Research Article |
Corresponding author: Róbert Kun ( rbert.kun@gmail.com ) Academic editor: Douglas Evans
© 2024 Róbert Kun, Dániel Babai, András István Csathó, Arnold Erdélyi, Judit Hartdégen, Attila Lengyel, Nikoletta Kálmán, András Mártonffy, Alida Anna Hábenczyus, Zsófia Szegleti, Ákos Vig, András Máté, Ákos Malatinszky, Tímea Tóth, Csaba Vadász.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kun R, Babai D, Csathó AI, Erdélyi A, Hartdégen J, Lengyel A, Kálmán N, Mártonffy A, Hábenczyus AA, Szegleti Z, Vig Á, Máté A, Malatinszky Á, Tóth T, Vadász C (2024) Effects of management complexity on the composition, plant functional dominance relationships and physiognomy of high nature value grasslands. Nature Conservation 55: 1-19. https://doi.org/10.3897/natureconservation.55.114385
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A significant proportion of Europe’s species-rich grasslands are semi-natural habitats. They have a long history of traditional management. Several studies have been carried out to conserve them, resulting in the establishment of subsidised conservation management schemes. On the other hand, many of these conservation management schemes have failed to provide locally adaptive solutions to maintain the diversity and functional status of species-rich grasslands. In addition, few studies have compared the conservation effectiveness of different levels of management complexity. The levels of management complexity in our study are based on how different management types (e.g. grazing and mowing etc.) and how different herbage removal intensities (e.g. lower and higher grazing intensities) are combined within and between years. To investigate this, we compared the overall effects of management complexity, herbage removal intensity and management type on plant diversity, plant functional type dominance relationships and plant physiognomy. Our field sampling was carried out in the sandy meso-xeric grasslands of the Turján Region of the Great Hungarian Plain (Central Hungary). We sampled nine 2 m × 2 m plots per grassland site (n = 12), recorded all the rooted plant species and estimated their percentage cover in each plot. High level of management complexity had significant positive effects on plant diversity, grazing had positive effects on plant diversity and phanerophyte density, while the studied levels of herbage removal intensity had no effect on diversity, plant functional types or plant physiognomy. In parallel, mowing and/or low levels of management complexity had some negative effects on conservation value (e.g. lower Shannon and Simpson diversity). In this landscape, the dominance of grazing and the more complex management is more optimal than relatively homogeneous mechanical mowing. The choice of management type and intensity is an important tool in the conservation management system of this landscape, but so too is its appropriate application in space and time. Through a detailed analysis of the effects of management complexity levels compared to management types and herbage removal intensity levels, we provide a new opportunity to make grassland management practices more effective for conserving biodiversity in this region, but it would be important to investigate these in different landscapes and conditions.
Grassland conservation system, management effects, management efficiency
A significant proportion of European landscapes are cultural landscapes that have been transformed and managed by humans (
Species-rich, semi-natural grasslands have been managed for centuries by small family farms to provide summer forage in pastures and to produce winter fodder in hay meadows for livestock (
As semi-natural grasslands have been developed and maintained by human management, active and adaptive nature conservation management should be implemented to maintain the species composition and vegetation structure of these habitats. Through a long learning process, nature conservation aimed to mimic the patterns and disturbance regimes of former non-intensive, traditional grassland management (
Conservation management of grasslands should also draw on the experience of local communities still practising traditional and adaptive grassland management (cf.
A list and an introduction to the management factors and their categories and sub-categories.
Management factor categories | Management factor subcategories |
---|---|
Type of grassland management (T) | Mowing (M): Mechanical mowing at the end of June or the first half of July with 10–15 cm of stubble. See details of management complexity later in this Table. |
Grazing (G): Pastures are mainly grazed by cattle from the end of April to the beginning of October each year. Shepherds often work with them. | |
Combined (C): Mowing and grazing are combined within the same year or between years. For more details, see management complexity later in this Table. | |
Herbage removal intensity (I) | Low: Grazing at < 0.5 Standard Livestock Units (SLU) per hectare or mown once a year. LUI value: 0.1 (Schneiders et al. 2011). |
High: Grazing at > 0.5 Standard Livestock Units (SLU) per hectare or mown once a year followed by grazing in the same year. LUI value: 0.2 (Schneiders et al. 2011). | |
Management complexity (C) | Low: Grazing with a standard sequence of two grazing units per year or one mowing with 10% uncut per year or one mowing per year combined with subsequent grazing. |
High: Mowing and grazing combined between years or grazing with different start times between years in a four-year rotation. |
In this study, we aim to reveal the effects of management complexity, management intensity levels and management types on plant diversity, plant functional type dominance relationships and plant physiognomy in species-rich meso-xeric, sandy grasslands of central Hungary. We hypothesise that high management complexity and low herbage removal intensity will positively affect plant diversity, plant functional state and physiognomy. We also hypothesise that grazing, in particular, has a positive effect on higher plant diversity and less graminoid (Poales) cover, more forbs and shrubs (Phanerophytes) cover. Our specific question is: How do low and high levels of management complexity affect plant diversity, vegetation physiognomy and plant functional type cover in relation to management type and herbage removal intensity?
The study sites are located in the Turján Region of the Great Hungarian Plain along the Danube in central Hungary, in the northern Kiskunság area. The study sites are relatively close to each other, within a circle of about 10 km diameter around the neighbouring villages of Kunpeszér, Tatársszentgyörgy and Kunadacs (Appendix
Most of the studied sites have been modified by local people in the past and present, through woodcutting and long-term grazing (
The surveys were conducted in June 2018 on 12 grassland sample sites, all of which were at least 5 ha and at most 10 ha in size (Appendix
The coordinates of the plots were recorded by GPS. Data were collected from 108 plots in the 12 grassland sites mentioned above, nine plots per site (see Appendices
We defined plant functional types (PFTs) as groups of species based on three growth forms: forbs including non-grassy herbs, graminoids (Poales) including grasses, sedges and rushes and phanerophytes including shrubs and small trees (
At each grassland site, we recorded three management factors at different levels, including intensity of herbage removal (I, with low and high levels), complexity of management (C, with low and high levels) and different types of management (T, including grazing, mowing and combined types) (Table
We calculated diversity measures, namely species number, Shannon index and Simpson index, from the plant species and estimated percent cover data recorded in each plot. The use of both diversity indices was important because the Shannon diversity index is more sensitive to the higher proportion of rare (often specialist) species, while the Simpson index is more sensitive to the balance of more dominant species. We built linear and generalised linear mixed effects models (with ‘lmer’ and ‘glmer’ functions from the ‘lme4’ package) to test the effect of management factors T, I and C as three fixed factors on plant diversity indices, on the abundance of PFTs and on vegetation physiognomy. Different families of distributions (Gaussian and Gamma) were used to treat each differently distributed dependent variable in the modelling (the ‘gamma_test’ function from the ‘goft’ package was used). In our analyses, site was a random factor in all models. To assess model fit, marginal R2LR was applied (
Management type, levels of herbage removal intensity and management complexity had similarly strong effects on species number based on model fits (R2 > 0.320, Table
Effects of different management factors, namely T: management type; I: herbage removal intensity of management; C: management complexity, on diversity measures in terms of model fit. Goodness-of-fit is expressed as R2LR values.
Management factors | Species number | Shannon diversity | Simpson diversity |
---|---|---|---|
R2 | R2 | R2 | |
T | 0.324 | 0.096 | 0.057 |
I | 0.325 | 0.023 | 0.011 |
C | 0.324 | 0.072 | 0.053 |
Significant differences in diversity and cover of phanerophytes in grasslands with low and high management complexity and different management types. Only models with minimum R2LR ≥ 0.100 fit (see Tables
With PFT categories as dependent variables, management type showed a strong relationship with graminoid and forb cover (Table
Effects of different management factors, namely T: management type; I: herbage removal intensity of management; C: management complexity in relation to forbs, graminoid and Phanerophyte cover. Goodness-of-fit is also presented in R2LR values.
Management factors | Forb species cover (%) | Graminoid species cover (%) | Phanerophyte species cover (%) |
---|---|---|---|
R2 | R2 | R2 | |
T | 0.368 | 0.430 | 0.121 |
I | 0.365 | 0.420 | 0.075 |
C | 0.368 | 0.415 | 0.097 |
Effects of different management factors, namely T: management type; I: herbage removal intensity of management; C: management complexity in relation to physiognomic factors in relation to grasslands. Goodness-of-fit is also presented in R2LR values.
Management factors | Litter cover (%) | Total plant cover (%) | Bare soil surface (%) | Average plant height in plots (cm) |
---|---|---|---|---|
R2 | R2 | R2 | R2 | |
T | 0.579 | 0.709 | 0.120 | 0.355 |
I | 0.572 | 0.705 | 0.090 | 0.298 |
C | 0.559 | 0.703 | 0.115 | 0.318 |
The two main components were presented in relation to forbs and graminoid (Poales) cover, based on principal component analysis. Higher graminoid cover was associated with mowing and higher forbs cover was mostly associated with grazing and combined management was intermediate between mowing and grazing (Fig.
Principal Component Analysis of diversity indices, plant functional type cover and physiognomic factors across management types. The diversity indices examined are species number (sp_num), Shannon (Sha) and Simpson (Sim) diversity. Plant functional type cover includes graminoids (Gram.), forbs and phanerophytes (Phanero.). Plant physiognomic factors are average plant height (height), total plant cover (full_cov), bare soil surface (bare_soil) and litter cover (litter_cov). Management types: mown, grazed and combined management. The direction, width and different colours of the ellipses in the figure show us the relationship between the samples of different management types. The length and direction of the arrows show the explanatory power and relationship of each variable studied with management types and other variables.
Principal Component Analysis of diversity indices, plant functional type cover and physiognomic factors across herbage removal intensity levels. The meaning of the abbreviations used in this Figure is given in the legend to Fig.
Different management types, mainly mowing and low and high levels of herbage removal intensity and management complexity, significantly affected the species composition and dissimilarity ratios of the grasslands studied (Fig.
Although different T choices played a less important role in influencing compositional diversity, the choice of the appropriate management type was also significant: grazing had a more positive effect on phanerophytes than mowing (Appendix
The increase of clonal, often highly competitive graminoid species with higher biomass production can reduce plant diversity (
Based on our results, special attention should be paid to the multiplicity of management factors (e.g. different management types or herbage removal intensity levels), including their spatio-temporal variability (
On the other hand, although we found that high management complexity is beneficial for grassland conservation, it may be difficult to apply such management complexity and the same methods in practice in other regions, for example, for several individual farmers. Our conclusions are most relevant in terms of the exact management complexity which we have investigated in our study. Each region is different in terms of management possibilities and environmental factors. It can be difficult to graze a site one year and mow it the next or to vary the intensity of management. It is also important to note that spatially and temporally complex management can be achieved in more ways than we have explored in our study. There are other and/or simpler ways, for example, mowing only every other year, mowing at the beginning of summer one year and at the end of summer the next. The use of different grazing animals and the leaving of uncut lines in different places on a grassland between years can also be effective tools for more complex management, depending on local conservation objectives and opportunities.
However, there are often practical difficulties in applying multiple aspects of management to the modelling of community diversity. Including more explanatory variables in a model requires larger sample sizes and a more balanced sample distribution (
Our aim was to collect, organise and compare the elements of the hard-to-compare, mosaic-like landscape of use according to various parameters, using systematic sampling and to quantify and generalise the treatment results obtained mainly through experience. We must emphasise as an important message to legislators and developers of support schemes that because each site is different, generalisation is limited.
High levels of management complexity and grazing as a management type are more positive and have a greater significance for grassland conservation (i.e. result in higher plant diversity, higher proportion of forbs etc.) than the intensity of herbage removal in our study area. At the same time, mowing and/or low levels of management complexity may have some negative effects on conservation value. These analyses can be used to identify what are the strong or direct and less strong or indirect effects in the conservation of high nature value grasslands. Further research is needed to verify these relationships across a wider range of different study systems in order to provide generalisable guidelines for conservation.
We would like to thank the following colleagues for their help with the fieldwork: Judit Deme, Zsolt Molnár, Abolfazl Sharifiyan, Gantuya Batdelger, Nikolett Pónya, Győző Haszonits and Dávid Schmidt.
The authors have declared that no competing interests exist.
No ethical statement was reported.
During the study, Róbert Kun received the ÚNKP-19-3-I-SZIE-37 grant, Attila Lengyel was supported by the National Research, Development and Innovation Office of Hungary (PD-123997), Dániel Babai was supported by the MTA Premium Postdoctoral Research Fellowship Programme of the Hungarian Academy of Sciences [grant number: PPD008/2017], and was supported by the MTA-Lendület program (Lendulet_2020-56).
All authors have contributed equally.
Róbert Kun https://orcid.org/0000-0002-9607-3110
Ákos Malatinszky https://orcid.org/0000-0001-6388-9191
All of the data that support the findings of this study are available in the main text.
Sampling design in the study area with factor combinations at each site and number of replicates.
Number of sites | Management type | Herbage removal intensity | Management complexity | Number of management factor combinations | Number of plots per site |
---|---|---|---|---|---|
1 | Mown | Low | Low | 1 | 9 |
2 | Mown | Low | Low | 1 | 9 |
3 | Mown | Low | Low | 1 | 9 |
4 | Grazed | Low | High | 2 | 9 |
5 | Grazed | Low | High | 2 | 9 |
6 | Grazed | High | Low | 3 | 9 |
7 | Grazed | High | Low | 3 | 9 |
8 | Grazed | High | Low | 3 | 9 |
9 | Grazed | High | High | 4 | 9 |
10 | Combined | Low | Low | 5 | 9 |
11 | Combined | Low | High | 6 | 9 |
12 | Combined | High | High | 7 | 9 |
Differences in PFT cover and diversity indices between different management types (mowing: M, grazing: G and combined: C) of semi-natural grasslands. The Table shows means and standard deviations of PFT groups and diversity indices. Significant differences in LMER Tukey post hoc tests between different management types are indicated by the letters ‘a’, ‘b’ and ‘c’.
MOWN | GRAZED | COMBINED | |
---|---|---|---|
Species number | 34.1±3.2a | 34.9±5.8a | 35.4±3.5a |
Shannon diversity | 1.6±0.4a | 1.9±0.3b | 1.8±0.3b |
Simpson diversity | 0.6±0.2a | 0.7±0.1b | 0.7±0.1b |
Forbs cover (%) | 19.0±11.9a | 24.4±15.2a | 31.1±20.2a |
Graminoid cover (%) | 82.2±11.3a | 74.3±18.2a | 57.8±22.2a |
Phanerophytes cover (%) | 2.2±1.7a | 5.1±3.8b | 4.0±4.0ab |
Mean plant height (cm) | 31.4±12.8a | 26.7±8.4a | 21.0±8.4a |
Total plant cover (%) | 94.0±3.3a | 95.6±3.1a | 87.6±8.7a |
Bare soil surface (%) | 0.6±0.4a | 1.6±1.6a | 1.8±1.9a |
Litter cover (%) | 5.7±3.3a | 3.8±2.7a | 10.9±7.7a |
Effects of herbage removal intensity of management on plant diversity and cover of PFTs. Table shows means and standard deviations of PFT cover and diversity indices. Results are based on LMER Tukey post hoc tests. Significant differences between different intensity levels are indicated by the letters ‘a’ and ‘b’.
LOW | HIGH | |
---|---|---|
Species number | 34.3±4.7a | 35.6±4.6a |
Shannon diversity | 1.7±0.4a | 1.8±0.4a |
Simpson diversity | 0.7±0.1a | 0.7±0.1a |
Forbs cover (%) | 26.4±19.1a | 22.5±11.3a |
Graminoid cover (%) | 66.9±21.3a | 79.5±15.1a |
Phanerophytes cover (%) | 4.4±3.9a | 3.7±3.3a |
Mean plant height (cm) | 25.9±10.8a | 26.5±9.2a |
Total plant cover (%) | 91.3±6.8a | 96.0±3.3a |
Bare soil surface (%) | 1.3±1.7a | 1.6±1.6a |
Litter cover (%) | 8.0±5.9a | 3.2±2.4a |
Differences between two levels of management complexity (low and high) on plant diversity and plant functional types. Table shows means and standard deviations of PFT cover and diversity indices. Results are based on LMER Tukey post hoc tests. Significant differences between different levels of management complexity are indicated by the letters ‘a’ and ‘b’.
LOW | HIGH | |
---|---|---|
Species number | 35.0±3.8a | 34.6±5.8a |
Shannon diversity | 1.7±0.4a | 1.9±0.4b |
Simpson diversity | 0.6±0.1a | 0.7±0.1b |
Forbs cover (%) | 24.2±17.2a | 25.6±15.3a |
Graminoid cover (%) | 76.3±21.2a | 66.3±16.2a |
Phanerophytes cover (%) | 3.4±2.9a | 5.2±4.3a |
Mean plant height (cm) | 29.7±10.2a | 22.2±8.6a |
Total plant cover (%) | 94.4±4.5a | 91.5±7.5a |
Bare soil surface (%) | 1.0±0.9a | 1.8±2.0a |
Litter cover (%) | 5.2±3.9a | 7.3±6.8a |
Differences in species composition dissimilarity between management types and levels of herbage removal intensity and management complexity, based on PERMANOVA analyses and Jaccard dissimilarity index.
Df | Sums of Sqs | Mean Sqs | F-test | R2 | Pr(> F) | |
---|---|---|---|---|---|---|
T | 2 | 3.126 | 1.563 | 5.792 | 0.099 | 0.001 *** |
Residuals | 105 | 28.332 | 0.270 | 0.901 | ||
Total | 107 | 31.458 | 1.000 | |||
I | 1 | 0.980 | 0.980 | 3.409 | 0.031 | 0.001 *** |
Residuals | 106 | 30.478 | 0.288 | 0.969 | ||
Total | 107 | 31.458 | 1.000 | |||
C | 1 | 1.783 | 1.783 | 6.368 | 0.057 | 0.001 *** |
Residuals | 106 | 29.675 | 0.280 | 0.943 | ||
Total | 107 | 31.458 | 1.000 |
Proportion of principal components expressed by eigenvalues, explained and cumulative proportions and their contribution to the variance.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | |
---|---|---|---|---|---|---|---|
Eigenvalue | 724.490 | 179.844 | 50.093 | 14.880 | 7.097 | 2.952 | 0.420 |
Explained share | 0.739 | 0.184 | 0.051 | 0.015 | 0.007 | 0.003 | 0.000 |
Cumulative share | 0.739 | 0.923 | 0.974 | 0.989 | 0.997 | 0.100 | 1.000 |