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Article

Short-Term Abandonment versus Mowing in a Mediterranean-Temperate Meadow: Effects on Floristic Composition, Plant Functionality, and Soil Properties—A Case Study

1
Research institute on Terrestrial Ecosystems, National Research Council, Via G. Moruzzi 1, 56124 Pisa, Italy
2
Institute for Information Science and Technologies “Alessandro Faedo”-National Research Council, Via G. Moruzzi 1, 56124 Pisa, Italy
*
Author to whom correspondence should be addressed.
Submission received: 23 November 2021 / Revised: 4 January 2022 / Accepted: 4 January 2022 / Published: 7 January 2022

Abstract

:
Hay meadows are secondary grasslands maintained by mowing, and their ecological importance resides in the inherent biodiversity and carbon stocking. We investigated the plant community and soil properties of a sub humid acid grassland near the Fucecchio marshes (Italy), managed as a hay meadow, mowed once a year, and not fertilized. Part of the meadow had been abandoned for three years. We analysed the soil properties (i.e., organic carbon and total nitrogen content, available phosphorus, pH, cation-exchange capacity, texture, and conductibility) and the plant community structure (composition, functionality, and species richness) of the two sides of the meadow (mowed and abandoned). Our aim was to highlight the changes in soil properties and vegetation community, and to find out to what extent abandonment can affect those dynamics. Our results showed that after short-term abandonment, soil pH, C and N increased; litter biomass and perennial forbs increased; and annual forbs decreased. New species colonising after abandonment, thus enriching the flora, may keep spreading and eventually hinder the growth of the specialists if mowing is not resumed. Certain valuable meadow habitats need constant human intervention to maintain their peculiar vegetation, most especially if they are a buffer zone in the proximity of natural protected areas.

Graphical Abstract

1. Introduction

Semi-natural (secondary) grasslands are considered as species-rich plant communities [1] and agro-pastoral activities, such as mowing or grazing, have been reported as positive drivers for biodiversity in terms of the total number of species, but not for rare species [2]. Grassland management also has a positive impact in terms of agronomic and ecological functioning [3]. In addition to the botanical and ecological importance of the hay meadows, economic aspects also inform these communities, as plant composition and richness are some of the key factors that affect forage quality [4].
Grasslands are usually dominated by grasses, and due to their heterogeneity and microtopography, they often hold a notable biodiversity of flora and fauna, especially pollinators [5]. Many of the living organisms of grasslands around the world are potentially affected by extinction debt because of land use changes [6] as well as the policies that give priority to afforestation [7].
As a result of land use change and abandonment, grasslands are among the most threatened ecosystems and are being studied in order to find ways to conserve nature as well as to create economic opportunities [8]. Conservation projects on grasslands and meadows with high botanical value vegetation require constant investigation since changes in soil properties due to management and climate can modify plant composition [9].
From an environmental point of view, grasslands are of great importance because of their capacity to stock 34% of the whole amount of organic carbon, most of which is in the soil (87%). Prolonged droughts and increases in temperature due to climate change, could turn grasslands from a sink to a source of carbon [10]. There are indications however, that managed grasslands, in a warmer world scenario, could counteract the carbon source effect exerted by unmanaged grasslands [11].
The factors affecting plant composition in meadows, at the field (e.g., mowing frequency and time, soil nutrients) and landscape scales (e.g., landscape heterogeneity, habitat fragmentation), have been widely investigated [3,12]. Extensive mowing of meadows (low-intermediate mowing and fertilization intensity) is a form of disturbance that enhances plant diversity since it prevents competitive exclusion and enables the co-existence of several species with different competition strengths in relation to light and soil resources [3,13]. However, although changing the mowing time can be either positive or neutral in terms of plant biodiversity [14], the moderate disturbance creates gaps in the vegetation and allows seedling recruitment and the vegetative spread of weak competitive species [15]. On the other hand, removing hay prevents the accumulation of litter and thus the nutrients in the soil [15].
Stopping the mowing of hay meadows may negatively impact species diversity and composition, as the natural secondary succession resumes in a very site-specific way [16]. Over the course of a few years, shrubs, tall grasses, and woody plants, which are strong competitors, gradually increase, whereas grassland specialists that are very sensitive to environmental changes decrease, resulting in a serious loss of biodiversity [3]. When meadow cultivation is abandoned, coverage by some unpalatable perennial weeds increases along with the soil nitrogen and carbon content [15].
The physical, chemical, and biological properties of the soil affect the vegetation in semi-natural grasslands [17]. Plant functional groups play an important role in species coexistence. A soil feedback mechanism affects the soil stock of C and N [3] and the variations in species composition and diversity more than the direct effect of mowing abandonment [18]. In fact, periodical mowing and the following harvest of plant biomass reduce the nutrient intake to the soil [19,20]. The cessation of mowing can cause eutrophication, leading to a decline in plant diversity due to the spread of species with fast growth rates as the greater nutrient availability leads to a stress reduction [21].
The ecological study of grasslands has been carried out in Northern and Central Europe and North America, as well as in the Mediterranean region. In particular, the dry-Mediterranean grasslands are important hot spots of biodiversity. However, Mediterranean grasslands in wet areas also have an important ecological value, creating buffer zones between arable land and wetlands [22]. The effects of haying or grazing have been mainly studied on dry grasslands and they are compared in terms of plant diversity, herbage quality and legumes presence, concluding that the three aspects are not compatible with only one form of management [23] and that plant traits are affected by the intensity of grazing and mowing [24]. Moreover, in the Mediterranean region, grasslands are strongly dependent on water availability in terms of vegetation and energy fluxes (productivity and carbon exchanges) [25].
Besides its important ecological role, such vegetation is related to the economic, cultural, and social heritage of the local population [26]. This makes its conservation a priority in many small-scale economies. Grasslands and meadows are important buffers when close to protected areas, as they tend to be managed in accordance with sustainable agriculture practices that are acceptable in such protected areas, and this proximity means that they have an important conservation value [22]. However, as semi-natural grasslands are likely to disappear in many geographical areas where the economy related to their cultivation is no longer profitable, the importance of carrying out the characterization of their vegetation has grown significantly, especially in wet areas ranging from small to large-scale, as the effects are strongly affected by the regional context [27].
The aim of the study was to find out how and to what extent the abandonment of yearly mowing affects species composition, plant functionality and the related soil properties of a sub humid hay meadow in a Mediterranean climate. This entailed discovering how the disturbance created by mowing helps to maintain grassland biodiversity. We thus investigated the soil properties and plant community structure, vegetation composition, richness, and functionality of two portions of the same meadow: one cultivated and one that had been abandoned for three years.

2. Materials and Methods

2.1. Study Site Description

The study site is located in Tuscany, central Italy, not far from the Padule di Fucecchio wetland nature reserve (43°47′ N, 10°43′ E). The area surrounding the reserve is farmed and there are lowland hay meadows [28]. The area is close to the boundary between the transitional Mediterranean-Oceanic climate and the semi-Continental Oceanic climates [29]. Total annual precipitation during the study was 1341 mm, (max 204 mm in January, min 3.8 mm in March) whereas mean minimum–maximum annual daily air temperatures were 4.2–29.6 °C.
Our study focused on a 3.5 ha grassland mowed to produce hay for horses. The management regime included a once-a-year mowing (May or June) without fertilisation. Three years before the sampling, part of the meadow had been abandoned as the owner decided to stop cultivation (Figure 1). The survey was conducted on the managed part of the meadow (hereafter hay meadow—HM), which was mowed in late June during the study, and on the part that had been abandoned (hereafter abandoned meadow—AM).

2.2. Data Collection

Vegetation and soil data were collected in May 2015 from four blocks (two in HM and two in AM), composed of eight plots (2 × 2 m), for a total of 16 plots per management type, avoiding meadow borders. The distance between plots in the same block was 2.5 m. Each plot was divided into four subplots (1 × 1 m): one subplot was designated in terms of species and plant functionality composition as well as light availability measurements, and the others in terms of destructive sampling (aboveground biomass and soil sample collections). The distance between blocks in HM and AM was 5 m (Figure 2).

2.3. Soil Analysis

Three soil samples (soil corer 2.5 cm Ø and 10 cm depth) were collected in each plot, both in HM and AM. The analyses were carried out to determine pH, total nitrogen (Ntot), organic carbon (Corg), available phosphorus (AP), cation-exchange capacity (CEC), texture, and electrical conductibility (EC) [30]. Corg and Ntot contents were determined by dry combustion using a Leco CHN Analyzer.

2.4. Vegetation Analysis

Each species was recorded in each plot and assigned to one of the following plant functional types (PFTs): annual forbs (AF), perennial forbs (PF), legumes (L), graminoids (G), and shrubs (S) [31]. The PFT composition was calculated by dividing the percentage canopy cover of each PFT by the total canopy cover of all the PFTs recorded [32]. Plant richness was evaluated by counting the number of species in each subplot. The aboveground biomass of plants rooted within a 0.2 m2 (two 10 × 100 cm) strips was collected in a subplot, and the biomass was separated into litter, graminoids, legumes, forbs and shrubs. The biomass was then dried at 60 °C until a constant weight [33].
Photosynthetically active radiation (PAR, µmol m−2 s1) was measured using a light meter (1-m length Decagon Ceptometer) on a cloudless day at solar noon (11 a.m. to 2 p.m.). For each subplot, PAR measurements were collected at ground level and above the vegetation. The light availability was calculated as the ratio of PAR below and above the vegetation.

2.5. Statistical Analysis

In order to evaluate the variance of plant biomass (Table 2) and soil properties (Table 3) in mowed and unmoved soil, we adopted ANOVA with nested factors to manage the random effect due to its division into four blocks.
In order to analyse the impact of soil and management factors on species composition we used redundancy analysis (RDA). RDA is a commonly used analysis method in the field of ecological environment and is very effective in the analysis of the impact of explanatory variables on response variables. The percentage cover of plant species was transformed using the Hellinger method [34]. In addition, species with a frequency of less than 5% (rare species) in the whole data set were excluded from the ordination analysis [18]. Data were reported as mean ± standard error. Statistical analyses were performed using R v. 4.1.1 (R Foundation for Statistical Computing, Vienna, Austria). In particular, the packages “nortest”, “stats”, “vegan” and “ggplot2” were used.

3. Results

3.1. Plant Community Composition

The species identified in the grassland are reported in Table 1. Twenty-seven species were common, 18 species were exclusive to the abandoned meadow (total = 45), and nine were found just in the hay meadow (tot n = 36). The average number of species per plot in HM was 15 ± 0.5, and 10 ± 0.9 in AM (p < 0.01), thus the plots in HM overall had fewer species, which were more uniformly distributed. In contrast, AM showed an overall higher number of plant species that were unevenly distributed among plots, and a lower number of species per plot. In terms of plant functionality (PFT), perennial forbs showed statistically significant differences, and were higher in AM compared to HM (p = 0.004), while annual forbs were higher in HM (p = 0.046). The most species rich PFT were graminoids and perennial forbs with 27 and 18 species, respectively, while annual forbs, legumes and shrubs accounted only for five, three and one species, respectively (Table 1; Figure 3).
Significant differences were detected in litter biomass, which was higher in AM (p < 0.001), while graminoids, legumes, forbs (both perennial and annual), and shrub biomasses did not differ significantly between the treatment and control plots (Table 2).
In HM, light availability was significantly higher, reaching 320, compared to AM, which reached 143 (p < 0.01).

3.2. Soil Properties and Plant Community

The soil was characterized by a sub-acid reaction and loam-sandy texture; CEC and EC were similar in the two sampled areas (Table 3); pH, Ntot and Corg were lower in HM than in AM (p = 0.001, p = 0.01, p = 0.01, respectively). The correlation reported in Table 4 shows a significant positive effect of soil carbon and nitrogen on the biomass of legumes, but a negative effect on species richness. Soil pH positively affected the biomass of the litter and negatively affected the biomass of graminoids and species richness. Litter biomass was negatively correlated to species richness.

3.3. RDA

Figure 4 reports the redundancy analysis which highlights the shared effects of the two explanatory variables, soil properties (pH, Corg, Ntot) and mowing, on the species composition of HM and AM. First, the AM and HM plots were well separated by the RDA, and AM plots showed a positive relationship with a higher pH. The RDA analysis revealed that the variation in species composition was significantly explained by pH (p = 0.01) and not by Ntot (p = 0.32) or Corg (p = 0.99). The RDA analysis also revealed that the variance cumulatively explained by the three constrained axes was 53%, of which 37% could be attributed to pH. The adjusted R-squared of the total RDA was 24%, while the R-squared value relating to pH was 28%. Species such as A. odoratum, D. carota subsp. carota, G. fragilis, L. ircutianum, P. lanceolata, S. flos-cuculi, T. campestre were found to be much more related to the HM plot. On the other hand, A. stolonifera, D. glomerata, H. perforatum, P. reptans and S. minor subsp. balearica were more related to the AM plots. Other species such as C. arvensis, H. lanatus and L. multiflora, were common to both plots, without being predominant in either.

4. Discussion

The short-term abandonment of the grassland mowing changed the soil properties, i.e., pH, total nitrogen, and organic carbon content, as well as the plant community in terms of species composition, richness, and functionality. The results of our study showed that mowing allows the conservation of a specific vegetation composition, rich in graminoids and annual forbs, while after three years of abandonment, new species started to colonise the grassland, especially perennial forbs, and the abundance of this already present functional type was increased. This led to differences in species composition, showing up species common and exclusive in relation to their management and a shift in functional groups from annual to perennial forbs, as well as changes in soil properties, with increases in pH, organic carbon, and nitrogen. Although the pH belongs to the moderately acidic category, the variation after abandonment showed a tendency towards a neutral reaction.
The vegetation analysed in the hay meadow included some species typical of the Arrhenatheretalia order of the Molinio-Arrhenatheretea class. This vegetation can be described as hygrophilous, meso-hygrophilous or mesophilous meadows which are found in the Mediterranean macro-bioclimate. These communities usually grow on soils that are mineral to variably rich in organic matter and include strongly manured to un-manured meadows [36]. The absence of species that are commonly found in the Arrhenatheretalia order, especially Arrenatherum eliatus (L.) P. Beauv. ex J. & C. Presl [37], could be related to the fact that the HM was not fertilized, as in fact Arrhenatherum elatius and the vegetation of the Arrhenatherion alliance is associated with fertilized meadows.
After the three-year abandonment, the sampled AM plots were much more uneven than the HM plots, with no clear dominating species in any of the sampled plots. On the other hand, in HM, A. odoratum showed high soil cover (abundance)in almost all the sampled plots.
Anthoxanthum odoratum is a perennial, low productive grass, which is tolerant to soil low nitrogen and acidic conditions, and with a noticeable scent. At low elevations, A. odoratum is often found in manured meadows belonging to the semi-natural class of Molinio-Arrhenatheretea Tüxen 1937.
Interestingly, other species such as G. fragilis, S. flos-cuculi, L. arundinaceum, D. carota subsp. carota, G. verum and T. campestre were much more related with the HM plot. The presence of G. fragilis highlights the alliance between Gaudinio fragilis and Hordeion bulbosi Galàn, Deil, Haug & Vicente 1997, S. flos-cuculi of the Molinion caeruleae koch 1926 alliance; L. arundinaceum of the Arrhenatherion elatioris Koch 1926 alliance, while the others were often abundant in the class Molinio-Arrhenatheretea.
The once-a-year mowing and subsequent removal of the above-ground biomass led to the development of small species (T. campestre) and late-flowering species (L. tuberosus) as well as non-clonal species (G. fragilis), due to the creation of gaps in the vegetation [38].
On the other hand, the three-year abandonment was not enough to create a mature and more homogeneous vegetation, although the possible small pedo-climatic differences among plots could have contributed to the floristic composition observed. In the AM, species of the Molinio-Arrhenatheretea class were less abundant, while other species of a different, species-poor vegetation class were subjected to a significant increase in their soil coverage in many AM plots. As regards D. glomerata subsp. glomerata, diagnostic species of the semi-ruderal and mesic alliance Convolvulo arvensis-Agropyrion repentis Görs 1966 which, in turn, belongs to the nitrofilous class Artemisietea vulgaris Lohmeyer, Preising & Tüxen ex Von Rochow 1951, were much more represented in AM. This could be consistent with the slightly higher Ntot content and, being a typical vegetation of disturbed habitats, with the fact that it had only been abandoned for three years.
Similarly, other species that are often found in Artemisietea vulgaris, such as A. stolonifera, were much more abundant in AM. The relatively abundant presence of P. reptans in AM, a species that is usually related to Molinio-Arrhenatheretea, could be explained by the subhumid nature of the studied area, as well as its suitability for disturbed habitats and the moderate non-homogeneity of the AM plots. In fact, P. reptans showed only a slight relation with D. glomerata in AM. Moreover, the presence of S. minor subsp. balearica may seem inconsistent, as this species grows in relatively dry environments. In any case, S. minor subsp. balearica in the RDA is not related to the species adapted to humid conditions, thus resulting in the unevenness of the AM. The prevalence of species that are related to species-poor vegetation suggests the beginning of a process leading to the competitive exclusion of species with a less competitive ability and in the long term to a sort of habitat homogenization in AM [39]. Species typical of wetlands found in the AM, e.g., L. salicaria, although in low abundance, suggest that an initial spread of such species in the wettest parts of the study area was able to increase as long as the meadow remained abandoned. Similarly, species, common near the edges where there was no mowing, such as A. eupatoria, F. vulgaris, started to spread after abandonment [40].
The differences in plant composition between AM and HM, highlighted by RDA, could also be related to the effects of litter accumulation after mowing had ceased. In fact, litter accumulation causes a shift in species composition as it negatively affects the seedling recruitment of many annuals, leading to a reduction of species richness in AM. This is due to the interception of light and the creation of a physical barrier, leading to a decrease in the number of seedlings per area unit. The same effect may also have been caused by the lower light availability (PAR) observed in AM [41]. Tolerance to litter accumulation, however, enhances the plant establishment of woody and clonal species because of better water conditions and reduced competition from other species [42].
In the hay meadow, the once-a-year mowing favoured the establishment of new individuals of some species, the persistence of others, and the faster growth of smaller species, such as A. odoratum, D. carota subsp. carota, G. fragilis, L. ircutianum, P. lanceolata, S. flos-cuculi, T. campestre, resulting in a high plant richness [13,43].
In both HM and AM, perennial forbs and graminoids were the most abundant, with the latter being mainly represented by perennials. The low percentage of annuals in HM and AM plots is consistent with the mowing period that did not allow an abundant growth as well as with the high presence of perennials in both Molinio-Arrhenatheretea and Artemisietea vulgaris classes. The cessation of mowing increased the percentage of perennial forbs, suggesting that the annuals found in the HM are tolerant to this management technique and are slightly favoured when allowed to colonize areas with a lower competition exerted by certain perennials related to the Artemisietea vulgaris class.
In the case of short-term abandonment, natural succession had begun, as demonstrated by the differences in the species compositions of the two plant communities, leading to a higher number of species in AM [44] The increase in species could be related to the higher abundance of species typical of the Artemisietea vulgaris, which came from the neighbouring area into the unmown meadow.
The effects of short-term abandonment on soil properties, i.e., pH, soil nitrogen and organic content, confirm the key role of soil chemical properties in influencing the vegetation in managed grasslands [45,46,47]. The vegetation also affects the soil properties with a feedback mechanism [3]. Some functional types, such as forbs, are effective in increasing the content of carbon and nitrogen in grassland soils [48]. In our study, pH, Corg and N were the soil properties that correlated negatively with species richness, as found in recently abandoned pastures [49], while other authors have reported that species richness is not influenced e.g., by pH values [50]. In AM, the increased litter deposition due to the lack of mowing and hay removal led to an increase in soil organic matter content and to higher values of Ntot and Corg, in Mediterranean grasslands, as land abandonment, leads to a quick increase in SOC [51]. The changes in soil properties correlated with the changes in species composition, suggesting that the cessation of mowing could be responsible for the short-term increase in some perennial forbs such as A. stolonifera, D. glomerata, H. perforatum, P. reptans and S. minor subsp. balearica [15].

5. Conclusions

Our results highlight the changes in the floristic composition and abundance of species together with the soil parameters and litter accumulation during the first phases of the abandonment of a sub humid meadow located at the border of Mediterranean and Oceanic climates. The hay meadow near the Fucecchio marshes, managed only by a once-a-year mowing, is a relatively species-rich habitat that is dependent on human agricultural activity. Mowing in May or June led to the persistence of hay species with different functionalities, increasing annuals, as well as to the modification of certain soil conditions with a slightly lower pH and nitrogen content, thus making it suitable for conservation and agriculturally/economically viable for the production of hay. In contrast, short-term abandonment increased the abundance of perennial and woody species, modifying the plant community structure in terms of species composition and functionality. In the early phase of vegetation succession, the species richness decreased in the abandoned meadow. At the same time, abandonment led to an increase in litter accumulation and a moderate increase in total nitrogen, soil organic matter and pH.
Further studies should investigate subsequent changes in the vegetation of the abandoned meadow together with the plant-soil relationships during the natural succession. The species that are most affected by mowing abandonment should be identified so that they can be used as indicators of vegetation homogenization in these particular pedo-climatic conditions. Since such ecosystems provide services to the local population and are influenced by field and landscape factors, an investigation into the relationship between local communities and small-scale economies based on grasslands and meadows could be of great interest in order to make grassland conservation a priority.

Author Contributions

Conceptualization, F.B. and F.V.; methodology, F.B., F.V. and V.L.; software, C.C.; validation, F.B., F.V. and V.L.; formal analysis, I.R., C.C. and M.S.; investigation, F.B., F.V., I.R. and M.S.; resources, F.B.; data curation, C.C.; writing—original draft preparation, F.B. and F.V.; writing—review and editing, F.B., F.V., C.C. and V.L.; visualization, F.B. and F.V.; supervision, F.B. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Informed Consent Statement

Not applicable.

Conflicts of Interest

The authors declare no conflict of interest.

References

  1. Feurdean, A.; Ruprecht, E.; Molnár, Z.; Hutchinson, S.M.; Hickler, T. Biodiversity-rich European grasslands: Ancient, forgotten ecosystems. Biol. Conserv. 2018, 228, 224–232. [Google Scholar] [CrossRef]
  2. Pykälä, J.; Luoto, M.; Heikkinen, R.K.; Kontula, T. Plant species richness and persistence of rare plants in abandoned semi-natural grasslands in northern Europe. Basic Appl. Ecol. 2005, 6, 25–33. [Google Scholar] [CrossRef]
  3. Petermann, J.S.; Buzhdygan, O.Y. Grassland biodiversity. Curr. Biol. 2021, 31, R1195–R1201. [Google Scholar] [CrossRef] [PubMed]
  4. Tallowin; Jefferson Hay production from lowland semi-natural grasslands: A review of implications for ruminant livestock systems. Grass Forage Sci. 1999, 54, 99–115. [CrossRef]
  5. Öckinger, E.; Eriksson, A.K.; Smith, H.G. Effects of grassland abandonment, restoration and management on butterflies and vascular plants. Biol. Conserv. 2006, 133, 291–300. [Google Scholar] [CrossRef]
  6. Kuussaari, M.; Bommarco, R.; Heikkinen, R.K.; Helm, A.; Krauss, J.; Lindborg, R.; Öckinger, E.; Pärtel, M.; Pino, J.; Rodà, F.; et al. Extinction debt: A challenge for biodiversity conservation. Trends Ecol. Evol. 2009, 24, 564–571. [Google Scholar] [CrossRef]
  7. Bremer, L.L.; Farley, K.A. Does plantation forestry restore biodiversity or create green deserts? A synthesis of the effects of land-use transitions on plant species richness. Biodivers. Conserv. 2010, 19, 3893–3915. [Google Scholar] [CrossRef] [Green Version]
  8. Haslgrübler, P.; Krautzer, B.; Tamegger, C.; Ševĉíková, M.; Tischew, S.; Rieger, E.; Kizekiva, M.; Golinski, P.; Scotton, M. SALVERE—Semi Natural Grassland as a Source of Biodiversity Improvement—A Central Europe Project. Grassl. Sci. Eur. 2011, 16, 526–528. [Google Scholar]
  9. Critchley, C.; Chambers, B.; Fowbert, J.; Sanderson, R.; Bhogal, A.; Rose, S. Association between lowland grassland plant communities and soil properties. Biol. Conserv. 2002, 105, 199–215. [Google Scholar] [CrossRef]
  10. Poeplau, C. Grassland soil organic carbon stocks along management intensity and warming gradients. Grass Forage Sci. 2021, 76, 186–195. [Google Scholar] [CrossRef]
  11. Chang, J.; Ciais, P.; Gasser, T.; Smith, P.; Herrero, M.; Havlík, P.; Obersteiner, M.; Guenet, B.; Goll, D.S.; Li, W.; et al. Climate warming from managed grasslands cancels the cooling effect of carbon sinks in sparsely grazed and natural grasslands. Nat. Commun. 2021, 12, 118. [Google Scholar] [CrossRef]
  12. Bretzel, F.; Vannucchi, F.; Romano, D.; Malorgio, F.; Benvenuti, S.; Pezzarossa, B. Wildflowers: From conserving biodiversity to urban greening—A review. Urban For. Urban Green. 2016, 20, 428–436. [Google Scholar] [CrossRef]
  13. Gaujour, E.; Amiaud, B.; Mignolet, C.; Plantureux, S. Factors and processes affecting plant biodiversity in permanent grasslands. A review. Agron. Sustain. Dev. 2012, 32, 133–160. [Google Scholar] [CrossRef] [Green Version]
  14. Humbert, J.-Y.; Pellet, J.; Buri, P.; Arlettaz, R. Does delaying the first mowing date benefit biodiversity in meadowland? Environ. Evid. 2012, 1, 9. [Google Scholar] [CrossRef] [Green Version]
  15. Kelemen, A.; Török, P.; Valkó, O.; Deák, B.; Miglécz, T.; Tóth, K.; Ölvedi, T.; Tóthmérész, B. Sustaining recovered grasslands is not likely without proper management: Vegetation changes after cessation of mowing. Biodivers. Conserv. 2014, 23, 741–751. [Google Scholar] [CrossRef]
  16. Pavlů, L.; Pavlů, V.; Gaisler, J.; Hejcman, M.; Mikulka, J. Effect of long-term cutting versus abandonment on the vegetation of a mountain hay meadow (Polygono-Trisetion) in Central Europe. Flora-Morphol. Distrib. Funct. Ecol. Plants 2011, 206, 1020–1029. [Google Scholar] [CrossRef]
  17. Cachovanová, L.; Hájek, M.; Fajmonová, Z.; Marrs, R. Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape. Folia Geobot. Phytotaxon. 2012, 47, 349–371. [Google Scholar] [CrossRef]
  18. Swacha, G.; Botta-Dukát, Z.; Kącki, Z.; Pruchniewicz, D.; Żołnierz, L. The effect of abandonment on vegetation composition and soil properties in Molinion meadows (SW Poland). PLoS ONE 2018, 13, e0197363. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  19. Venterink, H.O.; Kardel, I.; Kotowski, W.; Peeters, W.; Wassen, M. Long-term effects of drainage and hay-removal on nutrient dynamics and limitation in the Biebrza mires, Poland. Biogeochemisty 2009, 93, 235–252. [Google Scholar] [CrossRef]
  20. Pavlů, L.; Pavlů, V.; Gaisler, J.; Hejcman, M. Relationship between soil and biomass chemical properties, herbage yield and sward height in cut and unmanaged mountain hay meadow (Polygono–Trisetion). Flora-Morphol. Distrib. Funct. Ecol. Plants 2013, 208, 599–608. [Google Scholar] [CrossRef]
  21. Grime, J.P. Plant Strategies, Vegetation Processes and Ecosystem Properties, 2nd ed.; John Wiley & Sons Ltd.: Sussex, UK, 2001. [Google Scholar]
  22. Tardella, F.M.; Di Agostino, V.M. Vegetation of the “Altipiani di Colfiorito” wetlands (central Apennines, Italy). Plant Sociol. 2020, 57, 113. [Google Scholar] [CrossRef]
  23. Faria, N.; Peco, B.; Carmona, C.P. Effects of haying on vegetation communities, taxonomic diversity and sward properties in mediterranean dry grasslands: A preliminary assessment. Agric. Ecosyst. Environ. 2018, 251, 48–58. [Google Scholar] [CrossRef]
  24. Louault, F.; Pillar, V.D.; Aufrere, J.; Garnier, E.; Soussana, J.F. Plant traits and functional types in response to reduced disturbance in a semi-natural grassland. J. Veg. Sci. 2005, 16, 151–160. [Google Scholar] [CrossRef]
  25. Jongen, M.; Pereira, J.; Aires, L.M.I.; Pio, C. The effects of drought and timing of precipitation on the inter-annual variation in ecosystem-atmosphere exchange in a Mediterranean grassland. Agric. For. Meteorol. 2011, 151, 595–606. [Google Scholar] [CrossRef]
  26. Willig, M.R. Biodiversity and Productivity. Science 2011, 333, 1709–1710. [Google Scholar] [CrossRef] [PubMed]
  27. Socher, S.A.; Prati, D.; Boch, S.; Müller, J.; Baumbach, H.; Gockel, S.; Hemp, A.; Schöning, I.; Wells, K.; Buscot, F.; et al. Interacting effects of fertilization, mowing and grazing on plant species diversity of 1500 grasslands in Germany differ between regions. Basic Appl. Ecol. 2013, 14, 126–136. [Google Scholar] [CrossRef]
  28. Bartolini, A. Il Bosco di Chiusi e la Paduletta di Ramone. Indagini Naturalistiche e Attività di Ripristino Degli Habitat. Quaderni del Padule di Fucecchio n. 7; Centro di Ricerca, Documentazione e Promozione del Padule di Fucecchio; Università degli Studi di Pavia: Brescia, Italy, 2010. [Google Scholar]
  29. Blasi, C.; Michetti, L. The climate of Italy. In Biodiversity in Italy; Blasi, C., Boitani, L., la Posta, S., Manes, F., Marchetti, M., Eds.; Palombi Editori: Roma, Italy, 2007; pp. 57–66. Available online: http://www.pcn.minambiente.it/viewer/index.php?services=Fitoclima (accessed on 9 July 2021).
  30. ASA-SSSA. Methods of Soil Analysis, Part 1 and 3, Physical and Chemical Methods, 2nd ed.; ASA-SSSA: Madison, WI, USA, 1996. [Google Scholar]
  31. Duckworth, J.C.; Kent, M.; Ramsay, P.M. Plant functional types: An alternative to taxonomic plant community description in biogeography? Prog. Phys. Geogr. 2000, 24, 515–542. [Google Scholar] [CrossRef]
  32. Daubenmire, R.F. Canopy coverage method for vegetation analysis. Northwest Sci. 1959, 33, 43–64. [Google Scholar]
  33. Seabloom, E.W.; Borer, E.T.; Buckley, Y.; Cleland, E.E.; Davies, K.; Firn, J.; Harpole, W.S.; Hautier, Y.; Lind, E.; MacDougall, A.; et al. Predicting invasion in grassland ecosystems: Is exotic dominance the real embarrassment of richness? Glob. Chang. Biol. 2013, 19, 3677–3687. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  34. Legendre, P.; Gallagher, E.D. Ecologically meaningful transformations for ordination of species data. Oecologia 2001, 129, 271–280. [Google Scholar] [CrossRef]
  35. Bartolucci, F.; Peruzzi, L.; Galasso, G.; Albano, A.; Alessandrini, A.; Ardenghi, N.M.G.; Astuti, G.; Bacchetta, G.; Ballelli, S.; Banfi, E.; et al. An updated checklist of the vascular flora native to Italy. Plant Biosyst. Int. J. Deal. All Asp. Plant Biol. 2018, 152, 179–303. [Google Scholar] [CrossRef]
  36. Biondi, E.; Blasi, C.; Allegrezza, M.; Anzellotti, I.; Azzella, M.M.; Carli, E.; Casavecchia, S.; Copiz, R.; Del Vico, E.; Facioni, L.; et al. Plant communities of Italy: The Vegetation Prodrome. Plant Biosyst. Int. J. Deal. All Asp. Plant Biol. 2014, 148, 728–814. [Google Scholar] [CrossRef] [Green Version]
  37. Allegrezza, M.; Biondi, E. Syntaxonomic revision of Arrhenatherum elatius grasslands of central Italy. Fitosociologia 2011, 48, 23–40. [Google Scholar]
  38. Foster, B.L.; Gross, K.L. Species richness in a successful grassland: Effects of nitrogen enrichment and plant litter. Ecology 1998, 79, 2593–2602. [Google Scholar] [CrossRef]
  39. Smart, S.M.; Thompson, K.; Marrs, R.H.; Le Duc, M.G.; Maskell, L.C.; Firbank, L. Biotic homogenization and changes in species diversity across human-modified ecosystems. Proc. R. Soc. B Boil. Sci. 2006, 273, 2659–2665. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  40. Losvik, M.H. Plant species diversity in an old, traditionally managed hay meadow compared to abandoned hay meadows in southwest Norway. Nord. J. Bot. 1999, 19, 473–487. [Google Scholar] [CrossRef]
  41. Boob, M.; Elsaesser, M.; Thumm, U.; Hartung, J.; Lewandowski, I. Different management practices influence growth of small plants in species-rich hay meadows through shading. Appl. Veg. Sci. 2021, 24, e12625. [Google Scholar] [CrossRef]
  42. Galvánek, D.; Leps, J. The effect of management on productivity, litter accumulation and seedling recruitment in a Carpathian mountain grassland. Plant Ecol. 2012, 213, 523–533. [Google Scholar] [CrossRef]
  43. Klimešová, J.; Janeček, S.; Bartušková, A.; Lanta, V.; Doležal, J. How is regeneration of plants after mowing affected by shoot size in two specie-rich meadows. Folia Geobot. 2010, 45, 225–238. [Google Scholar] [CrossRef]
  44. Prévosto, B.; Kuiters, L.; Bernhardt-Römermann, M.; Dölle, M.; Schmidt, W.; Hoffmann, M.; van Uytvanck, J.; Bohner, A.; Kreiner, D.; Stadler, J.; et al. Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats. Folia Geobot. 2011, 46, 303–325. [Google Scholar] [CrossRef] [Green Version]
  45. Grime, J.P.; Thompson, K.; Hunt, R.; Hodgson, J.G.; Cornelissen, J.H.C.; Rorison, I.H.; Hendry, G.A.F.; Ashenden, T.W.; Askew, A.P.; Band, S.R.; et al. Integrated Screening Validates Primary Axes of Specialisation in Plants. Oikos 1997, 79, 259. [Google Scholar] [CrossRef]
  46. White, T.A.; Moore, K.J.; Barker, D.J. The importance of local processes to landscape patterns of grassland vegetation diversity. N. Z. J. Agric. Res. 2004, 47, 199–207. [Google Scholar] [CrossRef] [Green Version]
  47. Crawley, M.J.; Johnston, A.E.; Silvertown, J.; Dodd, M.; de Mazancourt, C.; Heard, M.S.; Henman, D.F.; Edwards, G.R. Determinants of Species Richness in the Park Grass Experiment. Am. Nat. 2005, 165, 179–192. [Google Scholar] [CrossRef] [PubMed]
  48. De Deyn, G.B.; Quirk, H.; Yi, Z.; Oakley, S.; Ostle, N.J.; Bardgett, R.D. Vegetation composition promotes carbon and nitrogen storage in model grassland communities of contrasting soil fertility. J. Ecol. 2009, 97, 864–875. [Google Scholar] [CrossRef]
  49. Diviaková, A.; Stašiov, S.; Pondelík, R.; Pätoprstý, V.; Novikmec, M. Environmental and Management Control over the Submontane Grassland Plant Communities in Central Slovakia. Diversity 2021, 13, 30. [Google Scholar] [CrossRef]
  50. Merunková, K.; Chytrý, M. Environmental control of species richness and composition in upland grasslands of the southern Czech Republic. Plant. Ecol. 2012, 213, 591–602. [Google Scholar] [CrossRef]
  51. Gabarrón-Galeote, M.A.; Trigalet, S.; van Wesemael, B. Effect of land abandonment on soil organic carbon fractions along a Mediterranean precipitation gradient. Geoderma 2015, 249–250, 69–78. [Google Scholar] [CrossRef]
Figure 1. Map of the site from 2012 to 2015, to highlight the cultivation before the abandonment of one portion.
Figure 1. Map of the site from 2012 to 2015, to highlight the cultivation before the abandonment of one portion.
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Figure 2. Map of the site with the position of the four blocks and the subplots, on the right HM (=hay meadow) on the left AM (=abandoned meadow).
Figure 2. Map of the site with the position of the four blocks and the subplots, on the right HM (=hay meadow) on the left AM (=abandoned meadow).
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Figure 3. Plant functional type composition (% PTF) in hay (HM) and abandoned (AM) meadow, as the ratio of the percentage canopy cover of each PFT to the total canopy cover of all the PFTs recorded. G = graminoid; PF = perennial forbs; L = legumes; AF = annual forbs; S = shrubs. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05; * = p < 0.05; ** = p < 0.01; ns = not significant.
Figure 3. Plant functional type composition (% PTF) in hay (HM) and abandoned (AM) meadow, as the ratio of the percentage canopy cover of each PFT to the total canopy cover of all the PFTs recorded. G = graminoid; PF = perennial forbs; L = legumes; AF = annual forbs; S = shrubs. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05; * = p < 0.05; ** = p < 0.01; ns = not significant.
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Figure 4. Ordination biplot of redundancy analysis (RDA) representing shared effects of two sets of explanatory variables (soil properties and management). C = total organic carbon; N = total nitrogen. (A) Circles in bold represent the HM plots, circles not in bold represent the AM plots, crosses represent the species. (B) the species reported are: A_odor = A. odoratum, A_stol = A. stolonifera, B_media, B_minor, C_divul = C. divulsa, C_mycon = C. myconis, C_arven = C. arvensis, C_spp = Cyperus sp., D_glom = D. glomerata, D_carota, L_aru = L. arundinaceum, G_verum, G_frag = G. fragilis, G_dis = G. dissectum, H_lan = H. lanatus, H_perf = H. perforatum, H_rad = H. radicata, K_spur = K. spuria, L_tuber = L. tuberosus, L_ircu = L. ircutianum, L_bien = L. bienne, L_mult = L. multiflora, P_lanc = P. lanceolata, P_rept = P. reptans, R_bulb = R. bulbosus, R_spp = Rubus sp., S_minor, S_flos = S. flos-cuculi, T_camp = T. campestris.
Figure 4. Ordination biplot of redundancy analysis (RDA) representing shared effects of two sets of explanatory variables (soil properties and management). C = total organic carbon; N = total nitrogen. (A) Circles in bold represent the HM plots, circles not in bold represent the AM plots, crosses represent the species. (B) the species reported are: A_odor = A. odoratum, A_stol = A. stolonifera, B_media, B_minor, C_divul = C. divulsa, C_mycon = C. myconis, C_arven = C. arvensis, C_spp = Cyperus sp., D_glom = D. glomerata, D_carota, L_aru = L. arundinaceum, G_verum, G_frag = G. fragilis, G_dis = G. dissectum, H_lan = H. lanatus, H_perf = H. perforatum, H_rad = H. radicata, K_spur = K. spuria, L_tuber = L. tuberosus, L_ircu = L. ircutianum, L_bien = L. bienne, L_mult = L. multiflora, P_lanc = P. lanceolata, P_rept = P. reptans, R_bulb = R. bulbosus, R_spp = Rubus sp., S_minor, S_flos = S. flos-cuculi, T_camp = T. campestris.
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Table 1. Species present in the meadow, family, and functional groups [35]. AM = exclusive after the abandonment; HM = exclusive to the hay meadow; C = common.
Table 1. Species present in the meadow, family, and functional groups [35]. AM = exclusive after the abandonment; HM = exclusive to the hay meadow; C = common.
SpeciesFamilyFunctional GroupSite
Agrimonia eupatoria L.RosaceaePerennial forbAM
Agrostis stolonifera L. PoaceaeGraminoidAM
Anthoxanthum odoratum L.PoaceaeGraminoidC
Avena sterilis L. subsp. sterilisPoaceaeGraminoidC
Briza maxima L.PoaceaeGraminoidHM
Briza minor L.PoaceaeGraminoidC
Carex divulsa StokesCyperaceaeGraminoidC
Carex flacca Schreb.CyperaceaeGraminoidHM
Carex hirta L.CyperaceaeGraminoidC
Carex spicata Huds.CyperaceaeGraminoidAM
Carex tomentosa L.CyperaceaeGraminoidAM
Centaurea nigrescens Willd.AsteraceaePerennial forbHM
Cichorium intybus L.AsteraceaePerennial forbAM
Clinopodium nepeta (L.) KuntzeLamiaceaePerennial forbAM
Coleostephus myconis (L.) Cass. ex Rchb. f.AsteraceaeAnnual forbC
Convolvolus arvensis L.ConvolvulaceaePerennial forbC
Convolvulus sepium L.ConvolvulaceaePerennial forbAM
Cynodon dactylon (L.) PersPoaceaeGraminoidAM
Cyperus spp.CyperaceaeGraminoidC
Dactylis glomerata L.PoaceaeGraminoidC
Danthonia decumbes (L.) DC.PoaceaeGraminoidHM
Daucus carota L. subsp. carotaApiaceaeAnnual forbC
Dianthus armeria L.CaryophyllaceaePerennial forbAM
Filipendula vulgaris Moench.RosaceaePerennial forbAM
Galium verum L.RubiaceaePerennial forbC
Gaudinia fragilis (L.) P. Beauv.PoaceaeGraminoidHM
Geraniumdissectum L.GeraniaceaeAnnual forbC
Holcus lanatus L.PoaceaeGraminoidC
Hypericum perforatum L.HypericaceaePerennial forbAM
Hypochoeris radicata L.AsteraceaePerennial forbC
Kickxia spuria (L.) Dumort.PlantaginaceaeAnnual forbC
Leontodon tuberosus L.AsteraceaePerennial forbHM
Leucanthemum ircutianum DC.AsteraceaePerennial forbHM
Linaria vulgaris Mill.PlantaginaceaePerennial forbAM
Linum bienne Mill.LinaceaePerennial forbC
Lolium arundinaceum (Schreb.) Darbysh.PoaceaeGraminoidC
Lotus conrniculatus L. FabaceaeLegumesAM
Luzula multiflora (Ehrh.) Lej.JuncaceaeGraminoidC
Lythrum salicaria L.LythraceaePerennial forbAM
Oenanthe pimpinelloides L.ApiaceaePerennial forbAM
Plantago lanceolata L.PlantaginaceaePerennial forbC
Potentilla reptans L.RosaceaePerennial forbC
Prunella laciniata L. (L.)LamiaceaePerennial forbC
Prunella vulgaris L.LamiaceaePerennial forbC
Prunella x intermedia LinkLamiaceaePerennial forbC
Ranunculus bulbosus L.RanunculaceaPerennial forbC
Rubus spp.RosaceaeShrubC
Sanguisorba minor Scop.RosaceaePerennial forbC
Silene flos-cuculi (L.) Clairv.CaryophyllaceaePerennial forbC
Sonchus oleraceus L.AsteraceaeAnnual forbAM
Trifolium campestre Schreb.FabaceaeLegumesHM
Urospermum delachampii (L.) F.W. SchmidtAsteraceaePerennial forbAM
Vicia sativa L.FabaceaeLegumesHM
Table 2. Plant biomass of functional types and litter in hay (HM) and abandoned (AM) meadow. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05; *** = p < 0.001; ns = not significant, calculated by ANOVA with nested factor (blocks have been considered as random effect).
Table 2. Plant biomass of functional types and litter in hay (HM) and abandoned (AM) meadow. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05; *** = p < 0.001; ns = not significant, calculated by ANOVA with nested factor (blocks have been considered as random effect).
Biomass DW (g)
HMAMp Values
Litter3.9 ± 0.8326.2 ± 2.83***
Graminoid46.0 ± 6.8933.8 ± 5.50ns
Legumes0.4 ± 0.090.5 ± 0.49ns
Forbs11.8 ± 1.4717.3 ± 3.80ns
Shrubs3.9 ± 1.915.6 ± 3.55ns
Table 3. Soil properties of hay (HM) and abandoned (AM) meadow. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05 (*); p < 0.01 (**); p < 0.001(***), calculated by ANOVA with nested factor (blocks have been considered as random effect). ns = not significant.
Table 3. Soil properties of hay (HM) and abandoned (AM) meadow. Data are means of 16 samples ± SE. Asterisks represent significance of p < 0.05 (*); p < 0.01 (**); p < 0.001(***), calculated by ANOVA with nested factor (blocks have been considered as random effect). ns = not significant.
Soil PropertiesHMAMp Values
pH5.6 ± 0.036.0 ± 0.05***
Corg%1.7 ± 0.092.2 ± 0.16**
Ntot%0.20 ± 0.020.26 ± 0.01*
AP0.66 ± 0.230.71 ± 0.21ns
CEC (cmol kg1)14.1 ± 1.2915.5 ± 0.68ns
EC (dS/m)0.20 ± 0.010.20 ± 0.01ns
Table 4. Pearson correlation between the soil properties and vegetation data (PFTs biomass and species richness). Coefficients of correlation in bold are significant for p ≤ 0.05 (*), p < 0.01 (**) and p < 0.001 (***). SR = species richness. Number of samples = 32.
Table 4. Pearson correlation between the soil properties and vegetation data (PFTs biomass and species richness). Coefficients of correlation in bold are significant for p ≤ 0.05 (*), p < 0.01 (**) and p < 0.001 (***). SR = species richness. Number of samples = 32.
pHECCECCorgNLitterGraminoidLegumesForbsShrub
pH
EC0.46
CEC0.360.36
Corg0.390.110.15
N0.390.140.250.67 ***
Litter0.54 **0.04−0.0010.31 *0.28
Graminoid−0.36 **−0.31 *−0.18−0.10−0.01−0.17
legumes0.110.080.220.58 *0.42 *0.08−0.07
Forbs0.430.230.110.120.300.11−0.0010.26
Shrub0.010.060.050.220.03−0.06−0.24−0.09−0.29 *
SR−0.32 *0.11−0.03−0.44 **−0.21 *−0.66 ***0.21−0.080.14−0.27
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Vannucchi, F.; Lazzeri, V.; Rosellini, I.; Scatena, M.; Caudai, C.; Bretzel, F. Short-Term Abandonment versus Mowing in a Mediterranean-Temperate Meadow: Effects on Floristic Composition, Plant Functionality, and Soil Properties—A Case Study. Agriculture 2022, 12, 78. https://0-doi-org.brum.beds.ac.uk/10.3390/agriculture12010078

AMA Style

Vannucchi F, Lazzeri V, Rosellini I, Scatena M, Caudai C, Bretzel F. Short-Term Abandonment versus Mowing in a Mediterranean-Temperate Meadow: Effects on Floristic Composition, Plant Functionality, and Soil Properties—A Case Study. Agriculture. 2022; 12(1):78. https://0-doi-org.brum.beds.ac.uk/10.3390/agriculture12010078

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Vannucchi, Francesca, Valerio Lazzeri, Irene Rosellini, Manuele Scatena, Claudia Caudai, and Francesca Bretzel. 2022. "Short-Term Abandonment versus Mowing in a Mediterranean-Temperate Meadow: Effects on Floristic Composition, Plant Functionality, and Soil Properties—A Case Study" Agriculture 12, no. 1: 78. https://0-doi-org.brum.beds.ac.uk/10.3390/agriculture12010078

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