©2017 Institute of Parasitology, SAS, Košice DOI 10.1515/helm-2017-0005 HELMINTHOLOGIA, 54, 1: 48 – 67, 2017 Soil nematodes in alpine meadows of the Tatra National Park (Slovak Republic) L. HÁNĚL Biology Centre CAS, Institute of Soil Biology, Na Sádkách 7, CZ-370 05 České Budějovice, Czech Republic, E-mail: [email protected] Article info Summary Received September 13, 2016 The assemblages of soil nematodes were studied at fi ve alpine meadow sites, 1763 – 2200 m. a. Accepted December 19, 2016 s. l., in the Tatra National Park in the Slovak Republic. A total of 110 species were distinguished, 19 species were recorded in the Slovak Republic for the fi rst time. The interesting new records are the occurrence of Sphaeronema alni at the elevation of 2003 m a . s. l. and the populations of Coomansus menzeli at four sites. The total numbers of species at individual sites varied from 45 to 72. The most abundant nematode genera were Aglenchus, Plectus, Acrobeloides, Paratylenchus, Eudorylaimus, Helicotylechus, and Aphelenchoides. The total mean abundance ranged from 704 to 2054 x 103ind.m-2 and the total mean biomass from 442 to 1531 mg.m-2. The lowest values of the Maturity Indices (∑MI, MI) were found at the highest elevation. The signifi cantly highest values of the Plant Parasite Index were at the lowest elevation. The mean values of the of the Enrichment Index varied from 16.3 to 38.4, the mean values of the Structure Index from 64.1 to 85.4. The Structure metabolic footprints were signifi cantly greater at the lowest elevation than at the highest elevation. Cluster Analysis and Principal Component Analysis performed on species presence and absence, genera abundance and genera metabolic footprints showed nematode assemblages at sites of high- er elevations different from those at sites of lower elevations. Keywords: soil nematodes; diversity, maturity; soil food web; alpine meadow Introduction Nematode faunas in grasslands are diverse and abundant (Wa- silewska, 1979; Boag & Yeates, 1998; de Goede & Bongers, 1998) Grassland (including shrub steppes, savannahs and prairies) is and may in fact control total grassland primary production (Scott the potential natural vegetation on 33 – 45 million km2 covering et al., 1979; Verschoor, 2001). Nematological studies of Central about a quarter of the Earth’s land surface (Lauenroth, 1979; European grasslands are numerous but relatively few alpine and Bardgett & Cook 1998; Bonan, 2008). Besides climatically de- subalpine meadows above timberline were investigated for nema- termined grasslands there are semi-natural and agricultural todes. Nevertheless, the available data showed a great variety of grasslands on land formerly covered by forests and mires. The nematode faunas in these habitats. total area of semi-natural grasslands in the Central and Eastern Since about 1940’s it is known that the nematode abundance in European countries is estimated at around 7 million hectares, a alpine meadows of Central Europe usually varies in order of sev- part of them was ploughed and is currently being restored (Jonge- eral millions of individuals per square metre, rarely is greater than pierová, 2008). ten millions or less than one million of individuals (Franz, 1950). 48 The Table 13 in his book deals with 42 species at 17 sites and it is of nematode taxa tends to be lower at higher elevations. Tempera- interesting that 14 species belonged to the order Dorylaimida and ture can be limiting factor for nematodes in cold climates, however this order had the greatest species richness at 14 sites. The large Hoschitz & Kaufmann (2004a) pointed out that the apparent tem- predator Coomansus zschokkei occurred at fi ve sites. Vinciguerra perature sensitivity may be the result of indirect effects, e.g. by (1988) found 31 species in three alpine meadows in Italian Alps, resource availability. 16 species belonged to the order Dorylaimida and C. zschokkei A drop in the soil moisture does not usually limit alpine and subal- occurred in two meadows. Gerber (1991) reported on 33 species pine meadows and the surplus of water leads to the formation of at alpine habitats in Austria but only three were from the order peat meadows and peat bogs accompanied by complex changes Dorylaimida. in nematode faunas (Háněl, 2015a). At lower elevations in grass- The extensive survey of nematode communities in mountain lands on soils susceptible to desiccation the total number of nem- grasslands from Romania carried out by Popovici (1998) showed atode taxa can be reduced, nematode communities differ in xero- much more diverse nematode faunas. However, a part of nema- philic and mesophilic grasslands (Ciobanu & Popovici, 2015) and todes could be determined to the genus level only. The synthe- a general trend of increasing nematode richness with decreasing sis of the data published by Popovici & Ciobanu (2000) therefore altitude can be reversed. gives the numbers of taxa (genera + species) as a measure of The behaviour of various community indices across alpine-subal- nematode taxonomic richness. They found that the alpine grass- pine grasslands is little known. Popovici & Ciobanu (2000) found lands (above 2000 m a. s. l.) had 46 – 50 nematode taxa. Aglen- that the Shannon’s index of generic diversity (H’) of the 36 sites chus agricola, Filenchus spp. and Paratylenchus spp. were the ranged between 2.38 and 3.47, without clear differences between dominant nematodes and plant feeders composed 62 – 66 % of in- the different nematode communities, and in eight alpine-subalpine dividuals. In the subalpine grasslands at the 1700 – 1850 m a. s. l. sites between 2.65 and 3.07. The ratios of hyphal to bacterial feed- (according to Popovici, 1998) the genera Filenchus, Acrobeloides, ing nematodes showed a constant preponderance of the bacterial Gracilacus, Paratylenchus, Plectus and Rotylenchus were preva- feeding group. It is consistent with results of Hoschitz & Kaufmann lent. Plant feeding nematodes composed 28 – 52 % of individuals (2004b). The minimum values of the Maturity index (MI) on the and the number of taxa varied from 60 to 88. The percentage of southern faces of the alpine summits were statistically signifi cant omnivores in subalpine and alpine stands varied from 3 to 12 % but did not seem biologically relevant (Hoschitz & Kaufmann, while at lower elevations they composed 5 – 41 % of individuals. 2004a). On the hand, multivariate analyses were effective in dis- Authors concluded that the environmental variables, such soil pH, tinguishing different nematode communities. total nitrogen, humus content, exchangeable bases and soil type This study gives a detailed survey of nematode assemblages at could explain the variations in the composition of nematode com- fi ve high-mountain meadows in the Tatra National Park (the west- munities in grasslands, but no single factor could be selected as ern part of the Carpathians) in the Slovak Republic. The main aims being of overriding importance. of the study are as follows: Hoschitz & Kaufmann (2004a) studied soil nematodes (families (i) To evaluate the species composition of nematode faunas. and trophic groups) of Alpine summits in Austria and especially (ii) To study the abundance of nematode genera and trophic differences between plots facing the four compass directions. The groups and the community indices based on abundance. results showed that the nematode abundances on the south or (iii) To estimate the biomass and metabolic footprints of nematode east facing side were signifi cantly higher than on the northern assemblages. and western sides. The aspects of the sites also affected nem- Preliminary results and methods were presented by Háněl (2015b) atode families and trophic groups and nematode diversity (num- at the 13th Central European Workshop on Soil Zoology and they ber of families) and abundance responded to differences in soil will be published in the Proceedings of the workshop. Therefore microclimate. Authors therefore concluded that nematode as- some topics such as selection of indices and statistical methods semblages are potentially good bioindicators of climate change. and problems with determination of some nematode populations The most abundant family at two Carex fi rma swards (2214 and are not discussed in detail in this paper. 2255 m a. s. l.) were bacterivorous Cephalobidae. This contrasts with the alpine sites studied by Popovici & Ciobanu (2000) where Material and Methods plant feeding nematodes dominated. In another study Hoschitz & Kaufmann (2004b) found that bacterivorous Acrobeloides were The research was carried out in alpine parts of the High Tatra most abundant nematodes at Caricetum curvulae sedge mat Mountains (the Tatra National Park), in the Slovak Republic, above (2595 m a. s. l.) while in an alpine pasture with dominant Poaceae the timberline. The areas studied has cold alpine climate, parent (1961 m a. s. l.) the most abundant nematodes were plant feeding/ rocks are mostly granites. Soil types are rankers, podzols and plant-associated Rotylenchus and Tylenchidae. This indicates that cambic podzols, acid and rich in organic matter. vegetation is also an important factor determining taxonomic and Nematodes were studied at fi ve meadow sites in two mountain trophic composition of nematode assemblages while the richness valleys of two lake catchments: 49 (i) the Furkotská dolina Valley of the Vyšné Wahlenbergovo Lake microscope Leica Leitz DMRB equipped with N PLAN 100x/1.25 (VWL) with soil-poor catchment. immersion oil objective and transmitted light interference contrast, V1: 49° 09’ 46.8’’ N, 20° 01’ 41.5’’ E; 2200 m a. s. l.; SW facing on maximum magnifi cation 1600x. As a basis for the determination of a 15° slope; Seslerietum distichae Krajina 1933 (dominant Ore- nematodes monographs written by Brzeski (1998) and Andrássy ochloa disticha (Wulf.) Link, Luzula alpinopilosa (Chaix) Breistr.) (2005, 2007, and 2009) were used plus publications dealing with (Matějka, 2015); soil pH(H2O) 4.7, soil carbon 10.4 %. individual taxonomic groups mostly cited in the books mentioned V2: 49° 09’ 22.3’’ N, 20° 01’ 40.4’’ E; 2003 m a.