2xI~l~A LUKEw I and JIf~i KOM.~REK2
1CT~choslovak Academy of Sciences, Institute of Soil Biology, Na Sgldkhch 7, CS-37005 0Jesk6
Bud~jovice, CT~choslovakia
2Czechoslovak Academy of Sciences, Botanical Institute, Dept. of Hydrobotany, DukeIskh 145,
CS-37982 Tl~ebofl,Czechoslovakia
Succession of Soil Algae on Dumps from Strip Coal-mining
in the Most Region (Czechoslovakia)
Keywords
Soil algae, Succession, Primary succession, Plant succession, Colonization of new biotoi~s,
Vegetation on dumps, C~echoslovakia
Abstract
LUKE.~OV~A. et KOMKREK,1. (1987): Succession of soil algae on dumps from strip c(ml-mining
in the Most region (Czechoslovakia). -- Foli~ Goober. Phytotax., Praha, 22: 355--362. --
In the years 1985-- 198ti, the primary succeSSion of algae on dumps from brown coal mining
was studied in the Most region (North Bohemia, Czechoslovakia). The colonization of sterile
clayic substr.atos by algae (several tectal species of chlorophytes and heteroconts) and mosses
(protonemata) starts before the first ccesis of higher plants (in deposits about 3 months old).
Diatoms and, later, cyanophytes accompany the algal community after 1 yeWr. Filamentous
types of hetcroconts and green algae occur first on 7 year old dumps. Green algae represent
the commonest group throughout the succession. The number of species continuously increases
with the age of the dumps and finally reaches about 40 species from the 18th to 30th year.
The species composition is similar to that in grassland biotopcs in Czechoslovakia.
INTRODUCTION
The large dumps from surface brown coal mining near the town of Most (North
Bohemia, Cz~echoslovakia) are suitable for studying plant colonization and succession
on new and sterile soils (comp. Ih~xcH 1982, 1984). In 1985 and 1986, we studied the
primary succession (colonization and species composition) of soil algae in this area, in
relation to the age of the dumps (from 3 months to about 30 years) and to the succession
of communities of vascular plants.
The dumps will cover about 400 km 2 (40 thousands hectares) in this region at the
final stage. They are partly artificially reeultivated, i.e., planted with trees and crop
plants. However, this presents many problems. The study of the natural primary
succession of microflora is important for the recognition of initial colonization by the
first pioneers, and for the study of humus formation and facilitation processes.
356 FOLIA GEOBOTAXICA ET PHYTOTAXO.KOMICA 22, 1987
The area is situated in the shelter of the I4ru~n6 hory mountains. It is, therefore, comparatively
dry (500--650 mm of precipitations per year, 300--350 mm in vegetation season)
and warm (annual mean 7 to 8.5 ~ (acc. to PltACr~ 1983). The substratum is composed almost
uniformly of miocene clay (about 80 %), and to a small extent of sand and gravels. Usually,
the substratum exhibits a scaled disintegration (cleavage) and is chemically characterized as
alumosilicates with Si02, AI2Oa and H20, and larger or smaller amounts of Fe, Mg, and
alkaline metals, pH varies slightly about 7 (TOBi~RI~'~.1973).
The higher plant succession on dumps passes through several characteristic stages (PRAc~
1982, 1983, '1984) (comp. Plate ll): (1) The initial period with solitary annual plants
(e.g., Polygonum nodo,~um, Senecio viscosus; up to 2nd--4th year), (2) Atriplex nitens community
(2--8 years), (3) community with dominant Carduus acanthoides and Sisymbrium loeselii
(4 -12 years), (4) community with characteristic Tanacetura mdgare (7--20 years; during this
period the soil starts to be covered continuously by plants and litter, and humus formation
starts; s~)litary shrubs of Sa~vbuvus ni~ra are characteristic of this stage), and (5) a grassland
community with dominant Arrhenatherum elatius and Calamagrostis epigeios with more soliVary
trees (Sarnbucus nigra, A cer pseudoplatanus, Betula pendula, Fraxinus excelsior, Sorbus aucupama,
etc.; 18--30 years), Tussilago ]ar/ara is the species occurring from about the 2nd year
throughout the succession period, particularly on slopes.
METHODS
The soil samples were collected at the end of May and in the middle of September in tile
years 1985 and 1986 from the surface layer to the depth of 1--2 cm, and cultured in Petri
dislms under laboratory conditions at room temI~rature and daylight intensity. The species
composition was verified by the method cf growth glasses according to Lv~n (1945) and by
th,~ cultivation of si~cies from suspended soil on 1.5 % agar plates with BBM.
The sampling sites were selected in places with typically developed succession stages of
higher plants as defined above. At th~ e~rly stages the different types of substratum (pure
clay or sand with gravel) we,re compared. From each sampling place several probes were
collected and mixed together before cultivation.
RESULTS
Altogether 46 species of soil algae were isolated from all localities studied. The
actual and cumulative numbers of species (MAoART~VR et WILSON 1967) are demonstrated
in dependence on the age of the dumps (Fig. 1). The actual numbers of species
30 ~ j"
20 I."
yeors 5 10 15 20 25 30
Fig. 1. Actual and cumulative numbers of soil algal species during plant succession on dumps
from coal mining.
LUKE~OV-~ AND KOM.~REK: SUCCESSION OF SOIL ALGAE ON DUMPS 357
found in cultured samples gradually increase approximately to the 18th year of existence
of a dump. In samples with the maximum species diversity 39 species were found,
but later the actual number decreases slightly. In dumps 30 years old, the number of
species decreased to 37 species on the average. The reason for this decrease is not clear;
deteriorating light conditions due to the denser cover of higher plants and of plant
litter on the soil surface, and a greater amount of consumers may play a certain role
here.
In Fig. 2 the changes in the development of species belonging to different taxonomic
groups (cyanophytes, diatoms, heteroconts and green algae) are presented, and
their occurrence is related to the developed higher plant communities. The first occurrence
of algal species was already recorded in the youngest successional stages of
dumps 3 to 6 months old. However, these algae were only sporadic solitary cells of
t~otrydiopsis sp. and of two representatives'of green algae, namely Chlamydomonas
thomassonii and Chlorella vulgar@. While Cb. thom~onii was found only in this first
Dominant
vascg!.ar ptants
Age of dumps
Cyanophyceae
~ .. |
"~,_,BacitiariophyceaeQJ
e
~- |
|1
Xanthophyceae
|
o
CNorophyceae
J JDesmidiates
U[otrichales
Chlorococca[es
Chtamydomon.
0
p~ifhout solitary
onnuo[
vegetalion
plqnts
3-6
months.
-9
r ~
Corduus. acon~h. Tanacetum Arrhenetherum
Sisymbrium toes. vutgore Colamagrostis
(Tussilago) (solffary trees )
1-2 years 7-8 years 18- Ig years *_30 years
| | |
| (9
| | u
.| | | |
I
1
.|
!iii!!i!i
oo,~176176
~xˇ
xxxxxˇ
ˇ215215215
~ x x ˇ
|
iiiii!:@iii!;i!i!iiil
| ::::::ii:
| ::i::::::
g~xˇ
|
9 ,~
9 1 7 6
D 9 1 7 6
,ooo~
!i ...... "xxxxx~
Fig. 2. Increase in the number of species in different algal groups in dependence on the ago
of dumps and on the succe~ion of communities of vascular plants.
358 FOLIA GEOP, OTANICA ET PHYTOTAXONO,~IICA 22, 1987
period of existence of the dumps (without higher plants), Botrydiopsis sp. and Ch.
vulffaris occurred more or less sporadically throughout the further succession. It is noteworthy
that protonemata of mosses were already found in samples from the first
successional stage (entirely without higher plants).
On dumps of about 1--2 years differences were found between sites on various
substrata, and between sites without vegetation and those at the initial Atriplexstage.
On pure clay without plants only two species were found, while at clayic localities
overgrown by Atriplex four species with dominant Botrydiopsis sp. were assessed.
From the clay mixed with sandy particles and gravel 10 species with dominant
Bolrydiopsis sp. and Tetracystis dissociata were cultured. This difference also coincides
with I%ACH'S (1983) statement that on sites with sandy deposits the first
annual plants germinate and settle remarkably sooner than in places with pure clay.
CHLOROPHYCEAE :
Chiamydomonas ca[tunae
Ch. macrosteitata
Ch. moewusii
Ch. thomossonii
Chtomydomonas sp. 1
ChIomydomonas sp. 2
ChIamydomonos sp. 3
ChIoromonas sp.
years
>
1 3 7 s 18 30
t 1 If I ~ ff I f$ ,-I
Chtoretia minutissima
Ch. mirabilis
Ch. saccharophi[a
Ch. vulgaris
Chlorel[a sp.
Chlorokybus at mophyticus
FernandineIla alpi na
I~yrmecia sp.
Neochtons cf. biIoba~a
Broc~eacoccus c~ minor
Pseudococcomyxo simplex
Radiosphaera sp
Tetracystis dissociate
I J I I J " J I 11 I I" ll I I I I " I l I ! " i l l " I l l l l l i i i l l I I I I
O e . . . ' H ' ' ' ' ' ' ' O ' ' ' ' ' ' ' ' ' O ' ' ' ' ' ' ' '
O 0 " ' ' ' ' ' ' ' ' ' ' ' ' ' ' ' ' ' ~ ~
. . . . . . . . . H . . . . . . ' ' . . . ~ 1 7 6 1 7 6
Gerr~ne[ia tar ricoJa
Chlorhormidiu m f [accidum
Leptosira sp.
Stichococcus baci[I.aris
Actinotoenium cucurbita
samples
Fig. 3. Course of the ocetwrence of soil algal species on dumps of different ages.
LUK~OV~ AN]) KO,~I~,RFK: 8UCCI'~SSION OF SOIL ALGAE O~ ]DUMPS 359
No such difference was recognized between the two substrata in later successional stages
in samples from sites with a more developed higher plant cover.
On dumps about 7 years old two types of communities of higher plants were studied.
One with dominant Tu~silago farfara occurring mainly on the slopes of dumps
and the other with dominant Carduus acanthoides and Si~mbrium loeselii. Also
in 30 year old dumps the species composition of soil algae was compared between
communities with dominant Arrhenatherum elat~u~on the one hand and Calamaqrostis
epiffeios on the other. In both community types, ahnost identical sets of algal species
were found, with dominant Chlorhormidium flaccidum and Pinnularia borealis.
The dependence of the algae on the substrate thus exists only at the initial stages of
algal succession. At later stages, the species composition and their total number de~
pend on the dump's age, without any noticeable relation to either the substratum
CYANOPHYCEAE :
Leptolyngbya edaphica
L nostocorum
Leptolyngbya sp.
Phor m~d~um~ulumnale
Nastoc sp
yea rs
1 3 7 9 18 30
I I {J I I f# I fl" t
b~bbDDppODD~
b~ DODIpppp~
BACILLARIOPHYCEAE :
Hanlzschia amphloxys
Navicu[a al0mus ------
N contenta
N mutico
N rn,jtica ssp nlvahs .........
N. neovent r~cosa .........
Nifzschia capite[tata ..........................................
N parvuta
Pinnularia boreahs
P obscura -- ~
XANTHOPHYCEAE.
Bot rydiopsis sp
He~erot hrix debi[is
Heterococcus sp
Pieurochloris magno
Vis steUo~a
. ~ 1 7 6 1 7 6 1 7 6 m D
~ 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6 1 7 6. . . . . . . ,~ . . . . . ~ 1 7 6
sarnptes
360 FOLIA GEOBOTANICA ET PHYTOTAXONOMICA 22, 1987
or the higher plant community. Chlorokybus atmophyticus (Plate 12a--d) was, however,
found only on sites with a developed community of Carduus and Sisymbrium.
The fact that the changes in species composition of soil algae on dumps are related
more closely to the age of the dumps than to the higher plant communities also coincides
with the statement of TOB~RN~' (1973), that the rapid development of the surface
layers of dump soils soon eliminates local deviations.
In Fig. 3 the start and maximum of the occurrence of different groups of algae
(differences in numbers of species) are clearly recognizable during the succession
of algal vegetation. Chlorophytes are the first phototrophic pioneers on the dumps
and generally the species-richest and most frequent (as the cell number could be
derived from the cultures). Altogether 26 species of soil green algae were found. As
for comparison with other biotopes, the chlorophytes are also commonly present in
other soils in Czechoslovakia. The species composition is not substantially different,
e.g., from cultivated meadows. Chlorella saccharophila and Tetracystis dissociata
seem to be characteristic species of the dumps. Four species of green algae (Chlorella
mirabili~v,Chlarokybusatmophyticus, Neochloris cf. bilobataand Tetracysti~ dissoclara)
were recorded in our samples for the first time in Czechoslovakia (comp. Plates
12, 13). The heterocontic alga Botrydiopsis sp., which occurs commonly in all later
stages, is also characteristic of the early stages of succession and is usually dominant.
However, heteroconts do not occur very commonly on dumps.
The next group are diatoms, the vegetation of which starts in about 1 year old
deposits. The majority of the 10 species found belong to commonly distributed soil
types with the exception of Navicula neoventricosa, which was found in two year
old sandy and gravel deposits for the first time in Czechoslovakia. Nitzschia parvula
is a characteristic diatom of the old dumps.
The latest-occurring group in dumps are cyanophytes, with the exception of two
species, namely Leptolyngbya edaphica and Nostoc sp., which were found only in one
year old dump. Cyanophytes are surprisingly rare.
:Figure 3 demonstrates the occurrence of all species in dumps of different ages.
The early start of green algae and Botrydiopsis is clearly expressed. Botrydiopsis sp.,
Chlorella saccharophila, Pseudoc~comyxa simplex, Tetracyslis dissociata and one
undetermined species of Chlamydomonas are the commonest species from one year
old deposits through all later stages. Several examples of rarely occurring species of
green algae found in the Most dumps, are documented in Plates 12--14.
DISCUSSION
The development (succession) of algal communities in different soil types is very
little known. The colonization of sterile deposits from coal mines by phototrophic
microorganisms provides a very rare opportunity to study this process under natural
conditions, j ust as in new volcanic areas.
Our results are comparable, e.g., with the results of ~Uw (1977) from the coal-mining
dumps in Kuzbas, of T~a~EVSKIJet ~TZNA(1967) from dumps of ashes near thermic power
station and deposits from metallurgic plants (both from the USSI4), or of BEHaEet SCHWABE
(1970), SC~IWAB~(1970) and SCa~CA~Eet BEn~aE (1971) from the volcanic Surtsey island
(Iceland). In spite of the distance between these areas the composition of pioneer algal species
LUKE~OV~ AND KOM.~REK: SUCCESSION OF SOIL ALGAE ON DUMPS 361
and genera with accompanying mosses is similar (Botrydiopsis, Chlorella, Chlamydomonas,
Chlorhorm'idium, Tetracystis and diatoms Hantzschia amphioxys, Pinnularia borecdis, Navictda
aromas%N. mutica). The early start of diatoms in the Most dumps corresponds more with the
data of SCHWAB~-et B~.trRE (1972) than with those of TXRS~.VSXIZet ~TrNA {1967), who found
the start of diatoms only with the first occurrence of perennial grasses. The high amount
of SiO2 in the Most deposits probably plays an important role in the development of diatom
vegetation.
The colonization of soils on Surtsey island is more rapid than on dumps. The greater
number of species from Surtsey is probably the result of the greater diversity of
microbiotopes; the studied localities on the dumps were much more uniform. Also odr
cultivation methods are, by necessity, selective, and may have eliminated some spe-
cies.
In comparison with all the authors cited (cf. also MARTINOVA1986), cyanophytes
were rarely found in the Most dumps. This situation corresponds with the low nitrogen
content in the Most dump soils; it increases very slightly during the years (ToBiIR~'X
1973). The reason for this difference from the data from other localities is not clear.
The early start of cyanophytes in Surtsey is probably mainly connected with the
warm steam outputs from the volcanic substrate, which do not exist in theMostdumps,
but may occur in other similar North Bohemian localities both natural and artificial
(cf. BRxmCz 1941, KOMX~K et Rosa 1957).
The algal communities studied are composed of special soil species, which are also
distributed in other soil biotopes; however, in each case it is impossible to describe
them as "mainly cosmopolitan and ubiquist" (MXRTI~OVA 1986).
In the dumps the contact of new sterile soils with the surrounding areas is closer
than in the case of new isolated islands (comp. HALPF~R)ret HARMON 1983). Yet the
succession of vascular plants as well as microphytes proceeds very regularly in our
localities. The original flora (including the microflora) is completely buried below the
very deep and sterile new deposits on over large areas. The new colonization, which is
fully dependent on the input of diaspores from the surrounding landscape (mainly
by wind and animals), is interestingly not as rapid, spontaneous and non-selective,
as one would have expected.
The number of species in 30 year old dumps is remarkably lower in comparison with
old dumps, described, e.g., by MARTINOVA(1986). The restricted anthropogcnic influence
and different chemical conditions of the substrate in our case are probably the
main reason for this difference.
The algae indicate the microecological situation very well (GoLLERBACH et ~TINA
1969). The primary succession in dumps can therefore serve as a good example for
recognition of the colonization process under given conditions.
LITERATURE CITED
BEHRE K. et SCHWABEG. H. (1970): Auf Surtsey/Island in Sommer 1968 nachgewiesene
nicht marine Algen. -- Schr./qaturw. Ver. Schleswig-Holst., Sonderb., p. 31-- 100.
BR~B~-Z R. (1941): Zur Kenntnis der Algenflora des Franzensbader und Sooser Quellenbereiches.
-- Beih. Bet. Centralbl. A, 61: 137--236.
GOLLE~BACHM. M. et ~TINAE. A. (1969): PoSvennyo vodorosli. [Soil algae.] -- A. N. SSSR,
Izd. ,,Nauka", Leningrad, 228 pp. [In Russian.]
362 FOLIA GEOBOTAXICA ET PHYTOTAXON())IICA 22. 1987
HALPERN C. B. et HAa~o~" M. E. (1983): Early plant succession on the nmddy river
mudflow, Mount St. tielons, Washington. -- Amer. Midl. Natur. 110 (1): 97--106.
KOM~REK J. et Rosa K. (1957): [Das Vorkommen der Blaualge Somˇierella cossyrensis (?)
in B6hmen.] -- Preslia, Praha, 29: 17--27. (In Czech.)
LU.~D J. W. G. (I945): Observations on soil algae I. The ecology, size and taxonomy of
British soil diatoms. Part I. -- New Phytologist 44 (2): 196--219.
MACARTHUg R. H. et WILSOh" E. O. (1967): The theory of island biogeography. -- Princeton
Univ. Press, Princeton, 203 pp.
MA~TINOVA O. A. (t986): Al'gougrupovann}a vidvaliv dolomitovich zbagaSuval'nieh fabrik
1)onee'koj oblasti. [Alg~l communities from dolomitic dmnps of the concentr~ating mills of
the l)oneek-region.] -- Ukr. bet. ˇ. 43/6: 83--85.
PRA(:H K. (1982): [Vegetation on substrates ~reated by mining]. -- Act,a Ecol. :Nat. Reg., Prague.
1982: 49--50. (In Czech.)
I)RACH K. (1983): PNsp~vek k otg~zkhm ekologick6 sukeese. [Remarks to the problem of the
ecological succession.]. -- Cand. dissert., CT~chosl. Aca~.l. Sci. inst. Bet., 237 pp. (In Czech.)
PRACH K. (1984): Selec~d results of the study of succession on dumps fi'om IJrown coal
mining (Most region, NW Bohemia). -- Acts bet. Slov. Acad. Sei. slov., ser. A, suppl. 1:
257--261.
PaACH K. (1985): [Succession -- one of the centrM terms in ecology.] -- Biol. Listy, Praha,
50 (3): 205--217. (In Czech.)
SCHWABE G. H. (1970): Zur 0kogenese auf Surtsey. -- Schr. Naturw. Ver. Sehleswig-Holst.,
Sonderb., p. 101-- 120.
SCHWABE G. It. et BE~Rb: K. (1971): 0kogenese der Insel Surtsey 1968 bis 1970. -- Naturwiss.
Rundsc*aau 24 (12): 513--519.
~U~UEVA M.~ G. (1977): Vodorosli na otvalach ugol'nych razrabotok v Kuzb~se i ich rol'
v poSvoobrazovanii. [Algem in coal-mining dumps in Kuzbas and their role in the soil formation.]
-- In: Razvitie i znuSenie vodoroslej v poSvach neSernozemnoj zony. Mat. me~vuz.
Konf. Kirov, Perm, p. 52--53. (In Russian.)
TAg(~EVSKIJ V. V. et ~TIN), E. A. (1967): Razvitie vodoroslej na promy~lennych otvalach.
[Development of algae in industrial dumps.] -- In: Sovr. sostojanie i perspektivy izuS.
posy. vodoroslej v SSSR; Mat. me~vuz. KonL Kirov, p. 146--150. (In Russian.)
ToB~aN.~ V. (1973): Fytocenologickh charakteristika vybran:~ch rostlinn:~,eh spoleSenstev n~kter~eh
v~sypek Mostecka. [Phytoeenological characteristics of selected communities of some dumps in
the Most region.[ -- Dissert., PNrodov. fak. Karlovy Univ., Praha. (In Czech.)
I~eeeived 14 January 1987
Plates 11-- 14
PLATE 11 LUKE~OV.~ A~'D KOM~REK: SUCCESSIOX" OF SOIL ALGAE OX DUI~IPS
,f
~ r ~ ' ~ 9. , ,, ~ .-.... 9
Dumps from coal-mining in the Mos~ region (Czechoslovakia): a -- 1 year old, b -- 8 years old,
c -- about 30 years old; (photo) PRAC~).
Folig Geobotanica et Phytot~xonomie,% Praha, 2'2, 1987
I'LATE 12 LIJKE.~OV:( AND KO3I,'~REK: 8VCCE.~SlON OF SOIL ALO-AE ON DITMI'S
l~are species of eoceal green algae found in dump soils: a - - d - - Chlorokybus atmophyticua,
e -- Fernandinella alpina, f--k -- Myrmecia sp.
Folla Oeobotanica et Phytotaxonomlca, Praha, 22, 1987
PLATE 13 LUKE,~OV][ AND KOM~{REK: SUCCESSION OF SOIL ALGAE ON DI:MPS
'.4
llm'e species of coccal grecn algae found in dump s,:ils (cont.): a--d -- Neochloris of. bilobat(l,
e-- h -- Tetracysti~" diesociata.
Folla fi-eobotanica et Phy~o~axonoinica. Praha, 22, 1987
PLATE 14 LUKE,~OV)[ A.N=]) KOM~{REK: SUCCESSION OF SOIL ALGAE ON DUMPS
::~" 9
!
0
-~Z'" ,r
,r
Characteristic species of algae in dump soils: a--e -- Bracteacoccus el. minor, f--h -- Geminella
terricola, i--k -- Heterothrix debilis (i -- with Botrydiopsis sp., libereting of autost)orcs ).
)Folia Geobot~nica et I hytotaxononiica, Praha, 22, 1987