1. Introduction
The ongoing socio-ecological processes are heightening the importance of environmental services not only within individual ecosystems but also across their entire complexes. “Eco-efficiency”, the intensification of production, with a simultaneous decrease in the impact on other habitats [
1], is becoming increasingly important in environmental management. On the one hand, there is a growing focus on the quality of habitats, the ecological well-being of populations, and the conservation of wildlife. On the other hand, it is essential to maintain the land area used for forests and crops to meet society’s needs [
2,
3].
The competing objectives outlined above are leading to more frequent clashes between foresters, farmers, and wildlife, particularly involving wild ungulates. In addition to providing compensation for crop damage, there is also a pressing need to take proactive measures to minimize such incidents. One potential approach is to establish dedicated areas with abundant forage to attract the animals’ attention away from farmers’ crops.
In the conditions of Central Europe, such lands can be mid-forest meadows. They play a specific role in the environment [
4]. Due to their characteristics, they have an impact on biodiversity (at all levels and approaches), nutrient flow in ecosystems, and the ecology of plants and animals. In many cases, glades can be characterized as ecotones, with all the advantages of ecological niches [
5]. Nevertheless, the open habitats within consistently managed forest complexes are not a significant element of the Central Europe landscape. Historically, since the 19th century, the focus on the productive aspect of rural areas has led to the neglect of potential benefits associated with maintaining mid-forest meadows [
5,
6,
7]. Currently, with sustainable approaches to the environment, where production is not the only factor for managed lands, the possible utilization of mid-forest meadows can be brought back to the attention. After all, the word “forest” in its original meaning did not refer to woods and production but to the places designated to preserve game species for hunting [
8].
In most places with a temperate climate, maintaining open habitats is strictly correlated with their exploitation by humans and livestock or by wild ungulates [
9,
10,
11,
12]. Neglected grasslands undergo a gradual decline in their forage potential (amount of biomass, herbage quality of harvested yield, and plant diversity), which ultimately leads to degradation, succession towards shrubs and forested areas, or the transformation of these lands into other types of crops [
13,
14,
15,
16,
17]. Since the 1930s, over 90% of semi-natural meadows in Europe have been transformed [
18]. It is especially important in ecotone habitats, where succession tends to be accelerated.
While in the right conditions, the sward of mid-forest meadows can be attractive for animals on its own, in many places (especially among the managed forests), those areas, like all permanent grasslands in Poland, are the subject of degrading factors, e.g., site and climate changes promoting unfavorable (in foraging perspectives) species [
19,
20,
21]. Decreasing yield and nutritional value can be reverted through proper management activities. Appropriate renovation activities, destructive or not for old swards [
20,
22], combined with an effective management strategy, are necessary to ensure the quality of produced biomass [
23,
24,
25]. The intensity of such treatments should be adjusted to the level of competition between all present plant species and desired ones.
Renovation should be tailored to the preferred goal, e.g., requirements of the main herbivores species that will utilize this area. Deer are more selective in comparison to cattle as they also look for the highest amount of available biomass and its “greenness” [
26,
27]. Deer prefer green grasses in the spring and forbs in the summer and tend to avoid patches with dead plant materials [
26]. However, offering the proper quality sward can influence their behavior throughout the seasons [
28]. Of course, patchy grazing by animals causes the uneven utilization of swards [
29], which impacts the efficiency of such areas [
30]. It is purposeful to connect the extensive foraging activity of wild animals with other techniques (e.g., mowing) to prolong the stability of sward productivity [
31].
Good-quality mid-forest meadows, as foraging places, can be a tool to control the populations of wild ungulates, with possible influences on the damage within forests and arable crops [
32,
33]. Besides providing feed, forest grasslands offer a sense of security due to their proximity to wooded areas, making them preferred even when nearby crops might provide slightly better forage.
The aim of the study was to evaluate the effectiveness of mid-forest meadow renovation using different methods in terms of both the quantity and quality of the sward yield. The renovated meadows were utilized as the foraging areas for the free-living deer population.
2. Materials and Methods
2.1. Experimental Site
The study was carried out from 2014 to 2017 on two selected mid-forest meadows in northwestern Poland (54°10′ N, 16°78′ E) by the Department of Grassland and Natural Landscape Sciences, Poznan University of Life Sciences, in cooperation with Polanów Forest District (Polish State Forest).
This study was conducted on the area managed by the Forest Wildlife Breeding Centre, characterized by a higher density of herbivore wildlife animal populations, in this case, red deer, compared to adjacent areas.
2.2. Weather Data
Meteorological data for Polanów (
Table 1), collected from the Visual Crossing platform [
34] and analyzed using the Vinczeffy Climatic Index (VCI) [
35], indicated that the meteorological conditions in the experimental area generally reflected the conditions across northwestern Poland [
36]. In 2013, the average VCI values classified the year as “medium-wet” (>0.151 mm/°C), with stable conditions observed during the vegetation season (April to October). In contrast, 2014 experienced extremely dry conditions (0.082 mm/°C) throughout most of the year, characterized by very low precipitation coupled with higher-than-average temperatures. Dry conditions persisted through the first half of the 2015 vegetation season (April to July). While meteorological conditions improved from August onward, the overall year was still classified as very dry (0.106 mm/°C). In comparison, 2016 and 2017 recorded notably higher VCI values (above 0.200 mm/°C), which placed them in the “rainy” category.
2.3. Experimental Design
Two experimental mid-forest meadows were established on two adjacent sites, located approximately 0.5 km apart. At these sites, each of the experimental treatments was established on an area of approximately 0.3 hectares. Such large areas were necessary to carry out the tested renovation methods using the full-scale machines available at the Forest Wildlife Breeding Centre. We also intended to check the feasibility of the applied renovation methods in practice. Two experimental mid-forest meadows were defined in the article as follows:
Object no. 1—Wnęka eng. Niche (object W)—54°6′16.761″ N, 16°47′30.403″ E, av. elevation 117 m: The renovation of this mid-forest meadow began at the beginning of the growing season in 2014 (sown 09.04.2014). While the entire site covered approximately 5.5 hectares, it was only partially utilized for the experimental purpose with five treatments designated for the study in one replication, with five large plots/research areas created (ca. 0.3 ha each). The reason for such decision was the terrain relief of object W because only about 1.5 ha was located on a flat terrain as in the case of the second mid-forest meadow, and the remaining part was hilly, which could have influenced the emergence, growth, and development of plants after sowing due to different exposures to light. The sward botanical composition before renovation was dominated by Dactylis glomerata (48.9%), Festuca arundinacea (18.9%), Festuca rubra (7.6%), Holcus mollis (10.3%), and other dicotyledonous species of low nutritional value.
Object no. 2—Topolowa Droga eng. Poplar Road (object TD)—54°6′5.925″ N, 16°46′55.418″ E, av. elevation 132 m: This mid-forest meadow underwent renovation in the beginning of autumn in 2013 (sown 28.09.2013). The entire object covered an area of 2.5 hectares and was fully adapted for the experiment. Within this meadow, four treatments were designated for this study, established randomly in two replications of approximately 0.3 hectares each (in total eight plots/research areas). The sward botanical composition before renovation was dominated by Festuca rubra (29.9%), Dactylis glomerata (14.2%), Festuca arundinacea (4.3%), Sonchus arvensis (4.0%), and other mono- and dicotyledonous species of low nutritional value.
The names of the objects (Wnęka and Topolowa Droga) are the common names of mid-forest meadows used in the Forest Wildlife Breeding Centre. Both objects are located on soils with a low class in the Polish Soil Classification system, indicating very poor agricultural suitability. These soils are typical brown soils with a granulometric composition dominated by sand with low fertility (
Table 2). Light clays were present in part of object W, but this area was excluded from the experiment [
32]. Overall, the habitats were defined by low nutrient content and limited water retention capacity. According to the Agricultural Drought Monitoring System [
33], the entire area of object TD falls within the “very susceptible to drought” category (the weakest one). About 50% of object W’s area is classified similarly, but due to the other environmental factors (slope, forest formation, altitude), the experiment was located on soils “susceptible to drought”.
On both objects, a comparison was conducted between control treatment (C), which did not undergo any renovation efforts, and treatments represented different renovation techniques. We tested the following methods:
Overdrilling using seeds of perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) (O): The overseeding technique included harrowing, which was used for sod preparation; then, the seeds in the rate of 20 kg/ha were sown using the Väderstad Rapid 300s Super XL seeder followed by rolling.
Full tillage usage for sowing the three types of seed mixtures M1, M2, and M3 (detailed composition is given in
Table 3): The full tillage technique included ploughing, then the harrowing operation for seedbed preparation and sowing seeds in the rate of 40 kg/ha using the Väderstad Rapid 300s Super XL seeder, followed by rolling. Additionally, cleaning cut was performed 6 weeks after sowing to prevent weed infestation.
Mixtures M1 and M2 were commercially available, and M3 was used by the authors for experimental purposes. Due to areas of both mid-forest meadow and seed mixture availability, the experimental design differed between the objects: TD included treatments C, O, M2, and M3 in two repetitions, and W included treatments C, O, M1, M2, and M3 in one repetition.
Both mid-forest meadows were previously utilized by the Forest Wildlife Breeding Centre as foraging areas for wildlife. The management schedule included fertilization with an application of 9 kg/ha of nitrogen, 30 kg/ha of phosphorus, and 45 kg/ha of potassium at the beginning of each growing season. The areas were grazed mainly by the free-living red deer population. The remaining non-grazed part of the sward was regularly mowed both in the beginning of summer and at the end of the growing season in autumn. The sward yield obtained from the mowing was subsequently conserved in the form of hay.
For eliminating the impact of animals on the sward, three 2 m × 2 m wire mesh grazing exclusion cages were placed on every experimental treatment. Grazing exclusion cages were used for the estimation of forage accumulation, which is a key response in grazing experiments on continuously stocked pastures. The management of experimental areas did not differ between the ungrazed area (exclusion cages) and grazed area, which was available for animals. The grazing exclusion cages were dismounted during the mowing events and removed with each growing season. The location of the grazing exclusion cages within the treatment areas was randomly changed during the following experimental years. This made it possible to carry out analyses on ungrazed and grazed areas.
2.4. Methods
For the assessment of habitat conditions, in which the effectiveness of the mid-forest meadows renovation was tested, soil moisture analysis was conducted using an ML3 ThetaProbe connected to an HH2 reader (Delta-T Devices Ltd., Burwell, UK). The device was configured with standard settings for mineral soil analysis (root depth—50 mm; field capacity—0.380 m3/m). Measurements were conducted on designated dates approximately every two weeks, from the beginning of April to the end of October in each study year between 2014 and 2017. Data were collected from six locations within each area. Additionally, on the same dates and similar regime of data collecting, the vitality of the sward dominant species was determined to assess the fertility of the habitat. The Soil Plant Analyses Development (SPAD) index, as a greenness indicator, correlated with the nitrogen status in plants recorded using Minolta SPAD-502Plus (Konica Minolta Optics, Tokyo, Japan) with standard settings. Due to the natural occurrence on all treatments, Dactylis glomerata was selected as the representative species.
Analysis of sward botanical composition was based on the yield proportion method [
37]. The sward samples, collected from each grazing exclusion cages using a square metal frame with three subsamples per cage (9 subsamples per plot per date), were separated into individual species, dried, and weighed. Based on the relation between the weight of each species to the total DM yield of the samples, the botanical composition was assessed. To unify the results between the experimental treatments, the individual results were summarized and presented as the proportion of plant functional groups (grasses, herbs, legumes).
Assessing the sward productivity of each treatment was based on the amount of sward biomass collected from the grazing exclusion cages (without grazing pressure). The timing of these assessments was synchronized with the standard management schedule for mid-forest meadows within the Polanów Forest District. In particular, sward harvest for hay production were conducted in June and October, before the end of the growing season. Sward samples were manually cut at an approximate height level of 5 cm from the ground within the area of each grazing exclusion cage (in 3 repetitions) using a square meter frame (0.5 m × 0.5 m) (9 repetitions per area per date). The collected herbage was dried in an oven at 60°C for 48 h for dry matter (DM) content determination. Based on the collected fresh biomass amount and DM content in the sward, the DM yield of each treatment (in kg DM per ha) was estimated.
The herbage quality of the renovated mid-forest meadows was evaluated using Wendee proximal feed ingredient analysis [
38] and dry matter digestibility (DMD). The analysis of the five basic components of dry matter for evaluation of nutritive value of forage (ash, crude protein (CP), ether extract (EE), crude fiber (CF), nitrogen-free extract (NFE)) was performed using near-infrared reflectance spectroscopy (NIRS) utilizing the FOSS DS 2500 Analyzer (Foss NIR Systems, Laurel, MD, USA). Sward samples from each experimental treatment were collected from grazing exclusion cages during the yield quantity sampling (covering all collection dates) and were prepared in accordance with the laboratory’s guidelines. The preparation protocol involved drying, grinding, and thoroughly mixing the samples, ensuring each had a minimum weight of 100 g. The dried samples taken during the period of investigations were then sent to the laboratory in two batches, one in 2015 and another in 2017. DMD was assessed in relation to acid detergent fibers (ADF) using the equation proposed by Di Marco [
39].
2.5. Statistical Analysis
The normality of the collected data was assessed using the Shapiro–Wilk test. Due to the instability of some of the habitat parameters, the Interquartile Range (IQR) method was used to detect and remove outliers from the data. Given the non-normal distribution of most data obtained from all experiments, the significance of differences between renovation methods was estimated using the Kruskal–Wallis test (one-way ANOVA on ranks). The statistical analysis was conducted using R Statistical Software (ver. 4.3.0) with the Agricolae package (function kruskal()). Groups that were statistically different were distinguished by a multiple comparisons test of rank sums, which is conceptually similar to Nemenyi’s post hoc test, and adjustments for multiple comparisons were made using critical values derived from rank distribution tables. Both procedures were applied within the used function.
4. Discussion
The objective of this study was to evaluate the most effective way to renovate the mid-forest meadows in terms of productivity. Environmental conditions noticeably influenced the experimental results observed at both meadows. Nevertheless, both sites presented challenging conditions, characterized by sandy soils, variable weather patterns, and unfavorable ecotone factors such as shading and uneven soil moisture levels due to the adjacent forest areas. In such conditions, two experiments were conducted in which different methods of renovation of mid-forest meadows were analyzed: overdrilling and full tillage methods using three mixtures with different botanical composition (M1, M2, and M3). In the research approach, it was very important to locate the experiments in the real habitat of mid-forest meadows, which are used by the free-living deer population.
Soil moisture stands out as a crucial factor influencing the potential success of grassland renovation and can be a strong predictor of meadow productivity [
40,
41,
42]. Grasslands are more tolerant of excessive water supply [
43] but are also more vulnerable to droughts compared to other ecosystems due to the shallow root system of most grasses. The availability of water is intricately linked to the quality and quantity of biomass produced, especially in short-term analyses [
44,
45], and water deficits can cause annual variations in yield. Soil structure, nutrient availability, and habitat patchiness show a direct correlation with soil water content [
46]. The ecosystemic interplay of soil, microbes, and plants is directly reflected in forage quantity and quality [
18,
47,
48,
49]. These relationships played an important role on the sites where the experiments were conducted. As indicated in
Figure 1, more difficult habitat conditions occurred on the object TD than on object W. Nevertheless, as a result of the renovation of the mid-forest meadows, satisfactory effects were obtained, measured by dry matter yield and herbage quality.
The habitat quality was also evident in the plant leaf greenness, as assessed using the SPAD index of
Dactylis glomerata as the representative species, because of its presence in the botanical composition of all analyzed swards. This parameter served to describe the overall vitality of the sward and the plant vigor [
50]. In controlled conditions, during the pot experiments, changing the level of soil water deficit caused the growth of the index in
Dactylis glomerata [
50,
51]. However, in field experiments, where environmental factors cannot be controlled, the SPAD level reflects the outcome of the more complex soil–plant feedback. As the leaf greenness is strictly attached to the chlorophyll content, the SPAD level can describe the photosynthetic capacity, as well as nitrogen content in the plant [
52]. Environmental factors influence the pigment composition in the plant tissues, making the SPAD index one of the key plant ecophysiological analyses [
53]. In the performed experiments, the significantly lower SPAD index was determined in the object TD in comparison to object W (
Figure 2). This poor condition also translated to the lower performance of the renovation methods, particularly at object TD, supporting the notion that site-specific factors play a crucial role in renovation success (49–52). The original sward’s yield in both mid-forest meadows was significantly influenced by habitat conditions. The total yield from control area in most experimental years, with the exception of object W in 2015, fell below the average for permanent meadows in Poland [
54]. On the other hand, mid-forest meadows are often located in less agriculturally valuable areas in the point of view of its soil fertility; hence, the obtained level of productivity should be assessed as satisfactory.
Overdrilling with Lolium perenne and Trifolium repens, chosen as an easy method to improve sward botanical composition, did not influence sward performance under the analyzed conditions. This underperformance is evident in the biomass produced, as the overdrilling treatment generally yielded slightly less than the control at both experimental objects. Furthermore, the limited representation of legumes each year suggests that the “new” seeds were outcompeted by the existing sward. The poor results of overdrilling the forest meadows at both sites were also due to difficult habitat conditions, especially soil moisture shortages in some periods, which turned out to be crucial for introducing valuable species to the existing turf.
The characteristics of the habitat patches directly impacted the survival and establishment of new plant species within the existing vegetation [
55,
56,
57]. Theoretically, better effects of the overseeding method were obtained on mineral soils [
58], but plant diversity, weed control, and overall performance remained unaffected by this technique, likely due to the persistence of existing vegetation and competition from established grasses [
48,
59,
60]. The lack of effects on yield quality or botanical composition suggests that habitat factors played a pivotal role, and increasing the seeding rate of any species used would not significantly affect the results [
61] or planting depth [
62]. Some studies suggested that the grazing can reduce sward competition [
55,
63], but for mid-forest meadows used as forage for the red deer population, such factors were insufficient to reduce competition.
The findings suggested that without additional measures, such as soil aeration techniques and targeted herbicide application, coupled with proper post-sowing care [
21,
64,
65,
66], overdrilling may have limited success in mid-forest meadow renovation. However, adding such operations would limit the main advantages of this procedure—the cost-effectiveness and minimalization of soil structure damages [
67].
The full tillage method effectively removed competition from old sward plant components, facilitating the establishment of new seedbeds. This approach proved to be a successful renovation strategy by creating an environment favorable for the growth of desirable sown species, particularly legumes and grasses. These species are crucial for enhancing the productivity and forage’s nutritional value [
14,
60,
68]. Additionally, full tillage positively influenced species diversity on the areas [
24,
66]. Although all treatments remained grass-rich (more than 70% of biomass) [
69], significant improvements in both yield quantity and quality were observed.
At object W, the highest overall yield was observed for the M3 mixture, followed by M2 and M1, with significant differences in yield quantities among all mixtures. Both M3 and M2 mixtures yielded sustainably throughout the experiment. In contrast, the M1 mixture, after an initial peak yield exceeding 9000 kg/ha, declined below the levels of the other two mixtures. This decline in M1 is attributed to a reduction in herbs (from 15% to 0.7%) and legumes (from 22% to 5.5%), changes not observed in the other mixtures. The performance loss was mainly due to heavy grazing pressure by wild red deer, which preferred this mixture in the first year of the experiment [
70]. Such forager impact can significantly alter biomass flow, plant species composition, growth rate, and nutrient flow, potentially damaging promising swards [
71]. The experimental areas were utilized as forage areas for game species without restricted access, considering this an environmental factor [
72].
Additionally, object W consistently demonstrated higher productivity measured by dry matter yield for all mixtures compared to object TD. For mixture M2, object W’s yield was 6712 kg/ha, almost double that of object TD’s 3691 kg/ha. Similarly, for M3, object W’s yield was 6415 kg/ha, more than double object TD’s 2995 kg/ha. These differences highlight the impact of object-specific conditions on yield, with object W’s more favorable environment (particularly soil moisture) supporting higher productivity.
Interestingly, while object W produced higher overall yields for both mixtures M2 and M3, object TD yielded forage with superior nutritional quality in several aspects. The forage from object TD exhibited higher crude protein (CP) and crude fat (EE) contents, lower crude fiber (CF) content, and slightly better dry matter digestibility. The impact of challenging environmental conditions, including the ecotone effect of tree canopies [
73], on various yield quality parameters is not fully understood and varies among species. However, increased forage quality under stress is supported by the literature [
74]. It appears that the functional group ratios in mixtures M2 and M3 responded better to stress factors (habitat, foraging, etc.) in mid-forest meadows. Additionally, the higher presence of weeds and herbs (e.g.,
Taraxacum officinale) can not only influence forage quality [
12] but also complicate predictions about forage quality [
75].
The notable effect of the full tillage method combined with sowing seed mixtures in enhancing both productivity and forage quality emphasizes the importance of selecting appropriate seed mixtures based on local conditions and desired forage outcomes [
22]. This was supported by Galvin [
57] who found that soil disturbance combined with the introduction of high-yield species significantly increases biomass production in semi-arid landscapes. The effectiveness of renovation techniques varies based on how they modify soil and plant interactions [
76]. Full tillage often resets these interactions more completely than overdrilling, providing a more favorable environment for the establishment and growth of desired species.