Invasion by Heracleum Mantegazzianum in Different Habitats - 711
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Journal of Vegetation Science 6: 711-718, 1995 © IAVS; Opulus Press Uppsala. Printed in Sweden - Invasion by Heracleum mantegazzianum in different habitats - 711 Invasion by Heracleum mantegazzianum in different habitats in the Czech Republic Pyšek, Petr1,3 & Pyšek, Antonín2 1Institute of Applied Ecology, University of Agriculture Prague, CZ-28163 Kostelec nad âern mi lesy, Czech Republic; Fax +42 203 97500; 2Husova 342, CZ-43982 Vroutek, Czech Republic; 3Present address: Botanical Institute, Academy of Sciences of the Czech Republic, CZ-25243 PrÛhonice, Czech Republic Abstract. Heracleum mantegazzianum, a tall forb from the the process itself represents a serious threat to native western Caucasus invaded several different habitats in the species, it also provides us with an unparallelled oppor- Czech Republic. The relation between invasion success and tunity for ecological studies (Vitousek et al. 1987). Many type of recipient habitat was studied in the Slavkovsk les of those studies have focused upon the history and hilly ridge, Czech Republic. The vegetation of 14 habitat types dynamics of invading species at various hierarchical occurring in an area of ca. 25 km2 was analysed using phyto- sociological relevés, and the invasion success of Heracleum levels. Rejmánek (1989) and KornaÊ (1990) provided (in terms of number of localities, area covered and proportion reviews of many of these studies and references to of available area occupied) was recorded separately in each of particular case studies. However, invading species and them. Site conditions were expressed indirectly using Ellenberg target communities cannot be studied independently indicator values. The hypothesis tested was that Heracleum because the species-community interactions are what spreads in the majority of vegetation types regardless of the determine the outcome of invasion. properties of the recipient vegetation. Recently, the most rapid progress in this field seems Community invasibility appeared to be affected by site to be associated with a shift in focus to the critical conditions and the composition of the recipient vegetation. interactions of invader and target community (Crawley The species is not found in acidic habitats. Disturbed habitats with good possibilities of dispersal for Heracleum seeds are 1987; Rejmánek 1989; Hester & Hobbs 1992; Lodge more easily invaded. Communities with a higher proportion of 1993; DeFerrari & Naiman 1994). Moreover, progress phanerophytes and of species with CS (Competitive/Stress- in the ecological study of invasions appears to be essen- tolerating) strategy were more resistant to invasion. The inva- tial for understanding what determines the dynamics of sion success was bigger in sites with increased possibilities of ecological communities (Lodge 1993). spread for Heracleum diaspores. Communities invaded by Invasive species are not only able to affect the struc- Heracleum had a lower species diversity and a higher indica- ture and function of an ecosystem (Versfeld & van tor value for nitrogen than not-invaded stands. It appears that Wilgen 1986; Witkowski 1991; Vitousek & Walker 1989; species contributing to community resistance against invasion Vitousek 1990) but can also reduce the biodiversity of of Heracleum, or capable of persisting in Heracleum-invaded stands, have similar ecological requirements but a different communities and landscapes, both at the local and re- life strategy to the invader. gional scales. They may cause considerable problems in landscape management (de Waal et al. 1994) and nature conservation (Usher 1988). Keywords: Alien plant species; Community invasibility; Land- The present study is aimed at analysing the effect of scape section; Plant trait; Site condition; Species diversity; the type of recipient vegetation on the invasion of Strategy. Heracleum mantegazzianum, an alien species from Asia and one of the most noxious European alien weeds (Py‰ek 1991, 1994; de Waal et al. 1994). The hypothesis Nomenclature: Tutin et al. (1964-1980). tested here is that, owing to its biological and ecological properties, Heracleum is capable of entering the major- ity of vegetation types in the landscape regardless of the Introduction properties of the recipient vegetation. Since Elton’s (1958) classic study, biological inva- sions have been receiving increasing attention (Py‰ek 1995) and in the last decades they have become one of the most attractive research fields in ecology (Drake & Mooney 1989; di Castri et al. 1990; Lodge 1993). While 712 Py‰ek, P. & Py‰ek, A. Material and Methods with willows forming the shrub layer. Road verges (0.5 %). Verges of smaller roads and paths Research area and habitats studied in the open landscape are included. Peat bogs (0.02 %). Acid, Sphagnum-dominated habi- The study area is located in the Slavkovsk les hilly tats. ridge (western Bohemia, Czech Republic) in the sur- roundings of the village of Prameny, 11 km east of the Study species town of Horní Slavkov (50° 09' N, 12° 48' E). The geo- logical substrate consists of Palaeozoic granite; the area Heracleum mantegazzianum (Apiaceae) is a mono- is situated at an altitude of 600 - 650 m a.s.l. and has a carpic perennial with a thick taproot, a stout stem attain- moderately moist climate with a mean annual tempera- ing 5.5 m height, large pinnate leaves (up to 3 m long) ture of 5 - 6 °C and an annual precipitation of 650 mm and usually 7 - 10 umbels bearing oval-elliptical, broadly (Veseck et al. 1957). Originally, the area was covered winged fruits 9 - 11 mm in size (Tutin et al. 1964-1980). by acidophilous beech and oak forests (Miky‰ka 1972). The species spreads exclusively by seed, and its seed At present, the landscape is managed moderately inten- production per plant may reach several tens of thou- sively; it is used mostly for silvicultural and agricultural sands (Neiland 1986; Brondegaard 1990). In the study purposes, including cattle breeding. The following habi- area, the species germinates in early spring (April), the tat types were distinguished in an area of ca. 25 km2 – flowering period starts in May, fruits appear in July and listed according to the proportion of the area they cover are shed from September onwards. which is given in parentheses. A summary of the species The species is native to the western Caucasus where composition is presented in App. 1. Ecological charac- it occurs in the upper forest belt of the southern slopes, teristics of the habitat types studied are given in Table 1. mainly in meadows, clearings and forest margins (Mladenova 1950). It was introduced in the 19th century Spruce forests (28.9 %). Forest plantations with spruce into the Czech Republic, as well as in other European as the main component of the tree layer. countries (Lundström 1984; de Waal et al. 1994). See Mown meadows (19.5 %). Mesophilous meadows used further Py‰ek (1991) for the historical dynamics of the for hay and mown twice a year. distribution of the species in the area, and Py‰ek (1994) Pastures (15.8 %). Used for cattle grazing. for an analysis of the ecology of its invasion. There are Moist grasslands (13.0 %). Sites in the vicinity of water two main reasons for the efforts in the Czech Republic courses, some of them periodically flooded. and elsewhere in Europe to eradicate the species from Fields (6.1 %). Arable land used for plantations of fod- infested areas (Williamson & Forbes 1982; de Waal et der and corn. This habitat type was excluded from al. 1994): (1) the replacement of native vegetation – the the data analysis because expressing its vegetation consequences of which have been discussed by Py‰ek characteristics in the way used for other habitats (1991) – and (2) injuries to the human skin caused by could be misleading due to the variety of crops phototoxic substances (Drever & Hunter 1970). In the cultivated. study area, Heracleum rapidly attains dominance in Urban sites (4.5 %). Most of the area of the village of invaded communities, with a cover of 50 -100 %. Part Prameny harbours vegetation of various succes- of the competitive superiority of Heracleum mante- sional ages occurring in man-made sites. gazzianum – the largest forb in central Europe – over Dry grasslands (3.7 %). Relatively thermophilous sites other plants is ascribed to its huge size and its ability to on south-exposed slopes, often on rock outcrops. shade the surrounding vegetation with its huge ground Road ditches (3.3 %). Representatives located in open leaves. The large seed-set (Neiland 1986; Brondegaard areas, further called ‘sunny ditches’ (1.7 %) and 1990) with dispersal promoted by water, wind and those shaded by neighbouring vegetation, called human-related factors (Jehlík & Lhotská 1970) also ‘shaded ditches’ (1.6 %) were treated separately. contributes to its rapid spread into various vegetation Water courses (2.3 %). Sites along the brook flowing types. through the village and along small streamlets in the open landscape. Sampling Forest margins (1.1 %). Ecotonal zones between forests and adjacent vegetation, mostly meadows, pastures Field research was conducted in 1992. Vegetation and arable land. was sampled using Braun-Blanquet’s seven-point scale Birch woodlands (0.7 %). Patches of deciduous or mixed 5, 4, 3, 2, 1, + and r (e.g. Westhoff & van der Maarel forests with birch and pine forming the tree layer. 1978). Five relevés were made in each habitat type. The Willow scrub (0.6 %). Moist sites, periodically flooded, size of the sample plot was 5 m × 5 m, except in those - Invasion by Heracleum mantegazzianum in different habitats - 713 Table 1. Characteristics of habitats in the study area and overview of their invasion by Heracleum mantegazzianum. Ellenberg indicator values were used to characterize site conditions: light (L), moisture (M), nitrogen (N), and soil reaction (R). Total number of species recorded (S-tot), mean species number per relevé (S-mean), and species diversity (Shannon index H') are presented as community characteristics. Habitats which are disturbed by human activities (DIST) and those in which dispersal of diaspores of Heracleum is probably promoted (DISP) are indicated by ‘yes’.