Effect of free-ranging cattle on mammalian diversity 885 Our findings indicate that high cattle abundance (relative
abundance ˃ 17) is incompatible with the presence of small mammals. Trampling and browsing by cattle reduce the het- erogeneity of the forest understorey for this group (Smith et al., 1996; Hayward et al., 1997). We could not identify small mammals to species level and the responses of indi- vidual species to cattle may vary (Schieltz & Rubenstein, 2016). Nevertheless, when present, small mammals had high relative abundance indices, so a prey base for medium-sized and small felids remains available. Small felids are indirectly affected by cattle, as indicated
by the human influence index. This index includes human population pressure, human land use and infrastructure and human access. Differential logging (i.e. logging of only par- ticular tree species and individuals of a certain size, resulting in impoverished species richness and affecting forest struc- ture) could also affect forest specialists such as the oncilla and margay. Therefore, we suggest direct hunting and asso- ciated activities, such as the presence of dogs (Perovic, 2002) and habitat transformation, are underlying explanations for the influence of humans on felids, not the presence of cattle directly.We did not record domestic cats, so these may not yet be a threat to small felids in the Yungas.We found that the Vulnerable oncilla had the lowest relative abundance and was the rarest of the felid group and therefore its con- servation status should be monitored. In agreement with Nanni (2015), we found brocket deer
only in areas with low cattle relative abundance. The human influence index did not affect these large herbivores, sug- gesting a direct effect of cattle on these two species. In agree- ment with Mazzini et al. (2018), we suggest biological inter- actions such as competition and dietary overlap between cattle and native herbivores are the cause of this effect. Species of conservation concern had low relative abun-
dance indices. Lowland tapir relative abundance was nega- tively influenced by distance to linear watercourses. Like the Malayan tapir Tapirus indicus and Baird’s tapir Tapirus baardii, which depend on proximity to water (Dudgeon, 2007; Reyna-Hurtado et al., 2019), lowland tapirs use water banks to browse and mate, and enter the water to take refuge from predators (Brooks et al., 1997). Lowland tapir relative abundance was not negatively influenced by cattle relative abundance index. Tapirs probably differ in feeding habits from cattle, browsing more on seedlings and fruits. Prey remains available for jaguars, but the ab- undance of wild prey is decreasing, which could provoke jaguars to start predating on cattle, with consequent escalations in human–predator conflict (Perovic, 2002; Cuyckens et al., 2014). The species of conservation concern considered here (lowland tapir, white-lipped peccary and jaguar) are large mammals, and body size is an indicator of extinction risk (Cardillo, 2005) as it determines suscepti- bility to hunting pressure and habitat selectivity. Our study was in areas that still have forest cover. Hence, the effects of
cattle and human activities such as habitat destruction and hunting for food (peccaries) and in retaliation (jaguars), are probably having negative, synergistic effects on species of conservation concern (Romero-Muñoz et al., 2020). These species depend on protected forests with extensive cover and protection against hunting. This is the first study based on an extensive camera-trap
survey to provide evidence that cattle affect the assemblage of mammals in the Austral Yungas, both directly and indir- ectly.Wehave provided guidelines for cattle abundance that should be implemented in protected areas where cattle rais- ing is allowed. However, we cannot provide guidelines for cattle abundance compatible with species of conservation concern (jaguar and white-lipped peccary), and their low abundances indicate their high risk of extinction in this region. Following Mazzini et al. (2018), we used directly measured cattle relative abundance at the local scale and complemented this with an indirect method at the regional scale (distribution modelling). Our work therefore provides both a method for future assessments of cattle impacts and an indicator of potential cattle abundance in unsurveyed areas in the Austral Yungas.
Acknowledgements We thank all field assistants and landowners who allowed us to conduct fieldwork, the Tinkunaku community, park rangers of the Secretariat of Environment and Sustainable Development of Salta province, the National Park Administration of Argentina; IDEA WILD for providing equipment; Leonidas Lizarraga for help downloading data; Agustin Abba for assistance with identification of ungulates; Tadeu de Oliveira for help with iden- tification of small felids; two anonymous reviewers for advice; and Martin Fisher and the teachers and classmates of the Conservation Leadership Programme Writing for Conservation Course in Colombia.
Author contributions Study conception: GAEC, LRM; study de- sign, fieldwork: GAEC, PGP; data input: GAEC, NGBT; data analysis: GAEC, writing: all authors.
Conflicts of interest None.
Ethical standards This research abided by the Oryx guidelines on ethical standards.
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Oryx, 2022, 56(6), 877–887 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321001538
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