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Mean difference effect size, g, and a mixed random effects model were used PR — plant species richness; AR — animal species richness; AA — animal abundance. Differences in species richness were most pronounced in the fungi Table 3. However, land abandonment also showed significant increases in animal and plant species richness. Among the taxa, we found significantly higher effect sizes for lichen and birds on abandoned land, while there was no global effect of land abandonment on arthropod and vascular plants Table 3 , Figure 4A. No clear patterns emerged for the extent of the study areas Figure 4C.

The number of years since abandonment did have a significant impact on effect size; but only studies that covered an abandonment period of thirty to forty-nine years not those with fifty or more years or less than thirty years of abandonment showed significant increases in species richness and abundance Table 3 , Figure 4D. Agroforestry, arable land, and pastures showed significantly different effect sizes between groups Table 3 , Figure 5A. On agroforestry and arable land, species richness and abundance increased after abandonment, while it decreased on pastures.

Permanent crops did not exhibit significant effects. D displays a bubble plot of the relationship between effect size and precipitation, with the size of the bubbles scaled according to the reciprocal of the standard deviation of the effect size. Effect sizes of studies performed in mountains and lowlands significantly differed from each other Table 3 , Figure 5B , with abandonment in lowland areas showing stronger increases in plant and animal species richness and abundance. Temperature did not show significant effects, whereas areas with high precipitation showed significant declines of plant and animal species richness and abundance after abandonment Table 3 , Figure 5D.

Land abandonment potentially has substantial environmental and socio-economic consequences [5]. This study presents the first formal meta-analysis that examines the particular impacts of land abandonment on biodiversity, using animal species richness, animal abundance, and plant species richness as indicators.

The analysis focused on the Mediterranean Basin, an area of comparable climate where land abandonment is prevalent. The meta-analysis revealed that land abandonment has been shown to slightly but significantly result in increases in plant and animal species richness and abundance. However, heterogeneity in responses to abandonment was high.

Among the empirical cases used in the meta-analysis, many pointed to increases, and others to decreases, in biodiversity after farmland abandonment. For example, when a simply structured vineyard in Israel was abandoned, the mean species richness values of perennial plants between vine rows increased from 0.

In contrast, mean plant species richness declined after abandonment from In some empirical studies, effect sizes went in different directions when different species groups [51] or different farmland habitats [52] were investigated. Using a diversity of indicators, a qualitative, global review of land abandonment came to similar insight, with 77 studies pointing to biodiversity losses, but another 39 studies reporting increasing biodiversity [3]. Thus, the responses of species richness and abundance are not consistent enough to support general conclusions about biodiversity trends on abandoned lands in the Mediterranean.

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Rather, these responses seem strongly mediated by the specifics of each case study, whether they pertain to spatial-temporal scale, land-use, landforms, climate, or other parameters. In regard to objective 1 , the diverging views on increases or decreases in plant and animal populations that result from land abandonment can be partly explained by the different taxonomic groups involved. All three kingdoms animals, fungi, plants showed an overall positive effect size after abandonment, but the strongest one was found for lichen remembering that all lichen cases were taken from one publication only.

Bird species richness also showed clear increases in response to land abandonment. Responses of vascular plant richness were heterogeneous, with some plant communities favored by agricultural management very likely those composed of ruderal, stress tolerant, and competitive farmland species and some very likely those composed of shrubland and woodland species favored by abandonment. A meta-analysis of land abandonment effects on bird distribution changes also found such heterogeneous differences, with decreasing occurrence of farmland bird species and increasing occurrence of woodland and shrubland species after abandonment [19].

As for objective 2 , our results showed that the temporal dimension of land abandonment studies is important [53]. Plant species richness often increases and exhibits strong dynamics in the first years after abandonment, but later species composition becomes more stable and species richness decreases.

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The intermediate disturbance hypothesis offers one potential explanation, as it predicts that plant competition has a greater influence on plant community development when it is not interrupted by disturbances such as cultivation, drought or grazing [50] , [54]. In a highly competitive environment, less successful competitors are often eventually suppressed. In our meta-analysis however, only studies considering an abandonment period of 30—49 years showed significant increases in species richness.

Obviously, several decades are needed until colonizers in the regional species pool trigger community succession. Therefore, our results highlight that comparatively long time periods are required before general increases in species richness and abundance can be detected. However, a non-significant decline in species richness after an abandonment period of 50 or more years may indicate that exclusion processes eventually follow colonization processes in many of the case studies. Substantially different outcomes were revealed for different agricultural systems [55] when objective 3 was investigated, confirming previous studies of the influence of farm-level attributes on biodiversity [56].

Species richness and abundance generally increased on cultivated habitats arable land, agroforestry after abandonment, and decreased in abandoned pastures. Cultivated habitats are generally more disturbed by agricultural activities and more distant in species composition from natural ecosystems than are pastures.

Effects on biodiversity values are also likely to vary within arable or pasture lands for the same reason. Biodiversity impacts after abandonment may differ between highly mechanized and simplified croplands for example, and traditionally grazed native pasture land where grazing may moderate competitive exclusion [59]. Regarding objective 4 , some effects of landforms, climate, and other contextual factors were revealed.

Particularly influential was the ecological region of the Mediterranean Basin where the study was carried out. Land abandonment impacts were more negative to species richness in areas of higher precipitation Table 3 , i. This pattern supports prevailing notions about non-equilibrium systems [60] ; in accordance with non-equilibrium concepts, in areas where abiotic factors do not limit competition as a major driver, human disturbance may favor greater species richness. Therefore, higher precipitation may lead to high levels of competitive exclusion when disturbance from agriculture ceases.

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When interpreting the results of our meta-analysis, several caveats need to be taken into account. Although meta-analysis is acknowledged as a straightforward method that yields robust quantitative results, relevant information reported in the empirical studies used may be lost, and some relevant studies may be missed in the selection process. Our search found that many papers could not be included because necessary summary statistics were not provided. This information could be gained from some, but by no means all, authors.

We tried to ensure comparability between primary studies by restricting our analysis to the Mediterranean Basin, and by adding standardized information temperature, precipitation, landform, protected area status from databases that covered the whole Basin or at least the European part of it. However, the strong variability that we found indicates that the relationship between land use and biodiversity in the Mediterranean may be too complex for general conclusions. Our analysis may be further limited because of publication bias, the idea that studies reporting significant differences are more frequently published than studies that do not find significant differences.

The distribution of effect sizes of our cases is rather symmetric and normal Figure 3 , so it does not indicate any obvious publication bias. Rather, most effect sizes are moderate, and only few are large. Before-after studies may be sensitive to random factors such good or bad years in terms of rainfall. However, only 2 out of the cases were before-after studies, so we believe that the influence of such factors on the outcome of our meta-analysis is low.

Another issue to be discussed is whether the inclusion of several cases per published study leads to pseudoreplication. Having the number of cases exceed the number of studies is very common in meta-analysis studies [61]. It can be addressed by randomly selecting one case per study and examining whether the confidence interval of the effect size for these selected cases is different from the confidence interval for all cases.

Consequently, the effect of pseudoreplication is not significant and our way of analysis is acceptable. Perhaps the most important limitation to address is the selection of biodiversity indicators. As have many other meta-analyses [57] , [63] , [64] , we focused our study on species richness. However, species richness can be an unreliable indicator of biodiversity [53] , and more sophisticated comparisons based on species composition would be more informative [65]. In addition to species richness, we considered species abundance for the assessment of the biodiversity impacts of land abandonment, as diminishing abundance may translate into reduced genetic diversity of populations [66].

Our approach did not allow us to consider studies of other dimensions of biodiversity, for example of differences in ecosystem diversity [67] , or of population changes in individual species [19].

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If these parameters were to be included, we suspect the overall impacts of land abandonment in Europe might more often lead to decreases in biodiversity [3] , [18]. In addition, our meta-analysis did not assess species composition. As a result losses in farmland biodiversity and especially agrobiodiversity that accompany many abandonment processes [27] might have been overlooked.

Our sample of primary studies was not distributed evenly across the Mediterranean Basin. In contrast, not a single one was assessed from the southern shore of the Mediterranean Basin. This might reflect a biased selection of study cases, but in large part can also be attributed to the fact that land abandonment is a particular phenomenon of the European Mediterranean, as land use pressure remains high in African and Asian regions [25] , [69].

Given that they represent regional-specific land-use systems, the dehesa and montado agroforestry systems of the Iberian Peninsula received a lot of attention in studies. In contrast, the biodiversity outcomes of land abandonment were comparatively little studied for arable land. Also, not all taxa received equal attention, and current studies do not allow identifying the specific kinds of plant and animal communities that are favored or hampered by land abandonment.

Future research studying land abandonment should strive to fill the gaps identified in this paper by focusing on neglected taxa and regions and by studying effects on species composition, turnover, and functional biodiversity rather than species numbers.

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  6. Other promising directions might be closer examination of biodiversity outcomes under different intensities of land management e. In particular, the landscape context of land abandonment and biodiversity needs much more attention. It is challenging to explain the contrasting impacts of a complex and spatially diverse process such as land abandonment [18]. Synthesizing the results of cases throughout the Mediterranean Basin, this meta-analysis indicates a slight increase in overall species richness and abundance after land abandonment.

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    The effects of land abandonment on biodiversity were mediated by a broad set of drivers.