- Open Access
Congruent responses of vascular plant and ant communities to pastoral land-use abandonment in mountain areas throughout different biogeographic regions
Ecological Processes volume 11, Article number: 35 (2022)
There is a long-term trend towards the abandonment of agro-pastoral activities in the mountain areas of Europe: the following encroachment process of semi-natural grasslands by shrubs is one of the main severe threats to the conservation of biodiversity in mountain environments. To better understand the impact of land abandonment, we analysed the reliability of plant functional groups, ant traits, and ant functional groups as indicators of land use changes. We carried out the research in Italy at four sites along a latitudinal/altitudinal gradient in three biogeographic regions (Mediterranean, Continental, Alpine). We identified three stages of a chronosequence at each site as representative of the plant succession in response to pastoral land-use abandonment.
As expected, both the plant and ant assemblages considerably differed across sites at the species level and, within each site, among the three stages. This trend was found also using ant traits, functional groups of ants, and plant functional groups. Ant and plant communities were related in terms of composition and functionality. Harvester ants and ants with collective foraging strategy were associated with annual legumes and grasses (Therophytes); ants with a strictly individual foraging strategy with Phanerophytes. Ant traits and plant functional groups indicated significant differences among the three stages of the chronosequence. However, ant functional groups could not discriminate between the stages represented by secondary grasslands currently grazed and shrub-encroached grasslands ungrazed.
Despite some limitations of ant functional groups in explaining the succession stages of land abandonment, our results suggest that ants are a good surrogate taxon and might be used as bioindicators of land-use changes and ecosystem functioning. Furthermore, our findings indicate that the functional group approach should be applied to other European ecosystems. Finally, reducing the taxonomic complexity could contribute to developing predictive models to detect early environmental changes and biodiversity loss in mountain habitats.
Agro-pastoral activities in mountain areas are crucial for maintaining the viability of rural traditions in Europe and are particularly relevant for resource and nature conservation (Gómez Sal and González García 2007). However, the abandonment of mountain environments due to socio-economic changes after the Second World War led to a drastic reduction of agro-pastoral activities and changed land uses (MacDonald et al. 2000; Tasser and Tappeiner 2002). Pastoral activities have shaped the landscape into a mosaic of plant communities supporting different levels of biodiversity and providing different ecosystem services (Yapp et al. 2010; Bagella et al. 2013; 2020; Pittarello et al. 2020). On the other hand, the cessation or the limitation of grazing management for several years has led to natural vegetation succession processes (Garbarino et al. 2013). Dwarf shrubs first, and trees later, were responsible for the encroachment of broad areas of semi-natural grasslands, which is considered one of the main threats to biodiversity conservation in mountain environments (Laiolo et al. 2004; Koch et al. 2015; Pittarello et al. 2016).
Identifying key taxonomic groups and possible relationships among them and land use patterns can help standardise efficient tools for biodiversity assessment and monitor the effects of land-use abandonment (Larrieu et al. 2018). Vascular plants are recognized as particularly suitable indicators in monitoring and conservation: they provide the physical structure for other organisms, including many habitat specialists distributed across broad environmental gradients, and are well known from a taxonomic point of view. For these reasons, vascular plants are traditionally considered one of the key surrogate taxonomic groups to select areas of concern in biodiversity conservation as they can reflect the diversity of other significant and less easily detectable taxa (Pharo et al. 2000; Burrascano et al. 2011; Bagella 2014; Bagella et al. 2014). However, many arthropod groups are considered good land-use change or abandonment indicators. Among them, ants are effective environmental indicators in almost all terrestrial habitats because of their wide range of climatic niches and their ability to cope with varying environmental conditions (Andersen and Majer 2004; Jiménez‐Carmona et al. 2020). Besides, they play multiple roles in food webs: herbivores, decomposers, predators, parasites, seed dispersers, and pollinators (Hölldobler and Wilson 1990). As they are influenced by a more complex suite of environmental variables than plants, ants can provide useful information on short-term environmental changes in highly dynamic and vulnerable ecosystems (Zina et al. 2021).
Simple metrics as species richness have proved to be unreliable tools in biodiversity monitoring (Andersen and Majer 2004; Wan Hussin et al. 2012). Indeed, ecosystem processes are affected by the functional traits of organisms involved, and the taxonomic identity is not suitable to make functional comparisons among different biogeographical regions (Westoby et al. 2002; Garnier et al. 2004; Segar et al. 2013). Since functional traits mediate the response of organisms to the environment and influence ecosystem functioning (Chelli et al. 2019), they may help predict how communities might respond to landscape changes. Likewise, functional groups can be helpful and cost-effective to detect congruent and robust patterns of community structure between regions by transcending the effects of historical factors on species composition (Hoffmann 2010; Laureto et al. 2015). The definition of functional groups, based on specific traits depending on the aims and scale of the research (Smith and Huston 1989; Wilson 1999), has a long history in ecology, especially for plants (e.g. Raunkiaer 1934). The impact of pastoral land-use abandonment on vegetation has been investigated, and previous studies showed that plant traits provide insight on key processes driving vegetation changes (Prévosto et al. 2011). Plant functional groups were also successfully used for measuring the recovery of vegetation after cropland abandonment (Heydari et al. 2020). Ant functional groups have a relatively recent history (Andersen 1995). Although Andersen's scheme (1995) is still questionable outside Australia, it could be a useful generalisation in broad studies of functional trait variation in Europe (Gómez et al. 2003; Azcarate and Peco 2012; Satta et al. 2012; Verdinelli et al. 2017).
Despite the importance of ant-seed dispersal in conserving semi-open habitats of European pastures (Travers et al. 2020), information on ant-plant relationships for the European mountain areas is minimal (Hevia et al. 2016; Heuss et al. 2019). In addition, a multispecies approach can provide a better assessment of the overall diversity of an area, reflect more accurately changes in diversity caused by habitat modification, and provide more complete information for halting the biodiversity loss (Hevia et al. 2016). Therefore, we aimed to investigate the possibility of using very different taxa as bioindicators by analysing the effect of land abandonment on ant and plant communities in Italian mountain areas. For this purpose, we followed the space for time substitution approach (Pickett 1989) in different sites, characterised by a shared history of land-use, we identified three different stages of a vegetation succession following a pastoral land-use abandonment. Moreover, to examine the effect of geographical variation, we considered a biogeographic gradient, i.e. Mediterranean, Continental, and Alpine (EEA 2016). The specific objectives of the research were: (i) to analyse vascular plant and ant community patterns in response to pastoral land-use abandonment; (ii) to evaluate if vascular plant functional groups, ant traits, and ant functional groups were reliable indicators of the main changes of community composition, regardless of the biogeographic region, and (iii) to verify a possible overall cross-taxon congruence between ant and vascular plant communities, at the level of species and functional composition, and investigating whether ants can be used as surrogates of biodiversity of the studied taxa.
Materials and methods
Study sites and experimental design
The research was carried out in Italy at four sites located along a latitudinal/altitudinal gradient, in three biogeographic regions (Fig. 1; Table 1). On a physiognomic basis, we identified three dynamic stages of a chronosequence at each site (Table 1), representing the plant succession from pastoral use to abandonment. Stage 1 was represented by secondary grasslands currently grazed (grasslands dominated by grasses and other herbaceous species, e.g. forbs, legumes, etc.). Shrub-encroached grasslands ungrazed for 20–30 years (grasslands dominated by shrub and fern species, with a sporadic presence of trees. The herbaceous layer was mainly dominated by grasses and other herbaceous species, e.g. forbs, legumes, etc.) were the dominating vegetation in stage 2. Stage 3 was represented by woodlands, abandoned for at least 50 years (woodlands with a close canopy cover). The overstorey layer was dominated by tree species, whereas the understorey by shrubs or ferns. The understorey herbaceous layer had a low cover.
A number of 3 to 9 sampling units (hereafter ‘plots’) were established for each stage of each site for sampling vascular plants and ants, for 60 plots (Table 1). In Val Vogna, being located above the treeline, only stage 1 and stage 2 were represented.
Surveys on vascular plants (hereafter 'plants') were performed from May to July 2019, at the peak blooming period, to identify all species more easily. Along with four transects corresponding to four perpendicular radii of each plot, we applied the vertical point-quadrat method (Daget and Poissonet 1971) to assess the frequency of occurrence of each plant species. Plot radius varied from 3 to 15 m, depending on the complexity of vegetation mosaic (Braun-Blanquet 1932). Plot size does not often influence patterns of vegetation composition (Otýpková and Chytrý 2006), and the variable sampling effort is considered suitable for analyses of changes that do not involve a comparison of alpha diversity (Chytrý and Otýpková 2003; Di Pietro et al. 2021). At regular intervals (12.5 cm to 50 cm, depending on the radius length), the plant species touching a vertical steel needle were identified and recorded (Perotti et al. 2018). Since the less frequent species are likely missed using this method, a complete list of plant species found within each circular plot was recorded. A schematic representation of the survey methodology is available in Additional file 1. Finally, in a 20 m radius buffer centred on the plot, the tree species percentage cover was detected from a photointerpretation of the most recent orthophotographs available from the Sardinian and Piedmont Region public repositories (Regione Piemonte 2018; Sardegna 2019). Nomenclature follows Pignatti et al. (2017).
The frequency of occurrence of each plant species recorded along the transects of each plot was converted to 100 measurements, i.e. species percentage cover (hereafter '%SC'). A %SC = 0.3 was attributed to all rare species (Pittarello et al. 2019).
Plant species were grouped into functional groups derived from their life form (Raunkiaer 1934) and forage type (Allen et al. 2011). Then, life forms following Pignatti (2005) (Fig. 2b) and forage types (Fig. 2a) were attributed to each plant species and combined to identify 22 plant functional groups (Fig. 2c). The sum of the %SC of the species belonging to each functional group was calculated afterwards for each plot.
Ant sampling was performed at the same time as the vegetation survey. Four pitfall traps (7-cm-diameter, 11.8-cm-deep polypropylene vials), sunk into the soil and partially filled with a solution of water and monopropylene glycol (10/1; v/v), were placed at the corners of a 10-m side square centred on each vegetation plot. A scheme of each ant sampling unit is available in Additional file 1. After one week, all traps were removed and the ants were identified at the laboratory. The abundance of ant workers for each group of four traps (sampling unit) was transformed in presence-absence for subsequent statistical analyses. The specimens were identified to species level (see Additional file 1: S3 for the list of dichotomous keys available in the literature and used in this work), and the nomenclature follows AntWiki (2021). All the ant species were scored for ten traits (Table 2 and Additional file 1: S4), to characterise better the different dimensions of their functional niche (Arnan et al. 2014). The traits were obtained from databases present in the available literature (Seifert 2007; Arnan et al. 2012, 2014, 2017; Retana et al. 2015; Scupola 2018; Boet et al. 2020) and personal observations (see Table 2 and Additional file 1 for full details). In addition to the trait-based approach, we followed the functional group approach to reduce the complexity of ant taxonomy further. Ants were then classified following a global model of ant community dynamics, based on functional groups in relation to environmental stress (factors limiting ant productivity) and disturbance (factors removing ant biomass), proposed by Andersen (1995) for Australian ant communities: Subordinate Camponotini (Cm), Hot-climate Specialists (HCS), Cold-climate Specialists (CCS), Cryptic Species (Cr), Opportunists (Op), and Generalised Myrmicinae (GM). Such a functional group scheme could be adapted to European ant assemblages and provide a good understanding of functional community structure and responses to land-use changes at a biogeographical scale.
For each plot, the number of species belonging to each qualitative trait and functional group and the average values of quantitative traits were computed.
Plants and ants as indicators
Non-metric multidimensional scaling (NMDS) was performed to examine the variation in the plant and ant community composition at the species level, across sites and chronosequence stage. Bray–Curtis measure was used to obtain all similarity matrices of plant %SC (Wildi 2010). Jaccard similarity index was used for ant presence absence data, as it is most suitable for the similarity calculation based on sparse binary data (Sanders et al. 2003).
Three NMDS analyses on plant and ant functional groups and ant traits were performed to test the hypothesis that community composition (and thus the placement of points on the ordination) is lacking in systematic structure across all study areas. Bray–Curtis similarity matrices were used for ordination analyses of all functional group data. As ant trait data derived from qualitative and quantitative traits, the dissimilarity matrix was computed with a Gower distance on standardised values (Pavoine et al. 2009). Distance matrices and NMDS were computed using 'vegdist' and 'metaMDS' functions, respectively, in Vegan R-package (Oksanen et al. 2019) of R ver. 3.6.0 (www.r-project.org).
Three Permutational Multivariate Analyses of Variance (PERMANOVA), each followed by multiple comparisons, were carried out based on 999 permutations using the 'adonis2' function in the Vegan R-package. PERMANOVAs were performed to test for differences among the three stages. Distance matrices and data transformation used in PERMANOVA were the same adopted in the NMDS of each functional group or trait. In addition, multiple comparisons with Bonferroni p-values correction were carried out with the 'pairwise.perm.manova' function of RVAideMemoire R-package (Hervè 2020).
Three Mantel tests with 999 permutations and Pearson correlation analyses were carried out on dissimilarity matrices to ascertain the multivariate correlation between ants and plants in terms of composition and functionality. The first Mantel test aimed at explaining the dependence of ants on vascular plant species. Jaccard distance and Bray–Curtis distance were used to compute the ant and vascular plants dissimilarity matrices, respectively. The second aimed at finding the relationship between ant traits and plant functional groups. As ant data derived from both qualitative and quantitative traits, the dissimilarity matrix was computed with a Gower distance on standardised values (i.e. the sum within each survey sum up to 1). The dissimilarity matrix for plant functional groups was calculated with Bray–Curtis distance. The third was carried out to test the dependence of ant functional groups on plant functional groups. Both dissimilarity matrices were computed with Bray–Curtis distance. Mantel tests were carried out using the 'mantel' function in Vegan R-package (Oksanen et al. 2019).
Plants and ants as indicators
Across the four study areas, 417 plant species (Additional file 1: S5) and 50 ant species (Additional file 1: S4) were found. NMDS provided a good ordination of plant and ant assemblages at the species level, as confirmed by low-stress values (0.14 and 0.10, respectively) (Fig. 3).
The four study sites were ordered and well separated along a biogeographic gradient (i.e. MDS1 axis), from the Mediterranean to Alpine. The three stages were well separated within the convex hull characterising each study site (i.e. along the MDS2 axis) from a taxonomic point of view.
The NMDS of plant functional groups proficiently discriminated amongst the three stages, and we also achieved consistent results when considering ant traits and functional groups (stressplot: 0.15, 0.21, and 0.18, respectively) (Fig. 4).
Furthermore, the PERMANOVA results confirmed the NMDS: the pairwise comparisons indicated significant differences amongst the three stages for both taxonomic groups, except for ant functional groups between stage 1 and stage 2 (p-value = 0.08) (Table 3).
Hemicryptophytes (grasses, legumes, and other dicotyledons) and Therophytes (both grasses and legumes) were mainly associated with stage 1 (Fig. 4A). Hemicryptophytes (both grasses and sedges) were associated with stage 2 as well, but a clear correspondence of Nanophanerophytes (other dicotyledons) was also evident. Phanerophytes were strongly associated with stage 3.
The discrimination of the three stages by ants, both in terms of traits and functional groups, fairly reflected vegetation patterns. Indeed, the sequence of the three stages, from 1 to 3, detectable through the convex hull centroids in the NMDS of vegetation, was observable in the NMDS plots of ant traits and functional groups as well (Fig. 4B). At the level of traits and functional groups for ants, data groupings were fairly well distinguishable within each phase, although the positions of plots in the multivariate space were partially overlapped. Stage 1 was characterised by seed harvester ants with a group and collective foraging strategy. Conversely, the woodlands of stage 3 harboured above all species with an individual foraging strategy. NMDS did not separate stages 1 and 3 from stage 2, which was characterised by an intermediate combination of ant traits. In terms of ant functional groups, the NMDS results showed a distribution of all sampling units similar to that obtained from the trait data (Fig. 4C). Cold-climate Specialists increased from stage 1 to 3, whereas Hot-climate Specialists followed the opposite trend. Stage 2 differed from stage 1 mainly for a higher contribution of Cold Climate Specialists and less importance of Hot Climate Specialists. Stage 3 was characterised by the absence of Hot Climate Specialists and the higher relative contribution of Cryptic Species.
In plants and ants, the patterns of species composition revealed by NMDS were characterised by a gradual change along the ordination axes from secondary grasslands over woodlands, i.e. across the chronosequence stages following the pastoral land-use abandonment. The comparison of community similarity matrices performed using the overall dataset showed a strong and significant relationship between the two taxa at the species level (Mantel r = 0.75, p < 0.001). However, such a relationship between community structures was moderate at the functional level. Mantel test performed on similarity matrices obtained from ant traits and plant functional groups showed moderate relationships between the two taxa (Mantel r = 0.36, p < 0.001). A direct correspondence between ants and vascular plants was also confirmed at the functional group level, although the cross-taxon congruence was weaker (Mantel r = 0.25, p < 0.001).
Plants and ants as indicators
At the species level, plant and ant communities showed a consistent pattern related to the biogeographic region. Stage 1 was the expression of secondary grasslands with a dominance of herbaceous species maintained by grazing livestock (Kahmen et al. 2002; Diaz et al. 2007). Secondary grasslands showed a strong differentiation, particularly at the opposite biogeographic positions, with different plant species: Dasypyrum villosum and Avena barbata in the Mediterranean site and Festuca rubra subsp. commutata and Poa alpina in the Alpine area above the treeline. At the two intermediate sites, species of secondary grasslands were mainly from the Festuco-Brometea class (Ciaschetti et al. 2020). The dynamic processes of grassland encroachment (stage 2) led to different communities depending on the biogeographical region of location (Bagella and Caria 2011). We found Pteridium aquilinum subsp. aquilinum in the Mediterranean, Rubus ulmifolius in the Continental, Prunus spinosa in the Alpine areas below the tree line and Rhododendron ferrugineum above the tree line. Woodlands vegetation (stage 3) was characterised by a floristic composition gradually more mesophilous (e.g. from Quercus ichnusae to Fagus sylvatica).
The ant communities in grasslands (stage 1) were typical of open habitats (Seifert 2007; Lentini and Verdinelli 2012; Scupola 2018; Verdinelli et al. 2017). In the Mediterranean region Tapinoma simrothi, Messor ibericus, Aphaenogaster spinosa commonly occurred in open grasslands, whereas in the Continental region Pheidole pallidula and other omnivorous species were frequent. In the Alpine regions, the community was represented by species seeking food both in the herbaceous and arboreal layers (Seifert 2007) and that were also well adapted to mountain area grasslands (Scupola 2018), with Lasius alienus and Formica fusca below the tree line and the boreal Formica lemani, Myrmica sulcinodis and Manica rubida above the tree line. At the stage 2 within the Alpine region, temporary parasitic and/or dulotic species (e.g. F. exsecta and F. lugubris) were sampled. Species commonly found under the trees or in shaded habitats of stage 3, such as Aphaenogaster ichnusa and Myrmecina graminicola, were shared with stage 2 as probably favoured by the abundance of shrubs and tall grasses (Verdinelli et al. 2017; AntWiki 2021).
In the Mediterranean area, annual legumes and grasses were dominant, as annual species are more tolerant to dry environments due to their fast growth rates and early and prolific seed set (Grime 1974). In Continental and Alpine areas Hemicryptophytes (mainly legumes, grasses, and sedges) are more abundant (Körner 2003) and grazing favoured their persistence (Illa et al. 2006). Stage 2 was defined by plant species typical of an early abandonment process, including tall and/or woody species (Pykälä 2005), which characterise the natural dynamic processes to dwarf shrub- and tree-dominated communities (Kesting et al. 2009). In the Mediterranean site we found tall Pteridophyte Hemicryptophyte species. Instead, in Continental and Alpine sites we found Nanophanerophyte dicotyledons. Stage 3 was entirely separated from stage 1 and stage 2, as the dominant species were Phanerophytes. Indeed, trees were sporadic in stages 1 and 2 as Phanerophytes are generally less frequent in grazing systems (Illa et al. 2006; Komac et al. 2013).
Also, ant traits and functional groups were good indicators of the time stages. Changes in ground cover may be important in determining the number of species able to use disturbed environments (Andersen 2019; Carvalho et al. 2020). These changes can have wide-ranging ecosystem effects given the role of ants as ecosystem engineers and their impact on many ecosystem functions (Folgarait 1998). The direct trophic relationship between ants and plants was clear in stage 1. Indeed, the seed diet, the collective foraging strategy, and other traits are associated with open habitats (Retana et al. 2015). Hot-climate Specialists include thermophilic taxa and specialist seed harvesters (Morton and Davidson 1988), whose occurrence was likely due to the combination of soil type and vegetation cover, which are the main drivers of ant assemblages (Bestelmeyer and Wiens 2001; Ríos-Casanova et al. 2006). Shrub-encroached grasslands offer moderate soil surface temperatures and substrate for potential prey and liquid food (i.e. extrafloral nectar and/or honeydew). These conditions favoured the Cold-climate Specialists and Cryptic Species (small to minute species, predominantly myrmicines and ponerines, that nest and forage primarily within soil, litter, and rotting logs), as a result of the decreasing insolation and increasing moisture (Arnan et al. 2012). The similarities between ant and plant communities' successional pathways are also remarkable in stage 3. Cold-climate Specialists and Cryptic Species increased their relative frequency in woodlands, where the thermal influence of canopy cover is stronger, and the insolation at ground level is minimal (Andersen 1995, 1997; Zina et al. 2021).
Even though the plant and ant community composition differed among sites, the two groups responded to pastoral land-use abandonment in a similar way, both at taxonomic and functional levels. This finding was coherent with the general principle that cross-taxon congruence is strengthened at a large scale (Westgate et al. 2014). Although climatic or biogeographic gradients generally favour a conformity of species numbers at large spatial scales, the strong congruence of community similarities between ants and plants was an interesting outcome of the present study. The impact of grazing abandonment on biodiversity has been investigated extensively in different ecosystems worldwide, but the effects on ants and plants have been usually treated separately. In Mediterranean grasslands, past evidence suggests that grazing abandonment harms biodiversity, and this view is mainly based on studies focused on vegetation (Peco et al. 2005, 2006). Mountain grazing abandonment in alpine areas can promote the establishment of vegetation characterised by few species and negatively affect biodiversity (Cislaghi et al. 2019). However, grazing involves different trophic levels (Filazzola et al. 2020), organisms and ecosystems (Maghniaa et al. 2017; Mannu et al. 2018, 2020), and the impact of grazing on biodiversity can be negative or positive. For example, grazing decreased the ant species richness in American and Australian rangelands (Woinarski et al. 2002; Boulton et al. 2005). In temperate rangelands, livestock grazing maintained grassland structure by suppressing woody encroachment and increased ant species richness (Schmidt et al. 2012). In Mediterranean grasslands, grazing abandonment increased the functional and species ant diversity (Azcárate and Peco 2012). In our study, we observed that ant and vascular plant community composition varied in a similar way in response to pastoral land-use abandonment. According to Radnan and Eldridge (2017), grazing in secondary grasslands favoured Hot Climate Specialists. However, the same authors showed that grazing had stronger effects on ants than any increase in shrub cover, as ant and plant communities can respond to pastoral land abandonment in an idiosyncratic way (Német et al. 2016). Although plants are considered poor surrogates of invertebrate biodiversity (Andersen and Majer 2004), our result shows that ants and plants can provide a congruent representation of biodiversity. This finding confirms what has been reported in recent studies involving plant communities, ant communities and other arthropods (Ford et al. 2012; Corcos et al. 2021; Zina et al. 2021). Ant communities might be classified according to structural attributes that parallel those adopted in vegetation science (Andersen 1995), and used as surrogates in biodiversity studies. However, the moderate level of cross-taxon congruence we found at the level of functional groups suggests the need for further investigations to develop new and comprehensive ant metrics suitable for pastoral mountain areas.
Our attempt to compare the community pattern of plants and ants in mountain areas of different biogeographic regions in response to the abandonment of pastoral land use provided encouraging results. Pastoral land use is one of the main driving forces that caused a change of species composition from secondary grasslands to woodlands in the two taxa: the abandonment promoted the woody species colonisation and the conversion of the original grassland ant community. Cross-taxa relationships confirmed this result. Although we did not test congruency within each chronosequence stage or biogeographic region, the analyses of the entire dataset showed that ant communities mirrored the plant species and functional groups at taxonomic and functional levels. The functional groups explain a good proportion of trait variation among species. They allow going beyond the taxonomic barriers and can be a valuable tool in cost-effective ecological surveys. Although the cross-taxon congruence was moderate at the functional group level, the results underline the reliability of ants as bioindicators of land changes and ecosystem functioning. This result is important because it could greatly facilitate biodiversity monitoring and conservation. Due to their role in food webs and ecosystem functioning, ants can provide a more fine-grained and dynamic view of ecosystems than plants. An in-depth study of the main drivers of ant community composition would certainly be a notable improvement in understanding how their structure and function vary in response to land use. However, simplified ant monitoring systems should be developed to meet the needs of land managers. Finally, our findings indicate that the functional group approach and the reduction of the taxonomic complexity could contribute to developing predictive models to detect early environmental changes and biodiversity loss in mountain habitats.
Data availability statement
The data supporting this study's findings are available from the corresponding author upon reasonable request.
Species percentage cover
Other Dicotyledons Chamaephyte
Other Dicotyledons Geophyte
Other Dicotyledons Hemicryptophyte
Other Dicotyledons Nanophanerophyte
Other Dicotyledons Phanerophyte
Other Dicotyledons Therophyte
Other Monocotyledons Geophyte
Other Monocotyledons Hemicryptophyte
Other Monocotyledons Therophyte
Not strictly Diurnal
Strictly group foraging strategy
Strictly collective foraging strategy
Both group and collective foraging strategy
Strictly individual foraging strategy
Both monogyny and polygyny
Monodomous and polydomous
Both dependent and independent
Mainly liquid feeder
Cold climate specialists
Hot climate specialists
Allen VG, Batello C, Berretta EJ, Hodgson J, Kothmann M, Li X, McIvor J, Milne J, Morris C, Peeters A, Sanderson M (2011) An international terminology for grazing lands and grazing animals. Grass Forage Sci 66:2–28. https://doi.org/10.1111/j.1365-2494.2010.00780.x
Andersen AN (1995) A classification of Australian ant communities, based on functional groups which parallel plant life-forms in relation to stress and disturbance. J Biogeogr 22:15–29. https://doi.org/10.2307/2846070
Andersen AN (1997) Functional groups and patterns of organization in North American ant communities, A comparison with Australia. J Biogeogr 24:433–460. https://doi.org/10.1111/1365-2656.12907
Andersen AN (2019) Responses of ant communities to disturbance: Five principles for understanding the disturbance dynamics of a globally dominant faunal group. J Anim Ecol 88:350–362. https://doi.org/10.1111/1365-2656.12907
Andersen AN, Majer JD (2004) Ants show the way Down Under: invertebrates as bioindicators in land management. Front Ecol Environ 2:291–298. https://doi.org/10.1890/1540-9295(2004)002[0292:ASTWDU]2.0.CO;2
AntWiki (2021) AntWiki: The Ants. https://www.antwiki.org. Accessed 28 Jul 2021
Arnan X, Cerdá X, Retana J (2012) Distinctive life traits and distribution along environmental gradients of dominant and subordinate Mediterranean ant species. Oecologia 170:489–500. https://doi.org/10.1007/s00442-012-2315-y
Arnan X, Cerdá X, Retana J (2014) Ant functional responses along environmental gradients. J Anim Ecol 83:1398–1408. https://doi.org/10.1111/1365-2656.12227
Arnan X, Cerdá X, Retana J (2017) Relationships among taxonomic, functional, and phylogenetic ant diversity across the biogeographic regions of Europe. Ecography 40:448–457. https://doi.org/10.1111/ecog.01938
Azcarate FM, Peco B (2012) Abandonment of grazing in a Mediterranean grassland area, consequences for ant assemblages. Insect Conserv Divers 5:279–288. https://doi.org/10.1111/j.1752-4598.2011.00165.x
Bagella S, Caria MC (2011) Vegetation series, a tool for the assessment of grassland ecosystem services in Mediterranean large-scale grazing systems. Fitosociologia 48:47–54
Bagella S (2014) Does cross-taxon analysis show similarity in diversity patterns between vascular plants and bryophytes? Some answers from a literature review. C R Biol 337:276–282. https://doi.org/10.1016/j.crvi.2014.01.003
Bagella S, Caria MC, Seddaiu G, Leites L, Roggero PP (2020) Patchy landscapes support more plant diversity and ecosystem services than wood grasslands in Mediterranean silvopastoral agroforestry systems. Agric Syst 185:102945. https://doi.org/10.1016/j.agsy.2020.102945
Bagella S, Filigheddu R, Caria MC, Girlanda M, Roggero PP (2014) Contrasting land uses in Mediterranean agro-silvo-pastoral systems generated patchy diversity patterns of vascular plants and below-ground microorganisms. C R Biol 337:717–724. https://doi.org/10.1016/j.crvi.2014.09.005
Bagella S, Satta A, Floris I, Caria MC, Rossetti I, Podani J (2013) Effects of plant community composition and flowering phenology on honeybee foraging in Mediterranean sylvo-pastoral systems. Appl Veg Sci 16:689–697. https://doi.org/10.1111/avsc.12023
Bestelmeyer BT, Wiens K (2001) Ant biodiversity in semiarid landscape mosaics, the consequences of grazing vs natural heterogeneity. Ecol Appl 11:1123–1140. https://doi.org/10.1890/1051-0761(2001)011[1123:ABISLM]2.0.CO;2
Boet O, Arnan X, Retana J (2020) The role of environmental vs biotic filtering in the structure of European ant communities, A matter of trait type and spatial scale. PLoS ONE 15:e0228625. https://doi.org/10.1371/journal.pone.0228625
Boulton AM, Davies KF, Ward PS (2005) Species richness, abundance, and composition of ground-dwelling ants in Northern California grasslands: role of plants, soil, and grazing. Environ Entomol 34:96–104. https://doi.org/10.1603/0046-225X-34.1.96
Braun-Blanquet J (1932) Plant sociology: The study of plant communities. McGraw-Hill, New York
Burrascano S, Sabatini FM, Blasi C (2011) Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning. Plant Ecol 212:829–841. https://doi.org/10.1007/s11258-010-9866-y
Carvalho RL, Andersen AN, Anjos DV, Pacheco R, Chagas L, Vasconcelos HL (2020) Understanding what bioindicators are actually indicating, Linking disturbance responses to ecological traits of dung beetles and ants. Ecol Indic 108:105764. https://doi.org/10.1016/j.ecolind.2019.105764
Chytrý M, Otýpková Z (2003) Plot sizes used for phytosociological sampling of European vegetation. J Veg Sci 14:563–570. https://doi.org/10.1111/j.1654-1103.2003.tb02183.x
Chelli S, Marignani M, Barni E, Petraglia A, Puglielli G, Wellstein C, Acosta A, Bolpagni R, Bragazza L, Campetella G, Chiarucci A, Conti L, Nascimbene J, Orsenigo S, Pierce S, Ricotta C, Tardella F, Abeli T, Aronne G, Bacaro G, Bagella S, Benesperi R, Bernareggi G, Bonanomi G, Bricca A, Brusa G, Buffa G, Burrascano S, Caccianiga M, Calabrese V, Canullo R, Carbognani M, Carboni M, Carranza M, Catorci A, Ciccarelli D, Citterio S, Cutini M, Dalle Fratte M, De Micco V, Del Vecchio S, Di Martino L, Di Musciano M, Fantinato E, Filigheddu R, Frattaroli A, Gentili R, Gerdol R, Giarrizzo E, Giordani P, Gratani L, Incerti G, Lussu M, Mazzoleni S, Mondoni A, Montagnani C, Montagnoli A, Paura B, Petruzzellis F, Pisanu S, Rossi G, Sgarbi E, Simonetti E, Siniscalco C, Slaviero A, Stanisci A, Stinca A, Tomaselli M, Cerabolini B (2019) Plant–environment interactions through a functional traits perspective: a review of Italian studies. Plant Biosyst 153:853–869. https://doi.org/10.1080/11263504.2018.1559250
Ciaschetti G, Di Musciano M, Pirone G, Di Cecco V, Pace L, Frattaroli AR (2020) A new pioneer association of detrital substrata of the hilly and low-mountain belts in Central Apennines (Italy). Plant Sociology 57:75–84. https://doi.org/10.3897/pls2020571/08
Cislaghi A, Giupponi L, Tamburini A, Giorgi A, Bischetti GB (2019) The effects of mountain grazing abandonment on plant community, forage value and soil properties: observations and field measurements in an alpine area. Catena 181:104086. https://doi.org/10.1016/j.catena.2019.104086
Corcos D, Lami F, Nardi D, Boscutti F, Sigura M, Giannone F, Pantini P, Tagliapietra A, Busato F, Sibella R, Marini L (2021) Cross-taxon congruence between predatory arthropods and plants across Mediterranean agricultural landscapes. Ecol Indic 123:107366. https://doi.org/10.1016/j.ecolind.2021.107366
Daget P, Poissonet J (1971) A method of plant analysis of pastures. Annales Agronomiques 22:5–41. https://doi.org/10.4081/ija.2019.1480
Diaz S, Lavorel S, McIntyre S, Falczuk V, Casanoves F, Milchunas DG, Skarpe C, Rusch G, Sternberg M, Noy-Meir I, Landsberg J, Zhang W, Clark H, Campbell BD (2007) Plant trait responses to grazing - a global synthesis. Glob Chang Biol 13:313–341. https://doi.org/10.1111/j.1365-2486.2006.01288.x
Di Pietro R, Fortini P, Misano G, Terzi M (2021) Phytosociology of Atractylis cancellata and Micromeria microphylla communities in southern Italy with insights on the xerothermic steno-Mediterranean grasslands high-rank syntaxa. Plant Sociology 58:133–155. https://doi.org/10.3897/pls2021581/07
EEA - European Environmental Agency (2016) Biogeographical regions. https://www.eea.europa.eu/data-and-maps/data/biogeographical-regions-europe-3. Accessed 16 Gen 2020.
Filazzola A, Brown C, Dettlaff MA, Batbaatar A, Grenke J, Bao T, Peetoom Heida I, Cahill JF Jr (2020) The effects of livestock grazing on biodiversity are multi-trophic, a meta-analysis. Ecol Lett 23:1298–1309. https://doi.org/10.1111/ele.13527
Folgarait PJ (1998) Ant biodiversity and its relationship to ecosystem functioning, a review. Biodivers Conserv 7:1221–1244. https://doi.org/10.1023/A:1008891901953
Ford H, Garbutt A, Jones DL, Jones L (2012) Impacts of grazing abandonment on ecosystem service provision: Coastal grassland as a model system. Agric Ecosyst Environ 162:108–115. https://doi.org/10.1016/j.agee.2012.09.003
Garbarino M, Lingua E, Weisberg PJ, Bottero A, Meloni F, Motta R (2013) Land-use history and topographic gradients as driving factors of subalpine Larix decidua forests. Landsc Ecol 28:805–817. https://doi.org/10.1007/s10980-012-9792-6
Garnier E, Cortez J, Billès G, Navas ML, Roumet C, Debussche M, Laurent G, Blanchard A, Aubry D, Bellmann A (2004) Plant functional markers capture ecosystem properties during secondary succession. Ecology 85:2630–2637. https://doi.org/10.1890/03-0799
Gómez C, Casellas D, Oliveras J, Bas JM (2003) Structure of ground-foraging ant assemblages in relation to land-use change in the northwestern Mediterranean region. Biodivers Conserv 12:2135–2146. https://doi.org/10.1023/A:1024142415454
Gómez Sal A, González García A (2007) A comprehensive assessment of multifunctional agricultural land-use systems in Spain using a multi-dimensional evaluative model. Agric Ecosyst Environ 120:82–91. https://doi.org/10.1016/j.agee.2006.06.020
Grime JP (1974) Vegetation classification by reference to strategies. Nature 250:26–31. https://doi.org/10.1038/250026a0
Hervè M (2020) RVAideMemoire, Testing and Plotting Procedures for Biostatistics. R package version 09–75. https://CRANR-project.org/package=RVAideMemoire.
Heuss L, Grevé ME, Schäfer D, Busch V, Feldhaar H (2019) Direct and indirect effects of land-use intensification on ant communities in temperate grasslands. Ecol Evol 9:4013–4024. https://doi.org/10.1002/ece3.5030
Hevia V, Carmona CP, Azcárate FM, Torralba M, Alcorlo P, Ariño R, Lozano J, Castro-Cobo S, González JA (2016) Effects of land use on taxonomic and functional diversity, a cross-taxon analysis in a Mediterranean landscape. Oecologia 181:959–970. https://doi.org/10.1007/s00442-015-3512-2
Heydari M, Zeynali N, Bazgir M, Omidipour R, Kohzadian M, Sagar R, Prevosto B (2020) Rapid recovery of the vegetation diversity and soil fertility after cropland abandonment in a semiarid oak ecosystem: An approach based on plant functional groups. Ecol Eng 155:105963. https://doi.org/10.1016/j.ecoleng.2020.105963
Hoffmann BD (2010) Using ants for rangeland monitoring, global patterns in the responses of ant communities to grazing. Ecol Indic 10:105–111. https://doi.org/10.1016/j.ecolind.2009.04.016
Hölldobler B, Wilson EO (eds) (1990) The Ants. The Belknap Press of Harvard University Press, Cambridge
Illa E, Carrillo E, Ninot JM (2006) Patterns of plant traits in Pyrenean alpine vegetation. Flora 201:528–546. https://doi.org/10.1016/j.flora.2005.10.007
Jiménez-Carmona F, Heredia-Arévalo AM, Reyes-López JL (2020) Ants (Hymenoptera, Formicidae) as an indicator group of human environmental impact in the riparian forests of the Guadalquivir river (Andalusia, Spain). Ecol Indic 118:106762. https://doi.org/10.1016/j.ecolind.2020.106762
Kahmen S, Poschlod P, Schreiber KF (2002) Conservation management of calcareous grasslands. Changes in plant species composition and response of functional traits during 25 years. Biol Conserv 104:319–328. https://doi.org/10.1016/S0006-3207(01)00197-5
Kesting S, Wrage N, Isselstein J (2009) Herbage mass and nutritive value of herbage of extensively managed temperate grasslands along a gradient of shrub encroachment. Grass Forage Sci 64:246–254. https://doi.org/10.1111/j.1365-2494.2009.00690.x
Koch B, Edwards PJ, Blanckenhorn WU, Walter T, Hofer G (2015) Shrub encroachment affects the diversity of plants, butterflies, and grasshoppers on two Swiss subalpine pastures. Arct Antarct Alp Res 47:345–357. https://doi.org/10.1657/AAAR0013-093
Komac B, Kéfi S, Nuche P, Escós J, Alados CL (2013) Modeling shrub encroachment in subalpine grasslands under different environmental and management scenarios. J Environ Manage 121:160–169. https://doi.org/10.1016/j.jenvman.2013.01.038
Körner C (2003) The alpine life zone. In: Alpine Plant Life, 3rd edn. Springer, Berlin.
Laiolo P, Dondero F, Ciliento E, Rolando A (2004) Consequences of pastoral abandonment for the structure and diversity of the alpine avifauna. J Appl Ecol 41:294–304. https://doi.org/10.1111/j.0021-8901.2004.00893.x
Larrieu L, Gosselin F, Archaux F, Chevalier R, Corriol G, Dauffy-Richard E, Deconchat M, Gosselin M, Ladet S, Savoie JM, Tillon L, Bouget C (2018) Cost-efficiency of cross-taxon surrogates in temperate forests. Ecol Indic 87:56–65. https://doi.org/10.1016/j.ecolind.2017.12.044
Laureto LMO, Cianciaruso MV, Samia DSM (2015) Functional diversity: an overview of its history and applicability. Natureza and Conservação 13:112–116. https://doi.org/10.1016/j.ncon.2015.11.001
Lentini A, Verdinelli M (2012) Formicidae. In: Vacante V, Gerson U (ed) Integrated control of citrus pests in the mediterranean region. Bentham Science Publishers, Sharjah. https://doi.org/10.2174/978160805294311201010231.
MacDonald D, Crabtree JR, Wiesinger G, Dax T, Stamou N, Fleury P, Gutierrez Lazpita J, Gibon A (2000) Agricultural abandonment in mountain areas of Europe: Environmental consequences and policy response. J Environ Manage 59:47–69. https://doi.org/10.1006/jema.1999.0335
Maghniaa FZ, Sanguin H, Abbas Y, Verdinelli M, Kerdouh B, El Ghachtouli N, Lancellotti E, Bakkali Yakhlef SE, Duponnois R (2017) Impact du mode de gestion de la subéraie de la Maâmora (Maroc) sur la diversité des champignons ectomycorhiziens associés à Quercus suber. C R Biol 340:298–305. https://doi.org/10.1016/j.crvi.2017.04.001
Mannu R, Pilia O, Fadda ML, Verdinelli M (2018) Variability of beetle assemblages in Mediterranean cork oak woodlands: does the higher taxa approach reliably characterize a specific response to grazing? Biodivers Conserv 27:3599–3619. https://doi.org/10.1007/s10531-018-1616-9
Mannu R, Pantini P, Sassu A, Verdinelli M (2020) A multidiversity approach to investigate the impact of mining exploitation on spider diversity in the abandoned mine district of Montevecchio-Ingurtosu (Sardinia, Italy). Environ Sci Pollut Res Int 27:32615–32627. https://doi.org/10.1007/s11356-020-09553-y
Morton SR, Davidson DW (1988) Comparative structure of harvester ant communities in arid Australia and North America. Ecol Monogr 58:19–38. https://doi.org/10.2307/1942632
Német E, Ruprecht E, Gallé R, Markó B (2016) Abandonment of crop lands leads to different recovery patterns for ant and plant communities in Eastern Europe. Community Ecol 17:79–87. https://doi.org/10.1556/168.2016.17.1.10
Oksanen J, Guillaume Blanchet F, Friendly M, Kindt R, Legendre P, McGlinn DR, Minchin PB, O'Hara RL, Simpson G, Solymos P, Henry H, Stevens M, Szoecs E, Wagner H (2019) vegan: Community Ecology Package. R package version 25–6
Otýpková Z, Chytrý M (2006) Effects of plot size on the ordination of vegetation samples. J Veg Sci 17:465–472. https://doi.org/10.1111/j.1654-1103.2006.tb02467.x
Pavoine S, Vallet J, Dufour A-B, Gachet S, Daniel H (2009) On the challenge of treating various types of variables: application for improving the measurement of functional diversity. Oikos 118:391–402. https://doi.org/10.1111/j.1600-0706.2008.16668.x
Peco B, de Pablos I, Traba J, Levassor C (2005) The effect of grazing abandonment on species composition and functional traits: the case of dehesa grasslands. Basic Appl Ecol 6:175–183. https://doi.org/10.1016/j.baae.2005.01.002
Peco B, Sanchez AM, Azcárate FM (2006) Abandonment in grazing systems: consequences for vegetation and soil. Agric Ecosyst Environ 113:284–294. https://doi.org/10.1016/j.agee.2005.09.017
Perotti E, Probo M, Pittarello M, Lonati M, Lombardi G (2018) A 5-year rotational grazing changes the botanical composition of sub-alpine and alpine grasslands. Appl Veg Sci 21:647–657. https://doi.org/10.1111/avsc.12389
Pharo EJ, Beattie AJ, Pressey RL (2000) Effectiveness of using vascular plants to select reserves for bryophytes and lichens. Biol Conserv 96:371–378. https://doi.org/10.1016/S0006-3207(00)00080-X
Pickett STA (1989) Space for time substitutions as an alternative to long-term studies. In: Likens GE (ed) Long-Term studies in ecology. Springer, New York
Pignatti S (2005) Indicator Values of the Vascular Plants of the Flora of Italy (Valori di bioindicazione delle piante vascolari della Flora d'Italia). Dipartimento di Botanica ed Ecologia dell'Università di Camerino, Camerino.
Pignatti S, La Rosa M, Guarino R (2017) Flora of Italy (Flora d’Italia), 2nd edn. Edagricole, Bologna
Pittarello M, Probo M, Lonati M, Lombardi G (2016) Restoration of sub-alpine shrub-encroached grasslands through pastoral practices: effects on vegetation structure and botanical composition. Appl Veg Sci 19:381–390. https://doi.org/10.1111/avsc.12222
Pittarello M, Probo M, Perotti E, Lonati M, Lombardi G, Ravetto Enri S (2019) Grazing Management Plans improve pasture selection by cattle and forage quality in sub-alpine and alpine grasslands. J Mt Sci 16:2126–2135. https://doi.org/10.1007/s11629-019-5522-8
Pittarello M, Lonati M, Ravetto Enri S, Lombardi G (2020) Environmental factors and management intensity affect in different ways plant diversity and pastoral value of alpine pastures. Ecol Indic 115:106429. https://doi.org/10.1016/j.ecolind.2020.106429
Prévosto B, Kuiters L, Bernhardt-Römermann M, Dölle M, Schmidt W, Hoffmann M, Van Uytvanck J, Bohner A, Kreiner D, Stadler J, Klotz S (2011) Impacts of land abandonment on vegetation: successional pathways in European habitats. Folia Geobot 46:303–325. https://doi.org/10.1007/s12224-010-9096-z
Pykälä J (2005) Cattle grazing increases plant species richness of most species trait groups in mesic semi-natural grasslands. Plant Ecol 175:217–226. https://doi.org/10.1007/s11258-005-0015-y
Radnan GN, Eldridge DJ (2017) Ants respond more strongly to grazing than changes in shrub cover. Land Degrad Dev 29:907–915. https://doi.org/10.1002/ldr.2911
Raunkiaer C (1934) The life forms of plants and statistical plant geography. Oxford University Press, London
Regione Piemonte (2018). AGEA 2018 - Ortofoto RGB. Geoservizi WMS e WMTS. https://www.geoportale.piemonte.it/geonetwork/srv/ita/catalog.search#/metadata/r_piemon:98fe6c87-2721-4193-a35a-5af883badce7
Regione Sardegna (2019). AGEA 2019 - Ortofoto RGB. https://www.sardegnageoportale.it/index.php?xsl=2419&s=40&v=9&c=14488&na=1&n=10&t=1&esp=1&tb=14401
Retana J, Arnan X, Cerdá X (2015) A multidimensional functional trait analysis of resource exploitation in European ants. Ecology 96:2781–2793. https://doi.org/10.1890/14-2326.1
Ríos-Casanova L, Valiente-Banuet A, Rico-Gray V (2006) Ant diversity and its relationship with vegetation and soil factors in an alluvial fan of the Tehuacán Valley, Mexico. Acta Oecol 29:316–323. https://doi.org/10.1016/j.actao.2005.12.001
Sanders NJ, Moss J, Wagner D (2003) Patterns of ant species richness along elevational gradients in an arid ecosystem. Glob Ecol Biogeogr 12:93–102. https://doi.org/10.1046/j.1466-822X.2003.00324.x
Satta A, Verdinelli M, Ruiu L, Buffa F, Salis S, Sassu A, Floris I (2012) Combination of beehive matrices analysis and ant biodiversity to study heavy metal pollution impact in a post-mining area (Sardinia, Italy). Environ Sci Pollut Res Int 19:3977–3988. https://doi.org/10.1007/s11356-012-0921-1
Schmidt AC, Fraser LH, Carlyle CN, Bassett ERL (2012) Does cattle grazing affect ant abundance and diversity in temperate grasslands? Rangel Ecol Manag 65:292–298. https://doi.org/10.2111/REM-D-11-00100.1
Scupola A (2018) The ants of Veneto (Le formiche del Veneto). World Biodiversity Association, Verona.
Segar ST, Pereira RA, Compton SG, Cook JM (2013) Convergent structure of multitrophic communities over three continents. Ecol Lett 16:1436–1445. https://doi.org/10.1111/ele.12183
Seifert B (2007) The Ants of Central and Northern Europe (Die Ameisen Mittel- und Nordeuropas), Lutra , Görlitz.
Smith TM, Huston MA (1989) A theory of spatial and temporal dynamics of plant communities. Vegetatio 83:49–69. https://doi.org/10.1007/BF00031680
Tasser E, Tappeiner U (2002) Impact of land use changes on mountain vegetation. Appl Veg Sci 5:173–184. https://doi.org/10.1111/j.1654-109X.2002.tb00547.x
Travers E, Pitz WT, Fichtner A, Matthies D, Härdtle W (2020) The role of semi-open habitats as dispersal corridors for plant species of woodlands and open habitats. Appl Veg Sci 24:e12526. https://doi.org/10.1111/avsc.12526
Verdinelli M, Yakhlef SEB, Cossu CS, Pilia O, Mannu R (2017) Variability of ant community composition in cork oak woodlands across the Mediterranean region: implications for forest management. Forest 10:707–714. https://doi.org/10.3832/ifor2321-010
Wan Hussin WMR, Cooper KM, Barrio Froján CRS, Defew EC, Paterson DM (2012) Impacts of physical disturbance on the recovery of a macrofaunal community: a comparative analysis using traditional and novel approaches. Ecol Indic 12:37–45. https://doi.org/10.1016/j.ecolind.2011.03.016
Westgate MJ, Barton PS, Lane PW, Lindenmayer DB (2014) Global meta-analysis reveals low consistency of biodiversity congruence relationships. Nat Commun 5:3899. https://doi.org/10.1038/ncomms4899
Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ (2002) Plant ecological strategies: some leading dimensions of variation between species. Annu Rev Ecol Syst 33:125–159. https://doi.org/10.1146/annurev.ecolsys.33.010802.150452
Wildi O (2010) Data Analysis in Vegetation Ecology. Wiley-Blackwell, New York. https://doi.org/10.1002/9780470664971
Wilson GB (1999) Guilds, functional types and ecological groups. Oikos 86:507–522. https://doi.org/10.1034/j.1600-0706.2003.12152.x
Woinarski JCZ, Andersen AN, Churchill TB, Ash AJ (2002) Response of ant and terrestrial spider assemblages to pastoral and military land use, and to landscape position, in a tropical savanna woodland in northern Australia. Austral Ecol 27:324–333. https://doi.org/10.1046/j.1442-9993.2002.01183.x
Yapp G, Walker J, Thackway R (2010) Linking vegetation type and condition to ecosystem goods and services. Ecol Complex 7:292–301. https://doi.org/10.1016/j.ecocom.2010.04.008
Zina V, Ordeix M, Franco JC, Ferreira MT, Fernandes MR (2021) Ants as bioindicators of riparian ecological health in Catalonian rivers. Forests 12:625. https://doi.org/10.3390/f12050625
The authors are grateful to Giovanni Reggiani (Società Agricola Montana), Donatella Acella (Allevamento Morag Fold 2009) and AGRIS Sardegna for hosting the experiments in their farms and providing the necessary technical assistance. Special thanks to Davide Barberis, Marco Cuboni, Alberto Tanda and Giovanni Rivieccio for their technical support to the fieldwork.
Research was carried out within the "iGRAL—Innovative beef cattle Grazing systems for the Restoration of Abandoned Lands in the Alpine and Mediterranean mountains" project (Ager—Agroalimentare e ricerca—research grant n. 2017–1164, Principal Investigator Giampiero Lombardi).
Ethics approval and consent to participate
Consent for publication
The authors declare no conflict of interests.
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Appendix S1. Schematic representation of surveys on vascular plants through the vertical point-quadrat method along four transects corresponding to four perpendicular radii of a circular plot. Appendix S2. Scheme of ant sampling design: four pitfall traps, 10 m distant from each other, were sunk into the soil and centred on each vegetation plot. Appendix S3. Ant dichotomous key reference list. Appendix S4. Ant species collected during the survey per each stage. Ant functional groups (FG): Subordinate Camponotini (Cm), Hot-climate Specialists (HCS), Cold-climate Specialists (CCS), Cryptic Species (Cr), Opportunists (Op), Specialised Predators (SP), and Generalised Myrmicinae (GM). Ant traits: Worker size (WS), Worker polymorphism (WP), Colony size (CS), Diet (D), Behavioural dominance (BD), Diurnality (Di), Foraging strategy (FS), Number of nests per colony (nN), Number of queens (nQ), Colony foundation (CF). For ant traits in species for which no data are reported in the literature, ten workers per species were randomly selected and measured for determining the WS; the mean thereof was used as a value for each species. Appendix S5. List of plant species found across the study sites associated with their Raunkiaer plant life form (Pignatti 2005), Forage type (Allen et al. 2011), Plant functional group, and the occurrence in the stages of the chronosequence. The latter is the combination of the life form and the forage type. Plant species nomenclature follows Pignatti et al. (2017).
About this article
Cite this article
Verdinelli, M., Pittarello, M., Caria, M.C. et al. Congruent responses of vascular plant and ant communities to pastoral land-use abandonment in mountain areas throughout different biogeographic regions. Ecol Process 11, 35 (2022). https://doi.org/10.1186/s13717-022-00379-9
- Biodiversity conservation
- Cross-taxon congruence
- Functional groups
- Land abandonment
- Mountain pastures