full text

COMPARATIVE ANALYSIS OF'SUCCESSIONALSTAGESOF'
SANDY VEGETATION - A CASE STUDY
K. Marq6czi
MaryiczL K (1993): Comparati'e a allsk of successionalnas6 af sandy wgetatian - 4 case
study. - Tiscia 27, 3-8.
different stagesof pimary
Abstrrct. The vegetationof sevenselectedsites, represenring
soheme,and to comparethe
suooession,
is aMlyzedin orderto esiablisha feasiblesuccessional
palhway
by
the
multivariate
methods
parhways
rhe
suggesled
succ€ssional
and
traditional
a nore or lesslinear successional
relev6ssuggests
The ordinationof micro-scalecoenological
wiih thetsditional theory.
s€quenc€
ofstudiedstandsconespoDding
Five g,?ical patchesw€re identifiedin eachstandby olassificaiionin order to repres€ntthe
Ordinalionof 6ese lypical patcheson the basisof species
intemalvariabilityof the vegetation.
frequ€ncies
shows,thatthe primarydynanic ofthe sandvegetationis moreooinploxThe linoar
suooessional
sequenceof studi€dstandsis uolikety, differeni kansitionsbetwoenstagesare
equiprobable,
andthe graphstructureofseriesis ratherreticular.
lype
Threegroupsofstandswere identifiedby diversityorderingi(i) an initial, oPengrassland
with mediumdiversity;the forestherblayeralsobelongsto
with low diversity;(ii) a grassland
andthe shrubbyhabitat.DiversityProfiles
this oategory;(iii) the mostdiverse,olosedgrassland,
of thesegroupswerecleariyseparated.
The m€thodsemployeddiffelenliateih€ vegetationof the studjedstandsaocordjngto their
sequence
doesnot necessarily
diversityandcomplexity,but the €stablished
speciescomposition,
reflectto therealhistorioaldevelop eutofthe standsin quesiion
Kewottlsi sancl! wgetatiotl, successioLntultituriate
ethah, di|e,sity ordering
Pi 659,H-6701Hungary
K. Marg6czi,DeparmenlofEcology,JATE Univ€rsity,Szeged,
Introductlon
On€ofthe typicalvegetationandflorl typesof
the HungarianPlain can be found on smdy areas
between the Danube and Tisza riv€|s. This
vegetationhas been shrdied by many authors,
amongthen such pioneersas Kerner(1863) and
Rapaics (1918). The successionalstases were
describedby Hargjtai (1940), zsolt (1943) and
Magyar (1960). Besides this olassical works,
Pr€cs€nyi(1981) studled the diversiry chaDses
dudns suocessionand there are severalpublications
phenomena
of
elucidatingcedainecophysiological
suchvegetation(e.9.Tubq 1984and Feketeet ai.,
1988). The spatial pattem as weli as the niche
belongingto the gnssland
r€lationsof somesp€cies
commlnity Festucetu raginatae lr.rc studied by
Molnir andNosek(1979)andFeketeet a1.(i980).
RecentlyFekete(1992)publisheda nelv conwich is differentfrom
oept of primary succession,
TISCIA 27
the traditional interpfetationby the Hungarian
phytosociologists.
He pointedoui that the primaD/
dynamicsof sandyvegetationis very complexrthe
vegetaiion-soil evolution is non-patali6l; ihe
suocession
is often deteminedby stoohastictran"
sitions between stages; the successionalgraph
structul€of the sand s€re is ratherrcticular than
The
linear;origin of climaxstageis hetercgeneous.
pioneersnssland do€s not accumulatesuffioient
humus fof the €srablislmentof steppe-m€adow
species,so the steppem€adow does not fit the
lin€. The standsof oak
pioneergmssiand-shrub
wood (the climax connuniq, in the kaditional
concept)arehabitat-dependent.
Although the classical phytocoenological
of sand)!egetation
successron
is given
oescr,prion
in the above meniioned papers, no detailed
nultivariate analysishas been made to describe
anongthe stnges.
vegetation
differ€nces
Classifications
and ordinationsarc suitablefor
analysis of succession,but the effectiveness of
djfferent methodsdep€ndson ihe natureof analyzed
dara(Mazzoleni,1991).
Unequivocal successional sequencesare not
necessarily self evident to critical observer
(Andersoq 1986). Ior example,computerbased
classification and ordination tecbdques d€fined a
successionalgradient that differed fion a theoretical or assumedsuccessionalclassification1nthe
case of a big sagebrusl/grasscomnuniry (Tueller
ald Platou,1991).
The aim of this studyis to analyz€the v€gelation of seleot€dhabitatsin a Hungariansandyarea
using multivariate methods and diversity
comparisons,
andto evaluatethe differert concepts
ofsuccession
accordingto th€r€sults.
The art comnuniry compositiona.]d epigeic
faunaofthe samestudyplotsanalysedin lhis paper
hasalsobeenstudied(Jlirdlinet al., 1993)
nesophilousand nesophilousspeciesbesidesthe
xerotolerant
ones.The phytocoenologicai
siatusof
thisstandis uncedain.
Plot 6. Shrubbyhabitatwith 50-60% covemge
of Crataegus nanagya, Juhipen$ connu,,i\.
Berbe s vulAaris andLigtsttun vulgarc sht\bs
Plot 7. Closedpoplar(Pop&lrsa/ba)forestwith
sama Robi ia pseudaacacia .Jees. Crutaegus
nonagtna and Junipetw cannunit give a sparse
The areasof the studyplotswereabout400 m2.
2. Field sanplins
A sinilar samplingprocedurex,asappliedthatr
desoLibed
by Szolletand Bartha(1991)with sone
modjfications.Long tnnseot of 200 contagious
smallplots, eachof 20x20 cm size, wereusedfol
Study area and Methods
samplingin eachstand.The total lransectof 40 m
lenghtwasbroken4-5 times,resultinga zig'zagline
L Site desctiption
and absenoe
"netling"the wholeplot. The presence
The field studieswer€ canied out in a naiure ofspeoi€irootingin the subplotsv,ererecorded.hl
leseNe in fhe southemparl of Hungary,betwe€n the shrubbyhabitaiandin the poplarforestonly the
of detectable
lhe rivers, Duna and Tisza near !o the village herblayerwassampled.The presenoe
w€rerecord€dandanalysed
K6leshalomrn June 1991.The study siLeis a lichenandmossspecies
complexof wind-blolvnsand dunes,SeveLalsuc- togetherwith thebiSherpLants.
cessionalsiagesof sandy vegetationoccur here
fiom bare sand to popl forcst For the preseni 3. Data ahalysis
The sevenstudysiteswere ordinatedaccording
study 7 plots were s€lect€d,representig these
ro rbe pooledfrequencyof occuningspeciesusing
stagesl
principal coordinateanalysis with Czekanowski
Plot 1. Almosi baresandwith sorneplant spe" index by the program package NuCoSA
cios belonging to the associationFr,ttrcalr,, (T6thlneresz,1991, 1993a).Virrgh (1986) havs
yaginatae.The bare sudacewithin the plot nlay found the sarneordinationrnerhodand similarity
index to be usetul for deiecting vegetation
derivefrofi an abandoned
sand-mine.
differences
in a similarscal€study.
In orderto represenr
the intemal variabilityof
Plot 2. Openperennialgrassland
with 30-40%
plant cov€rage,predonir\ ting by Festuca r.lgi ata the standsfive most abundantoharacterisiic
patch
(Festucetwn |aginatae danubiale). The ]|r,ossnnd types (10 contagious20x20 cm subplotof each)
were seleotedin eachstandby classificationof aU
lichenlayerwasalsoconsiderable.
200subplotsacoording
to theirspeci€scomposition,
similariq,indexby the NCLAS2
Plot 3. Similar to plot 2, but predominated
by usingthe Sorensen
Stipa borysthenica. (Fesh.tceh.t,nvaEinatae stipe- program of SYN-TAX lll prograF package
(Podani, 1988). These 20x100 cmz subplots,
tasutnborysthenicae)
represenlirglypical patches,were classificat€dby
Plot 4. Openperennialgrassland
predoninated NUCoSAusins Czekanowskisimiladty index and
by the species belonging to the Festucetun singl€ linkage soiting algorithm. Principal
laginatae comm]rr,ity ^ad Popuhts alba fomls a coordinate
analyses
(NuCoSA)ofthe samesubplols
shrubstoreyhere.Heightofit is s0 cn, about2s % were peLfomedusins asain Czekanowskiindex,
andthe resriltsof clusteringmethodand ordinalion
PIot 5. Closed grasslud with sone xero,
TISCIA 27
Analysis ol the diversity conditions were
performed by dive'siry o de iog rTodmeras?
1993a,b).This evaiuationdifferentiatesih€ diveBiry
sequedce of studied commurities based on
dominant lercus lare species. The Hill diversity
value is seisitive on rare species at low scale
parameter value and it is sensitive on dominant
speciesat high paranet€r value.
Ta$le 1- T[cpoolcd foqr...y values otth sldi€s in tlc 200 2ox2o 3n quadntctet shrdyplol Dah of22 rarc $ccies, whoscloolcd
f@qu;cy valucav.r. tclow l0 ae not indicaiod.Th€ uscd nonetrclahnc h aftor siron ( 1992).
SPECIES
Conplolh.cirn
lfi8eta
r
2
0
0
0
0
0
0
0
0
0
12
0 5
0 3
1
o
0
0
0
24
9
3
0
0
Equisetun rcn*istimrn
l
Euphorbia ctpaissi$
1
3
0
59
2
12
r24
10t
t1
5
3
0
5
0
0
0
9
STUDY ?LOTS
4
5
6
7
0
1
2
26
0
0
21
0
3
6
o
Ilt
0
0
1
2
t21
0
o
o
17
4
5a
l
0
0
0
0
0
0
0
5
0
29
0
63
0
3
2
1
r
3
0
a
2
0
0
2 3
0
0
0
18
o
24
55
0
2
t61
22
0 1 J
3
0
0
12
t3
3
PLllochdr.
sqrd,rdd
5
0
2
3
3
9
a
3
6
53
0
0
2
0
0
2
o
0
0
l
l
0
3 ?
t6
23
2
a
0
0
0
1
0
'1
A
12
51
5
l3
2
0
23
2
2
7
1
132
0
0
3
l5
NSCIA 27
t9
l5
33
1 6
2 5
23
R€sult
Ordi ation and cfassification of st dy
"ai
The ordinationolthe sevenstandsaccordingto
the pooledfrequencyof occuring speciesdefineda
-...\
successional sequence, rhat did not differ
\,
considembly from a theoreiical or assun€d
successionalorderj if the arch etrecl of the ordina'
tion methodis laken into coffideralion;or y the
plot 4 doesnot fit well into thesequence
(Fig. 1.)
The selected representaiive subplots of the
standswere sepantedrather well on the PCoA
scatiergran(Fig. 2.), and the classification
resulis
confirmthe togethemess
of thes€subplots(Fi8. 3.).
Thereare very few overlapsb€lweenthe diffefent
stands, consequentlybetween ihe successionaL
subplotr
stages.Ohly the subplotsof dre shrubbyhab(al r18.2.Pnncrpdcod'lDarcM(lyss ol Llie'eDresentative
o f r h cr J d . c d , k n d , L
. i n c se d c i ' c lrch c! u b p l o G
bcloncrns
(plot 6) form l1o separalecluster and they are
to lhc ,omochstcr,indiortcdby rta6 ir Fig,L Th€ d!,hcd
positionedfa. from eachotheron the cenlralpa|l of
linessisr lossibl. orddrsof rho samplcsco.sid.dng th.
rrchoffc.tof llr.ordin iotrmcLhod.
the scattergram.
The vegeiationofthis standis very
heierogeneous.
The identified patch types mny
rcpres€nidifferen!successional
stages,and can be area.Insidetb€ denoledclusterthe two siudy sites
(Fig 3.).
orderedin a feasiblesuccessional
sequence
that is areseparated
parallelwith the sequence
(Fig.
of tbe otherstands
Diversit,
2.t.
lig. L P ,cipalcoordlrar.nalysn oftlc shdy plolsaccodtug
ro thc poolcdfrc'$ancic,oftho occnirinsspccica.
ror tho
a.srirtion ofthc atudylioh indicatcd
by nuhlcs sc. thc
"Sludy d.a md Mctlods .
Altogether76 specieswere recordedin the 7
siands.The frequencies
of speciesare presented
on
Table 1. (Datfl of 22 rare specjes,whosesummed
fiequencyvalueswerebelow10 arenot indicaled.)
Three groups of standswefe identifLedby
diversityorderingr(i) an initial, opeograssland
type
with low diversity;(ii) a grasslandwith ftediuln
diversitytthe forestherb layer also belongsto this
cAtegory;(iii) the nrosl djverce,closedgrassland,
andthe shrubbyhabitat.Div€rcityprcfiles of these
groupswere.clearly
separaled.
In the first group the diversity of plot 1 is
unambiguouslylower, than that of the plot 4.
Howevernr the nediurn diveNily groupthe profile
ofplots 2,3 and7 crosseachother,this meanstha!
thesethreeco nuniiiescannotbe orderedsimply
by th€irdiversity.At lowerscalepalameter,ihat is,
regardingtherarespeci€sthe diversityofthe plot 7
is the highest,but the oppositeis right ai the higher
valueof scaleparameter- iegading the donr;nant
Discusslon
Some subplots do not join to any group; ihey
represent
unicalpatchrypes.On Fig. 2. the subplots
According to the tradilionai inte{pretations
ofplot 1 andplot 4. are encircledtogether,tlis lwo (Zsalr, 1943 and Magyar, 1960) the feasible
plots represent
the initial stageofsuccession
in this successional
sequenceof siudied stands is the
6
TISC]A 27
communiry,an amual grassland,
which is regarded
by thefaditional conceptls the firsr stageof sandy
successioncannotbe found in K6leshalomsite.
Festucdwgi ata is rhe rl]ost frequentspeciesm the
very open,pr€s'rmablypioneerstages(plot I and
plot 4). It is remarkable,that poplar can colonize
even in the very initial forn ol Festrcetut
vagrrdla?oommunitythe (plot4).
Fekete(1992)presenleda differentconoeptof
primary succession
on sandi the startingpoint is
Dsually Festucetun wgitlatae community. The
succession
terminatesar the Juniperc- Papuletun
along nany lines; this comnunity corresponds
to
the real forestsin this sere.The sleppemeadow
(closed grassland)does no! fit to the proneer
grassland'' shrub line and this makesthe graph
reticulaied
in thisxerosefies
The fesultof analysisreflectingto the intemal
Fis.3. DcndrogBboflh. Elrcscnhliv. sulploh oftho studied variabilityof the stands(Fig. 2.) suppo( rhis con$ande.Tlc a6 indi.rtc thc &lcvst clust rs,subflorsof
ceptfathef thanthe traditional,lin€ar successional
whicl oro cnciicl.dby lir$ in th. ordin ion diasmmD
sequefice.The starling point is undoubt€dlythe
complexoflhe plot I andplot 4; plot 2, 3, 5 and 7
e dpornis of successrol.
followirgalmosi baresand(plat 1) "- Ferhrenml could be nlrernarrv€
yaginatae danubiale (plot 2) - Fasfircetn
Besides this ar alternntiveexplanationof the
\)aginatae ltipetosrtl sabulosae (plar 3) - open scailergra is alsopossibler(I-a) - 3 - 2 - 5 - 7 plot
couldbe rccognizedaloflg an arch in the
grassland
with small poplartrees(plot 4) - closed sequence
Fig.2.
gLassland
(plot 5) - shrubbyhabital Glol 6) Tbe subplotsof plo! 6 may rcpresen!intermepoplarforest(plot 7). On the PCoAscattergram
of
standsthis soquenoe
oaDbe mofe ol lessrecogujzed diate stngesof altemativepathwaysleadingfrom
(plot 1,4)to the closedgrassland
if consideLing!h€ arch effect of the ordination the opeflgrasslnnd
nethod (Fig,1.).The arch effeci is strongerwhen Glot 5) andpoplarfol€st(plot 7). At th€ samerime
lypical patchesof plot 6
the ordinaredsampleshavefew speciesir conrnlon the subploisrepres€nling
(Mazzoleniet al., 1991) The Brcneho tcctonutt are ordered along aD arch, parallel with the
nlenlioned(1-zl)-. 3 " 2' 5 - 7 plot sequ€nce
in the
scattergran.That is, aherfiativeinteryretations
are
possiblewhefl t0,fu9 to identiry the successronaL
sequence
on the bas€of ihe ordinationresult.The
used methods are suitable to arange rhe
communitiesaccordhg to their complexjry,and
.n
speclescompositionbul it doesnot mean,that the
identifiedsequence
exactlycorresponds
to ihe r€al
,t
historical
de\,elopment
questior,
ofthe
stands
in
=
ln a hilly areatherearc considerable
diff€rences
in the physico-clremical
condiiionof the soil (i.e.,
humusandnitrogencontert,watercontentand soil
granulesize)ai ihe top ofthe sandhills, in the wind
groovesbetweenthem and at the relatively flat
afeis (K6rm6czi, 1983). Such environnenral
- - - - r l l
!
o.o
r.z
,.a
r.6
610 differences mry cause the development of
ai
altemativeerdpoinisofsuccession,
accordingto the
concept of Drake (1990), who resard rh€
Iis 4. Divc$iry ord€nngof thc ,tudy piots.lccoiins lo llic
environmentrlgrrdientsas | filter definug which
poolcd fEqnency valu$ of th. .ccunis sp€cies.Ior
setofspeciesis pemissibleto colonjze.
d€scriptionoftn. shdy plots indicaled!y dunbc.s scclxe
.studyd.a md M.tlods '.
The specles
diversityjs the highestin the closed
grassland(plot 5) and in the shrubbyhabitat(plot
T]SCIA27
6), but presumablebecauseof ditrerent reasons ln and L. call6 providedhelptul revjew of an earlier
the oaseof closedgrasslandconmunity $e nore venion of ihe manuscript.The work was founded
favourable environmental conditions of the slight by the OTKA V3 grantNo. 160?.
wind srove habitat (higher soil humidity, lower
of
wind effectandinsolation)allow the coexisrence
The shrubbyhabitatshaved€veloped
more species.
- In: Kikkava, J.
succession.
in the upper slope of a sand hil1, and lhe sbrtbs Andes6n,D.r. (1986):Ecolosical
D.J.(.ds.):CommunityE olosy,P.tic Md
md
Andcsn,
stand separatelyproviding a wide varieiy of
Proccs,Blrckw.ll sci. hbl., cdton, Austmlir,pp. 269'
microhabitats from the open, dry patchesbetween
235.
the shrubsto the shadedsitesunder their canopy, Drakc,J-A..(1990):Thc mechrnicsaf coDmuity asshbly ud
$cc.$ion - J.Th.or.Biol.l,/2,213-233.
were ihe soil is coveredby litter. Here the habrtat Iokete,6.
(1992):Thc I'olislicvi€w of$cccssionrconsidcred.causes
highernumberof speoies.
heierogeneity
P acsenyr rckcld,c.,Trba,Z, ald Mclk6,!, (1933):Backsroundlrocescs
Studyirgsimilar sandyvegeLalion
at tho populrtionl.vcl durinBsucc.ssionin Snssl ds o.
(1981) found that the speciesdiversiE,was the
sand.- Vcgclalio//,33 .ll.
is
iD
ihe
of
successiot,
that
lowestin the last stage
Fckere,C., Tnba,z. ud Picsanti, I. (1930):Niche studicsoD
forest.In presentcasethe diversityof foLestberb
sono phut sp.ciosol r srNslandconNnity. VU - Acla
B.L tl\ng, 26, 2At-297,
layer is high if resardingthe rare sPecies,bu!
lower if regardinglhe dominan!sPe- Hnrsitai,z, (1940)i NogykdrijsDijvaDyviligaIL A honoki
oonsiderably
(ll r lifc of NagykiJrais.
IL Plst
niiv6nyeiivclk@ctck,
cies (Fig. 4.). This communitydoes not evolve
coodrnitioson sand),- Bot$ikai. Ki'21.n6nyek.J7,205slage bur
aulomaticailylrom a pre"iousgrassland
2,44
the light shotage(theshadingeffeotofthe growing I5rd6n,C,,, G.ll{ L, Md Marg6cziK. (19t3): An! conmunity
qospositiotrin r Husr on $cc.$ion!l sanddunc uca, of
treecanopy)driv€sits developmenlSo,because
the considerable change ill environnenta] Komor, A.M. (1863): Pn$zorloboD d( Doraul.ndar' conditionsa different speciesPool is allowed !o
b.t*.on tha zonaiionof,Ndy
K6rn6czi,L, (r9S3)rCotrclarion
colorize(of. Drake,1990)
gRsElNdaDdthr physi.o-chcfiical
conditionoflhoi! soilin
The vegetaiionof plot 4 providesan eviderce
Bugrc,-Acir Biol. SzcAcd29, I l7-127
possrble
is
of opengrassland
rharlhe leforestation
^ l)I d l J L i ! . I i , , - ^ k r d { r n . r . K . d d 6 . B u t { e r l
M0srrr,P.Ll96O
but canno! be found cLear tfansitional slages MMzolctri,s., FLcnch,DD Nd Milos,J. (1991)rCo'nrralivc
sludyof chssification!trd odin ion mclhodson succet
and folest herb liyer
betweenthe open grass)and
riond d!t!, - Coonosc!
d,9l"l0l.
amongthe studiedPlots.
Molhir, E, N, lud Nosok,t,N,r Sidtial !6ce$c6 it r s6slmd
of
Pfesentpaper ref€rsto field investigations
cunnunit,l, -Aota Bol,Acad.sci. H!tr8,?J,339-348,
one single time Point. Direct evidecesabout the Podahi,J. (1988)rSYN"TAX lll User's h.nurl - ,{bstractr
L
Bot.niq /r, Supplomont
pathwayscan be drawnuP oDly aftel
successional
tuacs,hyi,l, (1981)iChdrsosin t[e divcEity of l[e legclalior
severalyears study period on Pe|m rehl plots
dldrg srcco$iotr-ach Bot,Acad.sci. Hung.27,139r93,
Resultof sucha long teffi sludycaMedout on the Ropaier,R.(ler8), Az Alliild ndvantffjldDjzij.lhm.. - Edesz€ti
vegetationof Hungariansandyarcashavenot beeD
l. (1992):A nrsy.loszasi .dany.s n6h hatiroz6F,
of the Sinon,
publjshedyet. Althoughbetterunderstanding
(D.t ihinlliotr book of tb. Hnhgdridr nsculd n66),
successionis very importnnt from the natue
T.nkdnyvkiad6, Budap.ri
consewatiorpoint of wiev as well, becausethe szollit, Cy, ud Brdl'a, S. (lr9l)r Paltetuft.ly* or dolonit.
sr.ssltrd coho!tritics u,inc nforhation thcory nodch '
of
consewation
activiryhaveio aim at naintainance
Abstrlct!Botrni.! /J, 4r-60.
stagesclose10 each other if it is 'L6tl1nfr6sz,a, (1993n):N!CoSA l.oi Nr'nbcr ctu"clri lor
all succ€sionftl
possible.BLtt ih€ nininal spalial scale of the
co'ntunitt !tudics and oll$r ecologicalarulicahons '
AbstactaBot ica I/,233-237.
rsno'known) errs dell
successionalprocesses
T6rllnaraszB. (19936.):Divoid 1,50:A ptogLaBfor diw6ity
Acknowledgem€nt
was
I an thanttul to L. Call6:this investigation
b) hin. r,d
carnedoul on rn€snrdysi'esselecred
joint io his projecton communityorganization
dufS. Bafha
on sand.B. T6thm6r€sz,
ing succession
orderina- Tisoia27,33-44.
Txellcr,P.T.lnd rlabu, K.A. (l9el):aphr succcsioDsrad'odt
9r,5?-63.
nr.6ts s.g.b&sl/gF$ ecosyst€n.Vegetatio
viiish, K. (1936): Th. cficct of hcrbicideson v.g.t tior
a Brltivarialcstudt- AbstractaBottrica 10,31?
dynamics:
340.
(Thc vcscZsoll,J. (1943):A ecnlendroi szisetn6v6nytrkamja.
tation oa szhtcrdrc Islmd). Indcx ltoni Bolarici Univ.
T]SCIA 27