Effect of road disturbance on ant diversity in a sector of the Central Biocenic Corridor located in the center of Arid diagonal of Argentina

doi:10.21203/rs.3.rs-2206207/v2

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Effect of road disturbance on ant diversity in a sector of the Central Biocenic Corridor located in the center of Arid diagonal of Argentina

https://doi.org/10.21203/rs.3.rs-2206207/v2

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published 03 May, 2023

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Linear infrastructures such as roads are among the most frequent generators of anthropogenic disturbances. Due to the expensive area that is affected along them, these great infrastructures represent a major conservation concern worldwide. Ants are an important component of natural ecosystems and are considered to be very sensitive to disturbance. The National Road Nº 150 is an important road of South America that forms part of the Central Bioceanic Corridor which will connect Brazil with Chile. In its trajectory, it crosses the Ischigualasto Provincial Park in Argentina. In order to evaluate the effects of this road on biodiversity in roadsides that run across this protected area, we compared species and functional group diversity of ants collected using pitfall traps between disturbed and undisturbed sites. In addition, we analyzed whether habitat variables, such as plant cover, gravel cover and soil hardness, explain ant species and functional group abundance in both site types. Twenty-one and 17 ant species were caught in disturbed and undisturbed sites, respectively. Roadside contained relatively more exclusives and rare species, while undisturbed sites harbored more common and dominant species. Hot climate specialists were more abundant at disturbed sites, whereas Opportunists at undisturbed sites. Ant species abundance seems to be positively influenced by gravel cover on roadside. In the present context of land use change, roadside could have an important function as reservoir and corridor for some ant species, and thus, plays a valuable role in the conservation of ant diversity in arid ecosystems.

Biodiversity

road edge effect

conservation

Monte desert

Disturbances lead to important change at various ecological levels of organization, disrupting ecosystem, community or population structure and changing resource, substrate availability, or the physical environment (Pickett et al. 1989). The most common forms of disturbance include herbivory, natural fires and human driven land use change (Lessard 2019). Among last disturbance, linear infrastructures, such as roads, are spread across nearly the entire world surface forming a fine-meshed network. An increase in development leads to an increase in the system of roads, because it facilitates the economic and social development of human populations (Arroyave et al. 2006). However, roads also generate many collateral problems for the conservation of ecosystems (Forman et al. 2002; Song et al. 2005; Delgado et al. 2007) and landscape integrity (Villemey et al. 2018).

Roads produce disturbance with a wide variety of abiotic and biotic effects on the surrounding landscape (Forman and Alexander 1998; Trombulak and Frissell 2000; Rotholz and Mandelik 2013). They transform the physical conditions adjacent to it, creating a new semi-natural habitat (or roadside), with consequences on biota that extend beyond the time of the road’s construction (Forman and Alexander 1998; Trombulak and Frissell 2000). Roadside represents the boundary between two contrasting habitats, road and natural habitat adjacent, over which significant ecological effects can be detected (Murcia 1995). Linear fragments or any fragments with high perimeter-to-area ratios as roadside, are “edge” habitats with features that can reduce their habitat value (Soulé and Gilpin 1991). Roadside habitats have different complexity, micro-climatic conditions, hydrology regimes and soil composition and density, high insolation and high exposure to wind compared to those in the nearby natural area (Delgado et al. 2007; Fruin et al. 2008; Kociolek et al. 2011; Rotholz and Mandelik 2013). As consequences, those greatly affect habitat quality, connectivity, dynamics of populations and communities, resulting effects on biodiversity (Lassau and Hochuli 2003; Gillies and St. Clair 2010; Zurita et al. 2012). Due to the large surface of natural habitat transformed by roads along thousands of kilometers, roadsides represent a major conservation concern worldwide (Forman 2000; Rotholz and Mandelik 2013).

The effect of roads and roadside on insects is varied. Frankel and Soulé (1981) consider that linear strips are likely to provide suitable habitat for this group because, in general, invertebrates have a low area requirement. Others underline the potential of roadside as conduits of gene flow and as reservoirs for many species of woodland invertebrates in highly fragmented landscapes (Zanden et al. 2013). In a recent global review of effects of roadside on this group, Villemey et al. (2018) concluded that abundance was higher on roadside than in surrounding habitats, but insect species richness was similar.

In order to evaluate the ecological effects of roadsides, it is necessary to gain a broad perspective on how communities and functional groups are affected. Among insects, ants (Hymenoptera: Formicidae) because are relatively sedentary and highly sensitive to long-term disturbance even occurring at relatively small scales in space and time (Kremen et al. 1993), change in habitat as those associated to roadsides can affect ant species richness and composition. It is so the ants have attracted considerable attention in disturbance studies because they are sometimes an useful bio-indicator in land management (e.g. Andersen and Majer 2004; Vanthomme et al. 2017). Habitat disturbances as change in structure, microclimatic conditions, resource availability and competitive interactions, are the most important indirect effects of disturbances on ant communities (Lassau and Hochuli 2003; Philpott et al. 2010; Andersen 2019). The road and trail impacts produced by edge effects on ant species diversity are also varied. They increase ant species richness in roadside compared to non-disturbed areas (Sanway et al. 1997; Azcárate et al. 2013; Kaur et al. 2019); roadside conditions improve the reproductive (Noordijk et al. 2009) and colony establishment of ant species (Farji-Brener 2000; Pirk et al. 2004; Vasconcelos et al. 2006; Itzhak 2008). Other studies found negative impacts, because roads and roadsides can create barriers to ants reducing movements (Laurance et al. 2004), decreasing richness species by edge effects and soil disturbances (e.g. Rivas-Arancibia et al. 2014; Palfi et al. 2017), affecting efficacy of seed dispersal mechanism (Parr et al. 2007), and facilitating the spread of invasive species which use the roadsides as invasion corridors (e.g. Walsh et al. 2004; Farji-Brener and Ghermandi 2008). Also, no change in richness and ant assemblages in the non-disturbed compared to disturbed areas of roadside were found (e.g. Beaumont et al. 2012; Pirk and López de Casenave 2017).

Although there are some studies that have evaluated the effects of disturbances caused by road on ants in Argentina, most of them have focused on some species in particular, as Acromyrmex in asphalt road in Patagonian (Farji-Brener 1996; Farji-Brener and Margutti 1997; Farji-Brener and Ghermandi; 2000, 2008), and Pogonomyrmex in dirt road in the Central Monte Desert (Pirk et al. 2004). While others studied on ant–seed interaction in the Patagonian steppe in areas near and far from a road (Pirk and López de Casenave 2017). So far, we do not have knowledge on any study that specifically assesses the consequences of disturbance produced by an asphalt road on the whole ant assemblage. For these reasons, we evaluated the diversity of ant assemblage associated with road disturbance, because they are an important environment modifier agent (Itzhak 2008; Palfi et al. 2017). This is a central concern for conservation of biodiversity and ecosystem management, principally when roads are crossed by protected areas.

The National Road Nº 150 (hereafter RN 150) is an important road of South America that forms part of the Central Bioceanic Corridor. This road will connect the Porto Alegre port in Brazil, on the Atlantic Ocean, with the Coquimbo port in Chile, on the Pacific Ocean, crossing the central sector of Argentina with an extension of 2,500 km. In view of the fact that this road runs across a protected area, the Ischigualasto Provincial Park (hereafter IPP), on its way through the San Juan province in Argentina (Borghi et al. 2012), it is important to assess biota associated with the roadside of this road. In this study we: 1) characterize species and functional group diversity and composition of ground-foraging ant assemblage in disturbed (roadside) and undisturbed (natural) sites, 2) evaluate whether presence and abundance of species and functional groups at the assemblage level respond to disturb caused by the road, and, 3) analyze which habitat variables better explain ant species abundances in both site types. Given that roadside represent opener habitats with lesser cover vegetation, due to recurrent maintenance and operation activities of road, compared to the nearby natural habitats, it has different and challenging micro-climatic conditions (such as high insolation and temperature), and soil with predominate of gravel; we predicted that ant diversity in term of species richness, common and dominant species would be greater in undisturbed natural habitats (natural site) than in disturbed habitat (roadside). Also, we predicted that abundance of arid-adapted ant species would be greater in disturbed habitats, because these species can be favored by the habitat opening on the roadside. Regarding functional composition, we expected that disturbance tolerant species and hot climate specialist species (Andersen 1995; Hoffmann and Andersen 2003) will be the most dominant group in disturbed areas.

Study area

The study was conducted in the IPP (30º 05’S, 67º 55’W) in the province of San Juan, Argentina. This protected area, together with the Talampaya National Park, was declared a World Heritage Site by UNESCO (United Nations Educational, Scientific and Cultural Organization). The Park comprises an area of 62,369 ha, with a mean altitude of 1,300 m a.s.l. and lies in the hyper-arid part of the arid diagonal Argentina, with predominance of the Monte of Mountains and Basins ecoregion. The climate is typically dry desert, with a mean temperature below 18°C within a range from 10°C to 45°C, and with a mean temperature in the warmest month reaching 22°C. Rainfall is concentrated from November to February, a mean annual rainfall of only 100 mm (Cortéz et al. 2005). The vegetation is xerophytic dominated by an open landscape with low plant cover (less than 30%), represented by shrubs (Larrea spp., Atriplex spp.), some trees (Prosopis chilensis, P. flexuosa), and a lower stratum and seasonal herbs (Márquez et al. 2005).

The RN 150 runs across the southern sector of the IPP. This road is 10 m wide of asphalt surface, a gravelly roadside of 4 m wide strip (2 m on each side) which is regularly disturbed by road maintenance activity, where to remove vegetation and to grade activity are carried out once and twice per year. While an adjacent strip 30 m wide (15 m on each side) with soil similar to natural environment is sporadically disturbed by remove vegetation.

Ant sampling

We sampled the ant communities of the study area in February 2013. A total of forty linear transects 50 m long parallel to RN 150 were established along 18 km long. Twenty transects were established at roadside (hereafter disturbed sites) located between 5 and 15 m from the road, and 20 transects in native (hereafter undisturbed sites) separated between 70 to 100 m from the road. Six pitfall traps separated 10 m away from each other were placed along each transect. All transects were separated from each other by 1 km to maintain their spatial independence and avoid pseudo-replication (Silva and Hernández 2015). Transect was taken as a sample unit. Pitfall traps consisting of plastic cups (6 cm diameter and 9 cm in depth) were buried so that its rim was flush with the ground, and was left in the field for 24 h. All traps contained 100 ml of water and ethylene glycol as a preservative, and one drop of detergent to break surface tension (Pérez-Sánchez et al. 2012).

All captured ants were preserved in 96% ethanol for identification. The ants collected were identified by E.A. and L.C. under a dissecting scope. Individuals were identified to genus and species level using available taxonomic keys (Kusnezov 1978; Palacio and Fernández 2003; Bolton 2013 for genus) and photographic material available in Antweb page. Ants were additionally assigned to functional groups following the classification used by Amatta et al. (2018) for the same sites, adapted from that proposed by Bestelmeyer and Wiens (1996) for Argentine Chaco. Voucher specimens were deposited at the ant fauna section of the Entomological collection of the IADIZA-CONICET in Mendoza, Argentina.

Habitat variables

On transects where ants were captured, we established 15 plots in order to register habitat variables. On each plot (50 m along the transect by 10 m wide), we established 5 subsamples (0.5 x 0.5 m) separated by 5 m intervals, where plant cover, gravel cover (both in percentages), and soil hardness using pocket penetrometer S-170 (10 recorded per each subsample) were recorded. The graves were categorized by size: small (< 1 cm); medium (2 to 3 cm); and large (4 to 5 cm). Plot was considered as a sample unit.

Statistical analysis

For the first objective, to quantify the species diversity in disturbed and undisturbed sites, we used Hill numbers (Hill, 1973), which represent the “effective number of species within a specific community” (Jost 2007). They are parameterized by a diversity order q, which determines the measures’ sensitivity to species relative abundances (Chao et al. 2014). For this study we applied the ⁰D, ¹D and ²D metrics, which are parameterized by the Hill numbers 0, 1 and 2, respectively (Jost 2006). When q = 0, the index is equivalent to species richness (⁰D); for q = 1, corresponds to the exponential of Shannon entropy (¹D); and for q = 2, the index corresponds to the reciprocal of the Simpson index (²D). The choice of multiple measures is important to express the complexity of the assemblages and detect the impact on rare (⁰D), common (¹D) and dominant species (²D) (Gotelli and Chao 2013; Chao et al. 2014). In order to examine visually ant species diversity in disturbed and undisturbed sites, and as a tool to evaluate sampling effectiveness, we building two types of integrated rarefaction and extrapolation curves (sample-size and coverage-based), with 95% confidence intervals, obtained by a bootstrap method based on 1,000 replications using iNEXT package (Hsieh et al. 2016). For both curves, in order to standardizing both site types to the same sampling effort (sample coverage 1.00), we used a reference of 6,574 individuals (i.e. the double the smaller reference sample size; Chao and Jost 2012). For comparisons of diversity between undisturbed and disturbed sites, the 95% CI were used at the same coverage level (the minimum one), and no overlap between CI values was indicative of significant differences (Cumming et al. 2007; Cultid-Medina and Escobar 2016). We determine how many times disturbed sites are more or less diverse than undisturbed sites because we expressing diversity as the effective number of species and making comparisons under the same sample coverage, then the replication principle is met and it is possible to calculate the magnitude of the difference in diversity (MD) among communities (Jost 2006). The MD was expressed as a percentage following to Cultid-Medina and Escobar (2016). Comparisons of diversity using 95% CI and use of MD analyses are recommended when effects of anthropic disturbances are assessment (Cultid-Medina and Escobar 2019). Evenness was estimated as ²Dα/⁰Dα (Jost 2010).

For the second and third objectives, we adjusted statistical models using in both cases as response variables, the abundance of each ant species. To evaluate disturbance effect, we included the disturbance situation as an explanatory variable (factor with two levels: disturbed and undisturbed). Rank abundance curves were used to visually understand how assemblages differ in number and relative abundances of species (Feinsinger 2003), and to illustrate differences over space of assemblage that would not be evident by changes in species richness (Avolio et al. 2015). Rank–abundance curves were performed using the Biodiversity R package (Kindt and Coe 2005) and the vegan package (Oksanen et al. 2019).

To assess the effect of habitat variables on abundance of ants (third objective), we used as explanatory variables: plant cover, soil hardness, and gravel cover considering different gravel sizes such as gravel smaller than 1 cm (small gravel), gravel size 2 to 3 cm (medium gravel) and gravel size 4 to 5 cm (large gravel). Prior to the analysis, we evaluate multicollinearity by performing a correlation analysis. We did not exclude variables because the r coefficients were in all cases < |0.7|. In both objectives, the statistical models were Generalized Linear Mixed models (GLMMs) with “transect” as a random effect, and Poisson error distribution (Crawley 2007); when error exhibited overdispersion (ĉ = 1.00), we adjusted a Negative Binomial error distribution. GLMMs were fitted using the glmer function from the lme4 package (Fox and Weisberg 2019), and the glmmadmb function from the glmmADMB package (Fournier et al. 2012). In each model, we used a backward elimination procedure to remove insignificant variables without losing important information. The significance of the factors was evaluated using LRT-test (Zeileis and Hothorn 2002). Results are presented as mean ± standard error (SE), and for null hypothesis, testing statistical tests were considered significant at α ≤ 0.05. All analyses were performed with R Software 3.6.1 (R Development 2019).

Species diversity and composition

A total of 8,872 ants were collected at disturbed and undisturbed sites, with an accumulated sampling effort of 5,760 hours (240 pitfall traps placed x 24 hours each one). Captured ants belong to four subfamilies, 10 genera and 21 species. The subfamilies richest in species were Myrmicinae (8 species) and Dolichoderinae (7 species), followed by Formicinae (5 species), and Dorylinae with only one species, whereas the richest genera were Camponotus and Forelius with four species each one (Table 1). A total of 5,585 workers belonging to 21 species were recorded at disturbed sites whereas 3,287 individuals of 17 species were found at undisturbed sites. Five of 21 species (Dorymyrmex exsanguis, Forelius nigriventris, Forelius rufus, Neivamyrmex diana and Crematogaster quadriformis) were caught exclusively in disturbed sites, while only one (Camponotus substitutus) of 17 species in undisturbed sites. Species that contributed more to dissimilarity between site types and counting with two or three time more workers were: Dorymyrmex planidens and Camponotus punctulatus in undisturbed sites, and Acromyrmex lobicornis and Pogonomyrmex brevibarbis in the disturbed sites. Thirteen species were the most numerically dominant in both site types, and they contributed with 98.2% and 99.5% of individuals captured in undisturbed and disturbed sites, respectively.

Table 1

Comparison of ant abundance and functional group recorded in disturbed and undisturbed sites across the RN 150 in the sector that runs through the IPP. LRTest was used to compare abundances between disturbed and undisturbed sites as worker number (mean ± ES) of each species ants and of each functional group (mean ± ES)
	Disturbed	Undisturbed	LRTtest
	Abundance (mean ± ES)		χ2	p
Subfamily Dolichoderinae
Dorymyrmex ensifer ^(Hcs)	10.2 ± 6.3	7.15 ± 2.22	0.28	0.59
Dorymyrmex exsanguis ^(Hcs)	0.5 ± 0.5	Absent	---
Dorymymex planidens ^(Hcs)	15.8 ± 8.4	18.4 ± 15.2	0.01	0.91
Forelius albiventris ^(Hcs)	30.3 ± 5.6	15.3 ± 3.9	3.27	0.07
Forelius chalybaeus ^(Hcs)	6.4 ± 2.9	6.1 ± 5.1	0.001	0.96
Forelius nigriventris ^(Hcs)	0.1 ± 0.4	Absent	---
Forelius rufus ^(Hcs)	0.3 ± 1.1	Absent	---
Subfamily Dorylinae
Neivamyrmex diana ^(Aa)	0.05 ± 0.05	Absent	---
Subfamily Formicinae
Brachymyrmex patagonicus ^(Op)	0.2 ± 0.1	0.7 ± 0.7	7.72	0.005
Camponotus mus ^(Sc)	0.2 ± 0.1	0.2 ± 0.2	0.14	0.7
Camponotus sp. ^(Sc)	9.8 ± 4.6	6.2 ± 2.8	0.36	0.54
Camponotus punctulatus ^(Sc)	4.7 ± 2.4	5.6 ± 2.3	0.07	0.77
Camponotus substitutus ^(Sc)	Absent	0.5 ± 1.6	---
Subfamily Myrmicinae
Acromyrmex lobicornis ^(At)	18.1 ± 13.2	3.9 ± 1.9	2.58	0.1
Acromyrmex striatus ^(At)	9.9 ± 3.6	3.9 ± 1.9	0.002	0.96
Crematogaster quadriformis ^(Gm)	0.5 ± 2.2	Absent	---
Pheidole bergi ^(Gm)	55.3 ± 19.9	22.7 ± 5.9	1.73	0.18
Pheidole triconstricta ^(Gm)	11.1 ± 4.7	8.6 ± 5.1	0.07	0.79
Pogonomyrmex brevibarbis ^(Hcs)	58.8 ± 12.3	18.0 ± 5.3	5.77	0.01
Pogonomyrmex cunicularius ^(Hcs)	36.3 ± 5.8	35.8 ± 4.1	0.003	0.95
Solenopsis sp. ^(Cr)	9.9 ± 4.6	4.2 ± 2.9	0.90	0.34
Indeterminate	1.0 ± 0.7	1.5 ± 0.9	2.39	0.12
Functional group
Army ants	0.05 ± 0.05	Absent	---
Attini	27.9 ± 13.9	13.4 ± 5.2	1.60	0.20
Cryptic	9.9 ± 4.6	4.2 ± 2.9	0.90	0.34
Generalized myrmicines	69.3 ± 27.9	33.5 ± 12.2	310.4	0.26
Hot climate specialists	158.8 ± 19.5	100.8 ± 17.7	6.59	0.01
Opportunists	0.2 ± 0.1	0.7 ± 0.6	7.72	0.005
Subordinate camponotini	14.6 ± 4.9	12.4 ± 3.2	0.13	0.71
^{Aa: Army ants, Hcs: Hot climate specialists, Gm: Generalized myrmicines, Op: Opportunists, At: Attini, Sc: Subordinate camponotini, and Cr: Cryptic}
^{Significant p−values are given in bold}

Using the number of individuals, the observed effective species richness (⁰D_(N)) was higher in disturbed sites, while the effective number of common species (¹D_(N)) and dominant species (²D_(N)) were significantly higher in undisturbed than disturbed sites (Table 2). Regarding species richness, the values observed were similar to those estimated for undisturbed and disturbed sites (Table 2). Diversity profiles were quite similar for undisturbed and disturbed sites, with decreasing scores from ⁰D_(N) towards ²D_(N) (Table 2, Fig. 1a). The completeness of the sampling was high in both situations, it reached 99.9% for disturbed, and 100% for undisturbed sites, indicating an effective sampling (i.e. all species expected to occur in each site type were effectively captured) for study area (Fig. 1b). When comparing the MD in effective species richness ⁰D_(N), effective number of common species (¹D_(N)) and the dominant species (²D_(N)) between disturbed and undisturbed sites, regardless of whether there are statistically significant differences, the disturbed sites were 1.23 times more diverse in term of effective species richness than the undisturbed sites (Table 2). Undisturbed and disturbed sites had a relatively similar effective number of common (¹D_(N)) and dominant species (²D_(N)), although disturbed sites was 0.9 and 0.89 times less diverse than undisturbed sites respectively. The undisturbed was a more equitable community (evenness factor = 0.52) than disturbed sites (evenness factor = 0.38). Disturbed sites contributed to the presence of uncommon species, singleton species (species represented by one individual) and doubleton (species represented by two individuals) with one species each one only registered in this site type.

Table 2

Observed diversities and the estimated asymptotes of diversities at disturbed and undisturbed sites in the RN 150 in the IPP in the sector that runs through the protected area
	Observed parameter	Estimated asymptote	Estimated SE	95% confidence interval
	Observed parameter	Estimated asymptote	Estimated SE	lower	upper
Disturbed sites
Species richness (⁰D_N)	21.0	21.5	1.32	21.03	29.44
Shannon diversity (¹D_N)	10.1	10.1	0.11	10.12	10.35
Simpson diversity (²D_N)	7.9	7.9	0.11	7.86	8.10
Species richness (⁰D_FG)	7.0	7.0	0.52	7.00	8.54
Shannon diversity (¹D_FG)	2.9	2.9	0.03	2.96	3.02
Simpson diversity (²D_FG)	2.4	2.4	0.03	2.42	2.49
Undisturbed sites
Species richness (⁰D_N)	17.0	17.0	0.13	17.00	17.00
Shannon diversity (¹D_N)	11.1	11.1	0.15	11.07	11.38
Simpson diversity (²D_N)	8.8	8.9	0.18	8.84	9.26
Species richness (⁰D_FG)	6.0	6.0	0.00	6.00	6.00
Shannon diversity (¹D_FG)	2.9	2.9	0.00	2.99	3.09
Simpson diversity (²D_FG)	2.3	2.3	0.04	2.29	2.38
^{Number of ants (N), functional group (FG), species richness (0D), diversity weighted by number of ants (qDN), and ±IC 95% for each qD}

Functional groups

Seven functional groups were identified, six of them were collected at both sites, but army ant group was exclusive at disturbed sites, although in low abundance (Table 1). The community was dominated by ant species belonging to Hot climate specialist (43%), followed by Subordinate camponotini (19%), Generalized myrmicines (14%), Attini (9%), and Opportunists, Cryptic and Army ants (5% each one). In the comparison of functional groups, effective functional group richness (⁰D_(FG)) and number of dominant functional groups (²D_(FG)) were significantly lesser in undisturbed than in disturbed sites, while did not statistical difference in effective number of common functional groups (¹D_(FG)) (Table 2). Comparing the MD in effective functional groups richness (⁰D_(FG)) and number of dominant functional groups (²D_(FG)), the disturbed sites were slightly more diverse than undisturbed sites (1.16 and 1.04 times, respectively). While diversity considering the effective number of common functional groups (1D_(FG)) was similar between disturbed and undisturbed sites. The disturbed sites were slightly lesser evenness compared to undisturbed sites (0.3 and 0.4, respectively

Road effect on species and functional group abundances

Numerically, the ant assemblage responds positively to disturbance caused by the road. Total abundance was significantly higher in disturbed sites (mean number of ants per transect 279.25 ± 42.71) than in undisturbed sites (164.35 ± 20.66 ants/transect; GLMM LRT-test χ2 = 6.49; p = 0.001). All exclusive species were registered with low abundance (they were rare species with ≤ 10 individuals). Slight difference was found between disturbed and undisturbed sites when abundance of each particular ant species was compared (Table 1); such as Brachymyrmex patagonicus that was significantly more abundant at undisturbed sites, and Pogonomyrmex brevibarbis and Forelius albiventris that were more abundant at disturbed sites; although for this last species the relationship was only marginally significant (Table 1). Inspection of the species abundance curves shows P. brevibarbis, Pheidole bergi and F. albiventris were 3.2, 2.4 and 2 times more abundant, respectively, in the disturbed than in undisturbed sites. Although Acromyrmex lobicornis was not as abundant in any of the sites, in the disturbed site its abundance was almost 4.5 times greater. Pogonomyrmex cunicularius was the most numerically dominant species in the undisturbed sites (Fig. 2).

Disturbed sites registered a higher number of ants in the majority of the functional groups than undisturbed sites (mean number ± SE of workers, 278.25 ± 42.85 and 162.80 ± 20.94, respectively; GLM, LRTtest χ2 = 2.93; p = 0.003). Hot climate specialists and Opportunists were the only functional groups whose abundance varied significantly between disturbed and undisturbed sites. Hot climate specialists were significantly more abundant at disturbed sites, whereas Opportunists at undisturbed sites. The other functional groups, Attini, Cryptic, Generalized myrmicines and Subordinate camponotini did not show a differential response to disturbance.

Effect habitat variables on abundance

Disturbed and undisturbed sites did not show differences for soil hardness and cover of large gravel (4 to 5 cm). However, undisturbed sites had the highest percentage plant cover, and a greater cover of combined medium and large gravel (> 2 to 5 cm); while the cover of small gravel (< 1 cm) was the highest in disturbed sites (Table 3). Plant species occurring at disturbed and undisturbed sites were overall similar, with the exception of some exclusive plant species for each site type (Table 4). Shrub was the life form predominant in both disturbed and undisturbed sites. One exotic plant species was recorded at the roadside (Portulaca oleracea), none in undisturbed site (Table 4).

In disturbed sites, the 52% of total ant abundance was explained by cover of different sizes of gravel (large, p < 0.001; medium, p < 0.001; and small, p = 0.01 gravel) (Marginal R_GLMM²), while the 98% was explained by these fixed variables (large gravel, p < 0.001; medium, p = 0.0003; and small, p = 0.0004), plus the random effect “transect” (Conditional R_GLMM²). All the variables had a positive effect, increasing abundance with increasing coverage of each size of gravel: large (estimate: 0.04 ± 0.01); medium (estimate: 0.07 ± 0.008), and small (estimate: 0.03 ± 0.01) gravel. The same fixed variables explained only a 24% (Marginal R_GLMM²) of total ant abundance recorded in undisturbed sites, while the combined effect of the fixed and random variables explained 98% of the variation (Conditional R_GLMM²). In these sites, habitat variables also had a positive effect on abundance: cover of large (estimate: 0.05 ± 0.007); medium (estimate: 0.03 ± 0.01), and small (estimate: 0.08 ± 0.02) gravel. The “transect” also had a significant effect on ant abundance in both disturbed and undisturbed sites (p < 0.0001).

Table 3

Comparison of habitat variables (mean ± SE) in disturbed and undisturbed sites in the NR 150 in the IPP, using a Generalized Linear Mixed Model (GLMM). Site was considered a random factor
Habitat variables	Disturbed sites	Undisturbed sites	LRT test	p
Soil hardness	3.49 ± 0.18	3.07 ± 0.27	1.72	0.18
Gravel < 1cm	65.76 ± 2.47	55.87 ± 5.52	4.24	0.04
Gravel (2–3 cm)	25.13 ± 3.03	32.09 ± 3.87	4.29	0.03
Gravel (4–5 cm)	9.11 ± 1.97	12.3 ± 2.5	1.09	0.29
Plant Cover	16.33 ± 3.0	25.66 ± 3.13	4.59	0.04

Table 4

Plant species recorded in disturbed and undisturbed sites in National Road N° 150, in the section that runs across Ischigualasto Provincial Park
Species	Undisturbed	Disturbed	Habit
Allenrolfea vaginata	X	-	Shrub
Allionia incarnata	X	-	Herb
Atriplex sp.	X	X	Shrub
Baccharis salicifolia	X	X	Shrub
Bulnesia retama	X	X	Tree/Shrub
Capparis atamisquea	-	X	Shrub
Cercidium praecox	X	X	Tree
Conyza bonariensis	-	X	Herb
Cyclolepis genistoides	X	-	Shrub
Cynodon dactylon	-	X	Grass
Deinacanthon urbanianum	X	-	Herb
Euploca mendocina	-	X	Herb
Geoffroea decorticans	X	X	Tree
Gymnocalycium castellanosii	-	X	Succulent
Heliotropium amplexicaule	-	X	Herb
Justicia lilloana	X	-	Shrub
Larrea cuneifolia	X	X	Shrub
L. divaricata	X	X	Shrub
Leptochloa crinita	X	X	Grass
Lycium boerhaviaefolium	X	X	Shrub
Lycium sp.	X	X	Shrub
Maytenus viscifolia	X	X	Shrub
Monttea aphylla	-	X	Shrub
Nicotiana glauca	-	X	Shrub
N. paa	-	X	Herb
Portulaca grandiflora	-	X	Herb
P. oleracea	-	X	Herb
Prosopis chilensis	X	X	Tree
P. flexuosa	X	X	Tree
P. torquata	X	X	Shrub
Salvia gilliesii	X	-	Herb
Sclerophylax kurtzii	X	X	Herb
Senna aphylla	X	X	Shrub
Senecio subulatus	X	X	Shrub
Solanum elaeagnifolium	X	-	Herb
Stipa speciosa	-	X	Grass
Tephrocactus aoracanthus	X	X	Succulent
Zuccagnia punctata	X	-	Shrub
Litter	X	X

Our results suggest that the roadside might have significant effects on ant assemblages. The effective species richness and the total abundance of ants have been positively affected by roads, as both measures were higher in disturbed sites. Though the higher total ant abundance in roadsides was due to the greatest contribution of the more common and dominant species, disturbed sites have rarer and exclusive species than undisturbed natural areas.

Tshiguvho et al. (1999) found that road borders were the more ant species-rich, and contained rarer species. In our case, disturbance may be responsible for the presence of uncommon species (singleton and doubleton) that was exclusive to the roadside. Thus, some fewer common species could be using roadside as refuge or as corridors to move to other areas, similarly as it was argued in others studies (Melis et al. 2010; Villemey et al. 2018). Then, roadside habitats might provide ants with suitable resources or environment conditions allowing the coexistence of more species than natural areas (Ribas et al. 2003).

It is important to mention that the species richness is overall a few informative measures to detect disturbance effects on ant assemblages, except when it is severe (Andersen and Majer 2004; Vanthomme et al. 2017). Assessments therefore typically focus on changes in species composition and abundance (Hoffmann and Andersen 2003; Calcaterra et al. 2010; de Castro Solar et al. 2016). In our study, when other metrics of diversity that include abundance were considered (¹D and ²D), the diversity calculated from the number of individuals per species revealed an increase in inequality in disturbed sites. However, this tendency was less marked when using the number of individuals per functional group, which can suggest that community function is maintained.

Habitat disturbance usually causes an increasing dominance by a few species in the ant assemblage (Andersen 1990; Majer et al. 1994). These species profit from disturbance generated by roads. The disparity observed in the roadside was mostly caused by the dominance of Pogonomyrmex brevibarbis, and probably by Forelius albiventris and Pheidole bergi. These species are considered generalists (Holldobler and Wilson 1990; Andersen 2000), and, as reported in others studies, they would be less sensitive to disturbance than the specialist ones (Leal et al. 2012; Pereyra et al. 2019). Positive association with roadsides was found in other Pogonomyrmex species. For instance, P. occidentalis occupies protected ditches in bare soil at the edge of roads (DeMers 1993), and their colonies occur in high density adjacent to linear disturbances, as dirt road and trail (Terranella et al. 1999). In the central Monte desert of Argentina, Pogonomyrmex pronotalis occurs in higher density in dirt roads probably because it finds favorable sites for nesting and establishment its colonies, as high bare soil cover, scarce litter, and more suitable temperatures (Pirk et al. 2004). Acromyrmex lobicornis, other species associated with roadsides, is considered a good colonizer of disturbed areas by its ability to repair mounds and relocate nests (Kusnezov 1978; Farji-Brener 2000). This Neotropical leaf-cutting ant species belonging to the group generalist herbivores, which proliferate in or benefit from human-disturbed habitats (Estes et al. 2011; Leal et al. 2014). Several studies report an increased density of leaf-cutting ants in disturbed areas with low vegetation cover generated by anthropogenic activities, as roads (Vasconcelos et al. 2006; Vieira-Neto et al. 2016; Siqueira et al. 2017).

A clear increase in the Hot-climate species group was observed in disturbed sites (roadside), while reverse was found in undisturbed sites with more occurrence of Opportunists group. Also, Generalized myrmicines showed a clear tendency to increase in roadside, but the difference was not statistically significant. The disturbed sites with low vegetation cover favored the occurrence of ant species characteristics of open habitats (e.g. Pheidole bergi). Indeed, species that showed a clear increase in roadside, belong to functional groups associated with open environments and disturbed areas, such as Hot-climate species group (Pogonomyrmex brevibarbis) and Generalized myrmicines (P. bergi), respectively. Hot-climate group comprises well arid-adapted species categorized as thermophilics with morphological, physiological and behavioral specializations (Bestelmeyer 2000). These attributes allow them to withstand high temperatures and low humidity, and to use open environments with scarce or low vegetation cover as roadside. The species included within the functional group Generalized myrmicines are characterized by world-wide distributions, relatively flexible habitat requirements, tolerance to high temperature, and ability to occupy different niches, such as they are highly efficient exploiting new resources. This functional group tends to predominate in environments experiencing moderate levels of disturbance (Andersen and Major 2004). These species can benefit from changing environments, such as roadsides, so they can use sides of roads as additional habitats and/or dispersal corridors (Vanthomme et al. 2017). On the other hand, species with opportunistic behavioral attributes and presumably poor competitive abilities, such as Brachymyrmex patagonicus, belong to the Opportunists group could take advantage of exploiting undisturbed sites, where more dominant species, such as Hot climate species are not so abundant (Andersen 1995). This species occurs naturally in most of Argentina, but in Chaco, a semiarid ecoregion (Pereyra et al. 2019), it is always associated with forest, as in the IPP (Amatta et al. 2018). Probably the harsh micro-environmental conditions of the disturbed areas do not allow it to inhabit the roadside. Occurrence of rare species exclusively captured in roadsides can be explain by attributes of Hot-climate group (Dorymyrmex exsanguis, Forelius nigriventris and F. rufus) and Generalized myrmicines (Crematogaster quadriformis). Camponotus belongs to the Subordinate camponotini group, which is known to be subdominant in disturbed areas (Hölldobler and Wilson, 1990). In this study, Camponotus species were scarce; C. punctulatus was twice higher in undisturbed sites, while C. substitutus was absent in roadside sites. These results agree with the idea of a negative association among Camponotus species and disturbed environments as roadsides. Surprisingly, Neivamyrmex diana, a species belongs to Army ant functional group, was captured exclusively on the roadside, although in very low abundance. The presence of this species could be explained by its nomadic behavior (Brady 2003). While the dearth of army ants in our samples reflects the fact that this group is primarily found in more structurally heterogeneous physiognomies, such as forests (Amatta et al. 2018), and are usually less diverse and abundant in open areas (Baccaro et al. 2015).

In arid or semi-arid regions, roadside may have a great impact on biodiversity, because they augment the available water resources in adjacent strips by rain run-off along roads and change in soil texture (Forman and Alexander 1998). In this study, the gravel cover was an explanatory variable of abundance of ants on both site types, but this effect was nearly twice stronger in disturbed sites than in undisturbed sites. Gravelly of roadside can produce changes in local microclimates by providing a substrate with high moisture content and well-drained site, suitable for ants adapted to desert environments to build their nests (Ettershank 1971) and foraging (Schumacher and Whitford 1976). For instance, soils very rich in clay and with little sand, harbor low numbers of ants (DeMers 1993). Pogonomyrmex species and some Pheidole species clear vegetation from around nest entrances and collect small stones to form a mound at the nest entrance (Ettershank 1971). Hypotheses to explain such behavior emphasize water conservation of desert ants (Ettershank 1971). Furthermore, since disturbed sites have a higher gravel cover, this substrate may represent a more heterogeneous habitat for ants, which could explain the higher richness, abundance and occurrence of exclusive species in roadside than in the natural areas (Ribas et al. 2003).

Two plant species exclusively registered on disturbed sites support our suggestion of changes in condition of substrate of roadside. Nicotiana paa is a native ruderal plant species, which selects disturbed of coarse-grained soil (Scarpa and Rosso 2011). Portulaca oleracea, is an exotic species abundant in warm and moist regions of the northern hemisphere (Chaube et al. 2014). This species was also found on the roadsides of other desert areas (Abd El-Ghani et al. 2013), and it shows tolerance to various metallic stresses in the roadside soil (Subpiramaniyam 2021). Also, roadside can offer a higher availability of food. In desert ant communities, where food is limited (Whitford 2002), an increased food availability can have a strong influence on abundance and diversity of ant species (Wynhoff et al. 2011). Road-kills animals, which are foraging targets for ants, may cause higher species richness and abundance on a roadside (Lassau and Hochuli 2003; Wynhoff et al. 2011). Although the availability of food was not quantified in present study, several times road-killed animals and carcasses on the roadside were found by us. Also, species such as guanaco (Lama guanicoe) and highly abundant domestic animals (cow and donkey) frequently use the roadside in this protected area (Cappa et al. 2020). This increases food availability not only by road-kills, but also by the feces and seeds dispersed through endozoochory (Campos et al. 2008). How changes in ant communities could happen by multiple variables, more studies are needed to confirm that the changes observed in the composition and structure of assemblage are effectively attributable to disturbance caused by the construction and maintenance of the road.

Even though the greatest impacts on flora and fauna is produced during the construction of the road, the resulting impacts of the operation and maintenance remain in the time (Muñoz et al. 2015), producing the so called chronic anthropogenic disturbances (sensu Singh 1998). Our findings indicated that changes of biotic and abiotic variables at roadsides, such as reduced plant cover and high gravel cover of roadside, increase species richness and abundance and promote the presence of rare species. Our results seem to indicate that the current management regime represents a chronic anthropogenic disturbance that produces only moderate disturbance for the ants, likely encourages niche diversification and co-existing more species (MacArthur 1972). A moderate disturbance level is expected in desert areas, such as disturbances have a greater impact on closed habitats than on open ones in correlation with the availability of annual precipitation, given that plant cover and structural complexity tend to increase with precipitation (Andersen 2019). According to Keals and Majer (1991), roadside can support many species of ant, particularly where there is native vegetation, and if it is wide enough to provide microhabitats for many species. Then the NR 150 seems to be appropriately wide to harbor a high richness of ant species.

In conclusion, roadsides with very little management as is the case of our study site seem to function as linear protected areas that support a higher richness and abundance of ant species than their adjacent natural areas. Though the higher ant abundance in roadsides corresponded to the most common and dominant species (which decreases equitability), roadsides allowed rare and exclusive species to persist in the harsh desert conditions. Thus, our results have important implications for the conservation of invertebrates, the provision of food for insectivorous species, and for the assessment of roadside environmental quality. Especially because this road will soon increase its traffic across the entire central area of Argentina, including the southern sector of a protected area (IPP). Fortunately, the current management of its roadsides would not compromise the conservation of, at least, their associated ant species.

Conflict of interest

The authors declare that there were no conflicts of interests for any of the authors regarding this study.

Acknowledgments

We thank the staff of Ischigualasto Provincial Park for providing all the necessary facilities during fieldwork. We are also grateful to INTERBIODES members.

Funding

This work was supported by Presidencia de la Nación Argentina and Gobierno de San Juan (Project Native Forest 2011–2021; Resolution N° 0441, SEAyDS).

Contributions

GSM conceived this research idea; study design was performed by GSM, AE and CLA; AE and GSM collected the field data; AE and CLA identified the ant specimens; AE and AN performed and written statistical analyses; the first draft of the manuscript was written by GSM and AE and all authors commented on previous versions of the manuscript, read, discussed the results, commented and approved the manuscript.

Abd El-Ghani MM, Salama FM, El-Tayeh NA (2013). Desert roadside vegetation in eastern Egypt and environmental determinants for its distribution. Phytologia Balcanica 192: 233-242.
Amatta E, Calcaterra LA, Giannoni SM (2018). Ant species (Hymenoptera: Formicidae) in the three forests of the Ischigualasto Provincial Park, a protected area of the Monte Desert, Argentina. Biodiversitas, Journal of Biological Diversity 19 (3): 831-839. https://doi.org/10.13057/biodiv/d190311
Andersen AN (1990). The use of ant communities to eval- uate change in Australian terrestrial ecosystems: a review and a recipe. Proceedings of the Ecological Society of Australia. 16: 347-357.
Andersen AN (1995). A classification of Australian ant communities, based on functional groups which parallel plant life-forms in relation to stress and disturbance. Journal of Biogeography 22: 15-29. https://doi.org/10.2307/2846070
Andersen AN (2019). Responses of ant communities to disturbance: Five principles for understanding the disturbance dynamics of a globally dominant faunal group. Journal of Animal Ecology 88(3): 350-362. https://doi.org/10.1111/1365-2656.12907
Andersen AN (2000). A global ecology of rain forest ants: functional groups in relation to stress and disturbance. In Agosti D, Majer JD, Alonso L, Shultz T, eds. Ants: standard methods for measuring and monitoring biodiversity. Smithsonian Institution Press, Washington DC, USA, pp. 25-34.
Andersen AN, Majer JD (2004). Ants show the way down under: invertebrates as bioindicators in land management. Frontiers in Ecology and the Environment 2: 291-298. https://doi.org/10.1890/1540-9295(2004)002%5B0292:ASTWDU%5D2.0.CO;2
Arroyave MD, Gómez C, Gutiérrez ME, Múnera DP, Zapata PA, Vergara IC, Andrade LM, Ramos KC (2006). Impactos de las carreteras sobre la fauna silvestre y sus principales medidas de manejo. Revista EIA 5: 45-57.
Avolio ML, La Pierre KJ, Houseman G, Koerner SE, Grman E, Isbell F, Johnson DS, Wilcox KR (2015). A framework for quantifying the magnitude and variability of community responses to global change drivers. Ecosphere 6: art280. https://doi.org/10.1890/ES15-00317.1
Azcárate FM, Seoane J, Castro S, Peco B (2013). Drove roads: Keystone structures that promote ant diversity in Mediterranean forest landscapes. Acta Oecologica 49: 107-115. http://dx.doi.org/10.1016/j.actao.2013.03.011
Baccaro FB, Feitosa RM, Fernandez F, Fernandes IO, Izzo TJ, Souza JP, Solar R (2015). Guia para os gêneros de formigas do Brasil. Manaus Editora INPA, Brazil.
Beaumont KP, Mackay DA, Whalen MA (2012) The effects of pre- scribed burning on epigaeic ant communities in eucalypt forest of South Australia. Forest Ecology and Management 271: 147-157. http://dx.doi.org/10.1016/j. foreco.2012.02.007
Bestelmeyer BT (2000). The trade-off between the thermal tolerance and behavioural dominance in a subtropical South American ant community. Journal of Animal Ecology 69: 998-1009. http://dx.doi.org/10.1046/j.1365-2656.2000.00455.x
Bestelmeyer BT, Wiens JA (1996). The effects of land use on the structure of ground-foraging ant communities in the Argentine Chaco. Ecological Applications 6: 1225-1240. http://dx.doi.org/10.2307/2269603
Bolton B (2013). AntWeb: ants of Bolton. World Catalog. www.antweb.org. Consultado el 20 de abril de 2022.
Borghi CE, Campos CM, Ortuño N, Beninato V, Andino N, Campos V, et al. (2012). Efeitos indiretos sobre a fauna do corredor bioceánico central em uma área protegida do deserto do Monte in P. P. Ischigualasto. In: A. Bager (Eds.), Ecologia de estradas: Tendencias e pesquisas Lavras. Universidade Federal de Lavras, Brazil, pp. 237-252.
Brady SG (2003). Evolution of the army ant syndrome: the origin and long-term evolutionary stasis of a complex of behavioral and reproductive adaptations. Proceedings of the National Academy of Sciences of the United States of America 100: 6575-6579. http://dx.doi.org/10.1073/pnas.1137809100
Calcaterra L, Cuezzo F, Cabrera S, Briano J (2010). Ground ant diversity (Hymenoptera: Formicidae) in the Iberá Nature Reserve, the largest wetland of Argentina. Annals of the Entomological Society of America 103(1): 71–83. http://dx.doi.org/10.1603/008.103.0110
Campos CM, Peco B, Campos VE, Malo JE, Giannoni SM, Suárez F (2008). Endozoochory by native and exotic herbivores in dry areas: consequences for germination and survival of Prosopis seeds. Seed Science Research, 18(2): 91-100. http://dx.doi.org/10.1017/S0960258508940344
Cappa FM, Giannoni SM, Ontiveros Y, Borghi CE (2020). Direct and indirect effects of roads on activity patterns of the largest South American artiodactyl (Lama guanicoe) in a hyper-arid landscape. Mammalian Biology 100(5): 453-461. http://dx.doi.org/10.1007/s42991-020-00035-9
Chao A, Jost L (2012). Coverage‐based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93(12): 2533-2547. http://dx.doi.org/10.1890/11-1952.1
Chao A, Chiu CH, Jost L (2014). Unifying species diversity, phylogenetic diversity, functional diversity, and related similarity and differentiation measures through Hill numbers. Annual review of ecology, evolution, and systematics 45: 297-324. http://dx.doi.org/10.1146/annurev-ecolsys-120213-091540
Chaube S, Mall S, Upadhaya PP, Rao GP (2014). Natural occurrence of 16SrI group phytoplasmas on purslane (Portulaca oleracea L.), a medicinal weed species in India. Medicinal Plants-International Journal of Phytomedicines and Related Industries 6(2): 150-153
Cortéz E, Borghi CE, Giannoni SM (2005). Plan de Manejo Parque Provincial Ischigualasto Fase I. San Juan, Argentina: Ente Autárquico Ischigualasto, Argentina: Gobierno de San Juan.
Crawley MJ (2007). John Wiley and Sons Ltd. Chichester, UK, pp 942.
Cultid-Medina CA, Escobar F (2016). Assessing the ecological response of dung beetles in an agricultural landscape using number of individuals and biomass in diversity measures. Environmental Entomology 46: 310-319. http://dx.doi.org/10.1093/ee/nvv219
Cultid-Medina CA Escobar F (2019). Pautas para la estimación y comparación estadística de la diversidad biológica (^qD). In: Moreno CE (Ed) La biodiversidad en un mundo cambiante: Fundamentos teóricos y metodológicos para su estudio. Universidad Autónoma del Estado de Hidalgo/ Libermex, Ciudad de México, pp. 175-202. http://dx.doi.org/10.1093/ee/nvv219
Cumming G, Fidler F, Vaux DL (2007). Error bars in experimental biology. The Journal of Cell Biology 177(1): 7-11. https://doi.org/10.1083/jcb.200611141
De Castro Solar RR, Barlow J, Andersen AN, Schoereder JH, Berenguer E, Ferreira JN, Gardner TA (2016). Biodiversity consequences of land-use change and forest disturbance in the Amazon: a multi-scale assessment using ant communities. Biological Conservation 197: 98-107. https://doi.org/10.1016/j.biocon.2016.03.005
Delgado JD, Arroyo NL, Arévalo JR, Fernández-Palacios JM (2007). Edge effects of roads on temperature, light, canopy cover, and canopy height in laurel and pine forests (Tenerife, Canary Islands). Landscape and Urban planning 81(4): 328-340. https://doi.org/10.1016/j.landurbplan.2007.01.005
DeMers M (1993). Roadside ditches as corridors for range expansion of the western harvester ant (Pogonomynnex occidentales Cresson). Landscape Ecology 8: 93-102. https://doi.org/10.1007/BF00141589
Estes JA, Terborgh J, Brashares JS, et al. (2011). Trophic downgrading of planet Earth. Science 333(6040): 301-306. https://doi.org/10.1126/science.1205106
Ettershank G (1971). Some aspects of the ecology and nest microclimatology of the meat ant, Iridomyrmex purpureus (Sm.). Proceedings of the Royal Society of Australia 84: 137-151.
Farji-Brener AG (1996). Posibles vías de expansión de la hormiga cortadora de hojas Acromyrmex lobicornis hacia la Patagonia. Ecología Austral 6: 144-150.
Farji-Brener AG (2000). Leaf-cutting ant nests in temperate environments: mounds, mound damages and nest mortality rate in Acromyrmex lobicornis. Studies on Neotropical Fauna and Environment 35(2): 131-138. https://doi.org/10.1076/0165-0521(200008)35:2;1-9;FT131
Farji-Brener AG, Margutti L (1997). Patterns of plant species in relation to Acromyrmex lobicornis nest-mounds on roadside vegetation in northwest Patagonia. International Journal of Ecology and Environment Sciences 23: 37-47.
Farji-Brener AG, Ghermandi L (2000). The influence of nests of leaf-cutting ants on plant species diversity in road verges of northern Patagonia. Journal of Vegetation Science 11:453-460. https://doi.org/10.2307/3236638
Farji-Brener AG, Ghermandi L (2008). Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas. Proceedings of the Royal Society B: Biological Sciences 275(1641): 1431-1440. https://doi.org/10.1098/rspb.2008.0154
Feinsinger P (2003). El diseño de estudios de campo para la conservación de la biodiversidad. Bolivia, Editorial FAN.
Forman RT, Alexander LE (1998). Roads and their major ecological effects. Annual Review of Ecology and Systematics 29: 207-31. https://doi.org/10.1146/annurev.ecolsys.29.1.207
Forman RT (2000). Estimate of the area affected ecologically by the road system in the United States. Conservation Biology 14: 31-35 https://doi.org/10.1046/j.1523-1739.2000.99299.x
Forman RT, Reineking B, Hersperger AM (2002). Road traffic and nearby grassland bird patterns in a suburbanizing landscape. Environmental management 29(6): 782-800. https://doi.org/10.1007/s00267-001-0065-4
Fournier DA, Skaug HJ, Ancheta J, Ianelli J, Magnusson A, Maunder M, Nielsen A, Sibert J (2012). AD Model Builder: using automatic differentiation for statistical inference of highly parameterized complex nonlinear models. Optimization Methods and Software 27: 233-249. https://doi.org/10.1080/10556788.2011.597854
Fox J, Weisberg S (2019). An R Companion to Applied Regression. Sage, Thousand Oaks CA. http://z.umn.edu/carbook
Frankel OH, Soulé ME (1981). Conservation and Evolution. Cambridge University Press: Cambridge, New York, pp 311.
Fruin S, Westerdahl D, Sax T, Sioutas C, Fine PM (2008). Measurements and predictors of on-road ultrafine particle concentrations and associated pollutants in Los Angeles. Atmospheric Environment 42(2): 207-219. https://doi.org/10.1016/j.atmosenv.2007.09.057
Gillies CS, St. Clair CC (2010). Functional responses in habitat selection by tropical birds moving through fragmented forest. Journal of applied ecology 47(1): 182-190. https://doi.org/10.1111/j.1365-2664.2009.01756.x
Gotelli NJ, Chao A (2013). Measuring and estimating species richness, species diversity, and biotic similarity from sampling data, pp. 195-211. https://doi.org/10.1016/B978-0-12-384719-5.00424-X
Hill MO (1973). Diversity and evenness: a unifying notation and its consequences. Ecology 54: 427. http://dx.doi.org/10.2307/1934352
Hoffmann BD, Andersen AN (2003). Responses of ants to disturbance in Australia, with particular reference to functional groups. Austral Ecology 28: 444-464. https://doi.org/10.1046/j.1442-9993.2003.01301.x
Hölldobler B, Wilson EO (1990). The ants. Belknap Press of Harvard University Press, Cambridge, MA, USA.
Hsieh TC, Ma KH, Chao A (2016). iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods in Ecology and Evolution 7: 1451-1456. https://doi.org/10.1111/2041-210X.12613
Itzhak MJ (2008). Seed harvester and scavenger ants along roadsides in Northern Israel. Zoology in the Middle East 44(1): 75-82. https://doi.org/10.1080/09397140.2008.10638291
Jost L (2006). Entropy and diversity. Oikos 113: 363-375.
Jost L (2007). Partitioning diversity into independent alpha and beta components. Ecology 88:2427-2439.
Jost L. (2010). The relation between evenness and diversity. Diversity 2: 207-232. https://doi.org/10.3390/d2020207
Kaur H, Torma A, Gallé-Szpisjak N, Šeat J, Gábor L, Gábor M, Gallé R (2019). Road verges are important secondary habitats for grassland arthropods. Journal of Insect Conservation 23: 899-907. https://doi.org/10.1007/s10841-019-00171-9
Keals N, Majer JD (1991). The conservation status of ant communities along the Wubin-Perenjori corridor. In: Saunders DA and Hobbs RJ (eds.) Nature Conservation 2: The Role of Corridors. Surrey Beatty and Sons, Sydney, pp 387-393.
Kindt R, Coe R (2005). Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies. World Agroforestry Centre (ICRAF), Nairobi.
Kociolek A, Clevenger A, St Clair C, Proppe D (2011). Effects of road networks on bird populations. Conservation Biology 25:241-249 https://doi.org/10.1111/j.1523-1739.2010.01635.x
Kremen C, Colwell RK, Erwin TL, Murphy DD, Noss RA, Sanjayan MA (1993). Terrestrial arthropod assemblages: their use in conservation planning. Conservation biology 796-808. https://www.jstor.org/stable/2386811
Kusnezov N (1978). Hormigas Argentinas: claves para su identificación. Miscelanea, Tucumán. N° 61.
Lassau SA, Hochuli DF (2003). Effects of roads on ant assemblages in the Sydney region: are patterns scale dependent. Records of the South Australian Museum 7: 283-290.
Laurance SG, Stouffer PC, Laurance WF (2004). Effects of road clearings on movement patterns of understory rainforest birds in central Amazonia. Conservation biology 18(4): 1099-1109. https://doi.org/10.1111/j.1523-1739.2004.00268.x
Leal IR, Filgueiras BK, Gomes JP, Iannuzzi L, Andersen AN (2012). Effects of habitat fragmentation on ant richness and functional composition in Brazilian Atlantic forest. Biodiversity and Conservation 21: 1687-1701. https://doi.org/10.1007/s10531-012-0271-9
Leal IR, Wirth R, Tabarelli M (2014). The multiple impacts of leaf- cutting ants and their novel ecological role in human-modified Neotropical forests. Biotropica 46: 516-528. https://doi.org/10.1111/btp.12126
Lessard JP (2019). Ant community response to disturbance: A global synthesis. Journal of Animal Ecology 88(3): 346-349. https://doi.org/10.1111/1365-2656.12958
MacArthur RH (1972). Geographical Ecology. Princeton University Press, Princeton 288 pp.
Majer JD, Delabie JH, Smith MR (1994). Arboreal ant community patterns in Brazilian cocoa farms. Biotropica 73-83. https://doi.org/10.2307/2389112
Márquez J, Martinez Carretero E, Dalmasso A, Pastrán G, Ortiz G (2005). Las áreas protegidas de la provincia de San Juan (Argentina) II. La vegetación del Parque Provincial Ischigualasto. Multequina 14: 1-27.
Melis C, Olsen CB, Hyllvang M, Gobbi M, Stokke BG, Røskaft E (2010). The effect of traffic intensity on ground beetle (Coleoptera: Carabidae) assemblages in central Sweden. Journal of Insect Conservation 14: 159-168. https://doi.org/10.1007/s10841-009-9240-3
Murcia C (1995). Edge effects in fragmented forests: implications for conservation. Trends in Ecology and Evolution 10: 58-62. https://doi.org/10.1016/S0169-5347(00)88977-6
Muñoz TP, Pascual Torres F, Gongález Mejías A (2015). Effects of roads on insects: a review. Biodiversity and Conservation 24: 659-682 https://doi.org/10.1007/s10531-014-0831-2
Noordijk J, Raemakers IP, Schaffers AP, Sykora KV (2009). Arthropod richness in roadside verges in Netherlands. Terrestrial Arthropod Reviews 2: 63-76. https://doi.org/10.1163/187498309x440085
Oksanen J, Blanchet FG, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin PR, O'Hara R B, Simpson GL, Solymos P, Stevens MH, Szoecs E, Wagner H (2019). Vegan: Community Ecology Package.
Palfi Z, Spooner PG, Robinson W (2017). Soil disturbance effects on the composition of seed-dispersing ants in roadside environments. Oecologia, 183(2): 493-503. https://doi.org/10.1007/s00442-016-3767-2
Palacio EE, Fernández F (2003). Claves para las subfamilias y géneros. In: Fernández F (ed). Introducción a las hormigas de la región Neotropical. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia.
Parr CL, Andersen AN, Chastagnol C, Duffaud C (2007) Savanna fires increase rates and distances of seed dispersal by ants. Oecologia 151: 33-41. https://doi.org/10.1007/s00442-006-0570-5
Pérez-Sánchez AJ, Lattke-Bravo JE, Viloria AL (2012). Composición y estructura de la fauna de hormigas en tres formaciones de vegetación semiárida de la península de Paraguaná, Venezuela. Interciencia 37: 506-514. https://www.redalyc.org/articulo.oa?id=33925376004
Pereyra M, Pol RG, Galetto L (2019). Ant community patterns in highly fragmented Chaco forests of central Argentina. Austral Ecology 44(4): 668-679. https://doi.org/10.1111/aec.12712
Philpott SM, Perfecto I, Armbrecht I, Parr CL (2010): Ant diversity and function in disturbed and changing habitats. In Lach L., Parr C.L. and Abbott K.L. (eds): Ant Ecology. Oxford University Press, New York, pp. 137–157.
Pickett ST, Kolasa I, Armesto II, Collins SL (1989). The ecological concept of disturbance and its expres- sion at various hierarchical levels. Oikos 54: 129-136. https://doi.org/10.2307/3565258
Pirk GI, López de Casenave JL, Pol RG (2004). Asociación de las hormigas granívoras Pogonomyrmex pronotalis, P. rastratus y P. inermis con caminos en el Monte central. Ecología Austral 14(1): 065-076.
Pirk GI, López de Casenave JL (2017). Ant interactions with native and exotic seeds in the Patagonian steppe: influence of seed traits, disturbance levels and ant assemblage. Plant Ecology 218(11): 1255-1268. https://doi.org/10.1007/s11258-017-0764-4
R. Core Team 2019. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. Vienna, Austria. URL https://www.R-project.org/
Ribas CR, Schoereder JH, Pic M, Soares SM (2003). Tree heterogeneity, resource availability, and larger scale processes regulating arboreal ant species richness. Austral Ecology 28(3): 305-314. https://doi.org/10.1046/j.1442-9993.2003.01290.x
Rivas-Arancibia SP, Carrillo-Ruiz H, Arce AB, Figueroa-Castro DM, Andrés- Hernánde AR (2014). Effect of disturbance on the ant community in a semiarid region of central Mexico. Applied Ecology and Environmental Research 12:703-716. https://doi.org/10.15666/AEER/1203_703716
Rotholz E, Mandelik Y (2013). Roadside habitats: effects on diversity and composition of plant, arthropod, and small mammal communities. Biodiversity and conservation 22(4): 1017-1031. https://doi.org/10.1007/s10531-013-0465-9
Scarpa GF, Rosso CN (2011). Etnobotánica del “coro” (Nicotiana paa, Solanaceae): Un tabaco silvestre poco conocido del extremo sur de Sudamérica. Bonplandia 2: 391-404. https://doi.org/10.30972/bon.2021422
Schumacher A, Whitford WG (1976). Spatial and temporal variation in Chihuahuan desert ant faunas. Southwestern Naturalist 21:1-8.
Silva PG, Hernández MI (2015). Spatial patterns of movement of dung beetle species in a tropical forest suggest a new trap spacing for dung beetle biodiversity studies. PLoS ONE 10(5): e0126112. https://doi.org/10.1371/journal.pone.0126112
Singh SP (1998). Chronic disturbance, a principal cause of environmental degradation in developing countries. Environmental Conservation 25:1-2. https://doi.org/10.1017/S0376892998000010
Siqueira FF, Ribeiro-Neto JD, Tabarelli M, Andersen AN, Wirth R, Leal IR (2017). Leaf-cutting ant populations profit from human disturbances in tropical dry forest in Brazil. Journal of Tropical Ecology 33(5): 337-344. https://doi.org/10.1017/S0266467417000311
Song IJ, Hong SK, Kim HO, Byund B, Gin Y (2005). The pattern of landscape patches and invasion of naturalized plants in developed areas of urban Seoul. Landscape and Urban Planning 70: 205-219. https://doi.org/10.1016/j.landurbplan.2003.10.018
Soulé ME, Gilpin ME (1991). The theory of wildlife corridor capability. Nature Conservation 2:3–8.
Subpiramaniyam S (2021). Portulaca oleracea L. for phytoremediation and biomonitoring in metal-contaminated environments. Chemosphere 280: 130784. https://doi.org/10.1016/j.chemosphere.2021.130784
Terranella AC, Ganz Lisa, Ebersole JJ (1999). Western harvester ants prefer nest sites near roads and trails. Southwestern Association of Naturalists 44 (3): 382-84.
Trombulak SC, Frissell CA (2000). Review of ecological effects of roads on terrestrial and aquatic communities. Conservation biology 14(1): 18-30. https://doi.org/10.1046/j.1523-1739.2000.99084.x
Tshiguvho TE, Dean WR, Robertson HG (1999). Conservation value of road verges in the semi-arid Karoo, South Africa: ants (Hymenoptera: Formicidae) as bioindicators. Biodiversity and Conservation 8: 1683–1695. https://doi.org/10.1023/A:1008911805007
Vanthomme H, Alonso A, Tobi E, Rolegha CL, Hita Garcia F, Mikissa JB, Alonso LE (2017). Associations of ant species to land cover types and human disturbance in south-west Gabon. African journal of Ecology 55(3):352-364. https://doi.org/10.1111/aje.12362
Vasconcelos HL, Vieira‐Neto EH, Mundim FM, Bruna EM (2006). Roads Alter the Colonization Dynamics of a Keystone Herbivore in Neotropical Savannas 1. Biotropica 38(5): 661-665. https://doi.org/10.1111/j.1744-7429.2006.00180.x
Vieira‐Neto EH, Vasconcelos HL, Bruna EM (2016). Roads increase population growth rates of a native leaf‐cutter ant in Neotropical savannahs. Journal of Applied Ecology 53(4), 983-992. https://doi.org/10.1111/1365-2664.12651
Villemey A, Jeusset A, Vargac M et al. (2018). Can linear transportation infrastructure verges constitute a habitat and/or a corridor for insects in temperate landscapes? A systematic review. Environmental Evidence 7, 5. https://doi.org/10.1186/s13750-018-0117-3
Walsh PD, Henschel P, Abernethy KA, Tutin CE, Telfer P, Lahm SA (2004). Logging speeds little red fire ant invasion of Africa. Biotropica, 36(4): 637-641. https://doi.org/10.1111/j.1744-7429.2004.tb00358.x
Whitford WG (2002). Ecology of desert systems. Academic Press, Elsevier Science Imprint, San Diego, California.
Wynhoff I., van Gestel R, van Swaay C, van Langevelde F (2011). Not only the butterflies: managing ants on road verges to benefit Phengaris (Maculinea) butterflies. Journal of Insect Conservation 15: 189–206. https://doi.org/10.1007/s10841-010-9337-8
Zanden EH, Verburg PH, Mücher CA (2013). Modelling the spatial distribution of linear landscape elements in Europe. Ecological Indicators 27:125–136. https://doi.org/10.1016/j.ecolind.2012.12.002
Zeileis A, Hothorn T (2002). Diagnostic checking in regression relationships, pp 5.
Zurita G, Pe’er G, Bellocq MI, and Hansbauer MM (2012). Edge effects and their influence on habitat suitability calculations: a continuous approach applied to birds of the Atlantic forest. Journal of Applied Ecology 49:503–512. https://doi.org/10.1111/j.1365-2664.2011.02104.x

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Effect of road disturbance on ant diversity in a sector of the Central Biocenic Corridor located in the center of Arid diagonal of Argentina

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