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Chronic Anthropogenic Disturbance As a Secondary Driver of Ant Community Structure: Interactions with Soil Type in Brazilian Caatinga

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Chronic Anthropogenic Disturbance As a Secondary Driver of Ant Community Structure: Interactions with Soil Type in Brazilian Caatinga Environmental Conservation (2017) 44 (2): 115–123 C Foundation for Environmental Conservation 2016 doi:10.1017/S0376892916000291 Chronic anthropogenic disturbance as a secondary driver of ant community structure: interactions with soil type in Brazilian Caatinga FERNANDA M. P. OLIVEIRA1,2, JOSÉ DOMINGOS RIBEIRO-NETO2 , ALAN N. ANDERSEN3 AND INARA R. LEAL4 ∗ 1Programa de Pós-Graduacão¸ em Biologia Animal, Universidade Federal de Pernambuco, Av. Prof. Moraes Rêgo s/n, Cidade Universitária, 50690–901, Recife, PE, Brazil, 2Programa de Pós-Graduacão¸ em Biologia Vegetal, Universidade Federal de Pernambuco, Av. Prof. Moraes Rêgo s/n, Cidade Universitária, 50690–901, Recife, PE, Brazil, 3CSIRO Land &Water, Tropical Ecosystems Research Centre, PMB 44 Winnellie, NT 0822, Australia and 4Departamento de Botânica, Universidade Federal de Pernambuco, Av. Prof. Moraes Rêgo s/n, Cidade Universitária, 50690–901, Recife, PE, Brazil Date submitted: 7 December 2015; Date accepted: 9 June 2016; First published online 19 August 2016 SUMMARY loss (Murcia 1995;Veldmanet al. 2009; Ahrends et al. 2010; Laurance et al. 2011). Much of the remaining habitat, Habitat loss is widely recognized as the major however, experiences chronic anthropogenic disturbance cause of global biodiversity decline, but remaining (CAD sensu Singh 1998) that does not result in habitat loss and habitat is increasingly threatened by chronic human fragmentation. Such disturbance includes firewood collection, disturbances. Using a multi-model averaging approach exploitation of non-timber products, hunting, overgrazing we examined the association between five chronic by livestock and intentional introduction of invasive plant disturbance surrogates and the richness and taxonomic species (Martorell & Peters 2005). Moderate levels of CAD and functional composition of ants in Brazilian can actually promote diversity (Connell 1978;Grime1979), Caatinga. Using pitfall traps in 47 plots near but high levels typically have negative impacts (Singh 1998). Parnamirim city (Pernambuco) across two soil types The ecological impacts of CAD have received far (sand and clay), we recorded 53 species from 27 less research attention than those associated with acute genera. Ant species richness on sand was slightly disturbances that lead to extensive habitat loss. CAD can higher than on clay, and was negatively related to have a major effect on vegetation structure (Sagar et al. most surrogates of anthropogenic disturbance. Soil 2003; Shaanker et al. 2004;Ribeiroet al. 2015), and can type and human population size were the main also disrupt key ecological processes such as seed dispersal predictors of ant species richness. Soil type was (Leal et al. 2014) and plant-protection against herbivores the most important predictor of functional group (Leal et al. 2015). Disruptions of such processes cause shifts abundance. Taxonomic and functional composition in the functional composition of trees, from long-lived, were influenced by soil type and disturbance, but this highly specialized species to disturbance-tolerant, short-lived relationship varied between clay and sandy soils. Ant generalized species (Sagar et al. 2003;Uniyalet al. 2010; functional composition showed a weak relationship Ribeiro et al. 2015;Ribeiroet al. 2016). However, there have with disturbance on sandy soils, but on clay soils been few studies of the impact of chronic disturbances on it showed predictable winner–loser replacement. We faunal diversity (Ribeiro-Neto et al. 2016). attribute the greater effect of disturbance on clay soils Ants have attracted considerable attention in disturbance to higher intensity of land use, and our study highlights studies because they are such a globally dominant faunal the importance of considering context dependence group and a popular bio-indicator in land management when evaluating biodiversity responses to disturbance. (Hoffmann & Andersen 2003; Andersen & Majer 2004). Keywords: ant diversity, ant functional groups, biodiversity, CAD can cause major shifts in ant species and functional human disturbance, semi-arid vegetation composition, even with little or no change in overall species richness (Hoffmann 2010; Franklin 2012). Such INTRODUCTION shifts can be directly caused by habitat change, but can also be mediated by changes in the abundance of Anthropogenic disturbance has severely reduced forest cover dominant species, which result in cascading indirect effects worldwide, and such habitat loss and associated fragmentation is widely recognized as the major cause of global biodiversity on subdominant and subordinate groups (Bestelmeyer & Wiens 1996; Hoffmann & Andersen 2003; Andersen & Majer 2004) and on ecological interactions (Leal et al. ∗Correspondence: Inara R. Leal Tel: 55-81-21267814 Fax: 55-81- 2014;Lealet al. 2015). Ant communities show predictable 21268348 e-mail: [email protected] replacement of disturbance ‘losers’ (disturbance-sensitive ∗Supplementary material can be found online at http://dx.doi. taxa) by disturbance ‘winners’ (disturbance-adapted taxa; see org/10.1017/S0376892916000291 Tabarelli et al. 2012), with disturbance typically favouring Downloaded from https://www.cambridge.org/core. University of Athens, on 01 Oct 2021 at 06:14:35, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0376892916000291 116 F. M. P. Oliveira et al. open-adapted taxa at the expense of highly specialized, Measurement of disturbance forest-associated functional groups (Andersen 2000; Hoff- We used the five measurements of disturbance that have been mann & Andersen 2003;Beaumontet al. 2012;Lealet al. previously described for the study plots (Leal et al. 2014), 2012). At a regional scale, disturbance is a secondary driver of based on surrogates related to population density and intensity variation in ant communities that is determined primarily by of land use, which tend to decrease with increasing distance soil type, and ant responses to disturbance can vary markedly from cities and roads (Ahrends et al. 2010). These surrogates with soil type (Hoffmann & Andersen 2003; Hoffmann 2010; were: the number of people (‘people’) and combined number Ribeiro-Neto et al. 2016). of cattle and goats (‘stock’) in the farms within 2 km of each We examined ant community responses to CAD and its plot; the distance to the nearest road (‘distance to road’); interaction with soil type in semi-arid Caatinga of north- the distance to the nearest farm (‘distance to farm’); and the eastern Brazil. We assessed the effects of CAD on both species distance to Parnamirim (‘distance to city’) (Leal et al. 2014; and functional diversity, focusing on three key questions. Ribeiro et al. 2015). People (W = 315, p = 0.824), stock (W First, what is the magnitude of effects of disturbance = 268.5, p = 0.466) and distance to road (W = 289.5, p = compared with those of soil type as the putative primary driver 0.366) did not vary with soil type (Wilcoxon matched-pair of local variation in ant communities? Second, to what extent tests), whereas distance to farm (W = 451, p < 0.001) and do the effects of disturbance vary with soil type? Third, to what distance to city (W = 199, p = 0.050) tended to be lower for extent does functional composition show predictable winner– sites with clay soils. loser replacement involving the loss of highly specialized, forest-associated functional groups? Ant sampling and functional classification MATERIALS AND METHODS Ants were sampled using six pitfall traps (13 cm height and Study site 12 cm diameter) with 10 m spacing in each plot. Traps were Caatinga vegetation consists of mosaics of seasonally dry partly filled with a 90% ethanol solution, and were operated tropical forest (sensu Pennington et al. 2009) and scrub (Velloso for a 48-hr period. Twenty-four of the plots (13 on clay and et al. 2002) covering 826 411 km² of north-eastern Brazil 11 on sand) were sampled from February to May 2010, and (Brasil-MMA 2011). It is one the world’s most biodiverse the remaining 23 (eight on clay and 15 on sand) were sampled and densely populated semi-arid regions, with more than from March to May 2013. All ants were sorted into species 23 million people at a density of 26 inhabitants/km² (INSA and, where possible, named. Unnamed species were assigned 2012). Nearly 10 million m3 year−1 of firewood and charcoal code numbers that apply only to this study. A complete set are obtained via exploitation of native vegetation (Gariglio of mounted voucher specimens is held in the ant collection et al. 2010), and there are more than 16 million goats (IBGE at Universidade Federal de Pernambuco (Pernambuco, 2010). Collectively, slash-and-burn agriculture, firewood Brazil). collection, overgrazing and hunting impose a continuum of For classifying ant species into functional groups we degradation varying from relatively minor biomass reduction adapted Andersen’s (1995) widely used scheme based on to complete desertification (Leal et al. 2005; Brasil-MMA global-scale responses of ants to environmental stress and 2011;Ribeiroet al. 2015). disturbance (see also Andersen 1997; Hoffmann & Andersen Our study was conducted in a continuous patch of Caatinga 2003) to be more specific to the regional fauna. Our groups vegetation near Parnamirim city (8°5’S; 39°34’W; 393 m were: Army Ants (species of Neivamyrmex and Nomamyrmex); asl) in Pernambuco state. The climate is semi-arid, with Dominant Dolichoderinae (primarily Dorymyrmex), Fire Ants an average temperature of 26°C and
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