3.1. Spontaneous Arable Weed Flora
Farming practice has been identified as the main factor explaining arable plant species diversity and abundance as several studies have stated [
22,
31,
32]. As an important factor for explaining segetal species richness, the management system was accessed concerning its impact on species diversity and species abundances. The management system is an indicator of the intensity of land use. Conventional farming represents a highly intensive farming system. Organic farms decrease inputs or do not use external inputs like fertilizers and can be considered as moderate to low intensive farming systems. The factors area, management and the interaction of area x management were analyzed concerning their impact on species numbers and diversity indices in current plant coverage. It was found that management was the only factor showing a significant impact on the species numbers in the dataset of the relevés.
Current vegetation data were analyzed concerning the impact of different factors on species numbers and on Shannon Diversity Index. Following
Table 1, the factors management and crop have a significant impact on species numbers with alpha = 0.001. However, the area was not found to be a significant explaining factor for species number variance. In general, species number and Shannon Diversity Index show similar patterns across the four different field trial variants (Gl-org, La-org, Gl-con, La-con). The factor crop is slightly less significant when testing its impact on Shannon Diversity Index (α = 0.05) compared to its impact on species richness (α = 0.01). Interaction effects were not significant in both species richness and Shannon diversity measurements indicating an independent experimental setup. Studies by Chamorro et al. [
4] have shown the potential for a conversion from conventional to organic farming practices affecting the recovery of a diverse arable plant community. Species abundance could be increased from 61 to 122 within a period of five years after the transition from conventional to organic farming system Chamorro et al. [
33].
The GLM found significant effects of all factors and all interaction effects. Species numbers in the region “Lahntal” are reduced compared to those of the other project region “Gladenbacher Bergland”.
All factors except the factor area La have positive effects on the species numbers per square meter in the fields. Concerning the variance of the data, area and management are the two factors explaining the main proportion of the variance of the data. For the spontaneous weed flora, average species numbers and Shannon Diversity Indices varied significantly among the four considered variants Gl.con, Gl.org, La.con and La.org as depicted in
Figure 2.
The organic variants Gl.org and La.org show significantly higher species numbers than the conventional variants. The number of species in the soil seed bank identified with the seedling-emergence method is higher than the number of species in current vegetation in three out of four variants. A reason for lower species numbers with the in-field identification might be the effect of competition between weeds and crops.
With the method of soil seed bank germinating, there is no crop, which will compete with the weeds for natural resources. This explains the higher number of wild herb species with the germinating method. In all cases of the seed bank method, the number of individuals is higher for the organic farming systems than in the conventional farming systems. The management system has a significant effect on species richness in the fields. In the investigation area of Gladenbacher Bergland there is a difference of 10 species in average between conventional and organic farming systems. The average species number in the conventional variant is eight whereas the average in the organic farming system is 18. In the other area (Marburg-Giessener Lahntal) the mean species number is lower (five species) and in the organic farming system the species number is 19. In both sites, organic farming systems show higher species richness and diversity in wild herbs. Conventional farming systems have selected a few species that do not react as sensitively as other species concerning the input of herbicides and fertilizers.
For all four groups except the conventional one in Gladenbacher Bergland, there are significant differences between the determination methods in species numbers. A reason for lower species numbers with the in-field identification could be the effect of competition between weeds and crops. With the method of soil seed bank germinating there is no crop, which will compete with the weeds for natural resources. This explains the higher number of wild herb species with the germinating method. In all cases of the seed bank method the number of individuals is higher for the organic farming systems than in the conventional farming systems.
The management system has a significant effect on species richness in the fields. In the investigation area of Gladenbacher Bergland there is a difference of 10 species in average between conventional and organic farming systems. The average species number in the conventional variant is 8 whereas the average in the organic farming system is 18. In the other area (Marburg-Giessener Lahntal) the mean species number is lower (5 species) and in the organic farming system, the species number is 19. In both sites, organic farming systems show higher species richness and diversity in wild herbs.
At the significance level of alpha = 0.05 only the factor crop was found to be a significant explanatory variable for differences in the individual and species numbers. However, area and management were not significant in the setting of the trial. For the relevé data, the effect of the factor year, area and management system were checked with a Generalized Linear Model, since the data have a Poisson distribution.
For the relevé dataset, the organic managed fields showed significantly higher average weed coverage as shown in
Figure 3.
The coverage numbers refer to the standardized relevé plot with the size of 100 m
2. The effect of the year on the average weed cover was significant, too. For the organic managed fields, variance of the data was much wider than those for the weed coverage of the conventional fields. Results of a Generalized Linear Model on the coverage of spontaneous weeds are compiled in
Table 2.
Nonmetric multidimensional scaling was used to analyze current vegetation data and to identify patterns in species distribution. NMDS was conducted in R package “vegan” and resulted in
Figure 4.
The best solution for the NMDS model concerning the increase in goodness of fit and the decrease in stress was found with a number of three dimensions. In this case, the correlation between the distance values and the observed dissimilarity expressed as R2 equals 0.97 and stress is reduced to a value of 0.178. The four crops winter rye (WR), winter barley (WB), winter wheat (WW) and spelt (SP) were identified as significant factors.
Most of the arable weed species were found in both areas, in the hill-sites around Gladenbach and the valley of river Lahn and in both of the management systems.
Some species show tendencies of a more specific occurrence concerning the factors area and management—here, Papaver dubium, Galeopsis tetrahit and Galium mollugo were some of the species showing higher abundance patterns in organic than in conventional farming systems.
In an experiment of Hyvönen & Huusela-Veistola [
1] some species with a high steadiness could be identified.
Galium aparine was detected on 41% of the fields followed by
Chenopodium album (68%),
Stellaria media (76%) and
Viola arvensis (84%) as the most frequent species in cereal cropping. This study could also identify some of these species as frequent species in cereal cropping.
Rumex crispus, Galium aparine and
Alopecurus myosuroides were more frequently found in conventionally managed fields. Moreover,
Vicia hirsuta and
Equisetum arvensis and
Cirsium arvense and
Stellaria media showed higher abundance patterns in more intensive cereal cropped fields.
The weed species Cirsium arvense, Centaurea cyanus, Vicia villosa and Rumex crispus were frequently associated with winter rye. In the crops winter wheat and winter barley, the weeds Trifolium pretense, Geranium dissectum, Tripleurospermum inodorum and Thlaspi arvense were frequently represented.
3.2. Soil Seedbank of the Arable Weeds
The reservoir of seeds of mostly annual arable plant species in the soil is an important parameter for analyzing the emerged arable weed flora of the past. Moreover, the soil seedbank allows forecasts regarding to the composition of futures’ arable plant communities and it provides information for conservation and regulation strategies for arable plant species [
34,
35].
The response variable in the soil seedbank dataset (individual numbers) is not normally distributed and follows a Poisson-distribution. For that reason, a Generalized Linear Model was chosen to conduct the analyses. A model with three fixed factors and two interaction terms was found to be the model with the best performance and predictive power. The results of the model are given in
Table 3.
Soil seedbank data was tested with a GLM concerning the impact of the factors area, management, soil depth and two two-way interaction effects between those mentioned factors. The Dredge-Function of the R-package “MuMIn” was used to find the model with the best performance and predictive power. Using the AIC and the AIC delta as a criterion for model selection, the model including all factors and two out three interaction effects was found to be the best one for modelling the individual numbers per square meter. For this model, AIC was 6149.2 and AIC delta was 0.17.
For the soil seed bank species numbers differed significantly from each other between the different farming methods (conventional and organic). Belowground species numbers were twice as high (18 species) as those in the conventional alternative (nine species) as depicted in the left part of
Figure 5.
A Kruskal–Wallis test was conducted with the software “R studio” to assess if organic farming period has a significant effect on the seed numbers of the soil seedbank. Testing results were significant, indicating significant differences in the data. Pairwise Wilcoxon test was followed up indicating significantly higher seed numbers of 18 and 27 years of organic growing, compared to 14 years of organic growing. Other groups did not differ significantly from each other. In the study, a maximum seed density is reached between 18 and 27 years of organic farming. After this period, the tendency of a decrease in species numbers per square meter can be observed.
As shown in
Figure 6, the number of weed seeds per square meter differs from organic management systems as affected by different time periods of organic growing. A Kruskal-Wallis Test was conducted to detect significant differences in the data. Results of this test are given in
Table 4, indicating significant higher seed numbers after 27 years of organic growing compared to 14 years, and also, the number of weed seeds in the soil per square meter is significantly higher in systems of 18 years organic farming, compared to those of 14 years. Evidence of an increase of the soils’ seed potential with an increasing time period of organic growing could be stated. This observation is applicable for the organic growing period until 27 years. Thereafter, however, the increase of seed numbers stagnates or fluctuates.
3.2.1. Analysis of Spring Vegetation
The seedbank data set was divided into three subgroups to access the effect of seasonal changes in the plant species communities. The first group includes species of the seedbank that germinated during a four-week period of April 2019, the second one contains those seedlings from June 2019 and the third one contains all additional seedlings from October 2019. Main germination of seeds was in June 2019. Way fewer seedlings emerged in autumn and spring. As shown in the graphical result of an NMDS in
Figure 7, Organic farming systems provide the highest species richness and species abundance. Species showing higher occurrences in conventional farming during spring are
Chenopodium polyspermum,
Galium aparine and
Juncus bufonius.In organic farming systems, Chenopodium album, Plantago intermediae and Rumex acetosella and Cirsium arvense showed higher abundances than in conventional farming systems.
Arable weeds of the soil seed bank samples were analyzed with a Nonmetric Multidimensional Scaling ordination (NMDS) as shown in
Figure 8. Resulting from this analysis, the management system has the main effect on explaining the variance in species abundance of the soil seedbank.
So, the impact of the farming practice could be stated in both, current vegetation and seedbank. There is a clear grouping of species concerning their occurrence pattern. Organic farming includes the highest proportion of total species numbers. There are a few species found more frequently in conventional farming systems. Tendencies found in current vegetation data are partially reflected by the soil seedbank.
Following seedbank data, the weed species Polygonum aviculare, Poa trivialis, Solanum nigrum, Stellaria media and Juncus bufonius appear as tolerant towards intensive farming methods, whereas the highest proportion of the total species number is only found in organic farming conditions at a higher abundance level. In current vegetation on arable fields, 49 segetal species were identified, which is a share of 60%. In total, 82 species that were identified in the soil seedbank over all fields were included in the experiment. Centaurea cyanus and Papaver rhoeas were mostly found under organic farming conditions which is reflected by both seedbank data and current vegetation. However, the impact of farming practice is considerably higher in the seedbank.
The current vegetation is more directly affected by environmental conditions like climate (rainfall) which results in a higher fluctuation. In contrast, the soil seedbank is reacting way slower and delayed to a change in environmental conditions. The seedbank represents the intensity level in farming over the last couple of years. The management system is the major factor whereas the area shows a lower significance for explaining the variance.
Figure 8 shows higher species numbers in organic cropping systems, indicated by the clustering of species along the factor organic management.
Species tend to have a lower distribution and spread along the factor of environment. Some species show tendencies to occur either in more extensive or more intensive farming. Species that are more likely found in intensive farming are Solanum nigrum, Matricharia chamomilla and Stellaria media. Under less intensive conditions, Fumaria officinalis, Centaurea cyanus and Holcus lanatus can be found more likely.
3.2.2. Indicator Species Analysis
Therefore, a grouping vector of sites was built on the basis of the four field trial variants consisting of two landscape areas (Lahn valley = La, Hillsites = Gl) and two management systems (organic = org, conventional = con). The assumption was met that both management system and landscape area could have an impact on the constellation of segetal species in arable fields.
For the area of Gladenbach and Lahntal under organic farming, the indicator species
Tripleuspermum inodorum and
Myosotis arvensis, Alopecurus myosuroides were the most significant indicators. Following the methodology from De Cáceres (2020) an indicator species analysis was conducted using the R package “indicspecies”. After running the analysis, 16 of 82 species in total were identified as indicator species. K-means algorithm was used to assign the species to four different groups. Following this procedure, four site group combinations were calculated as shown in
Table 5. Most of the indicator species are found in organic farming systems (group-nr. 1), way fewer under conventional farming conditions (groups 2 + 3 +4). A principal component analysis was conducted which resulted in an optimal number of four species groups. Within the four groups the proportion of explained variance in the data is around 76.7%. An indicator species analysis was conducted with the results shown in the table below. In organic farming, there are many species that can be used as indicators for organic managed sites; in contrast there are only two species as indicators for intensive farming methods of conventional agriculture, which are in this case
Daucus carota and
Anthemis arvensis. In this study,
Vicia hirsuta, Cirsium arvenese and
Rumex crispus were detected as indicator species for organic management systems. Intensive farming methods can be indicated by the species
Daucus carota and
Anthemis arvensis.
3.2.3. Impact of Crop Type on Species Richness
The effect of crop type has been stated by several studies with mainly a higher species richness and less geophytes in cereal fields than in root crops [
36]. As shown in
Figure 9 high variation in the weed individual numbers could be observed with significant differences between the crop types.
The highest species and individual numbers were found in the crops maize, red clover and spelt, whereas broad bean and winter wheat showed the lowest individual numbers per square meter.
To access the floristic similarity between the sites, Sorensen quantitative similarity index was calculated:
where:
A—the number of species in one of the two communities compared.
B—the number of species in the second community compared.
C—the number of common species in the compared communities.
In
Table 6, Sorensen’s Qualitative Similarity Index is given for each of the considered pairs of variants. Similarity between organic fields in two different sites was 65% and higher than those between organic and conventional sites (57.3%). Organic and conventional fields differ more in their species composition than organic fields among each other in different locations. However, the similarity between the organic fields is not considerably higher than those between conventional and organic farms, indicating that regional differences between arable weed communities might play a role although on a smaller regional scale.
As shown in
Figure 10, Shannon Diversity varied between spring and summer season. Significant higher Diversity values were found during the summer season. As Mennan & Ngouajio [
37] stated, there are seasonal cycles in the germination patterns of the weeds
Galium aparine and wild mustard (
Brassica kaber). Highest germination rates were stated during May with around 70% germination and a strong decrease in germination of around 30% in August.
Diversity of arable weed communities differs between spring and summer. Higher diversity is found during the summer months, since new species have germinated and contribute to a higher index.
Studies of Lososová et al. [
36] stated seasonal dynamics as a factor causing changes in arable plant community composition with higher species richness and beta diversity during the summer months. Species that were found in summer were mostly those also present in the spring germination.
There is no significant difference in species diversity between the two areas. There is no significant effect of the landscape area on the diversity of segetal species of arable fields. Studies by Swanton et al. [
38] suggest that there is an interaction effect between tillage system and soil type which may influence the distribution of soil seed bank vertically. The management showed significant effects on species diversity. Shannon Diversity Index is almost two times higher in organically managed fields than in the conventional ones. Several studies have stated the impact of an increasing farming intensity (herbicide application, plowing, fertilizing) on the diversity of arable plant communities. With higher intensity levels in farming, diversity of arable plant communities decreased significantly [
10,
39,
40,
41,
42].
The hypothesis was tested that there is a different accumulation rate of weed seeds in different soil depths resulting in different diversity indices. Therefore, a Kruskal-Wallis Test was conducted which has not become significant. Shannon Diversity Indices were plotted, as shown in
Figure 11. Though, species diversity has not been affected by the depth level of the soil sample. The farmers of the study fields have used a mixture of reduced tillage and ploughing. The arable fields show a homogenous mixture of seeds in the soil at a testing depth of 20 cm. A significant difference in the seed number and Shannon Diversity Index could not be found. This observation has also been stated by studies of Feledyn-Szewczyk et al. [
43] which showed the effect of the tillage system on the absolute weed seed numbers in the soil and their vertical distribution in the soil of arable fields. Feledyn-Szewczyk et al. [
43] showed that reduced tillage systems and ploughing systems tend to establish a mixture of seeds in the soil with higher average seed numbers in reduced tillage systems (4515 seeds m
−2 ) compared to ploughing systems (2080 m
−2).
Studies by Clements et al. [
40] analyzed the impact of different soil tillage methods on the seedbank composition of different weeds. It was found that deep soil tillage, like plowing, results in a more homogeneous distribution of the seedbank over depth. In contrast, in a system with minimal soil tillage, Clements et al. [
40] assumed that the upper 5 cm of the soil comprises over 60% of the seed bank. Research by Buhler et al. [
41] could identify grass seed accumulation in minimal tillage systems and found higher accumulation rates at the deeper soil levels in moldboard plowing systems.