Agroforest Syst DOI 10.1007/s10457-010-9359-y

Fungal fruitbodies and soil macrofauna as indicators of land use practices on soil biodiversity in Montado

Anabela Marisa Azul • Sara Margarida Mendes • Jose´ Paulo Sousa • Helena Freitas

Received: 22 January 2010 / Accepted: 28 October 2010 Ó Springer Science+Business Media B.V. 2010

Abstract The impacts of land use on soil biodiver- cutting practices followed by soil tillage (M), in sity are still poorly understood, although soil fungi and comparison with cutting practices with no soil tillage macrofauna are recognized to provide benefits to (Cu) and the control (C). The ECMF Laccaria laccata ecosystems. Here, we tested whether land use practices and subtomentosus exhibited a close used to control shrub density influences the fruiting relation with C and Cu plots while the saprobes, e.g., macromycetes (ectomycorrhizal-forming fungi— Entoloma conferendum, were associated to Ca and M ECMF—and saprobes) and soil macrofauna diversity plots. Most associated to Cu plots were present and abundance in Montado ecosystems. To address in C plots during the 2 years, but not in Cu after the this influence, we conducted a 2-years’ period mon- cutting practices (in the second year of sampling). itoring of fungi fruitbodies and macrofauna in sixteen Regarding soil macrofauna, higher values of taxa and experimental plots in Montado landscape in southern species richness were observed in C and Cu plots in the Portugal. A total of 4,881 frutibodies (57 taxa of first year of sampling. The species Aphaenogaster ECMF and 64 taxa of saprobic fungi) and 3,667 soil senilis and several Staphylinid morphospecies exhib- invertebrates (73 species and morphospecies) were ited a close relation with M plots, whilst most spider monitored in the experimental plots. There was greater families were directly associated to C and Cu plots. losses in sporocarps production and taxa composition, After the shrub cutting practices, higher values of taxa particularly the ECMF, in plots where shrub density and species richness of soil macrofauna were observed was controlled by permanent grazing (Ca) or involving in M and Ca plots; the presence of species with a high competitive ability to colonize disturbed areas faster might explain the results. Contrary to the frutibodies production and diversity, species richness and abun- dance within soil macrofauna were identical between Cu and C in 2004. Thus, fruiting macromycetes and A. M. Azul (&) H. Freitas soil macrofauna diversity and abundance in Monta- Department of Life Sciences, Centre for Functional do’s, appear highly sensitive to land use and somewhat Ecology, University of Coimbra, PO Box 3046, 3001-401 Coimbra, Portugal reflected a trend of severity to the current shrub e-mail: [email protected] management practices.

S. M. Mendes J. P. Sousa Keywords Fungal fruit-body Soil macrofauna Department of Life Sciences, IMAR-CMA, University of Coimbra, PO Box 3046, L. Ecosystems monitoring 3001-401 Coimbra, Portugal Land use Montado 123 Agroforest Syst

Introduction network that benefits plant communities by facilitating and influencing seedling establishment, by altering Quercus suber L. (cork oak) woodlands cover 23% of plant–plant interactions and by supplying and recy- the total forest area in Portugal (737 9 103 ha; DGRF cling nutrients (van der Heijden and Horton 2009), 2007). The cork oak woodlands are widespread in the while saprobic fungi are more specialized in decom- Mediterranean basin and cover about 2.2 million ha in posing dead organic plant material. In this context, a Europe (Portugal, Spain, France, and Italy) and North previous study reported that ECM fungal community Africa (Morocco, Algeria, and Tunisia). Most of cork associated to cork oak is quite diverse in both oak woodlands in Portugal are under agro-silvo- community structure and species composition, and pastoral exploitation, traditionally called Montado, affected by seasons, land use and cork oak mortality that is characterized by open oak formations of (Azul 2002; Azul et al. 2010). evergreen oaks (Q. suber and Quercus rotundifolia Soil fauna is also gaining importance in biodiversity L.), pastures and agricultural fields as undercover, assessment studies due to its active role on soil traditionally in a rotation scheme that includes fallows. processes and its sensitive response to changes in the The Montados are well adapted to the Mediterranean soil system (Bruyn 1997; Frouz 1999; Knoepp et al. environment and represent a good example of sustain- 2000; Rainio and Niemela¨ 2003; Sauberer et al. 2004). able agroforestry practice in Europe (Council of Soil organisms are representative of soil conditions Europe 1992) by combining two key aspects of land whether it’s physical, chemical or biological processes management: production and conservation, and due to we’re trying to assess (Blakely et al. 2002; Breure et al. their social and economic outcomes (Pinto-Correia 2005). This means that soil organisms could be used as and Vos 2004; Scarascia-Mugnozza et al. 2000). indicators of soil quality; soil invertebrates, one of the Changes in land management over the twentieth most abundant and diverse groups of soil organisms, century are thought to have contributed to Montado for example, often react very quickly to environmental landscape degradation (Joffre et al. 1999; Nunes et al. changes, with very sensitive responses (Bruyn 1997; 2005; Pinto-Correia 1993), with a significant decline Nickel and Hildebrandt 2003; Perner and Malt 2003; in cork oaks and plant and biodiversity (Da Rainio and Niemela¨ 2003). Some soil invertebrate Silva et al. 2008; Hector et al. 1999). Although indicators are already being used to assess certain successfully managed Montados remain, others are conditions (Beck et al. 2005; Hodkinson and Jackson increasingly being reforested with other tree species 2005;Ja¨nsch et al. 2005; Knoepp et al. 2000; Lavelle considered more lucrative by forest managers (e.g., et al. 2006; Nahmani et al. 2006; Rombke et al. Pinus pinea for the production of pine acorns; 2005; Ruiz Camacho et al. 2009; Sochova´ et al. 2006; Eucalyptus globulus, for the production of cellulose Souty-Grosset et al. 2005; Tischer 2005; Velasquez for the pulp industry), whilst other Montados are et al. 2007). abandoned and subject to shrub intrusion highly Several studies conducted in cork oak and holm- susceptible to fire (Nunes et al. 2005). In both cases, oak woodlands have been published (Cammell et al. a collapse of this highly-adapted and diverse Mediter- 1996; Da Silva et al. 2008; Deharveng et al. 2000; ranean-type ecosystem is observed. Rego and Dias 2000; Sousa et al. 1997) but none of Evaluating the impacts of management actions on these focused specifically on fungi and macrofauna as forests using bioindicators is widely recommended by indicators of the impacts of land use. European programs (Delbaere et al. 2002; EPBRS Understanding and predicting the consequences of 2002a, b). land use in soil biodiversity is emerging as one of the Plant mutualists, such as mycorrhizal fungi, and grand challenges for sustainable forest management, saprobes are widespread and are thought to maintain under climate change (European Environment Agency the structure and diversity of natural communities, 2004) and heavy mortality of evergreen oaks (Brasier influencing the performance of individual plants but 1996; Brasier and Scott 2008). In this work, we also altering plant community structure, plant produc- propose to investigate the performance of fruiting tivity, and nutrient cycling (Smith and Read 2008). macromycetes of ectomycorrhizal (ECMF) and sapro- Ectomycorrhizal (ECM) develop symbiotic structures bic fungi and soil macrofauna as indicators of land use on fine root tips and form a complex belowground in Montado ecosystems, an approach that can yield 123 Agroforest Syst insight into the link between management practices Montado landscape is dominated by Quercus and diversity descriptors of key soil components: fungi suber (40 to 60 trees per ha), but Quercus rotundi- and macrofauna. folia is also present, shrub strata (mainly Cistus salvifolius L., C. crispus L. and C. ladanifer L.) with plants with 4–5 years of age, occupying 65% of the Materials and methods total vegetation cover density. Cork is the main lucrative economical activity and it is harvested every Study site 9-year period; cattle breeding represent the second profitable activity. Land use is focused on practices to Field-work was conducted in a Montado landscape, control shrubs density to reduce the risk of fire. located in Foros de Vale de Figueira (Montemor-o- Novo, Portugal) (3884101000N, 882002300W) (Fig. 1). Experimental design The climate is typically Mediterranean, with a severe summer drought (2–4 months) and mild Sixteen experimental plots of 20 9 20 m were humid winters, with precipitation mainly from selected randomly among four Montado areas, each autumn to mid spring. The mean temperature ranges one with a different shrub management practice. The from 7.5°C (mean average in January) to 24°C four treatments include: the control (C), with no (mean average in July). Soils are classified as orthic shrub-cutting in the preceding 5 years, including the 2 luvisols with organic layers varying accordingly years (2003 and 2004) study period; the cut plots with land use and the year of perturbation; pH (Cu), with mechanized cutting practices that cut the ranges from 4.5 to 5.7. shoot of the plants and left it on the ground without till the soil afterwards; the cattle plots (Ca), with shrubs artificially maintained at low densities by permanent grazing of cows and sheep; and the mobilized plots (M), with mechanized practices with soil tillage that remove the completely the plants (Fig. 2). Shrub management practices were per- formed in Cu and M plots in the beginning of autumn 2004. Fruiting macromycetes of ECMF and saprobic fungi were monitored every 10 days during the peak fruiting period, from September to December, in 2003 and 2004. The plots were distributed erratically among four Montado areas and fruiting incidence per plot was assumed to be totally independent from the fruiting incidence of the neighbouring plots. Fungal fruitbodies were counted and mapped but not geo-referenced. Macromycetes were identified to species level to species level in most cases, according to Bon (1988), Courtecuisse and Duhem (1995) and Moreno et al. (1986); the unidentified species were also considered. The list of all taxa observed is presented in Appendix (Table 3). Taxa abundance of fungal fruitbodies was estimated as the cumulative number of individuals produced by a given taxon for each plot. The quanti- fication method for sporocarps was chosen instead of biomass analysis because of the limitations inherent to Fig. 1 Location of the Montado landscape in Montemor-o- sizes and biomass measurements among mushrooms, Novo: Alentejo region (southern Portugal) which frequently obscure fructification of smaller 123 Agroforest Syst

Fig. 2 Experimental areas selected in the Montado (C control, Cu shrub density controlled by cutting practices without soil tillage, Ca shrub density controlled by permanent grazing, M shrub density controlled by cutting practices followed by tillage of soil) individuals. Species frequency refers to the percentage Similarity) were performed based on the Bray–Curtis within the experimental areas in which a given species similarity index (Clarke and Gorley 2006). Data was log fruited at least once during the entire sampling period. transformed prior to analysis. This analysis was done on Sporocarps production was defined as the total number PRIMER 5 for Windows software (Clarke and Gorley of fruitbodies counted over the study period. 2006). Multivariate techniques were also used to Regarding soil macrofauna, sampling took place in associate taxa to stands: two correspondence analyses Autumn 2003 and Autumn 2004, for a period of 10 (CA), one per sampling year, were performed for fungi days. For each year, nine pitfall traps were settled on fruitbodies (stands were used as dummy environmental each plot, following a nested design. Samples were variables) and two principal component analyses brought to laboratory and biological material was (PCA), one per sampling year, were performed for soil sorted and identified to species level (when this was macrofauna data. Biological data used in these analyses not possible, morphospecies level was used). was also log transformed. The statistical significance of the canonical axes was evaluated by a Monte Carlo Data analysis permutation test. Both CA and PCA were performed in CANOCO 4.5 software (Ter Braak and Smilauer 1998). Fungi fruit bodies and soil macrofauna abundance and number of taxa at each stand were compared by an ANOVA, followed by a Newman–Keuls test when Results differences were found. In both cases, data was log transformed prior to analysis whenever normality or Fruiting macromycetes homoscedasticity criteria were not met (Zar 1996). For each zone, species diversity indices (Shannon–Wiener Abundance and biodiversity descriptors and Simpson) were calculated according to Magurran (1988). To evaluate differences between stands regard- A total of 4,881 fungal fruitbodies (57 ECMF taxa and ing community composition, ANOSIM (Analysis of 64 saprobe taxa; Table 3 in Appendix) were assessed 123 Agroforest Syst in the 16 experimental plots; 3,131 fruit bodies in 2003 Table 1 Fruitbodies abundance and diversity descriptors (114 taxa, 54 ECMF and 60 saprobes; Table 3 in 2003 2004 Appendix) and 1,750 fruit bodies in 2004 (55 taxa, 31 0 0 ECMF and 24 saprobes; Table 3 in Appendix). Fruitbodies S H D Fruitbodies S H D Overall, 33 families were observed in the 16 exper- C 1167 55 2.86 9.41 983 30 2.35 6.29 imental plots during the 2-year sampling period; being Cu 980 55 2.84 7.64 340 20 1.47 2.4 the families Russulaceae (20 taxa), Agaricaceae (15 Ca 668 25 2.23 5.39 395 13 2.07 6.52 taxa), Tricholomataceae (11 taxa), Thelephoraceae (9 M 316 38 2.88 12.31 32 5 1.42 3.88 taxa) and Amanitaceae (8 taxa) the best represented 0 (see Table 3 in Appendix). These five families S number of observed taxa, H Shannon diversity index, D Simpson diversity index, C control, Cu cutting practices using contributed with 52% of total taxa observed. Fruiting machinery with no soil tillage, Ca permanent grazing by cattle, ECMF fungi comprised 67% of total fruit bodies, M shrub management using machinery followed by soil tillage represented by 17 genera, mainly members of Russula,

Tomentella, Amanita and Lactarius (Table 3 in 1000 Appendix). The ECMF Astraeus hygrometricus (Pers.) ECMF 900 Morg. and Laccaria laccata (Scop.: Fr.) Berk. & 2003 2004 Broome were the most abundant with 1,008 and 721 800 fruit bodies, respectively, that corresponded to 35% of 700 the total fruiting macromycetes production (Table 3 in 2003 2004 Appendix). The saprobic community was represented 600 by 42 genera, mainly members of Clitocybe, Agaricus 500 and Coprinus (Table 3, in Appendix). The families

Agaricaceae (13 taxa) and Tricholomataceae (11 taxa) of fruit bodies 400 ° were the best represented, accounting for 57% of the N. 300 saprobic taxa (Table 3, in Appendix). Fruiting mac- romycetes community included 16 200 taxa (50% ECM) that produced up to 23% of total fruit 100 bodies (Table 3, in Appendix). No significant differences were observed on the 0 CCuCaM number of fruit bodies between the 16 plots, in 2003. However, in the second year sampling (after the shrub Fig. 3 Variation in ectomycorrhizal-forming fungal commu- cutting), significant differences were observed for the nity abundance as result of current techniques applied to four Montado areas on both abundance (F = 7.17, P\ control shrub density in the Montado areas (columns represent the mean and symbols the total abundance of fungal fruit 0.05) and taxa number (F = 6.85, P\0.05). In the first bodies, respectively; abbreviations as in Fig. 2) year sampling, the control (C) and the Montado areas with shrub density controlled by cutting practices with Changes in fruitbodies diversity and composition no soil tillage (Cu), both with plants with 4–5 years of among shrub management practices age and shrub strata occupying 65–75% of total vegetation cover, presented higher values in macro- Multivariate analysis clearly distinguished C and Cu mycetes richness and abundance (Table 1; Fig. 3). plots from Ca and M plots in 2003 (Fig. 5). Along Lower values in fruit bodies richness and abundance axis 1 (explaining 37.5% of the community compo- were observed in the plots where shrubs density and sitional variability) there’s a clear separation between growth are cut followed by soil tillage (M) and in the the—at the time—the C and Cu plots, and the Ca and plots where shrub management is conducted by M plots. In that sampling year, the ECMF Laccaria permanent grazing of cattle (Ca), respectively laccata and Xerocomus subtomentosus were closely (Table 1; Fig. 3). However, after the removal of related with C and Cu plots, whilst the saprobe understory shrubs in the second year sampling the Entoloma conferendum was associated to Ca plots. number of taxa and fruitbodies production decreased In 2004, the analysis performed (Fig. 6) separated all throughout the Cu and M plots (Table 1; Fig. 4). the four areas along Axis 1 (explaining 43.4% of the 123 Agroforest Syst

Saprobes Sco

500 1.0 2003 M 2004 400 2003

2004 Gsp 300

200 of fruit bodies °

N. Aph Lqu 100 Boa Lpe Cge Mme Hci Pez Lau Rcy Lch Rfo 0 C Lla CCuCaM Xsu Mqu Rfi Tve Fig. 4 Variation in saprobic fungi community abundance as Iri result of land use practices to control shrub density in the Ahy Montado areas (columns represent the mean and symbols the Cu Xhy total abundance of fungal fruit bodies, respectively; abbrevi- Eco ations as in Fig. 2) Mpr

Bae Cpu Ca Cru Lmo Pba Pca Man Pve -0.6 -0.4 0.8

Fig. 6 Correspondence analysis using the four experimental areas as dummy variables for fruit bodies data collected in

Gsp 2004 (abbreviations of experimental areas as in Fig. 2; see the Ca Fve Tbi Cfr codes in Table 3 in Appendix) Lmo Cru Cph Cpu Cge Cpi Pba Eco Cpl Aca Soil macrofauna Xsu M Ahy Iri Mqu CuHci Lpe Lla Abundance and biodiversity descriptors Tat LvoRso C Lch

Aph Ram Regarding soil macrofauna, 3,667 individuals were Ava Rcy Bas Rde caught in the traps: 1,799 individuals (53 species and Ctr Rfe Ame Lau Rkr Mme Rua Pse morphospecies) in 2003 and 1,868 individuals (34 Pez Rfo Bpl Rfi Rfr species and morphospecies) in 2004. Shrub manage- Pan Cmi ment techniques did affect significantly both the

-0.6 0.8 abundance (F = 4.82, P \ 0.05 for 2003 and F = -0.2 1.0 13.27, P\0.05) and the number of taxa observed per plot (F = 4.79, P\0.05 in 2003 and F = 7.87, P\0.05 Fig. 5 Correspondence analysis using the four experimental areas as dummy variables for fruit bodies data collected in in 2004). C and Cu plots presented the highest species 2003 (abbreviations of experimental areas as in Fig. 2; see the richness in 2003, followed by the M and Ca plots codes in Table 3 in Appendix) (Table 2). In 2004, a decrease in species richness and abundance was observed in all plots (with the excep- tion of Ca plot). However no significant differences community compositional variability), from the C were noticed in species diversity (Shannon) from one plots to the M plots. Most taxa associated to C and Cu year to the other. In the second year of sampling, plots before the cutting practices occurred in the C higher values of taxa abundance and species richness plots but no longer in the Cu plots in 2004. were observed in the M and Ca plots. 123 Agroforest Syst

Table 2 Soil macrofauna abundance and diversity descriptors ANOSIM (Global R = 0.839, P \ 0.01) corroborated 2003 2004 the differences in community composition observed and the pairwise tests performed revealed significant 0 0 Individuals S H D Individuals S H D differences between all plots (P \ 0.001 for every C 372 38 2.59 6.59 132 20 2.34 5.54 comparison made). In this first sampling period, ant Cu 519 41 3.01 13.95 132 19 2.6 11.65 species Aphaenogaster senilis and several Staphylinid Ca 242 29 2.77 12.01 1182 24 1.1 1.66 morphospecies were closely related to the M plots, M 666 33 2.44 6.76 420 27 2.21 4.69 whilst most of spider families were associated to less disturbed areas (Cu and C plots). Abbreviations as in Table 1 In the second year of sampling, the Principal Component Analysis performed (Fig. 8) clearly sep- Changes in community composition among shrub arated Ca and M plots from the remaining two stands management practices along Axis 1 (explaining 72.6% of the species- environment relation). Differences in community composition among areas were evaluated with an Multivariate analysis separated the plots with shrub P \ cut with machinery intervention followed by soil ANOSIM (Global R = 0.587, 0.01), which corroborated the differences observed. Pairwise tests tillage (M) from the remaining plots in 2003 (Fig. 7). revealed significant differences between all stands Along axis 1 (explaining 51.8% of the community P \ compositional variability) there’s a clear separation ( 0.001 for every comparison made), confirming that, even though C and Cu plots appear to be closer on between the M plots and the other plots, whilst Axis 2 the Principal Component Analysis, there were four (explaining 27.2% of the community compositional variability) separates the Ca plots from the other two different groups corresponding to the four plots in the study. In the second year of sampling, ant species shrub control techniques and C plots; using both axes, Aphaenogaster senilis was still closely related to the three different groups were observed—one with the, at the time, undisturbed stands (Cu and C), another with M plots as well as Staphylinid morphospecies and most Coleoptera. Species from some spider families, the Ca plots and the third one regarding the M plots. An like Lycosidae, and Thomisidae, were more related to the M and Ca plots. 1.0

Scy Ppa

Lc10 Cu Dip Gna Clg Lc13 Csc Gr2 Has Mtr Ca Gr1 Ppy Cha Lc1 Zoa Ta1 Xa4 Pti M Ppy Lin Pnt Zel St9 Ox2 Or1 Ase Iso OI2 C Cau Mst Aib Ap1 Cam Mss Mhi C Ld1 Chi Stg Opi Nem Col Xys Al1 Cur St6 Stg Cre Ld1 Lc7 Tes Cu Tet Pah Csc Fsa Or2 Hcl Agi Or4 Ths Ox2 OI4 Zor Aib Mei Pti OI1 Ox1 Bub Typ Hym Agi Ppa Lin Gas Sil St1 Ara Lc15 St9 Las Lc21 Ox1 Ozp Ap5 Ala Ase

Ca M -1.1 1.0 -1.0 -1.0 1.0 -1.0 1.5

Fig. 7 Correspondence analysis using the four experimental Fig. 8 Correspondence analysis using the four experimental areas as dummy variables for soil macrofauna data collected in areas as dummy variables for soil macrofauna data collected in 2003 (abbreviations of experimental areas as in Fig. 2; fauna 2004 (abbreviations of experimental areas as in Fig. 2; fauna codes are present in Table 4 in Appendix) codes are present in Table 4 in Appendix) 123 Agroforest Syst

Discussion Environment Agency 2004) and/or diseases (Brasier and Scott 2008). In this work we assessed soil macromycetes and Disturbances caused by cutting practices have macrofauna diversity descriptors in Montado land- been showed to have significant impacts in the scape to evaluate the influence of land use practices ECMF community diversity and composition (Byrd to control shrub density on soil biodiversity. Our et al. 2000; Hagerman et al. 1999; Smith et al. 2005). study revealed that shrub management influences the It is expected that the removal of understory species richness and abundance of fruiting macro- vegetation affect the ECM fungal community (Azul mycetes and soil macrofauna. 2002; Azul et al. 2010) with costs to the sporocarps Our results demonstrated that Montado’s under production. In our study, most of the ECMF taxa extensive silvopastoral exploitation comprise high associated to cutting practices with no soil tillage taxonomical diversity in fruiting macromycetes. Other (Cu) plots were present in the control (C) plots studies documented higher diversity in ECMF com- during the 2-years’ period, but not in Cu after the munity in old-growth forests dominated by Quercus shrub cutting practices were performed (2004), ilex L. (Richard et al. 2004) and coniferous (O’Dell indicating that ectomycorrhizal macrofungal compo- et al. 1999; Peter et al. 2001; Ferna´ndez-Toira´n et al. sition and fungal networks are indubitably affected 2006). The lower values in ECMF diversity may be by shrub management. The ECMF L. laccata and explained by our monitoring study limited to 2-years’ X. subtomentosus were closely related with less period. Richard et al. (2004) showed that in a Q. ilex disturbed plots (C and Cu in 2003; C in 2004) while old-growth forest fruiting macromycetes patterns was the saprobes were associated to Ca and M plots. This remarkably irregular, with 61.4% of the ECMF taxa corroborates the hypothesis that land use practices occurring one time during 3-years’ period. On con- allowing for autochthonous shrubs preservation, such trary, the saprobic community in Montado landscape as Cistus spp., are important to sustain ECMF (S = 64; Table 3 in Appendix) was identical to the old- diversity. However, the direct effects of shrub control growth Q. ilex forest (S = 68; Richard et al. 2004). This techniques in the specific mycobiont–phytobiont may be explained by the differential responses of associations and the activity of the interacting ECMF and saprobes to land use practices, and the mycelial systems of ECMF and saprobes and soil relatively low accumulation of favorable substrates to moisture conditions remain poorly understood but are decomposers. Also phenological patterns and physical extremely important in scenarios of drought-climate parameters, such climatic conditions are well known to change and oak mortality. influence fungi fructification. Cattle husbandry represents the second profitable The family Russulaceae covered 16.5% of the activity in Montado areas in study. Our results fruiting fruit bodies diversity (and 35% of ECMF). revealed that cattle grazing were rather efficient to A comparable tendency for the dominance of Russul- control shrub density and growth. Nevertheless, cau- aceae species was observed belowground in Montado tion in the interpretation of these results is recom- landscape (Azul 2002; Azul et al. 2010). The domi- mended since perceptible effects on fruitbodies nance of Russulaceae above- and below-ground was production and diversity were found, particularly on reported in Mediterranean (Bergemann and Garbelotto ECMF community (cumulative richness of ECMF 2006; Courty et al. 2008; Richard et al. 2004, 2005) taxa in C plots was 3 times higher than in Ca plots). and Temperate forests (Lilleskov et al. 2004; Tedersoo Cattle grazing even with restricted number of individ- et al. 2003), and it may be explained by the fact that uals imply multiple ecological consequences related to this ECMF family comprises a large range of species the plant species responses to herbivory (Ayres et al. and high diverse ecological requirements. The taxo- 2004) and to the nitrogen inputs provided by nomical diversity and cosmopolitan distribution (Avis et al. 2003; Edwards et al. 2004; Trudell et al. within Russulaceae further impose a better under- 2004), and may directly affect the sporocarp produc- standing of the putative role of these mutualist fungi in tion. The use of heavy machinery and livestock are soil processes in mediterranean ecosystems, e.g., also known to cause soil compaction, affecting soil stabilization following disturbance (Costa et al. water content and plant growth (Hamza and Anderson 2009; Pinto-Correia 1993), drought stress (European 2005). Our results call for further investigations to 123 Agroforest Syst evaluate the implications of livestock and nitrogen senillis, scutellaris and Staphylinidae inputs on ECMF community fruiting patterns. morphospecies 9) have very high abundances in the M Regarding soil macrofauna overall analysis showed and Ca plots; these taxa are known to be among the that different scenarios of shrub management cause opportunistic fauna (Andersen and Sparling 1997;da effects in soil macrofauna communities. The changes Silva et al. 2009; Dauber and Wolters 2005;Thorbekand observed were not only in terms of species richness and Bilde 2004), which take advantage of severe changes that abundance but also in terms of community composition. lead to the ‘‘disappearance’’ of dominant taxa in an Understory vegetation management appears to be one ecosystem after a disturbance (like shrub cut or tillage); of the factors causing a community composition shift in the absence of competition from these formerly (from 2003 to 2004), and its effect combined with the dominant groups, opportunistic fauna uses its ability to mobilization of the soil induce a more significant effect colonize the area that now presents different features. than the shrub cut alone. It is known that certain groups Overall, our study revealed that a considerable (or species within them), like Formicidae, Coleoptera or variation in soil biodiversity in Montado ecosystems Araneae tend to respond to changes in the structure of is explained by the current practices used to control their habitat, especially when its architecture is one of shrub density. This variation in soil biodiversity was the factors changing (Grill et al. 2005; Retana and Cerda´ observed in terms of species richness and abundance 2000). Our results are in agreement with previous but also in community’s composition of both fruiting studies, that showed that some groups, like , exhibit macromycetes and soil macrofauna. Our results a positive relation with the percentage of vegetation showed that shrub management based on cutting cover in Mediterranean systems (Retana and Cerda´ practices without soil tillage revealed to be less 2000). Other groups, like spiders, show a good corre- severe when comparing with permanent grazing or lation with the management intensity (Perner and Malt shrub management with tillage of soil afterwards. 2003) and its richness is strongly and negatively Although results cannot be generalized for other affected by an intensification of management. The forestry ecosystems, fruiting macromycetes and soil structure of the surrounding vegetation is also an macrofauna were measurable and reproducible in important factor when it comes to spiders (Grill et al. Montado’s, also to be sensitive indicators of soil 2005;McAdametal.2007;New2005)—web-building components population’s vulnerability use, and and active ground predator spiders are closely related somewhat reflected a trend of severity, to the current with a well structured understory vegetation cover (Cu techniques used to control shrub density. We believe plots in 2003; C plots in 2003 and 2004), but species that fruiting macromycetes and soil macrofauna may recognized as colonizers (like some from Thomisidae be useful indicators and extremely important tools family) are more associated with more open areas and, for explaining the ecological impacts of land use in therefore, are more associated with the Cu plots in 2004 forestry ecosystems. Our results imply further stud- and M and Ca plots in the 2-years period. Coleoptera ies to understand the impacts of these soil biodiver- families appeared to be associated to areas where sity changes regarding their role in ecosystems intensive management was adopted (soil mobilization functions and the plant physiological response to and cattle ranching). This is in agreement with the major aspects of cork oak sustainability in future: findings of Vanbergen et al. (2005), who showed the production and health. preference of some Coleoptera families for open areas. In the case of Scarabaeidae our results agree with Verdu´ Acknowledgments Financial support was provided by FCT- et al. (2007) that reported the strong association of this MCTES (Portuguese Foundation for Science and Technology) and European fund FEDER, project POCTI/AGG/42349/2001. group with grazed areas. Although, in the second year of AM Azul was supported by an individual grant from FCT- sampling, the highest values for species richness were MCTES (SFRH7BPD/5560/2001). observed in the M and Ca plots, this may have been an effect of opportunistic species, which were able to colonize the areas with new habitat configuration Appendix resulting from the management intervention. Some species (namely Aphaenogaster iberica, Aphaenogaster See Appendix Tables 3 and 4.

123 123 Table 3 List of taxa in the four experimental stands and information related to their fruiting pattern during the period 2003–2004 Taxon Family Edible Habit Host range C Cu Ca M CD AFRAFRAFRAFR

Agaricus augustus Fr. Agaricaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 2 1 (1, 0) Aau Agaricus campestris L.: Fr. Agaricaceae Edible Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 9 1 (1, 0) Aca Agaricus silvicola (Vittad.) Peck Agaricaceae Edible Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 3 1 (1, 0) Asi Agaricus xanthodermus Genev. Agaricaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Axa Amanita battarrae (Boud.) Bon Amanitaceae – ECMF Broad HR 7 2 (3, 3) 6 1 (3, 0) 0 0 (0) 0 0 (0) Aba Amanita caesarea (Scop.) Pers. Amanitaceae Edible ECMF Thermophilic 5 2 (2, 3) 0 0 (0) 0 0 (0) 0 0 (0) Ace Amanita franchetii (Boud.) Fayod Amanitaceae – ECMF Broad HR 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Afr Amanita muscaria (L.) Lam. Amanitaceae – ECMF Broad HR 2 1 (0, 2) 1 1 (1, 0) 0 0 (0) 0 0 (0) Amu Amanita pantherina (DC.) Krombh. Amanitaceae – ECMF Broad HR 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) Apa Amanita phalloides (Vaill. ex Fr.) Link Amanitaceae – ECMF Broad HR 20 2 (4, 3) 35 1 (4, 0) 0 0 (0) 0 0 (0) Aph Amanita rubescens Pers Amanitaceae Edible ECMF Broad HR 0 0 (0) 3 1 (2, 0) 0 0 (0) 0 0 (0) Aru Amanita vaginata (Bull.) Lam. Amanitaceae – ECMF Broad HR 5 1 (4, 0) 4 1 (3, 0) 0 0 (0) 0 0 (0) Ava Astraeus hygrometricus (Pers.) Morg. Diplocystidiaceae – ECMF Broad HR 241 2 (3, 3) 531 2 (3, 4) 236 2 (3, 1) 0 0 (0) Ahy Auricularia mesenterica (Dicks.) Pers. Auriculariaceae – Saprobe – 0 0 (0) 1 1 (1, 0) 0 0 (0) 12 1 (2, 0) Ame Bisporella citrina (Batsch) Korf and S.E. Carp. Helotiales – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Bic aereus Bull. Edible ECMF Thermophilic 14 2 (2, 3) 0 0 (0) 0 0 (0) 0 0 (0) Boa Bull. Boletaceae Edible ECMF Broad HR 2 2 (1, 1) 0 0 (0) 0 0 (0) 0 0 (0) Bed Boletus satanas Lenz Boletaceae – ECMF Thermophilic 1 1 (0, 1) 0 0 (0) 0 0 (0) 0 0 (0) Bsa Bovista aestivalis (Bonord.) Demoulin Agaricaceae – ECMF Broad HR 11 1 (2, 0) 17 2 (3, 2) 102 1 (0, 3) 0 0 (0) Bae Bovista plumbea Pers. Agaricaceae – ECMF Broad HR 0 0 (0) 0 0 (0) 0 0 (0) 10 1 (2, 0) Bpl Clitocybe fragans (With.) P. Kumm. Tricholomataceae – Saprobe – 0 0 (0) 0 0 (0) 14 1 (3, 0) 0 0 (0) Cfr Clitocybe geotropa (Bull.: Fr.) Que´l. Tricholomataceae Edible Saprobe – 66 2 (1, 3) 43 1 (2, 0) 22 1 (4, 0) 13 1 (2, 0) Cge Clitocybe gibba (Pers.) Kumm. Tricholomataceae Edible Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Cgi Clitocybe phaeophthalma (Pers.) Kuyper Tricholomataceae Edible Saprobe – 0 0 (0) 0 0 (0) 37 1 (2, 0) 0 1 (0) Cph Clitocybe squamulosa (Pers.) Fr. Tricholomataceae – Saprobe – 0 0 (0) 1 1 (1, 0) 0 0 (0) 2 1 (1, 0) Csq Collybia butyracea (Bull ex Fr.) Quel. Marasmiaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Cbu Conocybe pubescens (Gill.) Kuhn. Bolbitiaceae – Saprobe – 0 0 (0) 0 0 (0) 17 2 (3, 1) 0 0 (0) Cpu

Conocybe rubiginosa Watling Bolbitiaceae – Saprobe – 0 0 (0) 0 0 (0) 30 2 (3, 2) 0 0 (0) Cru Syst Agroforest Coprinus comatus (O.F. Mu¨ll.) Pers. Agaricaceae Edible Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 4 1 (1, 0) Cco Coprinus domesticus (Bolt.: Fr) S.F. Gray Agaricaceae – Saprobe – 0 0 (0) 9 1 (1, 0) 0 0 (0) 0 1 (0) Cdo Coprinus micaceous (Bull.) Fr. Agaricaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 14 1 (1, 0) Cmi gooetSyst Agroforest Table 3 continued Taxon Family Edible Habit Host range C Cu Ca M CD A FR A FR A FR A FR

Coprinus picaceus (Bull.) Fr. Agaricaceae – Saprobe – 0 0 (0) 5 2 (3, 1) 14 2 (4, 2) 2 1 (1, 0) Cpi Coprinus plicatilis (Curt.) Fr. Agaricaceae – Saprobe – 0 0 (0) 11 1 (2, 0) 34 1 (3, 0) 0 1 (0) Cpl Cortinarius amoenolens Henry ex Orton Cortinariaceae – ECMF Broad HR 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 1 (0) Cam Cortinarius trivialis Lge. Cortinariaceae – ECMF Angiosperms 96 1 (4, 0) 12 1 (2, 0) 0 0 (0) 0 1 (0) Ctr Crepidotus mollis (Bull ex Fr.) Kum. Inocybaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Cmo Crepidotus variabilis (Pers.) Kumm. Inocybaceae – Saprobe – 0 0 (0) 0 0 (0) 1 1 (1, 0) 0 0 (0) Cva Entoloma conferendum (Britzelm.) Noordel. Entolomataceae – Saprobe – 2 1 (1, 0) 0 0 (0) 284 2 (4, 2) 4 1 (0, 1) Eco Flammulina velutipes (Curt.) Sing. Physalacriaceae – Saprobe – 0 0 (0) 0 0 (0) 23 1 (1, 0) 0 0 (0) Fve Ganoderma applanatum (Pers.) Pat. Ganodermataceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Gap Gymnopilus penetrans (Fr.) Murrill Strophariaceae – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Gpe Gymnopilus spectabilis (Fr.) Sing. Strophariaceae – Saprobe – 0 0 (0) 0 0 (0) 72 2 (1, 1) 14 1 (0, 1) Gsp Hebeloma cistophilum Mre. Strophariaceae – ECMF Cistus specific 210 2 (2, 3) 113 1 (2, 0) 8 1 (1, 0) 0 0 (0) Hci Hebeloma sp.1 Strophariaceae – ECMF – 5 2 (2, 1) 0 0 (0) 0 0 (0) 0 1 (0) He1 Hygrocybe conica (Scop.) P. Kumm. Hygrophoraceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 2 1 (1, 0) Hco Hymenochaete rubiginosa (Dicks.) Le´v. Hymenochaetaceae – Saprobe – 0 0 (0) 0 0 (0) 2 1 (1, 0) 0 0 (0) Hru Inocybe rimosa (Bul.) P. Kumm. Inocybaceae – ECMF Broad HR 7 2 (2, 3) 53 2 (3, 4) 8 1 (3, 0) 1 1 (1, 0) Iri Laccaria laccata (Scop.) Cke. Tricholomataceae Edible ECMF Broad HR 640 2 (3, 4) 50 1 (3, 0) 13 1 (1, 0) 18 1 (1, 0) Lla Lactarius atlanticus Bon Russulaceae – ECMF Broad HR 0 0 (0) 0 0 (0) 3 1 (3, 0) 0 0 (0) Lat Lactarius aurantiacus (Pers.) Gray Russulaceae – ECMF Broad HR 41 2 (2, 3) 23 1 (4, 0) 0 0 (0) 0 0 (0) Lau Lactarius chrysorrheus Fr. Russulaceae – ECMF Quercus sp 39 2 (4, 2) 0 0 (0) 0 0 (0) 1 1 (1, 0) Lch Lactarius quietus (Fr.) Fr. Russulaceae – ECMF Quercus sp 16 1 (0, 3) 0 0 (0) 0 0 (0) 0 1 (0) Lqu Lactarius volemus (Fr.) Fr. Russulaceae Edible ECMF Quercus sp 7 1 (4, 0) 14 1 (4, 0) 0 0 (0) 1 1 (1, 0) Lvo Lentinellus cochleatus (Pers.) P. Karst. Auriscalpiaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Lco Lepiota josserandii Bon and Boiffard Agaricaceae – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Ljo Lepista nuda (Schumach.) Sing. Tricholomataceae – Saprobe – 0 0 (0) 0 0 (0) 1 1 (0, 1) 0 0 (0) Lnu Lepista sordida (Schumach.) Sing. Tricholomataceae – Saprobe – 0 0 (0) 4 1 () 0 0 (0) 0 0 (0) Lso Lycoperdon molle Pers. Agaricaceae – Saprobe – 0 0 (0) 0 0 (0) 50 2 (2, 3) 0 0 (0) Lmo Lycoperdon perlatum Pers. Agaricaceae – Saprobe – 14 2 (3, 2) 11 1 (3, 0) 4 1 (2, 0) 1 1 (1, 0) Lpe

123 Macrolepiota procera (Scop.) Sing. Agaricaceae Edible Saprobe – 5 1 (0, 4) 1 1 (0, 1) 30 1 (0, 3) 3 1 (3, 0) Mpr Marasmius androsaceus (L.) Fr. Marasmiaceae Edible Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 17 1 (1, 0) Man Marasmius quercophilus Pouz. Marasmiaceae – Saprobe – 112 2 (1, 3) 31 2 (2, 3) 0 0 (0) 60 1 (1, 0) Mqu 123 Table 3 continued Taxon Family Edible Habit Host range C Cu Ca M CD AFRAFRAFRAFR

Megacollybia platyphylla (Pers.) Kotl. and Pouz. Marasmiaceae – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 1 (0) Mpl Melanoleuca brevipes (Bull.) Pat. Tricholomataceae – Saprobe – 0 0 (0) 0 0 (0) 1 1 (1, 0) 0 0 (0) Mbr Melanoleuca melanoleuca (Pers.) Murr. Tricholomataceae – Saprobe – 27 2 (3, 2) 0 0 (0) 0 0 () 0 0 (0) Mme Meripilus giganteus (Pers.) P. Karst. Meripilaceae – Saprobe – 0 0 (0) 0 0 (0) 1 1 (1, 0) 0 0 (0) Mgi Mycena inclinata (Fr.) Que´l. Mycenaceae – Saprobe – 0 0 (0) 24 1 (1, 0) 0 0 (0) 0 0 (0) Min Omphalina sp. Hygrophoraceae – Saprobe – 22 2 (1, 1) 0 0 (0) 0 0 (0) 0 0 (0) Omp Omphalotus olearius (DC.) Sing. Marasmiaceae – Saprobe – 34 2 (1, 1) 0 0 (0) 0 0 (0) 0 0 (0) Ool Panaeolus antillarum (Fr.) Dennis Strophariaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 14 1 (1, 0) Pan Panaeolus semiovatus (Sow.) Lundell and Nannf. Strophariaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 43 1 (2, 0) Pse Panaeolus sphinctrinus (Fr.) Que´l. Strophariaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 2 1 (1, 0) Psp Psathyrella velutina (Pers. ex Fr.) Sing. Psathyrellaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 21 1 (1, 0) Pve Psathyrella candolleana (Fr.: Fr.) Psathyrellaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 17 1 (1, 0) Pca Peziza sp. Pezizaceae – ECMF Broad HR 21 2 (1, 1) 4 1 (1, 0) 0 0 (0) 0 0 (0) Pez Peziza badia Pers.: Fr. Pezizaceae – ECMF Broad HR 0 0 (0) 2 1 (1, 0) 26 2 (2, 1) 0 0 (0) Pba Phanerochaete sanguinea (Fr.) Pouzar Phanerochaetaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Psa Pisolithus arhizus (Scop.) Rauschert. Sclerodermataceae – ECMF Broad HR 1 1 (1, 0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Par Polyporus arcularius (Batsch.) Fr. Polyporaceae – Saprobe – 0 0 (0) 2 1 (0, 1) 0 0 (0) 0 0 (0) Poa Pulcherricium caeruleum (Schrad.) Parm. Phanerochaetaceae – Saprobe – 5 2 (1, 2) 4 1 (1, 0) 0 0 (0) 0 0 (0) Puc Rickenella fibula (Bull.) Raithel. Agaricomycete – Saprobe – 53 2 (3, 3) 50 2 (3, 4) 0 0 (0) 0 0 (0) Rfi Russula sp. 1 Russulaceae – ECMF – 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Ru1 Russula sp. 2 Russulaceae – ECMF – 4 1 (3, 0) 16 1 (3, 0) 0 0 (0) 0 0 (0) Ru2 Russula sp. 3 Russulaceae – ECMF – 5 1 (2, 0) 0 0 (0) 0 0 (0) 0 0 (0) Ru3 Russula sp. 4 Russulaceae – ECMF – 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) Ru4 Russula amoena Que´l. Russulaceae – ECMF Broad HR 0 0 (0) 16 1 (2, 0) 0 0 (0) 0 0 (0) Ram Russula amoenolens Romagn. Russulaceae – ECMF Broad HR 78 1 (4, 0) 32 1 (4, 0) 0 0 (0) 0 0 (0) Rua Russula cyanoxantha (Shaeff.) Fr. Russulaceae Edible ECMF Broad HR 86 2 (4, 3) 24 1 (4, 0) 0 0 (0) 0 0 (0) Rcy Russula delica Fr. Russulaceae – ECMF Broad HR 26 1 (4, 0) 3 1 (2, 0) 0 0 (0) 0 0 (0) Rde

Russula fellea (Fr.) Fr. Russulaceae – ECMF Broad HR 20 1 (3, 0) 5 2 (1, 1) 3 1 (3, 0) 0 0 (0) Rfe Syst Agroforest Russula foetens (Pers.) Pers. Russulaceae – ECMF Broad HR 78 2 (3, 4) 3 1 (2, 0) 0 0 (0) 0 0 (0) Rfo Russula fragilis Fr. Russulaceae – ECMF Broad HR 11 1 (4, 0) 2 1 (0, 2) 0 0 (0) 0 0 (0) Rfr Russula krombholzii Shaeff. Russulaceae – ECMF Broad HR 21 1 (3, 0) 25 1 (3, 0) 0 0 (0) 0 0 (0) Rkr gooetSyst Agroforest Table 3 continued Taxon Family Edible Habit Host range C Cu Ca M CD AFR AFR AFR AFR

Russula ochroleuca (Pers.) Fr. Russulaceae – ECMF Broad HR 1 1 (1, 0) 0 0 (0) 0 0 (0) 0 0 (0) Roc Russula sororia (Fr.) Romagn Russulaceae – ECMF Broad HR 43 1 (4, 0) 23 1 (2, 0) 0 0 (0) 4 1 (2, 0) Rso Russula xerampelina (Schaeff.) Fr. Russulaceae Edible ECMF Broad HR 6 1 (3, 0) 6 1 (2, 0) 0 0 (0) 0 0 (0) Rxe Schizophyllum commune (L.) Fr. Schizophyllaceae – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 6 1 (0, 3) Sco Scleroderma citrinum Pers. Sclerodermataceae – ECMF Broad HR 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) Sci Scleroderma polyrhizum (J.F. Gmel.) Pers. Sclerodermataceae – ECMF Thermophilic 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) Spo Stereum hirsutum (Wild.) Pers. Stereaceae – Saprobe – 5 1 (3, 0) 4 1 (1, 0) 0 0 (0) 4 1 (0, 2) Shi Stereum reflexum D. A. Reid Stereaceae – Saprobe – 2 1 (2, 0) 0 0 (0) 0 0 (0) 0 0 (0) Sre Tomentella atramentaria Rost., Bot. Tidsskr. Thelephoraceae – ECMF Broad HR 0 0 (0) 7 2 (3, 1) 0 0 (0) 1 1 (1, 0) Tat Tomentella brevispina (Bourd. and Galz.) M.J. Lar. Thelephoraceae – ECMF Broad HR 0 0 (0) 2 2 (1, 1) 0 0 (0) 0 1 (0) Tbr Tomentella sublilacina Ellis and Holw.) Wakef. Thelephoraceae – ECMF Broad HR 0 0 (0) 2 2 (1, 1) 0 0 (0) 0 0 (0) Tsu Tomentella subtestacea Bourdot and Galin, Bull. Thelephoraceae – ECMF Broad HR 0 0 (0) 2 2 (1, 1) 0 0 (0) 0 0 (0) Tos Tomentella stuposa (Link) Stalpers Thelephoraceae – ECMF Broad HR 0 0 (0) 3 2 (2,1) 0 0 (0) 0 0 (0) Tst Tomentella sp1 Thelephoraceae – ECMF – 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) To1 Tomentella sp2 Thelephoraceae – ECMF – 0 0 (0) 1 1 (1, 0) 0 0 (0) 0 0 (0) To2 Tomentella sp3 Thelephoraceae – ECMF – 0 0 (0) 3 1 (3, 0) 0 0 (0) 0 0 (0) To3 Tomentella sp4 Thelephoraceae – ECMF – 0 0 (0) 1 1 (0, 1) 0 0 (0) 0 0 (0) To4 Trametes hirsuta (Wulf.) Lloyd Polyporaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Thi Trametes versicolor (L.) Lloyd Polyporaceae – Saprobe – 2 1 (2, 0) 6 2 (1, 2) 12 1 (0, 3) 6 1 (0, 2) Tve Tremella encephala Pers. Tremellaceae – Saprobe – 0 0 (0) 6 2 (1, 2) 0 0 (0) 0 0 (0) Ten Tremella foliacea Pers. Tremellaceae – Saprobe – 1 1 (1, 0) 4 2 (1, 1) 0 0 (0) 0 0 (0) Tfo Tremella mesenterica Retz. Tremellaceae – Saprobe – 1 1 (1, 0) 0 0 (0) 0 0 (0) 2 1 (0, 1) Tme Trichaptum biforme (Fr.) Ryvarden Polyporaceae – Saprobe – 0 0 (0) 0 0 (0) 10 1 (1, 0) 25 1 (3, 0) Tbi Tricholoma sulphurum (Bull.) P. Kumm. Tricholomataceae – ECMF Broad HR 2 1 (0, 1) 0 0 (0) 0 0 (0) 0 0 (0) Tsu Volvariella speciosa (Fr.) Sing. Pluteaceae – Saprobe – 0 0 (0) 0 0 (0) 0 0 (0) 1 1 (1, 0) Vsp Xerocomus chrysenteron (Bull.) Que´l. Boletaceae – ECMF Broad HR 3 1 (0, 1) 0 0 (0) 0 0 (0) 0 0 (0) Xch Xerocomus subtomentosus (L.) Que´l. Boletaceae – ECMF Broad HR 14 2 (1, 4) 3 1 (2, 0) 5 2 (2, 3) 0 0 (0) Xsu Xylaria hypoxylon (L.) Grev. Xylariaceae – Saprobe – 0 0 (0) 51 2 (1, 2) 0 0 (0) 0 0 (0) Xhy

123 A fruitbody abundance determined as the number of the specimens over the two fruiting seasons in the 16 experimental plots, FR fruiting regularity determined as the number of seasons of fruitbodies occurrence, with the the total number of plots occupied in each fruiting season given in parenthesis, CD codes used in multivariate analysis Agroforest Syst

Table 4 List of soil macrofauna taxa collected in the study Taxon (Species/morphospecies) Order (sub-order) Family (sub-family) FC

Aphaenogaster gibbosa (Latreille, 1798) (Apocrita) Formicidae () Agi Aphaenogaster iberica Emery, 1908 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Aib Aleocharinae morphospecies 1 Coleoptera Staphylinidae (Aleocharinae) Al1 Alopecosa albofasciata (Brulle´, 1832) Araneae Lycosidae Ala Aphidinae morphospecies 1 Hemiptera Aphididae (Aphidinae) Ap1 Aphidinae morphospecies 5 Hemiptera Aphididae (Aphidinae) Ap5 Araneae morphospecies 1 Araneae n.d. Ara Aphaenogaster senilis Mayr, 1853 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Ase Bubas morphospecies 1 Coleoptera Scarabaeidae Bub Camponotus morphospecies 1 Hymenoptera (Apocrita) Formicidae () Cam Crematogaster auberti Emery, 1869 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Cau Chasmatopterus morphospecies 1 Coleoptera Scarabaeidae Cha Chilopoda morphospecies 1 Chilopoda n.d. Chi Calathus granatensis Vuillefroy, 1866 Coleoptera Carabidae Clg Coleoptera morphospecies 1 Coleoptera n.d. Col Crematogaster morphospecies 1 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Cre Crematogaster scutellaris (Olivier, 1792) Hymenoptera (Apocrita) Formicidae (Myrmicinae) Csc Curculionidae morphospecies 1 Coleoptera Curculionidae Cur Diplopoda morphospecies 1 Diplopoda n.d. Dip Formica sanguinea Latreille, 1798 Hymenoptera (Apocrita) Formicidae (Formicinae) Fsa Gastropoda morphospecies 1 Gastropoda n.d. Gas Gnaphosidae morphospecies 1 Araneae Gnaphosidae Gna Gryllidae morphospecies 1 Orthoptera (Ensifera) n.d. Gr1 Gryllidae morphospecies 2 Orthoptera (Ensifera) n.d. Gr2 Hahnia morphospecies 1 Araneae Hahniidae Has Carabus (Hadrocarabus) lusitanicus (Fabricius, 1801) Coleoptera Carabidae Hcl Hymenoptera morphospecies 1 Hymenoptera (Apocrita) n.d. Hym Isopoda morphospecies 1 Isopoda n.d. Iso Lasius morphospecies 1 Hymenoptera (Apocrita) Formicidae (Formicinae) Las Coleoptera larvae morphospecies 1 Coleoptera n.d. Lc1 Coleoptera larvae morphospecies 10 Coleoptera n.d. Lc10 Coleoptera larvae morphospecies 13 Coleoptera n.d. Lc13 Coleoptera larvae morphospecies 15 Coleoptera n.d. Lc15 Coleoptera larvae morphospecies 21 Coleoptera n.d. Lc21 Coleoptera larvae morphospecies 7 Coleoptera n.d. Lc7 Diptera larvae morphospecies 1 Diptera n.d. Ld1 Linyphiidae morphospecies 1 Araneae Linyphiidae Lin Meioneta pseudorurestris (Wunderlich, 1980) Araneae Linyphiidae Mei Messor hispanicus Santschi, 1919 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Mhi Messor morphospecies 1 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Mss Messor structor (Latreille, 1798) Hymenoptera (Apocrita) Formicidae (Myrmicinae) Mst Carabus (Macrothorax) rugosus Fabricius, 1792 Coleoptera Carabidae Mtr Nemesia morphospecies 1 Araneae Nemesidae Nem Caelifera morphospecies 68 Orthoptera (Caelifera) n.d. OI1 Caelifera morphospecies 71 Orthoptera (Caelifera) n.d. OI4 123 Agroforest Syst

Table 4 continued Taxon (Species/morphospecies) Order (sub-order) Family (sub-family) FC

Opiliones morphospecies 1 Opiliones n.d. Opi Caelifera morphospecies 1 Orthoptera (Caelifera) n.d. Or1 Caelifera morphospecies 2 Orthoptera (Caelifera) n.d. Or2 Caelifera morphospecies 4 Orthoptera (Caelifera) n.d. Or4 Oxytelinae morphospecies 1 Coleoptera Staphylinidae (Oxytelinae) Ox1 Oxytelinae morphospecies 2 Coleoptera Staphylinidae (Oxytelinae) Ox2 Ozyptila pauxilla (Simon, 1870) Araneae Thomisidae Ozp Pardosa hortensis (Thorell, 1872) Araneae Lycosidae Pah Laemostenus (Pristonychus) terricola (Herbst, 1784) Coleoptera Carabidae Pnt pallidula (Nylander, 1849) Hymenoptera (Apocrita) Formicidae (Myrmicinae) Ppa pygmaea (Latreille, 1798) Hymenoptera (Apocrita) Formicidae (Formicinae) Ppy Ptinidae morphospecies 1 Coleoptera Ptinidae Pti Scydmaenidae morphospecies 1 Coleoptera Scydmaenidae Scy Silvanidae morphospecies 1 Coleoptera Silvanidae Sil Staphylininae morphospecies 1 Coleoptera Staphylinidae (Staphylininae) St1 Staphylininae morphospecies 6 Coleoptera Staphylinidae (Staphylininae) St6 Staphylininae morphospecies 9 Coleoptera Staphylinidae (Staphylininae) St9 Steropus globosus (Fabricius, 1792) Coleoptera Carabidae Stg Tachyporinae morphospecies 1 Coleoptera Staphylinidae (Tachyporinae) Ta1 Tegenaria morphospecies 1 Araneae Theridiidae Tes Tetramorium morphospecies 1 Hymenoptera (Apocrita) Formicidae (Myrmicinae) Tet Theridion morphospecies 1 Araneae Theridiidae Ths Typhoeus morphospecies 1 Coleoptera Geotrupidae Typ Xantholininae morphospecies 4 Coleoptera Staphylinidae (Xantholininae) Xa4 Xysticus morphospecies 1 Araneae Thomisidae Xys Zelotes morphospecies 1 Araneae Gnaphosidae Zel Zodarion alacre (Simon, 1870) Araneae Zodariidae Zoa Zora silvestris Kulczynski, 1897 Araneae Zoridae Zor Taxa is alphabetically listed according to FC; FC fauna code used on multivariate analysis; n.d. not determined

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