chapter 23 - Part 1

 

Tab. 16. Between-year species similarity of the centiped community in MP 10
1992	57,14
1993	41,67	46,15
1994	50	53,85	87,5
1995	50	42,86	87,5	77,78
1996	50	42,86	87,5	77,78	100
1997	25	46,67	41,67	50,00	38,46	38,46
	1991	1992	1993	1994	1995	1996

Among the Isopods, only the mesohygrophilous Hyloniscus riparius and eudominant eurytopic Trachelipus rathkii were recorded in the first year of the monitoring. In the next years (Fig. 63), the number of recorded species increased. The dominance of H. riparius and of the stenotopic hygrophilous species Porcellium collicolum increased, while the dominance of T. rathkii suddenly decreased (Fig. 64). The mentioned changes indicate a slowing down or stopping in the locality drying (probably effect of the simulated flooding?), Tab. 17.

The carabid community in MP 10 has been monitored already since 1987. In that year the locality was flooded for a long time and, for a part, permanently swamped. Therefore the one-year sample from 1987 was taken as a standard for the original state of this community. Paludicolous Agonum moestum (15,95%,  Fig. 65), Patrobus atrorufus (13.79%) (Fig. 66), hygrophilous Platynus assimilis (4,96%,  Fig. 67) and the relatively tolerant species Pterostichus strenuus (29,31%) (Fig. 68), Carabus granulatus (7,76%) (Fig. 69) and Oxypselaphus obscurus (4,31%)(Fig. 70) predominated in the community. The high moisture or swamping was indicated by the presence of Europhilus fuliginosus, Europhilus micans  (Fig. 71) and Oodes helopioides.

In 1989, abundance of all hygrophilous species increased, except for Agonum moestum. Simultaneously, the abundance of the moderately hygrophilous Asaphidion flavipes increased (from 2 individuals in 1987 to 248 ind. in 1989, Fig. 72).

In 1990-1991 the whole catch was lower due to flooding of this locality (from 1348 individuals in 1989 the catch size decreased to 727 and 372 individuals respectively), but the community structure was considerably more similar to the original state of 1987 with a proportional similarity of 56,7-57,6%, and an abundance similarity of 39,6-40,5% (Fig. 73). Flooding affected also two hygrophilous, but not the expressively paludicolous species Patrobus atrorufus and Platynus assimilis.

Representation of the more tolerant or less hygrophilous species C. granulatus (35,6 and 22,5%), Pterostichus melanarius (6,5 and 14,2%), Pterostichus niger (13,5 and 22,3%) increased moderately in 1992, but in 1993 their increase become very strong. As a new species, the less hygrophilous Epaphius secalis (63 individuals, i.e. 3,8%) (Fig. 74) began to occur in this locality. After a decrease of 1991, the abundance of two less tolerant hygrophilous species increased (Platynus assimile from 18 ind. to 70 and 169 ind. respectively, and Patrobus atrorufus from 50 ind. to 94 and 111 ind. respectively), however the relative representation of Patrobus atrorufus decreased (from 13,4% to 8,8 and 6,7% respectively).

After a two-year interruption of sampling in 1994 and 1995, a striking decline of the cumulative abundance of all Carabides was recorded (from 1648 ind. in 1993 to 636 ind in 1996 and 396 in 1997). In 1996 and 1997 only three tolerant species predominated, viz. Pterostichus strenuus (30,1% respectively 36,36%), C. granulatus (16,82% respectively 19,19%) and Oxypselaphus obscurus (1993  10,1%). The relative and absolute abundance of these three species also strongly influenced the increase of the similarity indices between the whole-year samples from 1996 and 1997 with the sample from 1987, even in spite of the considerably lower abundance of some polyhygrophilous species. In 1996, an invasion of the xenocoenous species Trechus quadristriatus (Fig. 75) was observed, but in 1997 its abundance decreased a little.

The index of species similarity (in relation to 1987) exhibited a stabilised declining trend with small fluctuations (Fig. 73). The indices of proportional similarity and abundance similarity reflected the changes in the mutual proportion of species described above by sudden drops in 1989, 1992 and 1993.

In the frame of this locality, the changes of moisture and minute differences in altitude of soil surface (ranging from 50 to 60 cm) strongly influenced the spatial distribution of individual species [50]. After the drying of the locality and the retreat of the strongly hygrophilous species, the depressions were occupied by the more tolerant species which originally inhabited the small elevations. The driest elevated places were almost abandoned by the more tolerant carabides, but these species, having originally been concentrated in the elevations, were not substituted by other mesophilous species.

The amphibian taxocoenosis consisted of 5 species (Bombina bombina, Hyla arborea, Bufo bufo, Rana lessonae and Rana eculenta), but in individual years 1-4 species were recorded. While in 1991 only two species were fond, in 1993 there was only one (Fig. 76). An increase in the number of species was recorded in 1994 and 1995 when B. bufo and H. arborea appeared in this locality. In spite of certain changes in this taxocoenosis during the monitored period, (reflected also by the values of Jaccard´s index), the conclusions are ambiguous. However, the low number of species recorded in 1993 could result from the decrease of the groundwater level in the first half of 1993.

Tab. 18. Index of species similarity of amphibian taxocoenoses, MP 10

In the taxocoenosis of small terrestrial mammals, 9 species were recorded during the whole monitoring period, viz. Sorex araneus, S. minutus, Crocidura leucodon, Micromys minutus, Apodemus flavicollis, A, sylvaticus, Clethrionomys glareolus, Microtus arvalis, M. oeconomus. The catch size of A. flavicollis, Fig. 78, and A. sylvaticus evidently decreased after 1995. This obviously resulted from the artificial flooding of this area. C. glareolus was a dominant species without significant oscillations during the whole monitoring period (Fig. 77). The abundance of S. araneus and S. minutus oscillated greatly during the studied period, but it is not possible to identify a clear trend which could be considered as a response of these species to the changing conditions.

During a seven-year period altogether 32 butterfly species were recorded (in the period 1991-1992: 20 species, in the period 1993-1997: 29 species, Fig. 79). After the start of the Gabčíkovo structures operation, the species number increased strongly and in some species also their abundance increased. Among the mesophilous and hygrophilous species this included Erynnis tages, Ochlodes venatus, Leptidea sinaspis, Coenonympha glycerion and Araschnia levana. Among the xerophilous species Colias alfacariensis and the significant indicator of the xerothermic habitats Melitaea phoebe which invaded this monitoring plot, in 1996 and 1997, along the dikes from the close surroundings.

Taxocoenosis of birds (Aves) in MP 10 Kráľovská lúka was monitored in three partial plots - in an older willow-poplar stand with a rich stratum E₂  and in two poplar monocultures, one with an E₂ stratum, the other without it. In the willow-poplar stand, a number of species and individuals exhibited two contradictory trends during the monitored period. The numbers of species and individuals moderately increased in the breeding season, but out of the breading season decreased. In addition, the decrease in the number of individuals was greater.

In the mature poplar monoculture with a dense E₂ stratum over the whole partial plot, an average number of 9.3 species and 24.8 individuals occurred in the breeding season in 1992, while in the adjacent monoculture without an E₂ there were only 4.6 species and 7.7 individuals. Out of the breeding season, 6.7 species and 18.7 individuals occurred on the plot with an E₂ stratum, while on the plot without an E₂  stratum only 3.4 species and 6.6 individuals (Fig. 80, 81) occurred. When compared with the data from other flood-plain forests, e. g. with the monitoring plot Dunajské Kriviny, it is obvious that the poplar monocultures are inhabited by strikingly poorer ornithocoenoses.

In the course of monitoring in the breeding season lasting from June to July, the average number of species was 15 in 1989 and 1990, 17.5 in 1996, 18.6 in 1997, while the average number of individuals was 41 in 1989, 29 in 1990, 37.5 in 1996 and 39.3 in 1997. Out of the breeding season (data from February, April, August - October) the average number of species changed as follows: 14.3 in 1989, 12.6 in 1990, 9.8 in 1996, 11.2 in 1997, while the average number of individuals was 39.5 in 1989, 37.8 in 1990, 19.2 in 1996 and, finally, 28.2 in 1997.

It can be concluded that the bird communities did change significantly during the monitoring period. In the breeding season the average number of species per one observation was more or less stabile, but the number of individuals exhibited a generally decreasing trend. Out of the breeding season a moderate decrease was recorded in both the number of species and in the number of individuals. A continuation of the decreasing trend in size of bird population in future could result in changes in species composition. The changes observed in the bird communities were caused not only by the Gabčíkovo structures operation, but they are also resulted from another strong factors, mainly from the forest management system.

Evaluation

The present-day community structure of terrestrial animals reflects the hydropedological condition in this locality in recent years. Changes in the species composition indicate instability of the conditions in this locality. However, the general trend of decreasing species biodiversity (Oniscida, Coleoptera, Carabidae) and an increasing dominance of the mesohygrophilous and tolerant species (Mollusca, Coleoptera, Carabidae) continues, in spite of the fact that it slowed down or stopped in some taxocoenoses (Chilopoda, Coleoptera, Carabidae, Lepidoptera). This later fact indicates that the negative trends of succession have stopped or slowed down. However, it is obvious that this is a consequence of two considerably wetter vegetation periods resulting even in a longer flood in 1997. In general, the taxocoenoses turn toward the taxocoenoses typical of the drier types of the flood-plain forests. In the frame of the locality, the small local differences in moisture and altitude (in the range of 50-60 cm) resulted in changes of spatial distributions of species (Coleoptera). After drying off the locality and a retreat of the hygrophilous species, the more tolerant species originally inhabiting the small elevations moved into the depressions and there substituted for the hygrophilous ones. At the same time the elevation became less populated (Coleoptera, Carabidae). Changes observed in the ornithocoenosis could not be exclusively interpreted as a consequence of the Gabčíkovo structures operation, but they resulted to a considerable degree from the forest management system preferring the poplar lignicultures.

Some restoration measures were undertaken in this locality in 1996. Therefore, some positive changes in the animal community structure can be expected, similar to those already observed in the herbage stratum.

MP 14 Istragov

The area of this monitoring plot was covered by a soft-wood flood-plain forest (Salici-Populetum) and by groups of alders in some places. Closer to the original Danube stream, poplar monocultures were situated in an area in which an intensive clear-cutting took place in the last years of the monitoring. The sandy substrate in this locality reached a thickness exceeding 1 m. Soil moisture ranged from 25 to 35%, the ground water level moved around 2 m under the ground. The Danube damming and limited discharge in its original river bed resulted in a decrease of soil surface layers by approximately 15%. The extensive clear cutting of the poplar monocultures also contributed to a drying off and opening of this area. Up to 1992, the Istragov locality belonged to the strongly hygric habitats of the softwood flood-plain forests (Salici-Populetum myosotidetosum sensu Jurko, [15]). After the Danube damming and decrease of discharge in its original river bed, the ground water level decreased, the shallow depressions filled by water were reduced in number and extent and a xeroseries started in them [22, 32]. The construction of a structure to supply this locality with water has not still been finished. Therefore, succession tended toward communities of a drier variety of the softwood flood-plain forest (Salici-Populetum typicum) even in the lower situated places.

The forest stand as well as the malacocoenosis in the pre-dam conditions had a character similar to those on monitoring plot 10 (Kráľovská lúka). The strongly hygrophilous Zonitoides nitidus was eudominant (up to 77,6% of all individuals), also most other species were strongly hygrophilous (Carychium minimum, Oxyloma elegans, Pseudotrichia ribiginosa). Even the rare wetland species Euconulus alderi occurred in this locality after the Danube damming it was registered only once, in 1994. The number of recorded species of land snails is given in Fig. 19.

While in the recent year the strongly hygrophilous Z. nitidus (in 1993 even 77,6%) belonged to the eudominant species, later it was substituted by the hygrophilous C. minimum (55,8%). Other ripicolous species, P. rubiginosa (11,5%) and, after the Danube damming the euryoecious snail Cochlicopa lubrica (16,0%), belonged to the eudominant species. Changes of the between-year species similarity of the communities are presented in Tab. 19. While in 1994-1996 Cochlicopa lubrica was not found in the samples, it appeared in 1997. Its occurrence can indicate the beginning of a sequence of changes in the malacocoenosis structure which might result from an absence of floods and from reduced moisture in this monitoring plot. Also the hygrophilous forest species Vitrea crystallina appeared in the samples for the first time (as a subdominant species). In contrast to Z. nitidus and C. minimum, V. crystallina is not a typical wetland species. The typical wetland species Euconulus alderi (at present rare in the Podunajská nížina lowland) was found in the studied plot after the Danube damming only once, in 1994. Changes in the percentage of ecological groups of land snails in MP 14 are presented in Fig. 82.

The centipede taxocoenosis consisted of 10 species (Fig. 83), among which the eurytopic species Lithobius forficatus (Fig. 84) and L. mutabilis were eudominat. The typical mesohygrophilous species, L. crassipes, L. aeruginosus and L. curtipes and the hypogeic species Pachymerium ferrugineum (Fig. 84) reached a high dominance. Before the Danube damming, the hygrophilous species L. agilis was also recorded in this locality, in 1991 and 1992.

 

Tab. 20. Between-year species similarity of the community of chilopds in MP 14
1992	62,50
1993	55,56	66,67
1994	55,56	66,67	100
1995	40,00	50,00	60,00	60,00
1996	44,44	55,56	66,67	66,67	87,5
1997	33,33	41,67	50,00	50,00	63,64	70,00
	1991	1992	1993	1994	1995	1996

Drying and opening of this locality has not had a strong effect on the structure of the centipede community. In spite of this, the absence of the strongly hygrophilous species Lithobius agilis since 1993 is a signal that the locality is drying. The numbers of species recorded here after the Danube damming are presented in Fig. 83. The eurytopic L. forficatus and the mesohygrophilous species L. curtipes and L. crassipes (Fig. 84) maintained their dominant position. An increase in dominance of the hypogeic species Pachymerium ferrugineum (Fig. 84), which prefers sandy, but not moist or swampy sites, also indicated a shift in this monitoring plot toward the drier habitat types.

Changes in species similarity of the centipede taxocoenoses in individual years of the monitoring are presented in Tab. 20. The largest changes, as indicated by the lowest values of similarity index (33% and 41%), were between the pre-dam state and the last year of the monitoring (1997). They also confirm the slow shifting of this monitoring plot toward a drier habitat type.

The poor in species number community of terrestrial isopods (Fig. 85) indicated a slow process of drying in 1993-1996. The eurytopic Trachelipus rathkii (Fig. 86), able to tolerate a reduced soil moisture, was eudominant. In the last year of monitoring (1997), the hygrophilous species Porcellium collicolum appeared here reaching a high dominance. This might indicate stabilisation or even an increase in moisture conditions in this locality. It could be a possible positive effect of the artificially simulated floods. The between-year species similarity is presented in the Tab. 21.

In Istragov in 1989 two small hygrophilous carabids - Asaphidion flavipes (47,7%,  Fig. 87) and Bembidion femoratum (15,36%) strongly predominated. Other hygrophilous species Platynus assimilis (15,5%,  Fig. 88), Pterostichus strenuus (5,7%,  Fig. 89), Carabus granulatus (3,6%,  Fig. 90), Patrobus atrorufus (2,4%,  Fig. 91) have also reached a considerable dominance. The presence of the strongly hygrophilous species Europhilus fuliginosus and E. micans (Fig. 92) was of great ecological importance. A relatively high quantitative representation of the tolerant hygrophilous species, before the essential changes of the hydrological regime in this locality, reflected the predominance of strongly sandy soil on a great portion of the monitored plot.

In 1990-1992, the cumulative abundance of carabids decreased to one half of their original values (from 1738 individuals in 1989 to 672, 600 and 592). The largest decrease of abundance was recorded in Asaphidion flavipes (from 829 ind. to 170) and Bembidion femoratum (from 267 ind. to 59). The abundance of almost all other species also decreased to about 2/3 - 1/2 of the values recorded in 1989. In 1991, a sudden increase in abundance of Pterostichus niger (Fig. 93) was observed, which was recorded in other localities two years later.

During 1990-1992 a temporarily stabilised state was created which changed as late as in 1993 by a strong increase in abundance of C. granulatus (from 58 individuals in 1992 to 320 in 1993), P. atrorufus (from 11 ind. in 1992 to 211 ind. in 1993) and P. strenuus (from 24 ind. in 1992 to 222 ind. in 1993). At the same time the cumulative abundance of all carabids increased to 1432 individuals, i. e. more then twice.

After the interruption of sampling in 1994, the cumulative abundance of carabids in 1995 decreased to approximately the level of the year 1992. At the same time, the abundance of Asaphidion flavipes again increased suddenly, both absolutely and relatively (from 159 ind., i. e. 11,1% to 358 ind., i. e. 44,4%). The abundance and dominance of most other species decreased. Only the abundance of the tolerant hygrophilous species Epaphius secalis increased absolutely (from 12 ind. in 1993 to 40 ind. in 1995) (Fig. 94).

In the years 1996-1997, the quantitative proportion of species approximated the state of the period 1990-1991, but the cumulative abundance decreased. The flood in 1997, caused by the backwater from the Danube main river bed under the confluence with the by-pass canal, even made possible a temporal occurrence of some paludicolous species like Agonum moestum, Badister sodalis and Europhilus micans.

An index of the species similarity between the individual annual catches from 1990-1997 compared with the catch from 1989 fluctuated during the major part of the monitored period in the close limits of 32-36% (Fig. 95), with an exception in 1996 (25%) caused by the sudden decrease of the cumulative abundance and resulting in the absence of the rare species. The changes of indices of proportional similarity and abundance similarity were caused, first of all, by larger fluctuations in abundance of Asaphidion flavipes and by a high cumulative abundance in the years 1989 and 1995.

From the point of view of individual ecological groups of Carabids, the taxocoenosis in Istragov seemed to be the least affected of all monitored taxocoenoses. It is remarkable in this connection that, in spite of the general decrease of the ground water level by 1-1.5 m, changes of this level had fluctuation characteristics similar to before the start of the Gabčíkovo structures operation. In other localities the changes of the ground water level were considerably more moderate and fluent. On this basis it seems necessary to prove, in future, whether just sudden short term fluctuations are more significant for the survival of some hygrophilous species than the maintainance of the ground water table at a more or less stable level.

The amphibian communities were monitored in this locality in 1991, 1993, 1994 and 1996. In these years some 1-6 species of amphibians were recorded (Fig. 96). In 1993 only the species R. lessonae was recorded, the species recorded in 1991 did not occur in 1993. In 1994, all species recorded up to that time were observed, except for P. fuscus. In addition, two species of newts indicating the good quality and stability of the water were recorded. T. cristatus dobrogicus, T. vulgaris and B. bombina reached a high dominance. In 1996, 5 species of amphibians were recorded (Hyla arborea, Rana esculenta, R. lessonae, R. arvalis, B. bombina). Species similarity of amphibian community, expressed by means of Jaccard´s index, is presented in the Tab. 22. The results suggest that the community is in the process of stabilisation and has a predisposition to survive in this monitoring plot.

Tab. 22. Species similarity of the taxocoenosis of Amphibia in MP 14

During the seven-year monitoring, altogether 30 species of butterflies were recorded (in the period 1191-1992: 12 and 15 species, while in the period 1993-1997: 11-20 species, Fig. 97). Three xerothermophilous species (Pontia daplidice, Colias erate, Melitea phoebe) which were recorded in this monitoring plot after the start of the Gabčíkovo structures operation indicate a drying of this locality. As a consequence of drying of some parts of the flood-plain forest and timber exploitation, suitable conditions for an invasion of some mostly mesohygrophilous species from the surroundings were created (Zygaena filipendulae, Erynnis tages, Ochlodes vernatus, Papilio machaon, Leptidea sinapsis and Everes argiades).

In 1993, 1994, 1996 and 1997, the bird taxocoenoses were monitored in two partial plots. The first one was in an old willow stand almost without the E₂  stratum, and the second one included a poplar monoculture and the abutting willow stand. Because the monitoring plot was situated downstream, relatively near to the confluence of the original main river bed with the by pass canal, it was also flooded after the start of the structures operation. Before the start of its operation, the flood waves were often very high. In the close vicinity of the monitored plot continuous water tables surrounded by reed stands existed before the Danube damming. However, the character of the proper monitoring plot did not change visibly after the damming.

As follows from the diagrams of the average numbers of species and individuals per one observation in the old willow stand in the breeding season (Fig. 98) and out of the breeding season (Fig. 99), the average number of species decreased moderately and insignificantly, while the average number of individuals decreased more significantly. Species and individuals numbers in the breeding season stabilised in the last two years of the monitoring.

In the second partial plot the E₂ stratum was as well developed on the poplar monoculture as in the abutting old willow stand. Later, both parts were separated by an open strip, with a rich herbage layer which acquired a xerophilous ruderal character after the Danube damming. Before the damming a depression in the poplar monoculture was often filled by water, while after the damming only a little during the simulated floods. In the breeding seasons the average number of species was on the same level during whole monitoring period, but the average number of individuals decreased, Fig. 100. Out of the breeding season, the average number of species decreased very moderately, while the average number of individuals decreased relatively strongly, Fig. 101.

Evaluation

Up to 1992 the locality Istragov belonged among the typical hygric habitats of soft-wood flood-plain forest (Salici-Populetum myosotidetosum sensu Jurko, [14]). After the Danube damming and reduction of discharge in its original main river bed, the ground water level decreased and the shallow depressions filled by water were reduced in size and number. A xeroseries started in them. The planned structure for supplying this area with water has not been finished. Therefore, secondary succession in the lower places also tended to a variety of drier softwood flood-plain forest types (Salici-Populetum typicum). In the terrestrial communities a retreat of the species indicating waterlogged habitats or of more hygrophilous species (Mollusca, Coleoptera) was recorded. These species were substituted by the forest hygrophilous species not requiring a high moisture. This indicates the trend of drying of this locality. The observed changes also resulted from the extensive clear cutting and from the subsequent changes in the shrub and herbage strata. However, in 1996 the trend of aridisation and the retreat of strongly hygrophilous species moderated (Mollusca, Lepidoptera), and the adaptive succession changes were slower (Mollusca, Chilopoda) than the analogous changes in vegetation. The artificial floods probably contributed to the slowing down of these changes.

From the point of view of some ecological groups (Coleoptera, Carabidae), the taxocoenoses in Istragov seemed to be the least affected of all monitored taxocoenoses. It is remarkable in this connection that in spite of the general decrease of the ground water level by 1-1,5 m, the level changes fluctuated similarly to before the start of the Gabčíkovo structures operation. In other localities the changes of the ground water level are considerably more moderate and fluent. On this basis it seems necessary to prove in future, whether sudden short term fluctuations are more significant for the survival of some hygrophilous species than maintaining the ground water table at a stable level.

Area downstream from the by-pass canal not influenced by the project

MP 18 Sporná sihoť

MP 18  Kľúčovec - Sporná Sihoť - this monitoring plot was covered by an elm-ash stand mixed with poplar. The soil moisture at the ground surface ranges from 20 to 30%. Groundwater level oscillated in the depth of 2-3 m under the ground surface.

In summer 1991, thinning was carried out in monitoring plot MP 18. It was accompanied by a strong destruction of the soil surface by heavy machines and resulted in a large opening of the forest stand. This sudden anthropogenic intervention influenced the next evaluation of the malacocoenosis structure, especially from the view of small land snails which are closely bound to the litter and soil surface. In 1992, still before the Danube damming, the species of open landscape and species preferring open stands (Vallonia pulchella, Vallonia costata) began to invade the monitoring plot from the adjacent meadow. These species belonged among eudominant to dominant species in 1992-1993. Before the thinning the community had a mesohygrophilous character and was dominated by the tolerant forest species Alinda biplicata, Monachoides incarnatus, Trichia striolata and by the euryoecious species Punctum pygmaeum. Vallonia costata, a species preferring open forest stand, also occurred in the ecotonal parts of the monitoring plot even before the thinning. The malacocoenoses corresponded by its structure to the conditions of the transitional flood-plain forest.

This monitoring plot belonged to those areas which were flooded at the time of high discharges in the Danube. Because this locality was situated out of the zone with the bank drainage effect, no changes caused by the Gabčíkovo structures were expected here. Thinning of the stand in which the monitoring plot was situated considerably overlapped other influences which might potentially affect the malacocoenosis structure, When compared with the years prior to the thinning, besides the open landscape species, also the tolerant forest species Trichia striolata, Alinda biplicata, Arianta arbustorum began to substitute for the stenotopic forest species Aegopinella nites and Cochlodina  laminata. This was probably a consequence of the mentioned anthropogenic intervention. At the same time, the dominance of the euryoecious species Cochlicopa lubrica and Euconulus fulvus increased. Since 1996 it seems that after the regeneration of the herbage stratum, which was later accompanied by an impressive development of the shrub stratum, regeneration of populations of more stenotopic forest species (especially Semilimax semilimax) began. At present the studied malococoenosis has the character of a drier variety of the soft-wood flood-plain forest to a transitional type flood-plain forest. Thus, since 1990, the polyhygrophilous species (those which reflect also the larger annual fluctuation of soil moisture) were represented only as subrecedent species and they concentrated only in depressions. The moisture changes could also be indicated (but less reliably) by the forest hygrophilous species Vitrea crystallina, whose dominance culminated in 1992 and since that time decreased, until 1997 (perhaps as a consequence of drying processes after the thinning?).

Eight species of centipedes were recorded in the locality Sporná sihoť in pre-dam conditions (Fig. 102). In spite of the thinning and partial clear cutting of the monitoring plot, the centipede community was rich in representation of the mesohygrophilous species Lithobius crassipes and L. curtipes (Fig. 103), which are typical of this area. The eurytopic species L. forficatus maintained its eudominant position (Fig. 103), especially in the ecotonal strip between the forest and abutting meadow which represented a part of this monitoring plot. Another dominant species was L. aeruginosus (Fig. 103).

The locality was strongly influenced in 1991-1992 by a thinning, which was accompanied by the destruction of the soil surface by heavy machines for timber haulage. This intervention doubtless influenced changes in the centipede community in this monitoring plot. After the Danube damming, altogether 11 centipede species were recorded here. The number of species, however, strongly fluctuated in individual years (Fig. 102). The lowest number of species was recorded in the first year after the damming (1993). This low number of species was caused more by an extraordinarily warm and dry summer in 1993 rather then by the damming. Its connection with the Danube damming is, therefore, uncertain. The absence of the hypogeic species Necrophloeophagus flavus in the pitfall traps and sieving samples in 1993 and 1994 also resulted, with a high probability, from the extraordinarily dry summer in those years. The eurytopic species Lithobius forficatus and the mesohygrophilous species, characteristic of the flood-plain forests, L. crassipes, L. curtipes and Pachymerium ferrugineum, were eudominant. The between-year similarity of the centipede taxocoenosis in the years 1991-1997 reached the lowest values between the years 1993-1995 and 1993-1996 (Tab. 23). This was caused by a very dry and warm summer in 1993 and a very high species number recorded in 1995-1996. From the point of view of changes in the centipede taxocoenosis after the Danube damming, it can be concluded that the environmental conditions changed only a little and more or less invariable state was maintained during the whole period of the biota monitoring.

Altogether 6 species of the terrestrial Isopods were recorded in MP 18. Their number oscillated between 3 and 6 in individual years (Fig. 104). Since 1993, the relative abundance of the eurytopic species Trachelipus rathkii gradually decreased (Fig. 105) and, at the same time, the dominance of the mesohygrophilous species Hyloniscus riparius slowly increased. In the years 1996 and 1997, the species Armadillidum vulgare, preferring the drier habitats, disappeared and, contrarily, the hygrophilous species Porcellium collicolum began to occur in those years. These changes indicate a shift of this monitoring plot toward a moister habitat.

The amphibians were monitored in this locality in 1991, 1993, 1994 and 1996. In individual years 1-6 species were recorded: Bombina bombina, Hyla arborea, Rana ridibunda, R. lessonae, R. esculenta and R. arvalis (Fig. 106). The lowest number of species was recorded in 1993. The species composition was very similar in the years 1991 and 1994, as illustrated by the Jaccard index reaching 80%. In both years, the species H. arborea and R. esculenta were eudominant. Values of dominance of individual species, as well the values of the Jaccard index, indicate that the timber exploitation did not affected this taxocoenosis in the monitored period. Obviously, amphibians in this locality were not visibly influenced by the Gabčíkovo structures.

In the taxocoenosis of the small terrestrial mammals, altogether 7 species were recorded in the course of the monitoring, viz. S. araneus, M. minutus, A. flavicollis, A. sylvaticus, C. glareolus, M. arvalis and M. oeconomus. Among the species of the genus Apodemus, a stronger trend in the number of individuals of A. sylvaticus was observed, Fig. 107. Its number decreased, probably as a consequence of increasing humidity and floods. In contrast, A. flavicollis did not react to the condition changes and its abundance was stabile. Similarly, the abundance of C. glareolus did not change and it seemed that the moist habitat was favourable for this species. M. arvalis occurred only in a small number or not at all. This indicated that it did not inhabit this habitat continuously, and it moved here only in the drier periods which are more suitable for this species. Microtus oeconomus also occurred in this locality in the period 1993-1997, but without any trend in its abundance.

During the seven-year period, a total of 34 species of butterflies were recorded in the meadow in the centre of this locality (in the period 1991-1992: 10 and 13 species respectively;in the period 1993-1996: 11 – 22 species a year, Fig. 108). After the Danube damming, almost double the number of species were recorded than in the period 1991-1992. This difference was caused by a mowing of this relatively large monitoring plot, just at the time of occurrence of most butterfly species (turn of spring and summer), and by a lower frequency of collecting before the damming. Among the 17 species found after the Danube damming, xerothermophilous and mesophilous meadow species predominated. Also two hygrophilous species indicating moist swampy meadow - Heteropterus morpheus and Lycaena dispar - and one typical representative of the xerothermophilous habitats of a forest-steppe-like character - Melitea phoebe were found only after the Danube damming. Melitaea phoebe, from the southerly oriented slopes of the Little Carpathians and from surroundings of Štúrovo, probably invaded the area of protective dikes situated in the vicinity of this locality due to the aridisation of the Bratislava surroundings in the last years before the filling of the Čunovo reservoir.

Evaluation

No significant structural changes in the community of terrestrial animals were observed. The observed changes can be considered to be normal fluctuations within the limits of between year dynamics of flood-plain communities in conditions of a natural hydrological regimen. The communities were balanced and stabile. After the flood, which reduced the litter layer and in some places also the humus layer, population size of some epigeic species was reduced. However, the small hygrophilous species survived the floods especially well.

MP 23 Starý les

Monitoring plot MP 23 - Číčov - Starý les - represented a softwood flood-plain forest (Salici-Populetum) with admixed alders flooded every year. In many places, water stayed in small depressions all year round. After the start of the Gabčíkovo structures operation, the monitoring plot was flooded every year by water from the Danube main river bed and by the seeping ground water. The character of this monitoring plot did not change during the monitored period (1993-1997). The locality was situated downstream the confluence of the original Danube river bed with the tail race canal, thus out of the influence zone of the Gabčíkovo structures.

The studied malacocoenosis in a softwood flood-plain forest (as. Salici-Populetum phragmito-caricetosum) in the vicinity of the dead arm was characterised by a predominance of the hygrophilous species Carychium minimum, Zonitoides nitidus, Succinea putris and of the forest hygrophilous species Vitrea crystallina.

The land snail community was characteristic for this type of the softwood flood-plain forest also after the start of the Gabčíkovo structures operation. The development of this community was balanced during the whole monitoring period. Due to well functioning reparation mechanisms, the species of the wetland malacocoenosis are well adapted to the conditions of cyclical climax.

Monitoring of centipedes in this locality started as late as after the Danube damming in 1993. In the period 1993-1997, 12 species were recorded altogether (Fig. 109). The presence of the strongly hygrophilous species Lithobius agilis indicated the wetland character of this locality. The eurytopic species L. forficatus, mesohygrophilous species L. crassipes, L. curtipes and L. aeruginosus were eudominant (Fig. 110). The stability of the high soil moisture was indicated by the constant occurrence of L. agilis, whose dominance reached even 24% in 1997 (Fig. 110). Among other rare species, the presence of the termophilous hypogeic species Clinopodes linearis is to be mentioned. C. linearis occurred on elevated and drier margins of this locality. During the whole monitored period no significant changes were recorded in the centipede taxocoenosis (Tab. 25). This monitoring plot preserved a very natural character. It belongs to the most valuable remainders of the willow-alder flood-plain forest which have been preserved in the Slovak part of the Danube flood-plain.

Taxocoenosis of the terrestrial isopods was balanced. Except for the year 1993, the between-year species similarity reached 100% (Tab. 26). The mesohygrophilous species Hyloniscus riparius and the hygrophilous species Porcellium collicolum are characterised by a high dominance (Fig. 111). Values of dominance of the eurytopic species Tracheliplus rathkii continuously decreased from 1993 (Fig. 111). The presence of the hygrophilous species Porcellium collicolum, Porcelio scaber and Haplophthalmus mengiii confirm the invariably high soil moisture. The high dominance of Trachelipus rathkii in 1993 in both MP 18 (87%) and MP 23 (82%) probably resulted from the very dry and warm summer in those years.

Altogether 5 species of amphibians (Hyla arborea, Bombina bombina, Rana ridibunda, Rana esculenta, Rana lessonae) were recorded in this locality. Except for 1993, when only one species was registered, in all other years 4-5 species were observed (Fig. 113). The similarity of the species composition 1994-1997 was relatively high (Jaccard´s index 80%, Tab. 27). This indicated that no significant changes were in the taxocoenoses.

Tab. 27. Between-year species similarity of amphibia in MP Starý les.

During the five-year monitoring (1993-1997) altogether 36 butterfly species were recorded in this locality. During the investigation the species number gradually decreased due to the dike reconstruction, Fig. 114.. The low number of individuals of Pontia daplidice in the last years of the monitoring indicates a slow down of aridisation of this locality. A comparison of the present and the earlier state was not possible because the butterflies were not studied in this locality in pre-dam conditions.

Evaluation

The taxocoenoses of the terrestrial animals can be characterised as the typical communities of a regularly flooded softwood floodplain forest of the association Salici-Populetum with a minimal proportion of the eurypotent species invading into this monitoring plot in irregular periods from a nearby stand of poplar cultivars. The taxocoenoses with a high proportion of hygrophilous species and with a balanced state exhibit a highly original natural character and adaptation to the typical cyclic climax  of the floodplain forests. The recorded changes (Mollusca, Lepidoptera) lay within limits of usual fluctuations. The between -year fluctuations of abundance and dominance are minimal and are a natural manifestation of the dynamics of populations of individual species.

From the point of view of nature protection, the locality itself and its zoocoenoses deserve special attention because the species diversity of hygrobionts in this locality is the most similar to the natural conditions among all monitored localities. In addition, several species registered on red lists occur here. Because of the original character, this locality deserves increased protection and an extention of the protection zone of the Nature Reserve Lyon up to this locality. First of all it is necessary to prevent cutting the trees and other negative anthropogenic activities.

References

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Figures

Biological monitoring sites		Fig. 1. Species number of terrestrial Gastropods in the MP1 in the zone of increase of the ground water level.		Fig. 2. Fluctuation of Chilopod species number during the monitoring in MP 1
Fig. 3. Changes in Chilopod dominance in MP 1		Fig. 4. Fluctuation of species number of terrestrial Isopod in MP 1 in individual monitoring years.		Fig. 5. Changes in dominance of terrestrial Isopods in MP 1
Fig. 6. Changes in species number of Amphibia in MP 1		Fig. 7. Abundance changes in Apodemus sylvaticus in MP 1		Fig. 8. Abundance changes in Microtus arvalis in MP 1.
Fig. 9. Species number changes of Rhopalocera in MP 1		Fig. 10. Changes in species number of Rhopalocera in MP 1		Fig. 11. Changes in species number of Chilopods on MP3
Fig. 12. Changes in Chilopods dominance on MP 3		Fig. 13. Changes in species number of terrestrial Isopods during the monitoring on MP 3		Fig. 14.Changes in dominance of terrestrial Isopod in MP 3
Fig. 15 Changes in species number of Rhopalocera in MP 3		Fig. 16. Number of waterfowl population census in January		Fig. 17. Species number of waterfowl on the whole Slovak Danube section during the winter censuses
Fig. 18 Species number of waterfowl during the January census on the original Danube canal and the Gabčíkovo structures		Fig. 19. Number of terrestrial Gastropods on the monitoring plots in the by-pass zone		Fig. 20. Percentage of ecological groups of terrestrial Gastropods in MP 6
Fig. 21 Changes in species number of Chilopods in individual years in MP 6		Fig. 22 Changes in the chilopod dominance in MP 6.		Fig. 23. Changes in species number of terrestrial Isopods in MP 6
Fig. 24 Dominance changes of terrestrial isopods in MP 6		Fig. 25. Abundance of Agonum moestum in MP 6		Fig. 26. Abundance of Oxypselaphus obscurus in MP 6
Fig. 27. Abundance of Patrobus atrorufus in MP 6		Fig. 28. Abundance of Platynus assimilis in MP 6		Fig. 29. Abundance of Carabus granulatus in MP 6
Fig. 30. Abundance of Pterostichus niger  in MP 6		Fig. 31. Similarity indices of the carabid community in MP 6 in the period 1990-1997 compared with the state in 1989		Fig. 32. Changes in species number of Amphibians in MP 6
Fig. 33. Changes in abundance of Apodemus sylvaticus in the monitoring plot MP 6		Fig. 34. Changes in abundance of Sorex araneus and S. minutus in the monitoring plot MP 6		Fig. 35. Changes in species number of Rhopalocera in MP 6
Fig. 36 . Average number of species and individuals of birds in the breeding period in MP 6		Fig. 37. Average number of species and individuals in the MP 6 out of the breeding period		Fig. 38. Changes in species number of Chilopods in MP 7 during the monitoring period
Fig. 39 Changes in species number of terrestrial Isopods in MP 7 during the monitoring period		Fig. 40. Changes of dominance of terrestrial Isopods in MP 7		Fig. 41. Percentage of ecological groups of terrestrial Gastropods in the MP 9
Fig. 42. Changes of species number of Chilopods in individual years in MP 9		Fig. 43. Changes in Chilopod dominance in MP 9		Fig. 44. Changes in species number of terrestrial Isopods in individual years of monitoring in MP 9
Fig. 45 Changes in dominance of terrestrial Isopods in MP 9		Fig. 46 Abundance of Platynus assimilis in MP 9		Fig. 47 Abundance of Patrobus atrorufus in MP 9
Fig. 48 Abundance of Oxypselaphus obsucurus in MP 9		Fig. 49 Abundance of Asaphidion flavipes in MP 9		Fig. 50 Abundance of Carabus granulatus in MP 9
Fig. 51 Abundance of Pterostichus strenuus in MP 9		Fig. 52 Abundance of Pterostichus melanarius in MP 9		Fig. 53 Abundance of Pterostichus niger in MP 9
Fig. 54. Cumulative abundance of Carabids in the monitoring plots in the by-pass zone		Fig. 55. Similarity index of Carabids in monitoring plots in the by-pass zone		Fig. 56. Changes in species number of amphibians in MP 9
Fig. 57. Changes in abundance of A. flavicollis in MP 9		Fig. 58. Changes in abundance of C. glareolus in MP 9		Fig. 59. Changes in species number of Rhopalocera in MP 9
Fig. 60 Percentage of ecological groups of terrestrial Gastropods in the MP 10		Fig. 61. Changes in species number of Chilopods during the monitoring in MP 10		Fig. 62. Changes in dominance of Chilopods in MP 10
Fig. 63. Changes in species number of terrestrial Isopods during the monitoring in MP 10		Fig. 64 Changes in dominance of terrestrial Ispodos in MP 10		Fig. 65. Abundance of Agonum moestum in MP 10
Fig. 66. Abundance of Patrobus atrorufus in MP 10		Fig. 67. Abundance of Platynus assimilis in MP 10		Fig. 68. Abundance of Pterostichus strenuus  in MP 10
Fig. 69. Abundance of Carabus granulatus  in MP 10		Fig. 70. Abundance of Oxypselaphus obscurus  in MP 10		Fig. 71. Abundance of Europhilus micans in MP 10
Fig. 72. Abundance of Asaphidion flavipes  in MP 10		Fig. 73. Indices of similarity of the Carabid taxocoenosis in individual years compared with the year 1987 in MP 10 (in 1998, 1994 and 1995 the taxocoenoses were not monitored)
Fig. 74. Abundance of Epaphius secalis  in MP 10		Fig. 75. Abundance of Trechus quadristriatus  in MP 10		Fig. 76 Changes in species number of amphibians in MP 10
Fig. 77. Abundance changes in Clethrionomys. glareolus in MP10		Fig. 78. Abundance changes in Asaphidion flavicollis in MP 10		Fig. 79. Change in species number of Rhopalocera in MP 10
Fig. 80 Average number of species and individuals in the breeding season (TMB - poplar monoculture without E2 , TMS - poplar monoculture with E2 , LL - floodplain forest)
Fig. 81 Average number of species and individuals out the breeding season (TMB - poplar monoculture without E2 , TMS - poplar monoculture with E2 , LL - floodplain forest)
Fig. 82. Percentage of ecological groups of terrestrial Gastropoda in MP 14		Fig. 83. Fluctuation of species number of Chilopoda in MP 14 in the course of monitoring		Fig. 84. Dominance changes of individual species of Cilopoda in MP 14
Fig. 85. Fluctuation of species number of terrestrial Isopods in MP 14 in the course of monitoring		Fig. 86. Dominance changes of individual species of terrestrial Isopods in MP 14		Fig. 87. Abundance of Asaphidion flavipes  in MP 14
Fig. 88. Abundance of Platynus assimilis  in MP 14		Fig. 89. Abundance of Pterostichus strenuus  in MP 14		Fig. 90. Abundance of Carabus granulatus  in MP 14
Fig. 91. Abundance of Patrobus atrorufus  in MP 14		Fig. 92. Abundance of Europhilus micans  in MP 14		Fig. 93. Abundance of Pterostichus niger  in MP 14
Fig. 94. Abundance of Epaphius secalis  in MP 14		Fig. 95. Similarity indices of the Carabid taxocoenosis in MP 14 in 1990-1997 compared with the state in 1989 (in 1994 the locality was not monitored)
Fig. 96. Changes in species number of Amphibia in MP 14		Fig. 97. Changes in species number of Rhopalocera in MP 14		Fig. 98. Average number of species and individuals per one observation in the breeding season in MP Istragov - old willow stand
Fig. 99 Average number of species and individuals per one observation out of the breeding season in MP Istragov - old willow stand. (symbols as in fig. 98)
Fig. 100. Average number of species and individuals per one observation out of the breeding season in MP Istragov - poplar monoculture/willow stand. (symbols as in fig. 98)
Fig. 101. Average number of species and individuals per one observation out of the breeding season in MP Istragov - poplar monoculture/willow stand. (symbols as in fig. 98)
Fig. 102. Changes in species number of Chilopods in MP 18 in individual years of the monitoring		Fig. 103. Changes in dominance of Chilopods in MP 18		Fig. 104. Changes in species number of the terrestrial Isopods in MP 18 in individual years of the monitoring
Fig. 105. Changes in dominance of the terrestrial Isopods in MP18		Fig. 106. Changes in species number of Amphibia in MP 18		Fig. 107. Changes in abundance of A. sylvaticus in MP 18
Fig. 108. Changes in species number of Rhopalocera in MP 18		Fig. 109. Changes in species number of Chilopoda in MP 23 in individual years of monitoring		Fig. 110. Changes in dominance of Chilopods in MP 23
Fig. 111. Changes in dominance of terrestrial Isopods in MP 23		Fig. 112. Changes in species number of terrestrial Isopods in MP 23 in individual years of monitoring		Fig. 113. Changes in species number of Amphibia in MP 23
Fig. 114. Changes in species number of Rhopalocera in MP 23		Schissler dead arm (Hungary), September 1998