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Article

The Importance of Railway Lines for the Composition of Vegetation in Agricultural Landscapes: A Case Study

by
Jan Winkler
1,*,
Marta Smékalová
1,
Yentriani Rumeta Lumbantobing
2,
Jana Červenková
1,
Wiktor Sitek
2 and
Magdalena Daria Vaverková
2,3
1
Department of Plant Biology, Faculty of AgriSciences, Mendel University in Brno, Zemědělská 1, 613 00 Brno, Czech Republic
2
Department of Sustainable Construction and Geodesy, Institute of Civil Engineering, Warsaw University of Life Sciences-SGGW, Nowoursynowska 159, 02 776 Warsaw, Poland
3
Department of Applied and Landscape Ecology, Faculty of AgriSciences, Mendel University in Brno, Zemědělská 1, 613 00 Brno, Czech Republic
*
Author to whom correspondence should be addressed.
Land 2026, 15(4), 523; https://doi.org/10.3390/land15040523
Submission received: 31 January 2026 / Revised: 9 March 2026 / Accepted: 20 March 2026 / Published: 24 March 2026
(This article belongs to the Special Issue Species Vulnerability and Habitat Loss (Third Edition))
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Abstract

Railway corridors create persistent linear habitats embedded within intensively managed agricultural landscapes and can simultaneously support native biodiversity and facilitate the spread of undesirable taxa. We evaluated vegetation composition across five habitat types associated with railway line no. 250 (Havlíčkův Brod–Tišnov, Czech Republic): railway yard, railway embankment, railway land, field margin, and adjacent arable land. Vegetation was recorded using phytosociological relevés (10 m2) at four localities during three surveys in the 2021 growing season. In total, 83 plant taxa were identified, with pronounced differences among habitat types. Species richness and vegetation structure were highest in railway embankments, railway land, and field margins, whereas the railway yard and arable land exhibited lower diversity consistent with high disturbance intensity and substrate constraints. Canonical correspondence analysis distinguished habitat-affinity assemblages, indicating strong habitat filtering along the railway–agriculture gradient. Classification by origin and invasion status showed that non-native and invasive taxa were concentrated predominantly in railway embankments and adjacent habitats, suggesting elevated propagule pressure and potential spread into surrounding farmland. Colonization success (ICS) and colonization potential (ICP) indices indicated that railway-associated habitats can host taxa with high establishment capacity, contributing to successional stability within the corridor. These findings highlight railways as multifunctional elements of agricultural landscapes that require integrated vegetation management to balance biodiversity benefits with operational safety and invasive species risk.

1. Introduction

The development of railway transport began in the early 19th century and rapidly expanded across Europe and North America, becoming one of the most transformative infrastructure systems of the industrial era [1,2]. By the late 19th and early 20th centuries, railway networks had become key drivers of economic growth, urbanization, and landscape transformation worldwide [2]. Over time, railway systems evolved technologically and energetically, with substantial implications for global energy consumption and carbon emissions [3]. Railway transport continues to play a central role in global and regional supply chains [4].
In the Czech lands, railway construction intensified during the 19th century under the Austro-Hungarian administration and subsequently formed the structural backbone of regional connectivity [5]. The expansion of railway infrastructure substantially reshaped urban morphology, particularly in major cities such as Prague [6]. The long operational history of these corridors has resulted in persistent and spatially extensive anthropogenic habitats embedded within agricultural and urban landscapes. These long-established corridors therefore represent not only transport infrastructure but also structurally distinct ecological environments.
Railways represent a linear transport infrastructure that exhibits a number of shared habitat characteristics across geographical regions and climatic zones [7]. These habitats are exclusively anthropogenic in origin, as the original vegetation is removed and the substrate is subsequently technically modified for the construction of railway embankments and track installation [8,9]. Railway construction, as well as subsequent operation and maintenance, significantly alters environmental conditions, both through initial landscape disturbance and through long-term disturbances [10]. Understanding the processes of recovery and development of plant communities following disturbance therefore represents a long standing and fundamental topic of ecological research [10,11].
Railway environments create a wide spectrum of microhabitats suitable for plant species with differing ecological requirements [12]. These linear habitats are characterized by extreme environmental conditions, including dry, stony substrates exposed to intense solar radiation and often burdened by elevated levels of pollution, particularly heavy metals [13]. Rocky and exposed surfaces are typically colonized by xerophilous and thermophilous species, whereas mesophilous and hygrophilous species find suitable conditions in more shaded and moister microdepressions with less permeable surfaces [12]. These stress factors significantly limit the survival of many plant species [14,15].
Individual phases of railway construction and operation strongly influence the structure and distribution of vegetation. Plant communities in railway environments are additionally exposed to continuous disturbances resulting from intensive passenger and freight transport as well as from management interventions such as mowing or herbicide application, which are aimed at suppressing undesirable vegetation [16]. Vegetation control is necessary primarily to ensure railway traffic safety and to maintain the functionality of railway infrastructure. The main objectives include maintaining track quality and stability, preserving ballast bed stability, protecting wooden sleepers, ensuring worker safety, and preventing fires caused by dry vegetation [17]. Due to their linear nature and specific management regimes, railways also contribute to landscape fragmentation.
In agricultural landscapes, fragmentation is associated with a range of changes in biodiversity structure, community composition, abiotic conditions, and biotic interactions [18,19]. Fragmentation affects functional groups of organisms [20,21], alters biological community composition, and modifies competitive relationships among populations.
Changes in abiotic and biotic environmental characteristics induced by fragmentation may act as selective pressures on organism populations [22,23]. Fragmentation is often associated with reduced plant population sizes and limited availability of pollinators [24], which may lead to reduced genetic diversity and increased inbreeding depression [25]. Nevertheless, many studies indicate that organism responses to fragmentation can also be positive and in some cases contribute to increased population resilience across a wide range of environmental conditions [19].
Landscape fragmentation also creates habitats with differing living conditions, which is reflected in changes in vegetation structure and ecosystem functioning. Increased environmental heterogeneity may allow colonization by new plant species and lead to enhanced biodiversity. In contrast, areas subject to strong human influence are often characterized by homogeneous environmental conditions. Fragmentation of anthropogenic landscapes may, therefore, under certain circumstances, represent a tool for supporting biodiversity [26].
Transport infrastructure represents an important component of fragmentation and strongly influences both environmental conditions and landscape character [13]. Railway networks pass through various land use types, ranging from agricultural and urban areas to sites of high natural value or relatively undisturbed forests, thereby contributing to habitat fragmentation and facilitating the spread of nonnative species into more natural ecosystems. As a result, they can significantly affect the species composition of surrounding, particularly adjacent, communities [27].
Railway habitats are characterized by the co-occurrence of native and introduced ruderal plant species that colonize not only tracks but also embankments, sidings, and turntables. These habitats function as ecological corridors for plants with specific environmental requirements [28]. Transport infrastructure, including railways, also plays a major role in the dispersal of plant diaspores into new areas [29]. Railway corridors thus often facilitate the crossing of natural physical and ecological barriers and function as migration routes for plants [30,31].
The railway network represents a complex engineering infrastructure whose construction and operation require extensive landscape modification. The pronounced transformation of original habitats, combined with the linear character of railway lines, leads to fragmentation of habitat conditions. The heterogeneity of the resulting habitats arises from both single and recurrent anthropogenic disturbances, which simultaneously represent opportunities and constraints for the occurrence of individual plant species. Fragmented habitats along railway lines are characterized by different species composition and vegetation structure and promote the spread of plant taxa onto arable land in the surrounding agricultural landscape. The objectives of this study were to: (i) evaluate the species composition and vegetation structure of fragmented habitats associated with the railway network, (ii) identify plant taxa characteristic of these fragmented habitats, (iii) assess the representation of individual taxa in relation to vegetation succession, and (iv) evaluate the potential of railway lines as corridors for the spread of invasive plant species into the surrounding agricultural landscape.

2. Materials and Methods

2.1. Study Area

Railway line no. 250, in the section between Havlíčkův Brod and Tišnov, constitutes an important part of the rail connection between Prague and Brno crossing the Vysočina region in the Czech Republic (CR). This section traverses the dissected terrain of the Bohemian-Moravian Highlands and the South Moravian Highlands, which substantially influences its technical design and operational characteristics. The line follows a predominantly winding alignment with numerous curves and gradients, typically limiting operating speeds to 70–100 km/h. It is used for both long-distance passenger transport, particularly express services connecting Brno, Havlíčkův Brod, and Prague, and regional passenger transport serving towns and municipalities in Vysočina and South Moravia. Freight transport also plays a significant role, as this section functions as an important diversionary route during service interruptions on the main railway corridor routed via Česká Třebová. Historically, the line was constructed in the second half of the nineteenth century and progressively electrified during the twentieth century. At present, it is undergoing gradual modernization, particularly at railway stations and within signaling systems, while retaining the character of a technically demanding yet strategically important connection between Bohemia and Moravia [32].
The studied railway section is in the Žďár nad Sázavou District within the Vysočina Region and geomorphologically belongs to the Křižanov Highlands. The surrounding landscape forms part of a potato-producing agricultural region at an elevation of 525–580 m above sea level. Mean annual precipitation reaches approximately 640 mm and the mean annual air temperature is 7.1 °C. The parent material consists mainly of granites, syenites, and schists. Dominant soil types are Cambisols (Haplic, Calcaric, and Arenic), with smaller occurrences of Gleyic Luvisols and Gleyic Chernozems. These soils are generally non-skeletal to moderately skeletal, moderately to deeply developed, and show low susceptibility to waterlogging or drought, although they are vulnerable to acidification. The study area lies on gentle slopes of 3–7° with all-aspect exposure [33,34].

2.2. Selected Sites and Habitats

Within the monitored section of the railway line, four localities were selected to enable the assessment of five habitat types characterized by contrasting ecological conditions. The first locality is situated in the cadastral area of Velká Losenice (49.5563208° N, 15.8375858° E), the second in the cadastral area of Hamry (49.5688586° N, 15.9142503° E), the third in the cadastral area of Žďár nad Sázavou (49.5378403° N, 15.9493483° E), and the fourth in the cadastral area of Vatín (49.5291392° N, 15.9626936° E). At each locality, five habitats differing in the degree of railway influence, disturbance intensity, and conditions for vegetation development were identified. The selection of locations was guided by the effort to find locations in real conditions from the same climatic region and on the same railway line with comparable traffic intensity, which are directly adjacent to arable land. These habitats are shown in Figure 1 and Figure 2. Summary characteristics of the habitats are given in Table 1.
Railway yard
The habitat in question is situated near the tracks. The substrate underwent substantial modification during the construction of the railway, comprising predominantly stony and gravel-rich material with minimal humus content. The track’s structural design comprises layered components meticulously engineered to ensure stability, uniform load distribution, and effective drainage. The superstructure consists of steel rails mounted on sleepers, predominantly concrete, embedded in a ballast layer of crushed stone. The ballast contributes to the provision of elasticity, spatial stability, and facilitation of precipitation runoff. Beneath the ballast lies the substructure, consisting of a base layer of crushed aggregate or gravel-sand material that separates the superstructure from the formation layer and enhances load-bearing capacity. The formation layer represents a modified surface of the original soil or embankment material and may be stabilized using lime, cement, or reinforced with geotextiles. The subgrade comprises natural soil or artificially placed fill material. Additional structural components include geotextiles and geogrids that prevent mixing of layers and increase long-term stability, as well as drainage systems ensuring water removal. The entire subgrade system is designed to withstand high dynamic loads, adverse climatic conditions, and to minimize deformation. The substrate is strongly desiccated and intentionally drained. The width of the station on the single-track section was 3–4 m. Vegetation is controlled through irregular herbicide applications.
Railway embankment
This habitat is located adjacent to the tracks and represents a transition zone between the railway yard and the surrounding landscape. The embankment consists of compacted fill material, most commonly soil, gravel-sand mixtures, or crushed stone, forming the foundation for both the railway superstructure and substructure. Its construction must meet strict requirements for load-bearing capacity, stability, and drainage to withstand dynamic loads generated by train traffic and weather conditions. Embankment slopes are typically treated to reduce erosion and may be stabilized by vegetation, stone material, or technical measures. From an ecological perspective, railway embankments represent specific anthropogenic habitats characterized by dry, sun-exposed, and often nutrient-poor conditions. The width of the site ranged from 6 to 12 m, depending on local terrain and infrastructure conditions. Vegetation management is limited, with occasional and localized removal of woody plants.
Railway land
This habitat is located on the central slope of the embankment and forms a continuous inclined surface at all four localities. The area between the railway embankment and adjacent land is generally part of the railway protection zone. From a legal perspective, it represents land designated for railway operation and maintenance, including tracks, embankments, and adjacent areas required to ensure transport safety. This habitat was only moderately affected by terrain modifications during railway construction. In technical terms, it often includes toe ditches used for embankment drainage, as well as access or maintenance strips enabling infrastructure servicing. The width of the site ranged from 30 to 60 m, depending on local terrain and infrastructure conditions. Vegetation is not actively managed, and remnants of native vegetation occur locally.
Field margin
This habitat is located directly adjacent to arable land and represents the boundary between cultivated fields and neighboring land parcels. It is not used for crop production but is influenced by agricultural activities or sporadically affected by agricultural disturbances such as soil cultivation or herbicide drift. The impact of these disturbances is irregular and spatially limited.
Arable land
This habitat is situated on actively cultivated agricultural land. Different crops were monitored across the four localities. At the first locality, winter barley (Hordeum vulgare) was grown (sowing 15 September 2020, harvest 8 August 2021), followed by winter oilseed rape (Brassica napus var. napus). At the second locality, bread wheat (Triticum aestivum, sown 4 October 2020) was cultivated, followed by winter barley (Hordeum vulgare). At the third locality, triticale (×Triticosecale) was sown in autumn 2020. Due to a wet autumn and winter, the crop was damaged and resown with spring barley (Hordeum vulgare) on 12 April 2021, with harvest on 22 August 2021. In spring 2022, the field was planned to be sown with maize (Zea mays). At the fourth locality, maize (Zea mays, sown 9 May 2021, harvested 5 October 2021) was cultivated, with spring barley (Hordeum vulgare) planned for spring 2022. Vegetation on arable land is regularly disturbed by agricultural practices, including herbicide application.

2.3. Vegetation Assessment Methodology

The analyzed data were collected at individual sites using the phytosociological relevé method. At each of the four localities, five habitats were surveyed, with three relevés of 10 m2 recorded within each habitat. The size of the phytocoenological relevé was chosen in accordance with the commonly used methodology used for herbaceous vegetation. Total vegetation cover was first visually estimated for each relevé, after which individual plant species were identified, and their cover was estimated as percentages. Vegetation sampling was conducted three times during the 2021 growing season, in May, at the turn of July and August, and at the turn of September and October. During the field research, all safety rules set by the line operator were followed. Scientific names of plants were taken from the Pladias flora and vegetation database [36,37].
Recorded plant species were classified into groups according to their origin and invasion status using the classification framework of Pyšek et al. [38]. Taxa were assigned to the following categories: (i) invasive archaeophyte, (ii) transient archaeophyte, (iii) naturalized archaeophyte, (iv) transient archaeophyte or neophyte, (v) naturalized archaeophyte or neophyte, (vi) invasive neophyte, (vii) transient neophyte, (viii) naturalized neophyte, (ix) cultivated taxon, and (x) native taxon.
Colonization ability was assessed as an additional criterion. Indices describing the colonization ability of taxa within the Czech flora were derived from the framework developed by Prach et al. [39]. These indices were calculated using a database comprising 21 successional series, including both primary and secondary succession, originating from bare substrates. The database includes 1013 vascular plant taxa recorded in 2817 phytosociological relevés representing a wide range of habitats across the CR and successional stages spanning 1 to 150 years. Three indices were distinguished.
The successional age optimum represents the median time in years since disturbance at which a given taxon occurs during succession. Optimum values were defined within a range of 1 to 50 years. For taxa whose optimum exceeded 50 years and could not be estimated more precisely due to a limited number of successional stages, the value was set to 75 years.
The index of colonization success (ICS) expresses the frequency of occurrence of a taxon within the successional series database. ICS values were standardized to a scale ranging from 1, indicating absence, to 9, indicating a high frequency of occurrence across successional stages.
The index of colonization potential (ICP) integrates both the biological traits of taxa and their abundance in the landscape. ICP values range from 1, indicating low colonization ability, to 9, indicating high colonization ability.

2.4. Statistical Analysis

Vegetation cover data and associated railway habitats were evaluated using multivariate ecological data analyses. Species-specific cover values recorded at the studied localities were analyzed using multivariate statistical methods. The selection of an appropriate analytical approach was based on the length of ecological gradients estimated by detrended correspondence analysis (DCA). The gradient length obtained from DCA was 2.34 standard deviation units (SD), indicating that the response data have a compositional structure and that the use of a linear method is recommended. Based on this result, canonical correspondence analysis (CCA) was subsequently applied, using centering and standardization by species. The statistical significance of the ordination results was tested using a Monte Carlo permutation test with 999 permutations. All multivariate analyses and related calculations were performed using Canoco version 5.0 software [40].

3. Results

Across all monitored habitats and localities, a total of 83 plant taxa were recorded. Species richness differed markedly among habitat types.
Railway yard
A total of 23 taxa were recorded. The taxa with the highest cover were Equisetum arvense, Artemisia vulgaris, Tripleurospermum inodorum, Galium verum, and Aegopodium podagraria.
Railway embankment
A total of 40 taxa were recorded. The taxa with the highest cover were Festuca rubra, Arrhenatherum elatius, Calamagrostis epigejos, Aegopodium podagraria, and Carduus acanthoides.
Railway land
A total of 37 taxa were recorded. The taxa with the highest cover were Epilobium hirsutum, Arrhenatherum elatius, Festuca rubra, Aegopodium podagraria, and Anthriscus sylvestris.
Field margin
A total of 31 taxa were recorded. The taxa with the highest cover were Arrhenatherum elatius, Agrostis capillaris, Calamagrostis epigejos, Holcus lanatus, and Tanacetum vulgare.
Arable land
A total of 20 taxa were recorded. The highest cover was represented by cultivated crops: Hordeum vulgare, ×Triticosecale rimpaui, Triticum aestivum, Brassica napus, and Zea mays. Among weed species, the highest cover was observed for Poa annua, Polygonum aviculare, Galium aparine, and Galeopsis tetrahit.
Based on CCA, the spatial arrangement of plant taxa in relation to the railway-associated habitats was identified. The CCA results were statistically significant at α = 0.001, indicating a high explanatory power of the analysis. According to the ordination diagram (Figure 3), the taxa were grouped into eight distinct assemblages. The detailed assignment of individual taxa to these groups is provided in Table 2. The names and abbreviations of the plant taxa are listed in Table S1.
In Table 2, plant taxa are divided into eight groups based on the results of the CCA. The most taxa occur together in two habitats: railway embankment and railway land. The conditions of these two habitats are very similar for these taxa, so the listed species occur in both habitats.
On the contrary, the fewest taxa occur in the habitats railway embankment, railway land and field margin. Agricultural practices allow the occurrence of only a limited number of plant taxa.
From a dispersal perspective, taxa associated with the railway yard and adjacent railway habitats are relevant because they may spread along the corridor and into surrounding farmland. Conversely, taxa associated with arable land and the railway yard indicate potential exchange from cropland into the track zone.
With respect to origin and invasion status, native plant taxa were predominant across most of the studied habitats. Native taxa accounted for more than 50% of total cover in all habitats except arable land. An exception was the arable land habitat, where cultivated crops dominated, representing non-native introduced taxa and taxa grown in cultivation. In the railway embankment, railway land, and field margin habitats, archaeophyte and invasive taxa were also well represented in addition to native species. Neophyte invasive taxa were recorded exclusively at the railway embankment habitat. The distribution of plant taxa groups according to origin and invasion status is presented in Figure 4.
Based on successional age optima, taxa with optima of 21 years and older predominated in the railway embankment, railway land, and field margin habitats. In the arable land habitat, taxa with unknown successional optima, represented by cultivated crops, predominated, followed by taxa with optima of up to 5 years. In the railway yard habitat, the representation of taxa varied markedly among individual sampling periods, with taxa characterized by optima of up to 20 years being predominant. Especially at the railway embankment and railway land sites, the proportion of this group of taxa is higher than 60%. The distribution of plant taxa groups according to successional age optima is shown in Figure 5.
The index of colonization success in successional stages (ICS) reached values of 7, 8, and 9 for the majority of taxa across all habitats. Only in the arable land habitat did taxa with unknown ICS values predominate, represented by cultivated crops. The distribution of plant taxa groups according to ICS is shown in Figure 6.
With respect to the index of colonization potential (ICP), taxa with ICP values of 7, 8, and 9 were predominantly represented in the railway embankment habitat. In the railway land and field margin habitats, taxa with ICP values of 5, 7, and 8 predominated. In the arable land habitat, taxa with unknown ICP values, represented by cultivated crops, were dominant. The distribution of plant taxa groups according to ICP is shown in Figure 7.

4. Discussion

Human activities create a range of specific habitats that plants use to maintain viable populations. Railway lines represent synanthropic habitats that host characteristic plant assemblages [41,42,43]. The current extent of ecosystems in which human society has become the dominant ecological force is a defining feature of the Anthropocene [44,45]. Within this broader context, the interpretations presented below refer specifically to the studied railway corridor embedded in an intensively managed agricultural landscape of Central Europe.
Our results indicate that the least favorable conditions for vegetation occurred in the railway yard and arable land habitats. Both habitats are characterized by a high frequency of repeated disturbances, which was reflected in a lower number of recorded plant taxa. In addition, the gravelly track substrate is poorly suited for plant growth, which is consistent with the lowest vegetation cover. The railway yard habitat was characterized by taxa with very low cover (e.g., Avenella flexuosa, Potentilla erecta, Vicia hirsuta) and by taxa dispersing predominantly by wind (Epilobium adenocaulon, Taraxacum sect. Taraxacum). Taxa from adjacent habitats also penetrated into the railway yard (Equisetum arvense, Artemisia vulgaris, Tripleurospermum inodorum, Galium verum, Aegopodium podagraria). These taxa reach higher cover outside the railway yard, but may pose operational problems for railway infrastructure. According to Torstensson [17], plant residues and organic material can fill voids in the ballast bed and impair drainage. During winter, freezing water can then cause track displacement or deformation. Vegetation biomass on tracks also increases the risk of wheel slip and lengthens braking distance, thereby compromising operational safety.
The arable land habitat is primarily used for crop production (Hordeum vulgare, ×Triticosecale rimpaui, Triticum aestivum, Brassica napus, Zea mays), and their occurrence corresponded to their cultivation periods. In addition to crops, arable land supported weed species (Alopecurus pratensis, Chenopodium album, Convolvulus arvensis, Poa annua, Polygonum aviculare, Veronica polita) that were able to persist under repeated agricultural disturbances such as herbicide applications and soil cultivation.
The railway embankment and railway land habitats showed very similar vegetation composition and, according to the CCA results, their plant taxa formed a shared group. Dominant taxa were perennial grasses (Arrhenatherum elatius, Calamagrostis epigejos, Dactylis glomerata, Festuca rubra, Poa pratensis, Trisetum flavescens). Perennial dicotyledonous forbs were also strongly represented (Aegopodium podagraria, Anthriscus sylvestris, Artemisia vulgaris, Carduus acanthoides, Cirsium arvense, Epilobium hirsutum, Galium album, Solidago canadensis, Tanacetum vulgare, Urtica dioica).
The field margin habitat exhibited vegetation of a similar character to the two previous habitats, but with a higher representation of perennial dicotyledonous forbs (Fragaria vesca, Galium intermedium, Hylotelephium telephium, Linaria vulgaris, Verbascum nigrum, Veronica chamaedrys) and shrubs (Rubus sect. Rubus). The taxa present are capable of tolerating irregular agricultural disturbances. The railway embankment and railway land habitats provided more favorable conditions for plant growth. Similar patterns of increased functional and phylogenetic diversity along abandoned railway corridors have been reported from agricultural landscapes in Poland, where railway sites supported higher ecological heterogeneity than adjacent grasslands [46].
The railway embankment, railway land, and field margin habitats provided more favorable conditions for plant growth. The soil environment supported higher biomass production, and the intensity of repeated disturbances was lower.
Linear habitats associated with railway lines create conditions for vegetation of a more steppe-like character, although this is not a representative vegetation type for the studied locality. Habitats created by human society are a widely distributed component of Central European landscapes [47]. According to Woźnica et al. [48], plants have adapted to the specific conditions of railway lines, which has promoted the development of distinctive traits. Vegetation can also create additional problems, specifically the clogging of drainage systems by biomass, which leads to water retention and can cause failure of railway embankments or slopes [17].
Of particular interest are plant taxa capable of establishing across multiple habitat types, for which within corridor spread can be expected.
Taxa able to establish in both arable land and the railway yard include primarily annual weeds (Capsella bursa-pastoris, Myosotis arvensis, Viola arvensis) as well as some perennial species (Hypericum perforatum). Herbicides are applied in both habitats, which increases the risk of herbicide-resistant plant populations. Studies from Scandinavian railway systems have demonstrated that herbicide spraying from railway trains can result in drift deposition affecting adjacent non-target vegetation and drainage systems [49]. According to Torsterson et al. [17], herbicide resistance has been reported particularly in Equisetum arvense, Galium spp., and Conyza canadensis. Movement of plant populations between arable land and the railway yard is limited by the presence of additional linear habitats with more competitively dominant vegetation that prevents long-term persistence of annual species. Nevertheless, resistance risk should be systematically monitored and anti-resistance strategies implemented [50].
Arable land directly borders the field margin habitat, and taxa capable of establishing in both habitat types also occur in this interface. These are predominantly important arable weeds (Apera spica-venti, Galeopsis tetrahit, Galium aparine, Elymus repens). Field margins associated with railway lines may therefore function as sources of weed infestation in crops. This habitat is most often maintained by mowing, and regular vegetation management is essential to limit propagule production by economically problematic weed taxa.
Some taxa were able to establish in the railway embankment and railway land habitats as well as directly in the railway yard (Pimpinella saxifraga, Senecio viscosus, Tripleurospermum inodorum). For these taxa, spread from railway managed land into the track zone can be expected, where they may complicate railway operations. Tripleurospermum inodorum, a nonnative taxon, may have used railway infrastructure as a corridor for spread into new regions.
The perennial taxa Agrostis capillaris, Cerastium arvense, and Holcus lanatus were recorded in three habitat types (railway embankment, railway land, field margin). In the studied locality, railway-managed land functioned as a source habitat for several perennial taxa. However, whether similar patterns occur in other agricultural landscapes likely depends on local management intensity, connectivity, and propagule pressure.
The development of railway networks and associated transport has substantially contributed to the spread of nonnative and exotic plant species [28,51]. However, not all railway habitats are equally favorable for alien taxa. The highest occurrence of invasive neophytes was recorded in the railway embankment habitat, specifically Echinops sphaerocephalus, Lupinus polyphyllus, and Solidago canadensis. Invasive archaeophytes were most evident in the railway embankment, railway land, and field margin habitats and included Arrhenatherum elatius and Cirsium arvense. In our study system, habitats located between the track zone and arable land showed the highest occurrence of invasive taxa. Their role as sources of further spread should be evaluated in relation to regional landscape configuration and management regimes.
The specific spectrum of invasive taxa can vary substantially among regions. For example, railway lines in South Moravia (CR) have recorded Amaranthus retroflexus, Digitaria sanguinalis, and Epilobium adenocaulon [52].
Historically, railway transport enabled several taxa, such as Geranium purpureum, to expand into new regions across Europe and subsequently colonize additional urban habitats [53,54]. In our study, this taxon was recorded in the railway land habitat, although with low representation and low cover. Hutniczak et al. [15] point out that in areas surrounded by non-forest communities, succession on disused railways proceeds differently, with species belonging to the classes Artemisietea vulgaris and Molinio-Arrhenatheretea dominating.
Májeková et al. [55] emphasized two major factors influencing vascular plant species richness, mean annual temperature and freight train volume. Native species richness was higher at sites with temperatures below 8.65 °C, whereas nonnative species were more numerous in areas with intensive freight traffic (>7640 trains). Vegetation succession showed similar species composition in the railway embankment, railway land, and field margin habitats, where an optimum age of 21 years and older was identified. This pattern suggests the development of relatively stable grass–forb communities maintained under moderate disturbance regimes and limited soil turnover. In contrast, vegetation in the arable land and railway yard habitats represented earlier successional stages shaped by recurrent mechanical and chemical disturbances. Comparable successional trajectories have been documented in abandoned railway areas in Poland, where species composition was strongly influenced by surrounding land use and proximity to forest habitats [15]. These findings collectively indicate that railway-associated habitats may function as semi-stable successional systems whose structure depends not only on disturbance intensity but also on landscape context and connectivity.
Vegetation in the railway embankment, railway land, and field margin habitats showed high values of the colonization success index and the colonization potential index. This suggests that the present taxa have a strong capacity to establish not only within these habitats but potentially also on surrounding land, especially under pronounced changes in agricultural management.
Railway lines represent specific ecological migration corridors for plants under distinctive living conditions [43,48]. Conditions on railway infrastructure and adjacent land allow the persistence of very small populations of diverse taxa. Such small populations can serve as starting points for subsequent spread at the regional scale.
Propagules originating from various local and regional sources can establish in railway habitats. These taxa then enter ecological interactions, adapt to new environments, and can substantially influence biodiversity at broader spatial scales [16].
The results also indicate that railway infrastructure creates fragmented habitats that enable the long-term persistence of plant taxa that are not fully typical of the region’s original vegetation. Fragmentation can generate lasting and often difficult to predict consequences, including local increases in the abundance of certain species. Long time scales are required to fully understand the effects of landscape fragmentation [18,56]. Fragmentation also affects seed dispersal, particularly in wind dispersed taxa [57]. Our data indicate that wind dispersed taxa are more strongly represented in habitats exposed to stronger disturbance, particularly the railway yard (Epilobium adenocaulon, Taraxacum sect. Taraxacum). However, taxa with this dispersal mode also occurred in other habitats (Carduus acanthoides, Cirsium arvense, Senecio viscosus).
Although habitat fragmentation is often associated with negative impacts, many organism responses can be positive and can increase population resilience across a wide range of environmental conditions [19]. As reported by Hurajová et al. [58], habitat fragmentation can counteract homogenization in agricultural landscapes and enable the long-term persistence of species rich ecosystems. Vegetation can provide a number of ecological functions, including reducing the susceptibility of embankments to fire, stabilizing slopes, mitigating runoff, supporting biodiversity, and storing carbon [59,60], and investigating the interactions between land management practices, vegetation species composition, and climatic factors is also essential, as it can provide valuable insights for increasing the resilience of vegetative slope protection systems [61,62].
Respecting spontaneous vegetation can substantially support biodiversity [63]. Vegetation in agricultural landscapes also provides multiple ecosystem functions and increases the biological value of diverse anthropogenic habitats, including arable land [64,65], fallows and grasslands [66], vineyards [67], and landfills [68]. Conservation should therefore move beyond the traditional dichotomy of “wild” versus “managed” nature [65] and adopt hybrid conservation approaches that recognize high biodiversity in habitats often shaped by human society [69,70,71,72].
Railway lines have a strongly linear character, and within these lines habitat fragmentation occurs due to differing single event and repeated anthropogenic disturbances. Human society thus creates specific ecosystems composed of distinct plant taxa that provide specific ecosystem functions for other organisms and thereby substantially influence the structure of agricultural landscape ecosystems.

5. Conclusions

Railway lines fragment agricultural landscapes and create distinct linear habitats with contrasting vegetation composition and structure. The study leads to the following conclusions:
(i)
Vegetation composition and structure differed markedly among habitat types, reflecting variation in substrate properties and disturbance regimes. Species richness was highest in the railway embankment and railway land habitats, which exhibited highly similar species composition, whereas the lowest richness occurred in the railway yard and arable land habitats.
(ii)
Most taxa were restricted to a single habitat type, and only a limited number of species were able to establish and persist across multiple habitats, indicating strong habitat filtering along the railway–agriculture gradient.
(iii)
Vegetation associated with railway corridors displayed characteristics consistent with mid- to late-successional communities under moderate disturbance and introduced structurally and functionally diverse elements into otherwise homogeneous agricultural landscapes, thereby enhancing ecosystem-level heterogeneity.
(iv)
Railway embankment, railway land, and field margin habitats supported a higher representation of nonnative and invasive taxa, including Arrhenatherum elatius, Cirsium arvense, Echinops sphaerocephalus, Geranium purpureum, Lupinus polyphyllus, and Solidago canadensis.
Under specific management and landscape conditions, these habitats may act as secondary sources for the spread of undesirable taxa into adjacent farmland. Certain plant taxa may directly influence railway operation and track stability, including Equisetum arvense, Artemisia vulgaris, Tripleurospermum inodorum, Galium verum, and Aegopodium podagraria. Other species, such as Capsella bursa-pastoris, Myosotis arvensis, and Viola arvensis, may spread into arable land and function as agricultural weeds, including potentially herbicide-resistant populations. Overall, railway corridors function simultaneously as biodiversity-supporting elements and as potential dispersal pathways for problematic taxa. Effective management therefore requires an integrated approach that balances ecological value with operational safety and agricultural protection.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/land15040523/s1, Table S1. Plant taxon name and abbreviation.

Author Contributions

Conceptualization, J.W. and M.S.; methodology, J.Č. and J.W.; validation, M.D.V. and J.W.; formal analysis, J.W.; investigation, J.W. and Y.R.L.; resources, W.S. and J.W.; data curation, M.S. and J.W.; writing—original draft preparation, J.Č. and J.W.; writing—review and editing, M.D.V. and M.S.; visualization, W.S. and J.W.; supervision, J.Č. and W.S.; project administration, J.W.; funding acquisition, J.W. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The data are not available to the public in order to preserve the originality of the data, for the successful completion of Marta Smékalová’s studies.

Conflicts of Interest

The authors declare no conflicts of interest.

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Land 15 00523 g001
Figure 1. Definition of the monitored railway section (modified image) [35].
Figure 1. Definition of the monitored railway section (modified image) [35].
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Figure 2. Location of vegetation relevés within the Žďár nad Sázavou study area.
Figure 2. Location of vegetation relevés within the Žďár nad Sázavou study area.
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Figure 3. Ordination diagram illustrating the relationships between recorded plant taxa and selected habitats based on CCA results (pseudo-F = 1.9; p = 0.001). Explanation of abbreviations: AegPoda—Aegopodium podagraria, AetCyna—Aethusa cynapium, AgrCapi—Agrostis capillaris, AchMill—Achillea millefolium, AloPrat—Alopecurus pratensis, AngArch—Angelica archangelica, AntSylv—Anthriscus sylvestris, ApeSpic—Apera spica-venti, ArrElat—Arrhenatherum elatius, ArtVulg—Artemisia vulgaris, AveFlex—Avenella flexuosa, BarVulg—Barbarea vulgaris, BraNapu—Brassica napus, Bryophy—Bryophyta, CalEpig—Calamagrostis epigejos, CapBurs—Capsella bursa-pastoris, CarAcan—Carduus acanthoides, CenCyan—Centaurea cyanus, CenScab—Centaurea scabiosa, CerArve—Cerastium arvense, CirArve—Cirsium arvense, ConArve—Convolvulus arvensis, DacGlom—Dactylis glomerata, EchSpha—Echinops sphaerocephalus, EchVulg—Echium vulgare, ElyRepe—Elymus repens, EpiAden—Epilobium adenocaulon, EpiHirs—Epilobium hirsutum, EquArve—Equisetum arvense, EroVern—Erophila verna, FalConv—Fallopia convolvulus, FesRubr—Festuca rubra, FraVesc—Fragaria vesca, GalTetr—Galeopsis tetrahit, GalAlbu—Galium album, GalApar—Galium aparine, GalInte—Galium intermedium, GalVeru—Galium verum, GerPurp—Geranium purpureum, GerRobe—Geranium robertianum, HolLana—Holcus lanatus, HorVulg—Hordeum vulgare, HylTele—Hylotelephium telephium, HypPerf—Hypericum perforatum, CheAlbu—Chenopodium album, LeoHisp—Leontodon hispidus, LeuVulg—Leucanthemum vulgare, LinVulg—Linaria vulgaris, LupPoly—Lupinus polyphyllus, MelOffi—Melilotus officinalis, MyoArve—Myosotis arvensis, MyoRamo—Myosotis ramosissima, OenBien—Oenothera biennis, OnoVicii—Onobrychis viciifolia, PimSaxi—Pimpinella saxifraga, PoaAnnu—Poa annua, PoaPrat—Poa pratensis, PolAvic—Polygonum aviculare, PotErec—Potentilla erecta, RubSect—Rubus sect. Rubus, RumAcet—Rumex acetosa, SenVisc—Senecio viscosus, SenVulg—Senecio vulgaris, SilLati—Silene latifolia, SolCana—Solidago canadensis, TanVulg—Tanacetum vulgare, TarSect—Taraxacum sect. Taraxacum, TraOrie—Tragopogon orientalis, TriHybr—Trifolium hybridum, TriInod—Tripleurospermum inodorum, TriFlav—Trisetum flavescens, TriAest—Triticum aestivum, TusFarf—Tussilago farfara, UrtDioi—Urtica dioica, VerNigr—Verbascum nigrum, VerArve—Veronica arvensis, VerCham—Veronica chamaedrys, VerPoli—Veronica polita, VicCrac—Vicia cracca, VicHirs—Vicia hirsuta, VioArve—Viola arvensis, ZeaMays—Zea mays, ×TrRimp—×Triticosecale rimpaui).
Figure 3. Ordination diagram illustrating the relationships between recorded plant taxa and selected habitats based on CCA results (pseudo-F = 1.9; p = 0.001). Explanation of abbreviations: AegPoda—Aegopodium podagraria, AetCyna—Aethusa cynapium, AgrCapi—Agrostis capillaris, AchMill—Achillea millefolium, AloPrat—Alopecurus pratensis, AngArch—Angelica archangelica, AntSylv—Anthriscus sylvestris, ApeSpic—Apera spica-venti, ArrElat—Arrhenatherum elatius, ArtVulg—Artemisia vulgaris, AveFlex—Avenella flexuosa, BarVulg—Barbarea vulgaris, BraNapu—Brassica napus, Bryophy—Bryophyta, CalEpig—Calamagrostis epigejos, CapBurs—Capsella bursa-pastoris, CarAcan—Carduus acanthoides, CenCyan—Centaurea cyanus, CenScab—Centaurea scabiosa, CerArve—Cerastium arvense, CirArve—Cirsium arvense, ConArve—Convolvulus arvensis, DacGlom—Dactylis glomerata, EchSpha—Echinops sphaerocephalus, EchVulg—Echium vulgare, ElyRepe—Elymus repens, EpiAden—Epilobium adenocaulon, EpiHirs—Epilobium hirsutum, EquArve—Equisetum arvense, EroVern—Erophila verna, FalConv—Fallopia convolvulus, FesRubr—Festuca rubra, FraVesc—Fragaria vesca, GalTetr—Galeopsis tetrahit, GalAlbu—Galium album, GalApar—Galium aparine, GalInte—Galium intermedium, GalVeru—Galium verum, GerPurp—Geranium purpureum, GerRobe—Geranium robertianum, HolLana—Holcus lanatus, HorVulg—Hordeum vulgare, HylTele—Hylotelephium telephium, HypPerf—Hypericum perforatum, CheAlbu—Chenopodium album, LeoHisp—Leontodon hispidus, LeuVulg—Leucanthemum vulgare, LinVulg—Linaria vulgaris, LupPoly—Lupinus polyphyllus, MelOffi—Melilotus officinalis, MyoArve—Myosotis arvensis, MyoRamo—Myosotis ramosissima, OenBien—Oenothera biennis, OnoVicii—Onobrychis viciifolia, PimSaxi—Pimpinella saxifraga, PoaAnnu—Poa annua, PoaPrat—Poa pratensis, PolAvic—Polygonum aviculare, PotErec—Potentilla erecta, RubSect—Rubus sect. Rubus, RumAcet—Rumex acetosa, SenVisc—Senecio viscosus, SenVulg—Senecio vulgaris, SilLati—Silene latifolia, SolCana—Solidago canadensis, TanVulg—Tanacetum vulgare, TarSect—Taraxacum sect. Taraxacum, TraOrie—Tragopogon orientalis, TriHybr—Trifolium hybridum, TriInod—Tripleurospermum inodorum, TriFlav—Trisetum flavescens, TriAest—Triticum aestivum, TusFarf—Tussilago farfara, UrtDioi—Urtica dioica, VerNigr—Verbascum nigrum, VerArve—Veronica arvensis, VerCham—Veronica chamaedrys, VerPoli—Veronica polita, VicCrac—Vicia cracca, VicHirs—Vicia hirsuta, VioArve—Viola arvensis, ZeaMays—Zea mays, ×TrRimp—×Triticosecale rimpaui).
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Figure 4. Distribution of plant taxa groups according to origin and invasion status. TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
Figure 4. Distribution of plant taxa groups according to origin and invasion status. TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
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Figure 5. Distribution of plant taxa groups according to successional age optima. TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
Figure 5. Distribution of plant taxa groups according to successional age optima. TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
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Figure 6. Distribution of plant taxa groups according to the index of colonization success (ICS). TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
Figure 6. Distribution of plant taxa groups according to the index of colonization success (ICS). TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
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Figure 7. Distribution of plant taxa groups according to the index of colonization potential (ICP). TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
Figure 7. Distribution of plant taxa groups according to the index of colonization potential (ICP). TI. first evaluation date, TII. second evaluation date, TIII. third evaluation date.
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Table 1. Summary characteristics of the monitored habitats.
Table 1. Summary characteristics of the monitored habitats.
HabitatInfluence of RailwayVegetation DisturbanceConditions for Vegetation
Railway yardStrong influence during railway construction and operationHerbicide applicationUnfavorable conditions and strong disturbance pressure
Railway embankmentStrong influence during construction and limited influence during operationOccasional and localized removal of woody vegetationModerately unfavorable conditions and limited disturbance pressure
Railway landLimited influence during construction and operationNo regular vegetation managementFavorable conditions without significant disturbance pressure
Field marginNo direct influenceLimited influence of soil cultivation and herbicide applicationFavorable conditions under low disturbance pressure
Arable landNo direct influenceSoil cultivation and herbicide applicationFavorable conditions under strong disturbance pressure
Table 2. Habitat-affinity groups of plant taxa based on CCA.
Table 2. Habitat-affinity groups of plant taxa based on CCA.
Preferred HabitatPlant Taxa (Abbreviations)
Railway yardAvenella flexuosa (AveFlex), Epilobium adenocaulon (EpiAden), Potentilla erecta (PotErec), Taraxacum sect. Taraxacum (TarSect), Vicia hirsuta (VicHirs)
Railway yard
and railway embankment,
railway land		Pimpinella saxifraga (PimSaxi), Senecio viscosus (SenVisc), Tripleurospermum inodorum (TriInod)
Railway embankment,
railway land		Aegopodium podagraria (AegPoda), Aethusa cynapium (AetCyna), Achillea millefolium (AchMill), Angelica archangelica (AngArch), Anthriscus sylvestris (AntSylv), Arrhenatherum elatius (ArrElat), Artemisia vulgaris (ArtVulg), Barbarea vulgaris (BarVulg), Bryophyta (Bryophy), Calamagrostis epigejos (CalEpig), Carduus acanthoides (CarAcan), Centaurea cyanus (CenCyan), Centaurea scabiosa (CenScab), Cirsium arvense (CirArve), Dactylis glomerata (DacGlom), Echinops sphaerocephalus (EchSpha), Echium vulgare (EchVulg), Epilobium hirsutum (EpiHirs), Equisetum arvense (EquArve), Erophila verna (EroVern), Fallopia convolvulus (FalConv), Festuca rubra (FesRubr), Galium album (GalAlbu), Galium verum (GalVeru), Geranium purpureum (GerPurp), Geranium robertianum (GerRobe), Leontodon hispidus (LeoHisp), Leucanthemum vulgare (LeuVulg), Lupinus polyphyllus (LupPoly), Melilotus officinalis (MelOffi), Myosotis ramosissima (MyoRamo), Oenothera biennis (OenBien), Onobrychis viciifolia (OnoVicii), Poa pratensis (PoaPrat), Rumex acetosa (RumAcet), Senecio vulgaris (SenVulg), Silene latifolia (SilLati), Solidago canadensis (SolCana), Tanacetum vulgare (TanVulg), Tragopogon orientalis (TraOrie), Trifolium hybridum (TriHybr), Trisetum flavescens (TriFlav), Tussilago farfara (TusFarf), Urtica dioica (UrtDioi), Veronica arvensis (VerArve), Vicia cracca (VicCrac)
Railway embankment, railway land and field marginAgrostis capillaris (AgrCapi), Cerastium arvense (CerArve), Holcus lanatus (HolLana)
Field marginFragaria vesca (FraVesc), Galium intermedium (GalInte), Hylotelephium telephium (HylTele), Linaria vulgaris (LinVulg), Rubus sect. Rubus (RubSect), Verbascum nigrum (VerNigr), Veronica chamaedrys (VerCham)
Field margin and arable landApera spica-venti (ApeSpic), Galeopsis tetrahit (GalTetr), Galium aparine (GalApar), Elymus repens (ElyRepe)
Arable landAlopecurus pratensis (AloPrat), Brassica napus (BraNapu), Chenopodium album (CheAlbu), Convolvulus arvensis (ConArve), Hordeum vulgare (HorVulg), Poa annua (PoaAnnu), Polygonum aviculare (PolAvic), Triticum aestivum (TriAest), Veronica polita (VerPoli), Zea mays (ZeaMays), ×Triticosecale rimpaui (×TrRimp)
Arable land, railway yardCapsella bursa-pastoris (CapBurs), Hypericum perforatum (HypPerf), Myosotis arvensis (MyoArve), Viola arvensis (VioArve)
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Winkler, J.; Smékalová, M.; Lumbantobing, Y.R.; Červenková, J.; Sitek, W.; Vaverková, M.D. The Importance of Railway Lines for the Composition of Vegetation in Agricultural Landscapes: A Case Study. Land 2026, 15, 523. https://doi.org/10.3390/land15040523

AMA Style

Winkler J, Smékalová M, Lumbantobing YR, Červenková J, Sitek W, Vaverková MD. The Importance of Railway Lines for the Composition of Vegetation in Agricultural Landscapes: A Case Study. Land. 2026; 15(4):523. https://doi.org/10.3390/land15040523

Chicago/Turabian Style

Winkler, Jan, Marta Smékalová, Yentriani Rumeta Lumbantobing, Jana Červenková, Wiktor Sitek, and Magdalena Daria Vaverková. 2026. "The Importance of Railway Lines for the Composition of Vegetation in Agricultural Landscapes: A Case Study" Land 15, no. 4: 523. https://doi.org/10.3390/land15040523

APA Style

Winkler, J., Smékalová, M., Lumbantobing, Y. R., Červenková, J., Sitek, W., & Vaverková, M. D. (2026). The Importance of Railway Lines for the Composition of Vegetation in Agricultural Landscapes: A Case Study. Land, 15(4), 523. https://doi.org/10.3390/land15040523

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