pe nor th o f the A lps

( d o ma in e Cen t r e - Eu r o p éeen ,

sensu

OZE ND A , 1994) by the end of the late-

Holocene . Three forest fo rmat ions dominated : (1 ) mixed broad- leaved

Plants

Tracheophytes

Bryophyte s

A lgae ( incl. blue-greens)

Lichens

Fungi

Total

2000

1000

6000

1000

30000

40000

Animals

Marrunals

Birds

Reptues

Fishes

A m ph i b i a ns

Inver tebrates

Total

70

170

10

50

20

40000

40300

Table 1. Estimated numb ers of indigenous species in C zechia. Data are from variou s sources.

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PIRINEOS 151-152

woodlands in the lowland and coll ine belt , (2) closed beech and f ir forests in

the lower montane bel t , and (3) monodominant spruce ta iga in the upper

montane belt . The forest formations comprised about 35 tree species, 300

vascu lar herbs , and cor responding ly low numbers o f consumers a t the lower

trophic levels. A ctual and pot entia l compo sition of these forests is describe d

in detail in the phytosocio logical (e.g. , OBERDORFER, 1977-1992; ZLA TNÍK ,

1959;

ELLENB ER G, 1 99 6; MU C I NA etal, 1993 ; MORA VEC et al, 1983) and

palyn ologi cal l i terature (FIRBA S, 1949; 1952; HU NTLE Y & BIRKS, 1983). The

Pleistocene glacial periods inhibited migration and the f lora and fauna

becam e re la t ive ly imp over ish ed in C en tra l Europe in con trast to the m ore

species- rich Eastern Forests o f No r th A mer ica .

The f loodplain woodlands and colhne oak-hornbeam woods at lower and

warmer alt i tudes have been reported to contain 50 vascular plant species per

relevé (about 400 m^ in size) , however, most of the zonal vegetation in the

foothil ls of Hercynian mountains consists of oligotrophic oak forests, beech

forests a nd spruce taiga, w ith < 20 vascular plan t species per relevé. There is a

high beta diversity in azonal woodlands along deeply-cut r iver valleys and on

l imestone (kars t ) . On h i l l s ides , the nor th fac ing s lopes tend to host

psychrophi lous beech

Fagus sylvatica)

comm uni t ies , whi ls t the w arm southern

slopes on l imestone host durmast -oak

(Quercus

pubescens) wo o d s wh er e p l an t

and animal species of a diverse geographical origin occur together .

The b iod ivers i ty of the C en tra l Europea n landscape is h igher in pa tchy

and t ree less azonal hab i ta ts , where p lan ts and an imals do no t face the

adversi ty o f c losed-canopy shade and roo t compet i t ion by t rees i s reduced .

The c losed wo odl and s of C en tra l Europe become patchy natura l ly in

ex treme environments such as the dry and ho t sou th- fac ing s lopes a long the

river valleys, in the karst areas and on exposed cliffs of volcanic hil ls. These

xero-thermic habitats are found along the large r ivers, such as the Berounka,

Danube, Sázava , Saale , Vl tava and Weser , on l imestone , e .g . : Swabian Ju ra ,

Bohemian Karst , Morav ian Karst , in the basa l t ic Doupov Hi l l s and on the

neovolcan ic cones of NW Bohemia (Ceské St redohor í ) in the ra in shadow of

the Ore Mounta ins . The t ree less pa tches are dominated by

Festuco-Brometea

grasslands, con ta in ing a mix ture o f re l ics and forerunner species , inc lud ing

some continental elements lef t from the Late Glacial when this biome was

wid esp r ead in Cen t r a l Eu r o p e .

Water logging , par t icu lar ly in depressions and areas o f permanent spr ings

and m ires p roduc es a var ie ty o f we t lands wh ich m ay occur locally in fo rests .

There is a variety of l ife forms present in wetlands which interact with the

physica l env ironment and in f luence microhydro logy and micro topocl imate ,

and thus create favourable conditions for enhanced beta diversity. Soligenous

mires of the

Scheuchzerio-Caricetea fuscae

class (fens) and ombrogenous mires





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of the

Oxycocco-Sphagnetea

class (bogs) m ake rem arka ble cont r ibutio n to

b iod ivers i ty in C en tra l Europe , par t icu lar ly wi th regard to b ryo phyte s a nd

inver tebra tes . For example , on the C ervené Bla to bog in South B ohemia ,

SPITZER & JA ROS (1993) identif ied 570 species of butterf lies over an area of

less than 3 km^. The drying out of these wetlands creates favourable

conditions for tree growth, and thus a high reduction in beta diversity. The

adverse effects of waterlogging on trees may combine with effects of low

tempera ture in the montane and subalp ine bel ts o f the Hercynian mounta ins .

This is the case for mires and f lushes in the High Sudetes (particularly in the

Gian t Mts . ) , where numerous arc t ic -a lp ine p lan t and an imal species have

survived the pressures of forest succession in the Holocene and contr ibute to

today's biodiversity.

Co ld wind-exposed hab i ta ts , par t icu lar ly a t h igh a l t i tude wi th a shor t

growing season can prevent the successful establishment of trees by delaying

the hard en in g of the xy lem t issues . C om mo n in the ne ighb our ing A lps an d

Wes tern C arpa thian s, this treeless bald or alpine tun dra is a rar i ty in the

Hercynian middle-mounta ins . They most ly occur as smal l t ree less i s lands on

top of some r idges and peaks, in snowbeds and corries. Some of them can be

viewed merely as azonal islands within the montane belt : e.g. Plesné Lake,

C erné Lake, C er tovo Lake and Gross A rber-See corr ies in Bohemian Forest, the

Zechengrund in the Ore Mts . , o r the summit ba ld of the Harz . However , the

larger treeless areas o n top of the Vosges, Black Forest , Giant M ts., Snow Mts .

and Jesenik Mts. form a true alt i tudinal belt , sometimes referred to as

subalpine or even alpine. Their diversity is best ref lected by the number of

habitats and communities. For example, in the three ranges of the High

Su d e te s , ab o u t 7 0 p l an t co mmu n i t i e s h av e b een d i s t i n g u i sh ed in 1 1

phytosocio log ica l c lasses (hab i ta ts) :

Asplenietea trichomanis

(crevices) ,

Thlaspietea rotundifolii

(scree),

Juncetea trifidi

(summit tundra) ,

Salicetea herbaceae

( snow beds) ,

Mulgedio-Aconitetea

(tall-herb communities) ,

Montio-Cardaminetea

(f lushes) ,

Scheuchzerio-Caricetea fuscae

(fens),

Oxycocco-Sphagnetea

(bogs) ,

Nardo-Callunetea (mat-grass and heath communi t ies) , Betulo carpaticae-Alnetea

virids (subalp ine scrub) , and Vaccinio-Piceetea (krum mho lz) . The species

richness of these habitats is well documented in f lorist ic and faunistic works;

SOUREK (1969) enumerated 1150 vascular species in a single range of the

Giant M oun tain s, which equ als half the total estim ated for C zechia (Table 1) .

4. The hum an vegetat ion history of the Hercynian m ountains

Human- induced f ragmenta t ion of the con t inuous forest cover in the warm

low lands of C en tra l Europe bega n in the Neol i th ic , about 5,000 B.P. resu lt ing





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PIRINEOS 151-152

in today ' s mosaic o f na tura l , semi-natura l and cu l t iva ted forests , pastures ,

meadows, arab le land and patches o f t ree less c lear ings ( fo r a summary see

ELLENBERG, 1996). Forest clearing and the expansion of agriculture have

progressive ly m oved in to the mo ntan e bel t and to the r idges of the He rcynian

mounta ins . Pastur ing a t the summits o f the Vosges has been ment ioned as

early as the 9* century (SCHMITT, 1963), whilst the uplands of the High

Sudetes w ere no t exp lo i ted un t i l abou t the 16* cen tury (LOKVE NC , 1978) .

Us ing the evidenc e of archives and old charts, JENÍK & HA M PE L (1992)

poin ted ou t some d i f ferences in the exp lo i ta t ion of summ it tun dra on va r ious

estates in the Jesenik Mts., where the 16th century, too, appeared to be the

beginning of pasturing. The application of ar t if icial forest regeneration,

cu l t iva t ion of monodominant s tands of Norway spruce

(Picea abies

(L.)

Karsten) and Scots pine

{Pinus sylvestris),

and the introduction of exotic tree

species such as

Pseudotsuga menziesii, Pinus strobus, Robinia pseudoacacia

for

example have caused large changes in the foo th i l l s o f Hercynian mounta ins

over the past 200-300 years . The in tensive ma nag em ent has no t caused forest

loss in the monta ne bel t in most o f the Hercyn ian rang es, bu t species d ivers i ty

has decl ined main ly th rough the expansion of con iferous s tands.

A l t i tud in al zonat ion in the He rcynia n mou nta i ns d i ffers in the we st f rom

that in the east . In the Vosges and in the Black Forest , European beech

(Fagus

sylvatica)

and silver fir

{Abies alba)

h ad p r imar i ly d o m in a ted mu ch of

uppermost montane bel t up to the t ree- l imi t , however fo l lowing t ree

cu t t i n g , b u r n in g an d a i r p o l lu t io n i t h a s b een t r an s f o r med in to

monodominant p lan ta t ions. In con trast , in the Bohemian Forest , the Ore

Mts .

and Sudetes , beech forests se ldom reached an a l t i tude of > 1000 m and

co n i f e r o u s t a ig a wi th No r way sp r u ce p r ed o min a ted u p to ab o u t 1 3 0 0 m .

Ho wev er , f o r e s t c l ea r in g an d lo g g in g , f o l lo wed b y e i th e r n a tu r a l

r eg en e r a t io n o r p l an ta t io n in t h e l a s t 3 0 0 y ea r s h av e r e su l t ed in

mo n o d o min an t sp r u ce s t an d s o v e r mo s t o f t h e se mo u n ta in s t o d ay . Th e lo ss

of b road leaf s tands caused an impover ishment o f so i ls and reduced bo th

b e lo w- an d ab o v eg r o u n d sp ec ie s r i ch n ess ( p r e l im in a r y d a t a o n f u n g i an d

bacter ia are avai lab le a t C har les Univ ers i ty ) . Wind th row s fo l lowed b y ba rk

beet le ou tbreaks were another fac tor in changing forest composi t ion wi th

prominent ef fec ts on b iod ivers i ty .

Norway spruce , European beech and s i lver f i r fo rm a krummholz in the

t ransi t ion towards the t ree l ine in many Hercynian ranges where there i s no

natura l ly occur r ing dwarf p ine

{Pinus mugo).

Monodominant s tands of P.

mugo

form a krummholz on ly in the Gian t Mounta ins wi th smal l pa tches a lso

in the Bohemian Forest and Iser Mts. Remarkably, the Jesenik Mts. were never

co lon ised by dwarf p ine , though i t i s widespread to the west , in the Gian t

Mts. and to the east , in Babia Góra ( the we sternm ost C arpath ian ran ge w i th

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Pinus mugo). The are no alt i tudinal l imits to tree growth in the Ore Mts. and

thei r species- r ich Zechengrund is p robably on ly a smal l i s land in the snow-

rich leeward side (see below).

The position of the treeline and the presence or absence of the competitive

dwarf pine have played an impo rtant role in the deve lopm ent of biodiversity in

the Hercynian mountains (JENÍK & LOKVENC, 1962). Not only alt i tude, but

genera l re l ie f and topocl imate ( i r rad ia t ion , wind ac t ion , dura t ion and

distr ibution of snow, snow creep and avalanches) have diversified the pattern of

the alpine-like belt with its wide range of plant and animal species from either

the A lps and the borea l/subar ctic region (SOUKU POVÁ et

al,

1995). In the m id-

Holocene period, marked by the expansion of Central European woods above,

their current alt i tudinal l imits, only the highest summits, steep f lanks and

corries remained treeless. Treelessness has been a prerequisite for the

development and maintenance of fragile plant populations and communities of

alpine character. In a similar context, C A RBIENER (1966) prefers to write abo ut

subalpine vegetation in the Vosges, and BOGENRIEDER

et al.

(1982) have

coined the term subalpine island for the highest area of Feldberg in Black Forest.

The long-term ma intenan ce of the local alpine-like areas with ou t relict forest

species and their continued treelessnes remain a puzzle for the palaeo-

ecologist . Palynological data have suggested that during the climatic optimum,

ab o u t 5,000 years ago , forest vitali ty in C entral Europe increased and the alpin e

timberline a scend ed 200 to 400 m above i ts prese nt-day posit ion (FIRBA S, 1952;

de VA LK, 1981; HÜ TTEM A NN & BORTESCHLÀGER, 1987) . This means tha t

a l l Hercynian ranges would have been covered by forests and the i r

helioph ilous arctic-alpine populatio ns shou ld have become extinct. A lso, there

would have been no place for the microevolutionary processes which resulted

in the differentiation of some non-forest plant taxa in the Holocene period (for

details see JENÍK, 1983).

5. The causes of the distribution pattern of biodiversity

A s a wh o le , t h e Her cy n ian mo u n ta in r an g es l ack h ig h a l t i t u d e , t h e

d ivers i ty o f rocks, la rge areas o f l imestone , and complex i ty o f relief, they

only excep t ional ly surpass the na tura l t ree- l imi t , and the i r ac id , nu t r ien t -

p o o r p a r en t r o ck s se ld o m c r ea t e a f e r t i l e su b s t r a tu m f o r eu t r o p h ic

ecosystems. This i s ref lec ted in the species-poor zonal vegeta t ion wi th a

smal l number o f t ree species and few compet i t ive ferns and grasses ,

er icaceous shrubs and a modest c ryp togamic f lo ra . The fauna is s imi lar ly

sp ec ie s p o o r . Ho wev e r , t h e r e a r e so me lo ca l i sed cen t r e s o f h ig h

b io d iv e r s i ty i n ce r t a in lo ca l i t i e s wh ich h a r b o u r en h an ced n u mb er s o f

91





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PIIUNEOS 151-152

sp ec ie s , man y in f r a sp ec i f i c t ax a , an d cu r io u s h y b r id s b e tween p a r en t s

der ived f rom eco log ica l ly d is tan t hab i ta ts .

A n exam ple of locali t ies w hic h repeate dly ap pea r in floristic and faunistic

lists is the three ranges of the High Sudetes. The corries (or cirques) of KoteLné

Jámy, Labské Jámy, Sniezne Kotly, Kotly of Great and Litt le Staw in the Giant

Mts, or Velká Kotlina corr ie in the Jesenik Mts. have several hundreds of

vascu lar p lan t species , whi ls t the ^species-poor communi t ies on neighbour ing

slopes are composed of only a dozen vascular species (JENÍK et al., 1983a; b).

The above cor r ies also have a h igh nu mb er of mosses , hepat ics , mol luscs and

insects . Enviro nme nta l a nd phytosocio log ica l analyses ( JENÍK

1961;

JENÍK

et

al.,

1980) have identif ied some common features in the topographical posit ion

of these species-r ich sites. They are mainly ( i) on east-facing slopes at the

eastern marg in of la rge summit p la teaux or on the eastern s ide of p rominent

saddles , ( i i ) across the summit p la teaux on the adverse s ide of a p rominent

funnel-shaped valley, ( i i i) in the hollows of corr ies or snowbeds, ( iv) si tuated

under p rominent corn ices , covered regular ly by deep snow dr i f t s and

affected by frequent avalanche action (JENÍK, 1990).

The driving force behind this special biodiversity pattern is the prevail ing

wester ly winds, a fea ture wel l documented by summit meteoro log ica l

s ta t ions on Snezka Peak and Szren ica (MÍGALA

et al.,

1995). The westerlies

are s t reamlined and accelera ted in funnel shaped windward val leys , and

sweep across the cor responding summit p la teau or saddle areas . This wind

action is documented by f lag-trees, unilaterally eroded turf and ablation

forms in the snow surface in winter . Over the sheltered lee slopes, the laminar

air f low changes to turbulent eddies, a process clearly detectable by the

movement of clouds, airborne snow crystals and by the deposit ion of drif t

snow (STURSA

et al.,

1973). In summer, deposit ion of other airborn particles,

such as so il , l i tte r and seeds has a lso been do cum ented . Simi lar topogra phica l

configuration and related air currents appear in the summit areas of all

Sudetic rang es and there are 10 pro mi ne nt areas of enh ance d species div ersity

re la ted to them. JENIK (1961) has descr ibed a model ca l led anem o-orogra phic

system, which consis ts o f th ree in terdependent components : (1 ) windward

funnel-shaped val ley , (2 ) accelera t ion summit zone , and (3) leeward

turbulence zone. The validity of this model has later been confirmed in the

Vosges (C A RBIENER, 1966; 1969; de VA LK, 1981), in the Boh em ian Forest

(SOFRO N & STEFA N, 1971) and in the Feldberg region in Black Forest

( JENÍK, unpubl ished) . I t i s the leeward tu rbu lence zone tha t co incides wi th

the cen t res of h igh b iod ivers i ty . C ur ren t we ather in g and local ly ou tcro pping

base-r ich rocks, which are kept bare by the secular action of avalanches and

water erosion improve nu tr ien t avai lab i l i ty a t these s i tes . In the summer ,

d issec ted tu r f and exposed so i l a re invaded by seed and spores t ranspor ted

92





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sometimes from afar , thus further increasing the r ichness of f lora and fauna.

Treelessness has been main ta ine d by avalanches, whic h even occur red d ur in g

the c limat ic op t imu m of the mid-H olocene d ue to snow sw ept f rom the la rge

su mmi t p l a t eau x .

Desp i te the avalanches and erosion by mel t water the leeward hab i ta ts o f

th e an emo - o r o g r ap h ic sy s t ems h av e , ma in ta in ed a s t ab le m ic r o h ab i t a t

diversity and, in the course of millenia, accumulated a genetically diverse

assemb lag e f r o m r a th e r d i f f e r en t mo u n ta in an d so me t imes lo wlan d

ecosystems. Relicts from the Late Glacial and early-Holocene grow together

wi th s ub- therm ophi lou s species of the mid-H olocene . These s ites funct ion as

micro-evolu t ionary workshops where gene recombinat ion and hybr id iza t ion

produce new infraspecif ic taxa or microspecies (JENIK, 1983). The corries of

the Sudetes, for example, have been the speciation centres for the apomictic

Hieracium

genus (20 endemic species) . The Velká Kotlina in the Jesenik Mts.,

i s in the lee o f a double anemo-orographic system, and has accumula ted

about 500 vascular plant species (some of them have recently become extinct) .

I t also has a r ich vertebrate and invertebrate fauna, which makes this place

the most species r ich locali ty in the whole of the Hercynian mountains of

C entra l Europe ( JENIK et al., 1983a; b) . Similarly, a prominent anemo-

o r o g r ap h ic sy s t em mak es p o ss ib l e t h e p r e sen ce o f b io g eo g r ap h ica l ly

ou tsta ndi ng species in the back wa ll of the corr ie nea r Great A rber Lake in the

Bohemian Forest ( JENÍK

et al,

1998).

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A p p e n d i x

Brief characterist ics of selected ranges in the He rcyn ian m ou nta ins ; A ,

highest peaks with alt i tude, B, extent, C, climate, D, l i thology of the summit

area, E,

georelief,

F, posit io n in relation to imm igratio n, G, eviden ce of

microevolution, Fî, disturbance factors impacting on biodiversity, and J,

selected references.

The Vosges

A:

Ho hne ck 1362, Kastelberg 1346

B: about 100 km in length and 40 km in width

C: annual precipitation > 1900..mm,.._eu=marit ime pluviométrie regime,

suboceanic thermal reg ime

D: main ly porphyro id gran i te

E: f lat- topped summit area, gentle slopes towards west, deeply cut corr ies

on the east-facing slopes

' F: ra ther so l i ta ry wi th in the Hercynian A rch , ca lcareous Ju ra to the sou th

G: numerous apomictic microspecies, e.g. in the

Sorbus

g en u s

H: graz ing in the summ it area , wo od cu t t ing , ai r po l lu t ion

J: JA EGER (ED.), 1963; C A RBIENER, 1963; 1966; 1969; DE VA LK, 1981.

Black

Forest

A : Feldb erg 1493

B: about 160 km in length, 50 km in width

C : ann ual p rec ip i ta t ion > 2000 mm , d if ference betw een w ind exposed and

lee slopes

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D:

main ly gneiss and hunter sandstone , smal l a reas o f g ran i te

E: most ly smooth and undula t ing sur face , wi th a number o f cor r ies and

s n o w b e d s

F: close to the calcareous J ura and the A lps

G: infraspesific taxa, e.g., within

Gnaphalium supinum,

differing from

popu la t ions in the A lps

H: woo d cu t t ing , con iferous p lan ta t ions , a ir po l lu t ion

J : BA RTSC H & BA RTSC H, 1940; BOGENRIEDER

et al,

1982.

Bohemian F orest

A : Great A rber 1456

B:

about 130 km in length and 70 km in width

C: annual p rec ip i ta t ion > 1400 mm, warmer NE f lanks

D : mainly gneiss, smaller summit areas of granit ic rocks

E: modera te s lopes , a la rge up land p la teau wi th few prec ip i tous wal ls in

corries

F: m em be r of a chain of rang es of the Boh emia n Massif wi th the A lps lyin g

across the Danube val ley

G: infraspecific varieties of Picea excelsa (Sumava proven ance) , en dem ic

Gentiana bohémica su b sp . gabretae

H: woo d c u t t ing for g lass indust ry , spruce p lan ta t ions of a l ien prov enanc e ,

insect popula t ion explosions, and peat exp lo i ta t ion

J: KUNSKY, 1968; SOF RON & STEFA N, 1971.

Giant Mountains

A: Snezka Peak 1602

B: about 35 km in length and 10 km in width and connected to a massif of

the Jizera Mts.

C: cool and oceanic climate; high proportion of precipitation is as snowfall

D:

granite in the main Silesian r idge, gneiss and phylli te in the Bohemian

ridge

E: two para l le l r idges connected by two large up land p la teaux wi th

polygonal so i ls and numerous cor r ies and snowbeds

F: highest range of the Sudetes stretching along W to S direction

G: 30 endemic vascular taxa, mainly microspecies of the

Hieracium

genus, a

popula t ion of Sorbus sudetica der ived f rom two van ished paren ts (S.

chamaemespilus, S. aria)





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H: cutting of wood, catt le grazing, tourism, air pollution

J: JENÍK, 1961; SOUREK, 1969; SYKORA et al, 1983 ; J A HN , 1985 .

Jeseník Mountains

A : Prad id 1492

B: about 25 km in leng th an d 4 km in w id th

C : co o l su b o cean ic c l ima te wi th h ig h max imu m win d sp eed s an d

abundance of snow

D : phyl l i te , amphibol i te , gneiss

E: a funnel- shaped r idge wi th smooth summits , a few snowbeds and a

prominent corr ie (Velká Kotlina)

F: easternmost range of the Sudetes connected with a few low Sudetic

r idges

G: infraspecif ic endemics of vascular plants

(Plantago atrata

ssp .

sudetica

,

Dianthus carthusianorum

ssp .

sudeticus

A U T H ) a n d

Hieracium

m icrospecies .

H: g rass cu t t ing , wo od c u t t ing , monospeci fic sp ruce p lan ta t ions

J: JENÍK, 1961; JENÍK & HA MPE L, 1992 .

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Montañas Hercínicas

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