Mastozoología Neotropical ISSN: 0327-9383 [email protected] Sociedad Argentina para el Estudio de los Mamíferos Argentina
Cuevas, M. Fernanda; Ojeda, Ricardo A.; Jaksic, Fabian M. ECOLOGICAL STRATEGIES AND IMPACT OF WILD BOAR IN PHYTOGEOGRAPHIC PROVINCES OF ARGENTINA WITH EMPHASIS ON ARIDLANDS Mastozoología Neotropical, vol. 23, núm. 2, 2016, pp. 239-254 Sociedad Argentina para el Estudio de los Mamíferos Tucumán, Argentina
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Sección Especial MAMÍFEROS EXÓTICOS INVASORES
ECOLOGICAL STRATEGIES AND IMPACT OF WILD BOAR IN PHYTOGEOGRAPHIC PROVINCES OF ARGENTINA WITH EMPHASIS ON ARIDLANDS
M. Fernanda Cuevas1, Ricardo A. Ojeda1 and Fabian M. Jaksic2
1 Grupo de Investigaciones de la Biodiversidad (GiB), IADIZA, CCT Mendoza CONICET, CC 507, 5500 Mendoza, Argentina. [Correspondence:
ABSTRACT. Wild boar is an invasive species introduced to Argentina for sport hunting purposes. Here, this spe- cies is present in at least 8 phytogeographic provinces but we only have information in four of them (Pampean grassland, Espinal, Subantarctic and Monte Desert). We review the ecological strategies and impact of wild boar on ecosystem processes in these different phytogeographic provinces and identify knowledge gaps and research priorities for a better understanding of this invasive species in Argentina. We observe that foraging strategies of wild boar consist in consuming mainly plant species rich in energy, especially those with bulbs and fuits having high concentrations of carbohydrates and lipids, and the use of habitat is closely associated with food resource availability. Wild boar generate a broad variety of impacts reflected in plant species composition, structure and biomass, and changes in soil properties such as increased soil degradation in the Monte Desert. We conclude that the impacts of wild boar in Argentina are mostly negative, demanding more interaction among relevant players (scientists, government officials, managers of protected areas, and landowners) to plan the population control strategies needed to mitigate damage to native ecosystems and agricultural production in the country.
RESUMEN. Estrategias ecológicas e impacto del jabalí en provincias fitogeográficas de Argentina con énfasis en las tierras áridas. El jabalí es una especie exótica introducida en Argentina para la caza deportiva. Aquí, esta especie está presente en al menos 8 provincias fitogeográficas aunque solo tenemos información para cuatro de ellas (Pampeana, Espinal, Subantártica y Desierto del Monte). Se hace una revisión de las estrategias ecológicas y del impacto del jabalí sobre el ecosistema invadido en diferentes provincias fitogeográficas de Argentina, identificando los vacíos y necesidades de investigación para un mejor entendimiento de esta especie invasora. Observamos que la estrategia de forrajeo del jabalí consiste en consumir principalmente especies de plantas ricas en energía, con un alto contenido de carbohidratos y lípidos presentes en bulbos y frutos; y que el uso del hábitat está principalmente asociado con la disponibilidad de los recursos alimenticios. El jabalí en Argentina genera una amplia variedad de effectos relacionados con la composición, estructura y biomasa de la vegetación, y con cambios en las propiedades del suelo, donde genera un aumento de la degradación del mismo, ejemplifi- cado en el Desierto del Monte. Concluimos que el impacto generado por el jabalí en Argentina es mayormente negativo, por lo que es necesaria la interacción de diferentes actores (científicos, gobierno, administradores de
Recibido 17 julio 2015. Aceptado 10 marzo 2016. Editor invitado: RA Ojeda 240 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br
áreas protegidas y propietarios de campos) para planificar una estrategia de gestión y control de las poblacio- nes de jabalí, y por lo tanto mitigar su daño sobre el ecosistema nativo y los sistemas productivos del país.
Key words: Argentina. Disturbance. Foraging. Habitat. Sus scrofa.
Palabras clave: Argentina. Disturbios. Forrajeo. Hábitat. Sus scrofa.
INTRODUCTION distribution of wild boar (Jedrzejewska et al., 1997; Acevedo et al., 2006). For example, The wild boar (Sus scrofa) is native to Eurasia rainfall generates an increase in the number of and northern Africa (Long, 2003), but now has pregnant females during rainy years (Fernán- one of the widest geographic distributions of dez-Llario and Mateos-Quesada, 2005). Hunt- all exotic mammals (Oliver et al., 1993). Due ing pressure also affects its spatial behavior by to the damage it causes in natural ecosystems modifying choice of resting sites (Scillitani et and agricultural areas, it is considered one of al., 2009). Heterogeneous landscapes (i.e., high the 100 most harmful invasive species of the diversity of food resources and high availability world (Lowe et al., 2000). The wild boar is an of shelters) also favor high densities of wild omnivorous species with a diet dominated by boar as compared to homogeneous habitats plant material (between 87 and 99%) and a (Fernández-Llario, 2004; Acevedo et al., 2006). smaller representation of animal matter (Schley The wild boar is considered an ecosystem and Roper, 2003). It has a high reproductive engineer due to its rooting behavior, which capacity due to characteristics such as preco- transforms the physical state of biotic and cious sexual maturation (from 5 to 12 months), abiotic materials (Jones et al., 1994; Crooks, its relatively short gestation time (120 days), 2002). To forage, wild boars overturn extensive and large litter size (5-7 piglets) (Gethöffer et areas of soil, leaving them bare of vegetation al., 2007; Herrero et al., 2008). It also has a (Hone, 1988). This rooting behavior creates a high tolerance to different climatic conditions, complex mosaic of disturbed patches of differ- reflected in its wide geographic range (Oliver ent ages and sizes (Cuevas et al., 2012). This et al., 1993). For all these traits, wild boar has type of activity sometimes generates benefits expanded successfully to all continents except to native and exotic flora, such as increasing Antarctica (Long, 2003). Nowadays this species plant richness and cover of perennial grasses occurs in different parts of the world either in (Tierney and Cushman, 2006). But it also causes pure wild or barely-modified feral form due negative effects such as mixing soil horizons, to its crossing with domestic pigs (Baber and reducing vegetative cover and litter, accelerating Coblentz, 1986; Merino and Carpinetti, 2003). the leaching of ions from litter and soil (e.g., In its native range, this species occupies dif- Ca, P, Zn, Cu, and Mg), increasing nitrate con- ferent types of habitats such as forests, shrub- centrations and soil respiration, and decreasing lands, mangroves, grasslands, and wetlands the abundance of soil arthropods (Singer et al., (Rosell et al., 2001). In spite of that, wild boar 1984; Mohr et al., 2005; Risch et al., 2010). prefers those habitats that offer high energy Here we review and summarize the literature food, such as acorns, and have high vegetation on wild boar ecology and ecosystem impacts cover for protection from predators (includ- in different phytogeographic provinces of ing hunters), as well as those close to water Argentina, identifying knowledge gaps and resources (Kurz and Marchinton, 1972; Massei research priorities toward a better understand- and Genov, 1995). ing of this invasive species in Argentina. The Several climatic and ecological factors have aims of our contribution involve an overview been described to affect the abundance and of scientific studies of wild boar ecology and A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 241 impact in Argentina, with a particular focus in which the species uses available resources in their ecological strategies to survive in semi- the invaded habitat, such as diet, habitat use, arid conditions. resource selection, that permit wild boar to We performed searches in specialized jour- survive in it) and impact on soil properties nals and data bases (e.g. Blackwell, Elsevier, and plant species composition and structure. Google Scholar, Scielo and Scopus), using dif- ferent combinations of the following keywords: THE WILD BOAR IN ARGENTINA Sus scrofa, feral pig, wild boar, ecology, feeding habits, diet, habitat use, impact, Argentina. We The wild boar was first introduced to Argentina also reviewed doctoral and Master’s theses. in 1906 in San Huberto ranch, La Pampa for We set the search for studies between 1970 hunting purposes (Daciuk, 1978). After that, and 2015 and included studies referring to wild boar reintroductions occurred several wild boar and feral pigs. The literature search times in different parts of the country, such yielded 234 studies of which 17 were relevant as in Collun-có ranch, Neuquén in 1917 and to the objective of this study (Table 1), from Huemul ranch, Río Negro in 1924 (Daciuk, which we found 13 scientific articles, 3 doc- 1978). Furthermore, the continuous installation toral theses and 1 technical report related to of hunting grounds involves the introduction wild boar ecological strategies (i.e., way in of new populations of this species around the
Table 1 Summary of wild boar studies in the phytogeographic provinces of Argentina.
Phytogeographic Impact Ecological traits References province
Pampean grassland Potential impact Population abundance Merino and Carpinetti (2003); on Pampas deer Perez Carusi et al. (2009) (Ozotoceros bezoarticus celer) populations
Espinal Seed predation of Diet, habitat use Govetto (1999); Ballari (2013); Butia yatay Ballari et al. (2015b)
Habitat use, distribu- Schiaffini and Vila (2012); tion range expansion Gantchoff and Belant (2015); Pescador et al. (2009)
Subantarctic Impact on vegetation Barrios-García (2012); and soil properties Barrios-García and Simberloff (2013); Barrios-García et al. (2014)
Seed predation of Sanguinetti and Kitsberger (2010) Araucaria araucana
Monte Desert Impact on vegetation Cuevas et al. (2010); Cuevas (2012); and soil properties Cuevas et al. (2012)
Diet, habitat use Cuevas et al. (2013a); Cuevas et al. (2013b) 242 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br country. In a recent study, Ballari et al. (2015a) about 700 individuals at the beginning of the evaluated the current status of wild boar in study to more than 2000 at the end, including Argentina’s system of protected areas. They several peaks reaching over 4000 individuals. showed that wild boars are present in at least A second study conducted by Perez Carusi et 10 ecoregions belonging to 8 phytogeographic al. (2009) used the same methodology of count- provinces (High Andean, Monte Desert, Chaco, ing, but also compared the spatial distribution Paranaense, Pampean grassland, Espinal, Pa- and abundance of feral pigs with the sympatric tagonian and Subantarctic provinces) (Fig. 1). pampas deer (Ozotoceros bezoarticus celer), a While in the rest of the provinces (Puna, native species with endangered conservation Prepuna, Yungas) they did not find any record status (Ojeda et al., 2012). They reported in- of this species. They also found novel ecore- direct evidence of a potential negative impact gions being occupied by this species, like High of feral pigs on pampas deer: compared to Andean, Parana Flooded Savanna and Iberá previous years there was a decrease in pampas Marshes, which indicates that wild boars are deer abundance and spatial distribution while continuously expanding their geographic range feral pigs increased on both counts. The authors in Argentina. Nevertheless, studies about their also suggested that the decrease in abundance ecology and ecosystem impacts in different of pampas deer could be related to other fac- biomes of Argentina are scarce. Information is tors such as the outbreak of FMD (foot and available in only four phytogeogeographic prov- mouth disease) and/or the effect of poaching. inces: Pampean grassland, Espinal, Subantarctic Still, Perez Carusi et al. (2009) found a negative region, and Monte Desert (Fig. 1, Table 1). correlation between abundances of pampas deer and feral pigs, but did not assess the habitat Pampean grassland or food use of these two species. Although The climate of this province ranges from tem- pampas deer is a strict herbivore and pigs are perate to warm, having rainfall throughout the omnivorous, the bulk of feral pig diet is plant whole year that decreases from north to south matter (90%) (Schley and Roper, 2003; Ballari and from east to west (range: 600-1100 mm and Barrios-García, 2014). Further research on rainfall annually). Dominant vegetation is the niche axes (e.g., food, habitat) of these spe- grasslands with a lower presence of halophyte cies may provide a more thorough assessment steppes, marginal forests, and several types of of their coexistence mechanisms. hydrophilic shrubs (Cabrera, 1971). Pérez Carusi et al. (2009) observed a 400% For this biome, there are two studies about increase of pig population during the period Sus scrofa, in which area occur feral populations of study. However, this finding should be taken of domestic pigs, both in Bahía Samborombón cautiously, because at the end of Merino and (35°26’ S, 57°47’ W) where feral pigs were re- Carpinetti’s (2003) study, pig population was ported in 1980 (Merino and Carpinetti, 2003; over 2600 individuals, while in Pérez Carusi et Pérez-Carusi et al., 2009; Ballari et al., 2015a) al.’ (2009) study it was 2690 individuals. Still, (Table 1). This area has an annual precipita- comparing both studies, there was a substantial tion of 1000 mm and small patches of “tala” increase in pig density, from 1.59 ind/km2 to forest (Celtis tala) surrounded by humid and 7.78 ind/km2. It should be noted that these salty grasslands (Cabrera, 1971). This area has studies differed in number of sighting trips a special significance as a refuge for several na- and transects, and the size of surveyed area. tive species that have disappeared in other parts Espinal of the Pampas region (Merino and Carpinetti, 2003). Between 1995 and 1998, Merino and The Espinal phytogeographic province has a Carpinetti (2003) assessed feral pig populations diverse climate that changes latitudinally from using aerial counts. They found that pig abun- warm and wet in the northern part to temper- dances showed an accelerated increase, from ate and dry in the central and southern areas. A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 243
Fig. 1. Argentine phytogeographic provinces showing the presence of wild boar (white circles). Numbers indicate provinces with ecological and ecosystem impact information of the species. Map extracted from Katinas et al., 2007. 244 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br
Accordingly, the vegetation also varies, showing may cause wild boar to compensate for lack of deciduous dry forests, palm groves, grasslands, protein by eating more animal matter (Schley savannahs and shrub steppes (Cabrera, 1971). and Roper, 2003). The southern part of this region was home to Regarding habitat use, Govetto (1999) found the first introduction of wild boar in Argentina a high density of wild boar signs in Yatay palm (San Huberto ranch, La Pampa province). De- forest during the masting period (February spite having the oldest wild boar population in and March), and also Ballari (2013) found that the country, there is no scientific information wild boar prefers habitats with a dominant tree evaluating its interactions with native species canopy, e.g., Yatay palm forest and forest of or the impact upon the invaded ecosystem. exotic xerophytes (white cedar Melia azedarach, Instead, the ecology and impacts of wild boar Gigg’s firethornPyracantha atalantoides, honey in this region have been conducted in an area locust Gleditsia triacanthos, broad-leaf privet that differs markedly from the introduction site: Ligustrum lucidum, Chinese privet Ligustrum El Palmar National Park (EPNP), Entre Rios sinense). Agricultural lands that surround the province (31°50’ S, 58°17’ W) (Govetto, 1999; park were not preferred by wild boars, possibly Ballari, 2013; Ballari et al., 2015b) (Table 1). due to abundant food resources present in the The landscape is characterized by a hetero- park, including the supplemental feeding (corn) geneous mosaic of vegetation patches that for hunting practices (Ballari et al., 2015b). includes gallery forests, shrublands, grasslands Subantarctic and savannahs, with Yatay palms (Butia yatay) in highlands (Movia and Menvielle, 1994). The The climate of this phytogeographic region climate is warm (annual mean temperature is temperate and wet with mean temperature 28.9 °C) and wet throughout the year with 9.5 °C in the northern portion and 5.4 °C in the no dry season (annual mean precipitation south. Annual precipitation goes from 2000 mm 1300 mm; Papadakis, 1974). Wild boars have on the west bordering with Chile to 750 mm been reported in this protected area since 1950 to the east of this region. Dominant vegetation (Ballari et al., 2015a). types are pure or mixed forests of conifers like Based on stomach contents, Ballari et al. cordilleran cypress (Austrocedrus chilensis, Fam. (2015b) found that 81.2% of the wild boar diet Cupressaceae) and alerce (Fitzroya cupressoides, at EPNP is plant material and almost 18.8% is Fam. Cupressaceae), evergreen species like coi- animal matter. They observed that during the hue (Nothofagus dombeyi, Fam. Nothofagaceae), fruiting of Yatay palm (an endemic and protect- and deciduous species of southern beech such ed species) wild boars eat those fruits to reach as ñire (N. antarctica, Fam. Nothofagaceae) approximately 50% of their diet in summer. and lenga (N. pumilio, Fam. Nothofagaceae). But during winter/autumn and spring, when These forests are mixed to a lesser extent with those fruits are not available on the ground, grasslands and peatlands (Cabrera, 1971). Wild boar fed mainly on corn (a supplemental feed- boars have been introduced to Patagonian forest ing used in EPNP for controlling the species, since 1917 (Daciuk, 1978). and to promote hunting), this item comprising Within this region, wild boar studies have between 40 and 50% of the diet. Only during focused on habitat use, distribution range and spring, the bulk of the diet is corn (42%) and impact on soil properties and vegetation (Pes- animal matter (27%). So, it appears that dur- cador et al., 2009; Sanguinetti and Kitsberger, ing the masting period of Yatay palm, boars 2010; Schiaffini and Vila, 2012; Barrios-García, prefer to eat it over the supplemental corn. The 2012; Barrios-García and Simberloff, 2013; relatively high dietary content of animal matter Barrios-García et al., 2014; Gantchoff and Bel- in EPNP could be related with the ingestion ant, 2015) (Table 1). of corn and fruits of Butia spp., whose species Regarding habitat use, Schiaffini and Vila are high in carbohydrates but low in protein (2012) registered the presence of wild boar (Schley and Roper, 2003; Hoffman et al., 2014). signs through transects along an altitudinal So their gain of caloric requirement from corn gradient (from 300 to 1200 m elevation) at A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 245
Los Alerces National Park (LANP, 42°50’ S, Results showed that wild boar rooting generates 71°52’ W). They found that between 600 and a 60% reduction in aerial plant biomass, this 700 m there was the highest abundance of wild negative effect being stronger inNothofagus boar signs and that at elevation 1200 m there forests. Cover of herbs and grasses was lower was no evidence of its presence. Nothofagus in rooting patches, herb cover being least in dombeyi and N. antarctica forests were used Austrocedrus and Nothofagus forests while grass by this species more than forests of N. pumilio cover was also lower in Nothofagus forest. Shrub and grasslands. The authors concluded that the cover was also negatively affected by rooting, increased presence of wild boar in intermediate but it was in shrubland that the authors noticed elevations was associated with the dense under- the major impact. Regarding litter decomposi- story vegetation of Nothofagus forests, which tion rate, this was lower in rooting patches. provides warmth and moisture conditions that At soil level, wild boar rooting only modified boars need to meet thermal requirements. Also, soil compaction, making this attribute lower the high canopy (40 m) provides shelter from in both forests (Table 2). Finally, the authors hot summer temperatures (~30 °C). Further, concluded that impact caused by wild boar is the dense understory of bamboo (Chusquea greater aboveground than belowground. culeou) affords protection from frost during In a second enclosure experiment Barrios- cold days, allowing boars to find food under it. García and Simberloff (2013) evaluated the Gantchoff and Belant (2015) evaluated the effect of rooting on non-native seedling estab- influence of environmental and anthropogenic lishment and plant growth, and wild boar’s role factors on wild boar occurrence in a tourist site in seed dispersal of native and exotic plant spe- of Nahuel Huapi National Park (NHNP, 40°57’ cies. Rooting patches showed higher non-native S, 71°33’ W) using camera traps. Similarly as seedlings and biomass of seedlings, biomass in Schiaffini and Vila’s (2012) study, they found being greatest in shrublands. All exotic plant that wild boar frequently uses Nothofagus species except for sweet brier (Rosa rubiginosa) dombeyi and N. antarctica forests compared and elmleaf blackberry (Rubus ulmifolius) with N. pumilio forests. They found that lon- showed higher establishment in rooting patches. ger distance to human settlements and closer The authors also found that both soil samples distances to roads were the most important and feces showed equal composition of seed anthropogenic variables influencing the oc- species, but in fecal samples there were fewer currence of wild boar in the park. That result non-native species of seed compared with soil plus the boars’ nocturnal activity indicates that samples. Lastly, they found a positive effect boars use roads as corridors to move across (invasional meltdown) in the establishment of forests while avoiding human due to hunting non-native seedlings in rooting patches and a pressure (Gantchoff and Belant, 2015). negative effect in non-native seed dispersal. Regarding distribution range, Pescador et Sanguinetti and Kitzberger (2010) evaluated al. (2009) assessed the presence and the rela- the impact of wild boar on seed survival and tive abundance of wild boar in 1985 and then seedling establishment of Araucaria araucana repeated it in 2005 along transects in Lanín in LNP through an experiment at different National Park (LNP, 39°34’S, 71°27’W). They distances from a female tree of araucaria. found that after 20 years wild boar increased The authors observed that wild boars pre- its range with a spread rate of 3500 ha per year. ferred mixed A. araucaria–N. pumilio over Studies about impact of wild boar on vegeta- A. araucaria–N. antarctica forests for feeding. tion structure, seed predation, and soil proper- They found that wild boar consumed between ties were recently conducted in LNP and NHNP. 10 and 30% of available seeds. Predation was At NHNP, Barrios-García et al. (2014) studied greater in places with low plant cover and close the impact of wild boar on vegetation and soil to seeding trees. When they excluded wild properties through an enclosure experiment in boars, they found that the number of surviving three different plant communities:Austrocedrus seeds increased, resulting in higher seedling chilensis, Nothofagus dombeyi and shrublands. establishment during non-masting years (boars 246 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br
Table 2 Effects of wild boar’s rooting in soil properties in both Monte Desert and Subantarctic phytogeographic provinces.
Monte Desert Subantarctic Forest Soil Properties Disturbed Undisturbed Disturbed Undisturbed
Physical Hardness – + – + Moisture + – No change Temperature ------No change Texture Silt – + ------Clay – + ------Sand + – ------
Chemical Total Nitrogen No change No change Mineral Nitrogen + – No change Nitrate + Nitrite + – ------
NH4 No change ------Organic Carbon No change ------C/N ratio + – ------Organic matter No change ------pH No change No change Total Carbon ------No change Extractable P ------No change
Microbiological Soil respiration – + No change Ammonifiers No change ------Cellulolytics No change ------N fixers No change ------Nitrifiers No change ------
ate proportionally more seeds during such peri- acterized by high temperatures, water deficit ods than during masting). Finally, the negative and low plant productivity, generating a great effect of wild boar was reflected at individual challenge to the survival of plants and animals tree level, but not at population scale. in these environments (Cloudsley-Thompson, To sum up, wild boars in the Subantarctic 1975; Brown et al., 1979; Polis, 1995). region more often use Nothofagus forests and South American aridlands have played an those habitats are the most affected by rooting important role in the evolution of the temper- behavior. The disturbance by wild boar affects ate biota of the continent, with high biological properties not only at community level but diversity that contains a large percentage of also at ecosystem scale, changing plant com- endemic genera and families (Ojeda et al., munity composition and structure, decreasing 1998). In Argentina, aridlands are undergoing decomposition rates, and promoting invasive rapid habitat conversion as a result of human plant establishment and growth. activities (agriculture, grazing, logging, etc.), desertification, and salinization (Ojeda and Monte Desert Mares, 1982). To these challenges, environ- Aridlands are one of the most extensive terres- mental changes driven by climate change are trial habitats on the planet, occupying about a added, as well as changes caused by invasive third of the Earth’s surface. Aridlands are char- species (Boulanger et al., 2007; Cuevas et al., A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 247
2012). Considering that the temperate biomes Wild boar studies within this phytogeograph- of Argentina concentrate the highest numbers ic region have been conducted in the Man and of invasive mammals of South America (Ojeda Biosphere (MaB) Reserve of Ñacuñán (34°02′ S, et al., unpublished data), the study of invasive 67°58′ W), Mendoza province. The landscape species in the process of expansion is an in- is characterized by a heterogeneous mosaic teresting opportunity to assess the conditions of vegetation patches. Dominant habitats are and constraints that these species face within Prosopis woodland or algarrobal (Prosopis the dynamics of aridland invasibility. flexuosa, Fam. Fabaceae), Larrea shrubland or In Argentina, 57% of the territory consists of jarillal (Larrea cuneifolia, Fam. Zygophyllaceae), aridlands (Verbistk et al., 2010). Overall, there and sand dunes. The climate is semiarid and are 23 exotic species of mammals (including strongly seasonal, with hot, humid summers feral populations of domestic species), with and cold, dry winters. Mean annual precipita- at least six of them found in this region: wild tion and temperature are 326 mm and 15.6 ºC, boar (Sus scrofa), European rabbit (Oryctolagus respectively, with a maximum annual mean of cuniculus), European hare (Lepus europaeus), 23.8 ºC and a minimum annual mean of 7.6 ºC blackbuck (Antilope cervicapra), donkey (Equus (Estrella et al., 2001; Labraga and Villalba, asinus), red deer (Cervus elaphus) (Novillo and 2009). Wild boar was first sighted in this area Ojeda, 2008). in the 1980’s (Cuevas et al., 2010). Monte Desert is a subtropical to warm tem- Cuevas et al. (2013a) studied habitat use by perate desert and semidesert located in western wild boar through signs such as tracks and Argentina (Abraham et al., 2009) (Fig. 1). The rooting. Considering tracks at the habitat level, climate is dry and warm in the northern por- the authors did not find any difference among tion and dry and cold in southern part of this the three available habitats, which means that phytogeographic province. The precipitation at least for displacement (moving from one varies between 80 and 250 annual mm, and place to another) wild boars used the differ- temperature from 48°C to -17°C (Labraga and ent habitats in proportion to their availability. Villalba, 2009). Dominant vegetation types are Regarding rooting activity they found that wild shrubland steppes of xerophytes, psammophytes boar positively selected Larrea shrubland and or halophytes, as well as marginal Prosopis avoided Prosopis woodland. At the microhabi- woodlands (Cabrera, 1971). tat level, herb cover was the most important In this region, wild boar has been studied factor affecting wild boar presence, showing from a great variety of aspects, including habi- a positive association between this and the tat use, diet, climatic influence, and impacts abundance of signs. on vegetation composition and on physical, Based on wild boar feces, Cuevas et al. chemical and microbiological soil properties (2013b) found that the diet consisted of 96% (Campos and Ojeda, 1997; Cuevas et al., 2010; plant matter and 4% of animal matter. Herbs Cuevas, 2012; Cuevas et al., 2012; Cuevas et were the most frequently consumed food item al., 2013a; 2013b) (Table 1). Although wild (~50%) followed by woody species. Aerial parts boar is not physiologically adapted to arid en- were consumed more frequently during the vironments (Baber and Coblentz, 1986), they dry season, whereas during the wet season, have successfully colonized them worldwide, fruits and animal tissue were more frequent. such as in the deserts of USA, Australia, and Regarding trophic selection, herbs were the Argentina (Barrett, 1978; Saunders and Giles, only food item selected by wild boars, while 1995; Cuevas et al., 2010). For that reason, trees like algarrobo dulce (Prosopis fleuxosa) studies that help us understand the ecologi- were consumed as available only during the cal strategies that boars use in environments wet season, which is the season where this quite different from their native range may tree species bears fruit. They also found that yield insights about the traits and factors wild boar used food resources according to that constraint or facilitate the expansion of seasonal availability, observing a broader tro- invasive species. phic niche with higher plant diversity in the 248 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br wet season. Finally, the most consumed food vironmental temperatures (Rosell et al., 2001; items (fruits of Prosopis flexuosa, leaves of Dexter, 2003). In the Monte Desert, Cuevas malvisco [Sphaeralcea miniata], and bulbs of et al. (2013a) found that wild boars showed a papilla [Pitraea cuneato-ovata]) had high forage behavioral response related with daily move- quality and high carbohydrate contents, which ments patterns to increased environmental means immediate energy for the organism. temperature, but they did not find a strong Cuevas et al. (2013a) found that wild boars association with free water. used the habitat as a function of food avail- To sum up, ecological strategies of wild boar ability. This is because Larrea shubland, which in aridlands of Argentina where water resource was the only habitat selected by boars through is scarce and exposure to sun is high, shade rooting sign (their main way of finding food), could be essential for surviving. Therefore, it is associated with high herb cover, and herba- is necessary that wild boars minimize the ex- ceous plants were the most frequent food item posure to high temperatures and maximize the as well as preferred in their diet (Cuevas et al., food intake of high quality forage to maximize 2010; Cuevas et al., 2013b). High carbohydrate their energy input. input is considered important in the diet of an According to Campos and Ojeda (1997), wild individual as it is an essential component in boar causes damages by chewing nearly 100% of keeping the body in good physical condition ingested seeds of Prosopis flexuosa. While that and also for the accumulation of reserves to study was based on only 3 samples of feces, a be used during more critical periods (food more recent study (Cuevas, unpublished data) scarcity) and/or periods of highest energy has observed that of a total of 1618 seeds (39 demand (reproduction) (Abaigar, 1993). Fur- fecal samples), 30% had the entire (apparently thermore, around this protected area there are healthy) seed coat while the remainder 70% no croplands so the ingestion of energy-rich was damaged, either by boar chewing (17.3%), food is crucial for boar survival, particularly bruchid insects (31.7%), or other causes (21%). in arid conditions where the majority of plants Nevertheless, of all the “healthy” seeds (250), have high fiber content and low nutritional the author observed that only one had ger- value (Noy-Meir, 1973). This foraging strat- minated after one and a half months. Future egy enables wild boar to maximize energy studies are needed to understand the role of budget through food selection. Besides being wild boar in the life cycle of such a key spe- a generalist species (Rosell et al., 2001), in the cies as Prosopis fleuoxa in the Monte Desert. semiarid environment of the Monte Desert the Regarding the effect of wild boar rooting, wild boar appears as a species that selects both Cuevas et al. (2012) observed that disturbed space (habitat use) and food (herbs) (Cuevas patches were modified in physical, chemi- et al., 2013a; 2013b). cal and microbiological soil properties in Regarding climatic influences on wild boar a short-term (fresh disturbance) (Table 2). activity at local scales, Cuevas et al. (2013a) Regarding the impact on vegetation, those found a positive association between the authors found that wild boar activity through number of days with low temperature and the rooting produced a decrease in plant richness number of wild boar signs recorded in the and diversity, generating a negative effect Reserve. This means that the seasonal activ- on perennials such as tomillo (Acantolippia ity and/or daily movements of wild boars in seriphioides), jarilla (Larrea cuneifola) and periods or seasons of high temperature were llaullín (Lycium sp.), and on annuals such reduced. Thus, temperature could be a limit- as verbena (Glandularia mendocina), papa ing factor for wild boar activity, especially in del quirquincho (Heliotropium mendocinum), aridlands, because boars lack sweat glands or malvisco (Sphaeralcea miniata) and llantén other cooling physiological mechanisms for peludo (Plantago patagonica). These impacts maintaining hydric and thermal balance. They could occur either by mechanical action when require free water, shade, a diet rich in water, boars forage and many plants remain with their and/or a behavioral response to increased en- roots exposed, or by consumer action, because A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 249 many of these species were found in the diet in community composition and ecosystem (Cuevas et al., 2012; Cuevas et al., 2013b). functioning, resulting in many cases in the The only species that was favored by rooting disappearance of native species through pre- was Pitraea cuneato-ovata, which is an annual dation, competition for resources, spread of native species with high water requirements diseases, alteration of genetic diversity, habitat that grows in waterlogged and disturbed soils destruction, increased soil erosion, changes in (Stasi and Medero, 1983). This positive effect hydrology and nutrient cycles, disruption of on P. cuneato-ovata establishment could be due soil regimes, among other effects (Brown, 1989; to the change of soil properties by wild boar. Mack and D’Antonio, 1998; Byers et al., 2002; The high C/N ratio found in disturbed soils Lockwood et al., 2007). The study of invasive indicated that nitrogen mineralization was species in invaded habitats is necessary not faster in these soils (Cuevas et al., 2012). This only as a good opportunity to address topics could be because of the high soil moisture and such as basic processes in ecology but also the oxygenation (lower compaction) found in those invasion process, how ecosystems function, patches, or the incorporation of litter into the and to evaluate the effectiveness of population soil, which was found to be lower there (Cuevas management plans (Sax et al., 2007). et al., 2012). It should be pointed out that the In this review we could observe that although longer the time between processes of miner- wild boar is an omnivorous species whose alization and requirements of new vegetation, diet consists mainly of plant matter, it prefers the lower the efficiency of nutrient uptake and items rich in energy such as bulbs and fruits use (Abril, 2002). So when mineral nitrogen is of Pitraea cuneato-ovata, Prosopis flexuosa, released during periods without vegetation, it Araucaria araucana, and Butia yatay. These is subjected to loss by leaching or volatiliza- resources represent immediate energy for boars. tion (Abril, 2002). Thus, the high contents of Hence their main food strategy, especially mineral nitrogen found in disturbed soils could in arid and semi-arid ecosystems where the be lost due to rains, leaving the soil without majority of plants have low nutritional value the nitrogen needed for future plant growth due to their high fiber contents (Noy-Meir, (Cuevas et al., 2012). To sum up, Cuevas et al. 1973). Extreme conditions in arid and semi- (2012) concluded that the physical alteration of arid environments involve seasonal and spatial soil due to wild boar rooting has consequences variation of resources (van Horne et al., 1998) on its chemical properties. And these new which can have significant consequences on the soil characteristics could be responsible for a population dynamics of species, especially in reduced plant cover and less soil bulk density, periods of scarcity (Ostfeld and Keesing, 2000). which could increase soil degradation by wind High dietary intake of carbohydrates (e.g., in erosion. Even though this impact is at the mi- the fruits mentioned above) is expected to be crosite scale, disturbance by wild boars could compensated with high intake of protein from be another factor contributing to accelerating animal matter for proper nutrition (Schley and the desertification process in the Monte Desert Roper, 2003). For that reason wild boar would (Cuevas et al., 2012). be increasing the consumption of animal mat- ter in periods of fruiting when they eat more DISCUSSION AND CONCLUSIONS food rich in energy (Schley and Roper, 2003). This was observed in both Espinal and Monte Globally, invasive species have a significant provinces. This strategy was also observed in effect on both economic and environmen- places where the species is native and where tal systems (Vitousek et al., 1997). In many it is introduced (Barrett, 1978; Abaigar, 1993; countries, economic losses due to biological Massei et al., 1996; Schley and Roper, 2003). invasions have been and continue to be in the Regarding the Subantarctic region, future millions (Pimentel et al., 2001). At an ecologi- studies evaluating wild boar feeding habits, cal level, the establishment of new species to including their seasonal variation, could help us new environments has led to major changes understand why this species prefers Nothofagus 250 Mastozoología Neotropical, 23:239-254, Mendoza, 2016 MF Cuevas et al. http://www.sarem.org.ar - http://www.sbmz.com.br forests to other types of environments. Those While soils of Patagonian forest are derived forests likely offer not only shelter but also food, from volcanic ashes with high capacity of sta- because others studies have shown that wild bilizing soil organic matter, buffering pH, and boar habitat use is a function of food availability retaining P and water which confers high resis- (Barrett, 1982; Welander, 2000; Cuevas et al., tance to nutrient loss (Diehl et al., 2003), soils 2013a). Similar information on diet and habitat in Monte Desert have an inherent tendancy use of wild boar is also needed in the Pampean (fragility) to desertification attributable to an grassland province, where niche interactions interaction between the system’s own fragility and possible competition between this species due to aridity, erosive forces from water and and Pampas deer require evaluation. wind, salinization processes, and anthropic ac- The mesquite Prosopis spp. plays an important tions such as livestock pressure, logging, and role in the organization of animal and plant fire regime modification (Villagra et al., 2009). communities (Mares et al., 1977). In Monte Therefore, soils in the Subantarctic region may Desert, wild boar consume Prosopis flexuosa, a be better buffered to short-term disturbances key species for its provision of shelter, shadow, in comparison with Monte Desert soils, where and fruit to many native animals such as the presence of a new disturbance factor (wild the zorro gris or South American gray fox boar rooting) could have consequences such as (Lycalopex griseus), mara or patagonian hare increasing desertification. (Dolichotis patagonum), vizcacha or plains As we mentioned above, temperature and vizcacha (Lagostomus maximus), and domestic the availability of free water are two important animals such as cows and horses (Campos factors for wild boar population distribution and Ojeda, 1997). In another study, Lynes and and abundance. In several cases, when the Campbell (2000) reported that 70% of seeds of temperature is high wild boars are restricted American carob (Prosopis pallida) in the feces to areas of dense vegetation cover and close to of wild boar germinated. Future studies on the water resources (Dexter, 1998; Acevedo et al., viability and germination of seeds of Prosopis 2006). Cuevas et al. (2013a) observed that in the flexuosa in feces of wild boar are thus needed Monte Desert, daily movements of wild boars to understand the role of boars in the recruit- in periods or seasons of high temperature were ment of this key tree species in Monte Desert. reduced. Thus, at local scale, temperature could In addition, the impact of wild boar upon be a limiting factor in wild boar activity. At Yatay palm requires additional investigation. regional scale, wild boar could be also affected Boars may serve dual roles as possible seed dis- by temperature: in areas where temperature is persers (they defecate whole seeds upon eating low there is an increased presence of wild boar its fruit) and as predators upon Yatay seedlings, (Cuevas, unpublished data). Studies focused where during non-masting periods they dig on movements, activity pattern, home range, around the plant, leaving their roots exposed and reproductive capacity in different climatic and causing it to die (Ballari, 2013). Therefore, conditions are needed to understand why this wild boar may in fact reduce the recruitment species is so successful. of Yatay palm in EPNP. Future studies about Regarding management strategies of wild its role in the predation or dispersal of Yatay boar in Argentina, Ballari et al. (2015a) found palm are crucial to determine this. that 54% of the surveyed protected areas apply Regarding the impact of wild boar rooting some control method. Hunting was the most behavior, we found evidence suggesting no commonly used technique of wild boar control, substantial changes in soil properties in the a method that managers of protected areas (e.g., Subantarctic region, whereas in the Monte EPNP, Islas de Santa Fe National Park, Laguna Desert there were modifications of physical, de Llancanelo Natural Wildlife Reserve and chemical, and microbiological properties caused Campos del Tuyú National Park) have reported by rooting, and leading to wind erosion of soil. to be effective for reducing the boar population. These differences in the impact of rooting may However, the authors found that hunting and be tied to soil characteristics and resilience. in combination with others methods—such as A REVIEW OF WILD BOAR’S ECOLOGY AND ITS IMPACT IN ARGENTINA 251 traps—were actually ineffective and did not (scientists, government officials, managers of reduce the abundance of this invasive species. protected areas, landownwers) to plan a strat- In EPNP, the method of baiting the species with egy to control its populations, thus mitigating supplemental feeding (corn) could in fact have damage to native ecosystem and the productive the unintended consequence that boars more systems of the country. frequently use the protected area, rather than the private agricultural lands that surround ACKNOWLEDGEMENTS the park, due to the supplemental food being Special thanks to Nate Upham for assisting in the English available the whole year. A similar situation version of the manuscript and for his comments, to Lili- was found in Europe, where landowners bait ana Katinas for helping us with the figure and to Cecilia the wild boar to keep them in woodland areas Scoones and Roberto Kiesling for the assistance with the and protect their crops (Cellina, 2008). While common names of several herbs species. the current baiting strategy in EPNP benefits LITERATURE CITED landowners, it appears to be detrimental to the park’s objectives regarding the conservation of ABAIGAR T. 1993. Régimen alimentario del jabalí (Sus endemic species such as Butia yatay. Although scrofa) en el sureste Ibérico. Doñana Acta Vertebrata 20:35-48. the control method has been effective according ABRAHAM EM. 2001. Geomorfología y suelos. Pp: 123- to managers of the protected area (Ballari et al 126. In: El desierto del Monte: La reserva de Biosfera 2015a), it keeps wild boar population within de Ñacuñán (S Claver and S. Roig-Juñent, eds.). the park. 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