Abstract:

Among the ecoregions that comprise the high-Andean zone of the Tropical Andes, the Puna and the Yungas stand out for covering a large part of the Peruvian Eastern Andes. Located in the ecotone of these two ecoregions, the Pampa Hermosa National Sanctuary (PHNS) houses one of the priority areas for conservation in Peru. However, the biodiversity of the high-Andean zone of the sanctuary and its surroundings remains poorly studied. Thus, through camera traps and transects, we sought to inventory for the first time the medium and large mammals from the high-Andean region of the PHNS and its buffer zone. We recorded 11 native and three domestic species of medium and large mammals. The richness of native mammals sampled reached 91.7% of the estimated richness (S est.= 11.99 ± 1.85). Among the native species Odocoileus virginianus had the highest relative frequency (56%). We recorded three endemic mammals from the Tropical Andes, Cuniculus taczanowskii, Tremarctos ornatus, and Pudu mephistopheles. The observed richness was higher than most surveys of medium and large mammals carried out in the Puna-forest ecotone in Peru, where the reported richness ranged from 4 to 13 species. In addition, our records of Leopardus pardalis and Eira barbara are the highest for all distribution of these two carnivores. Our results showed that more than 90% of the species recorded were found in it, demonstrating that the entire high-Andean region of the PHNS and its surroundings has an important value for the local mammal community.

Keywords
Camera-trap; Eira barbara; Leopardus pardalis; Odocoileus virginianus; Puna; Tropical Andes

Resumo:

Dentre as ecorregiões que compreendem a zona alto-andina dos Andes Tropicais, a Puna e as Yungas se destacam por abranger grande parte dos Andes Orientais peruanos. Localizado no ecótono entre estas duas ecorregiões, o Santuário Nacional Pampa Hermosa (SNPH) abriga uma das áreas prioritárias para conservação no Peru. No entanto, a biodiversidade da zona alto-andina do santuário e seu entorno permanece pouco estudada. Dessa forma, através de armadilhas fotográficas e transectos, buscamos inventariar pela primeira vez os mamíferos de médio e grande porte da região alto-andina do SNPH e de sua zona de amortecimento. Nós registramos 11 espécies nativas e três espécies domésticas de mamíferos de médio e grande porte. A riqueza observada dos mamíferos nativos atingiu 91,7% da riqueza estimada (S est.= 11,99 ± 1,85). Dentre as espécies nativas, Odocoileus virginianus foi a que apresentou a maior frequência relativa (56%). Registramos três mamíferos endêmicoss dos Andes Tropicais, Cuniculus taczanowskii, Tremarctos ornatus, e Pudu mephistopheles. A riqueza observada foi superior que a maioria dos levantamentos de mamíferos de médio e grande porte feitos em ecótono Puna-bosque no Peru, onde a riqueza reportada variou de 4 a 13 espécies. Além disso, os registros de Leopardus pardalis e Eira barbara são os mais elevados para toda distribuição destes dois carnívoros. Nossos resultados demonstraram que mais de 90% das espécies registradas foram encontradas nela, demonstrando que toda região alto-andina do SNPH e seu entorno tem um importante valor para a fauna de mamíferos local.

Palavras-chave:
Andes Tropicais; Armadilhas fotográficas; Eira barbara; Leopardus pardalis; Odocoileus virginianus; Puna

Introduction

Peru is considered as one of the most megadiverse countries in the world (Noss 1990NOSS, R. F. 1990. Indicators for monitoring biodiversity: a hierarchical approach. Conserv. Biol. 4(4):355-364. https://doi.org/10.1111/j.1523-1739.1990.tb00309.x
https://doi.org/10.1111/j.1523-1739.1990... , Shanee et al. 2017SHANEE, S., SHANEE, N., MONTEFERRI, B., ALLGAS, N., PARDO, A.A. & HORWICH, R.H. 2017. Protected area coverage of threatened vertebrates and ecoregions in Peru: Comparison of communal, private and state reserves. J. Environ. Manag. 202:12-20. https://doi.org/10.1016/j.jenvman.2017.07.023
https://doi.org/10.1016/j.jenvman.2017.0... ) and much of this biological diversity is due to the climatic and geographic influence created by the Andes Mountains and the Humboldt Maritime Current (Fajardo et al. 2014FAJARDO, J., LESSMANN, J., BONACCORSO, E., DEVENISH, C. & MUNOZ, J. 2014. Combined use of systematic conservation planning, species distribution modelling, and connectivity analysis reveals severe conservation gaps in a megadiverse country (Peru). PLoS One 9(12):e114367. https://doi.org/10.1371/journal.pone.0114367
https://doi.org/10.1371/journal.pone.011... ). The diversity of ecosystems and habitats created by these factors give rise to a scenario where the neotropical fauna finds countless opportunities for speciation and radiation (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ).

Among the countries with the greatest diversity of mammals, Peru is in the fourth place, with 569 species (Pacheco et al. 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ), only behind Brazil within the neotropical region (SERFOR 2018SERFOR- Servicio Nacional Forestal y de Fauna Silvestre. 2018. Libro Rojo de la Fauna Silvestre Amenazada del Perú. 1 ed. SERFOR, Lima. https://www.serfor.gob.pe/portal/wp-content/uploads/2018/10/Libro-Rojo.pdf (Last access on 07/05/2021).
https://www.serfor.gob.pe/portal/wp-cont... ). Of the 10 large ecoregions existing in Peru (Brack-Egg 1986BRACK-EGG, E. 1986. Las Ecorregiones del Perú. Bol. de Lima 44:57-70.), the Selva Baixa and the Yungas (or Selva Alta) have the highest rates of mammalian diversity and endemism, respectively (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ). However, with a significant mammal diversity (63 spp.) (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ), the Puna is the dominant ecoregion of the Peruvian high-Andean zone, rising from 3,000-3,500 m to 4,500-5,000 m (Rolando et al. 2017ROLANDO, J.L., TURIN, C., RAMÍREZ, D.A., MARES, V., MONERRIS, J. & QUIROZ, R. 2017. Key ecosystem services and ecological intensification of agriculture in the tropical high-Andean Puna as affected by land-use and climate changes. Agric., Ecosyst. Environ. 236:221-233. https://doi.org/10.1016/j.agee.2016.12.010
https://doi.org/10.1016/j.agee.2016.12.0... ). The Puna is characterized by high-altitude grasslands, and it is subdivided into two ecosystems based on rainfall: (1) Dry Puna, which spans the high plateaus of southern Peru; and (2) Wet Puna, which is found from the center-south to the center-north of the Peruvian Andes (Josse et al. 2009JOSSE, C., CUESTA, F., NAVARRO, G., BARRENA, V., CABRERA, E., CHACÓN-MORENO, E., FERREIRA, W., PERALVO, M., SAITO, J. & TOVAR, A. 2009. Ecosistemas de los Andes del Norte y Centro. Bolivia, Colombia, Ecuador, Perú. Intercooperation, CONDESAN-Proyecto Páramo Andino, Programa BioAndes, EcoCiencia, NatureServe, IAvH, LTA-UNALM, ICAEULA, CDC-UNALM, RUMBOL SRL, Lima. http://www.saber.ula.ve/handle/123456789/39336 (Last access on 10/05/2021).
http://www.saber.ula.ve/handle/123456789... ). In the ecotone regions between the Wet Puna and the Cloud Forests of Yungas, a long transition zone is formed between grassland and forest environments, where there is a great sharing of the native fauna and flora of these two ecoregions (Simpson 1983SIMPSON, B. B. 1983 An historical phytogeography of the high Andean flora. Ver. Chilena Hist. Nat. 56:109-122. http://rchn.biologiachile.cl/pdfs/1983/2/Simpson_1983.pdf (Last access on 11/05/2021).
http://rchn.biologiachile.cl/pdfs/1983/2... , Buytaert et al. 2011BUYTAERT, W., CUESTA‐CAMACHO, F. & TOBÓN, C. 2011. Potential impacts of climate change on the environmental services of humid tropical alpine regions. Glob. Ecol. Biogeogr. 20(1):19-33. https://doi.org/10.1111/j.1466-8238.2010.00585.x
https://doi.org/10.1111/j.1466-8238.2010... ).

Because they are regions with extremely rugged terrain, adverse climatic conditions, and difficult access (Jiménez et al. 2010JIMÉNEZ, C.F., QUINTANA, H., PACHECO, V., MELTON, D., TORREALVA, J. & TELLO, G. 2010. Camera trap survey of medium and large mammals in a montane rainforest of northern Peru. Rev. Peru. Biol. 17(2):191-196. http://sisbib.unmsm.edu.pe/BVRevistas/biologia/biologiaNEW.htm
http://sisbib.unmsm.edu.pe/BVRevistas/bi... ), these high-Andean zones of the Peruvian Eastern Andes are poorly studied (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , Medina et al. 2012MEDINA, C.E., ZEBALLOS, H. & LÓPEZ, E. 2012. Diversidad de mamíferos en los bosques montanos del Valle de Kcosñipata, Cusco, Perú. Mastozool. Neotrop. 19(1):85-104. https://www.redalyc.org/pdf/457/45723408008.pdf
https://www.redalyc.org/pdf/457/45723408... ) and, consequently, the status of biological diversity remains with several local gaps in most of these Andean areas. In this sense, the Protected Areas are essential tools for the conservation of this Andean biodiversity to be explored and, therefore, have a fundamental role in the scientific and ecological development of the country (Shanee et al. 2017SHANEE, S., SHANEE, N., MONTEFERRI, B., ALLGAS, N., PARDO, A.A. & HORWICH, R.H. 2017. Protected area coverage of threatened vertebrates and ecoregions in Peru: Comparison of communal, private and state reserves. J. Environ. Manag. 202:12-20. https://doi.org/10.1016/j.jenvman.2017.07.023
https://doi.org/10.1016/j.jenvman.2017.0... , Bax & Francesconi 2019BAX, V. & FRANCESCONI, W. 2019. Conservation gaps and priorities in the Tropical Andes biodiversity hotspot: Implications for the expansion of protected areas. J. Environ. Manag. 232:387-396. https://doi.org/10.1016/j.jenvman.2018.11.086
https://doi.org/10.1016/j.jenvman.2018.1... ).

In the Peruvian Andes only 36% of the biological diversity existing is protected by the Protected Areas system (Fajardo et al. 2014FAJARDO, J., LESSMANN, J., BONACCORSO, E., DEVENISH, C. & MUNOZ, J. 2014. Combined use of systematic conservation planning, species distribution modelling, and connectivity analysis reveals severe conservation gaps in a megadiverse country (Peru). PLoS One 9(12):e114367. https://doi.org/10.1371/journal.pone.0114367
https://doi.org/10.1371/journal.pone.011... ). This becomes even more worrying given the fact that Peru is the country with the largest extension of the Tropical Andes, one of the main hotspots of global biodiversity and holder of the highest rate of endemism in the world (Myers et al. 2000MYERS, N., MITTERMEIER, R.A., MITTERMEIER, C.G., DA FONSECA, G.A. & KENT, J. 2000. Biodiversity hotspots for conservation priorities. Nature 403(6772):853-858. https://doi.org/10.1038/35002501
https://doi.org/10.1038/35002501... , Josse et al. 2011JOSSE, C., CUESTA, F., NAVARRO, G., BARRENA, V., BECERRA, M.T., CABRERA, E., CHACÓN-MORENO, E., FERREIRA, W., PERALVO, M., SAITO, J. & TOVAR, A. 2011. Physical geography and ecosystems in the tropical Andes. In Climate Change and Biodiversity in the Tropical Andes (S. K. Herzog, R. Martínez, P. M. Jørgensen & H. Tiessen, eds.). Inter-American Institute for Global Change Research (IAI) and Scientific Committee on Problems of the Environment (SCOPE), São José dos Campos, p. 152-169. http://www.saber.ula.ve/handle/123456789/39237 (Last access on 12/05/2021).
http://www.saber.ula.ve/handle/123456789... ).

Located in the Peruvian Central Andes, the Pampa Hermosa National Sanctuary (PHNS) houses an important transitional area of the Puna and Yungas ecoregions of the department of Junín (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). Despite being considered a priority area for the conservation of Andean-Amazonian biodiversity (SERNANP 2009, 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... , Arias et al. 2016ARIAS, E., PACHECO, V., CERVANTES, K., AGUILAR, A. & ÁLVAREZ, J. 2016. Diversidad y composición de murciélagos en los bosques montanos del Santuario Nacional Pampa Hermosa, Junín, Perú. Rev. Peru. Biol. 23(2):103-116. http://dx.doi.org/10.15381/rpb.v23i2.12381
http://dx.doi.org/10.15381/rpb.v23i2.123... ), few expeditions sought to systematically inventory the local biological diversity, especially in the high-Andean zone (> 2,500 m) from the sanctuary, due to its difficult access. Historically, the fauna of the region was first studied in the mid-19th century by naturalist Johann Jakob von Tschudi in the Montaña de Vitoc (Tschudi, 1844aTSCHUDI, J.J. 1844a. Mammalium conspectus quae in Republica Peruana reperiuntur et pleraque observata vel collecta sunt in itinerere. Archiv für Naturgeschichte 10: 244-255., 1844bTSCHUDI, J. J. 1844b. Untersuchungen über die Fauna Peruana: Therologie. Scheitlin und Zollikofer, St. Gallen.). In this pioneering expedition, fish, amphibians, reptiles, birds and mammals were collected (Serrano-Villavicencio et al. 2020SERRANO-VILLAVICENCIO, J.E., BARTOLETTI, T. & BUENO, C. 2020. Mammals collected by Johann Jakob von Tschudi in Peru during 1838 1842 at Muséum d’histoire naturelle de Neuchâtel. Bol. Mus. Para. Emílio Goeldi. 15(3):905-931. https://doi.org/10.46357/bcnaturais.v15i3.249
https://doi.org/10.46357/bcnaturais.v15i... ), mainly in the Yungas ecoregion, about 20km away from the PHNS. In the recent decades, among the groups already systematically inventoried in the PHNS and its surroundings, there are insects, birds (Silva et al. 2016SILVA, F.Z., VALLADARES, N.A., FLORES, M.A., CARBONEL, D.C.U., MAURICIO, N.N., BENAVIDES, F.P., RAMOS, K.P., ÑAUPA, A.Q., AVILA, C.S., HUAMANHUILLCA, S.T. & HUAMÁN, R.Y. 2016. Línea base de la flora y fauna del Santuario Nacional Pampa Hermosa - Sector San Damián. Rev. Cienc., Tecnol. Humanid. 7(2):11-38. http://revistas.unprg.edu.pe/openjournal/index.php/revistacientifica/article/view/57/37 (Last access on 05/05/2021).
http://revistas.unprg.edu.pe/openjournal... ), bats (Arias et al. 2016ARIAS, E., PACHECO, V., CERVANTES, K., AGUILAR, A. & ÁLVAREZ, J. 2016. Diversidad y composición de murciélagos en los bosques montanos del Santuario Nacional Pampa Hermosa, Junín, Perú. Rev. Peru. Biol. 23(2):103-116. http://dx.doi.org/10.15381/rpb.v23i2.12381
http://dx.doi.org/10.15381/rpb.v23i2.123... ) and plants (La Torre-Cuadros et al. 2007LA TORRE-CUADROS, M.Á., HERRANDO-PÉREZ, S. & YOUNG, K.R. 2007. Diversity and structural patterns for tropical montane and premontane forests of central Peru, with an assessment of the use of higher-taxon surrogacy. Biodivers Conserv. 16:2965-2988. https://doi.org/10.1007/s10531-007-9155-9
https://doi.org/10.1007/s10531-007-9155-... , Silva et al. 2016SILVA, F.Z., VALLADARES, N.A., FLORES, M.A., CARBONEL, D.C.U., MAURICIO, N.N., BENAVIDES, F.P., RAMOS, K.P., ÑAUPA, A.Q., AVILA, C.S., HUAMANHUILLCA, S.T. & HUAMÁN, R.Y. 2016. Línea base de la flora y fauna del Santuario Nacional Pampa Hermosa - Sector San Damián. Rev. Cienc., Tecnol. Humanid. 7(2):11-38. http://revistas.unprg.edu.pe/openjournal/index.php/revistacientifica/article/view/57/37 (Last access on 05/05/2021).
http://revistas.unprg.edu.pe/openjournal... ). However, all these surveys were carried out in the lower part (< 2,000 m) of the sanctuary and its buffer zone.

Thus, the biodiversity of the high-Andean zone of the sanctuary and its surroundings remains poorly studied, especially with regard to the terrestrial mammals community. In the PHNS buffer zone, there are several reports of conflict between the communities in the San Pedro de Churco village and the Andean bear (Tremarctos ornatus) (Rojas-Vera Pinto & Butrón 2016ROJAS-VERA PINTO, R., & BUTRÓN, R. 2016. Diagnóstico de la interacción humano-oso andino en el Santuario Nacional Pampa Hermosa. Informe de trabajo. SERFOR, Lima., Rojas-Vera Pinto 2019), signaling the presence of important endangered species in the sanctuary area. For this reason, in this present study, we sought to inventory, for the first time, the medium and large mammals from the high-Andean zone of the PHNS and its buffer zone. In addition, we compared our observed richness with that of other surveys carried out in similar altitudinal ranges in the Peruvian Eastern Andes.

Material and Methods

1. Study area

The study was carried out in the PHNS (75º35’09” - 75º24’43”W; 10º58’53” - 11º06’30”S) and in its buffer zone, concentrating on the west side of sanctuary, where the S.P. Churco village (75º33’24”W; 11º02’13”S) is located (Figure 1). The site is situated on the eastern flanks of the Tropical Andes, in the department of Junín, central Peru (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). The area of the PHNS is 11,543 ha and its buffer zone is 20,292 ha, totaling 31,835 ha of direct and indirect protection of the Andean-Amazonian Peruvian mountain ecosystems along a wide altitudinal range (1,130 to 4,080 m) (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). The study area was concentrated in the high-Andean zone of the sanctuary and its surroundings, covering an area of approximately 2,500 ha, between 3,200 and 3,700 m.

Figure 1
a) Location of Peru in South America and the study site; b) Location of department of Junín (light green) in Peru and the study site; c) Ecoregions covering the Pampa Hermosa National Sanctuary (PHNS) and its buffer zone and location of the S.P. Churco village; d) Ecosystems and land cover of the high-Andean region of the PHNS and its buffer zone and location of the 16 sampling units in the study area.

The study area is located in an ecotone, where two ecosystems belonging to distinct ecoregions are found (Pulgar-Vidal 1987PULGAR-VIDAL, J. 1987. Geografía del Perú: las ocho regiones naturales: la regionalización transversal: la micro regionalización. Peisa, Lima.): (1) the Wet Puna of the Puna ecoregion, extending from 3,250 to 5,000 m, characterized by high-Andean grasslands with herbaceous-shrubby vegetation padded with mosses and grasses arranged in clumps (Brack 1986BRACK, A. 1986. Ecología de un país complejo. Gran geografía del Perú. Nat. y Hombre 2:175-319., López 2010LÓPEZ, R. P. 2010. Los Pisos de vegetación y la flora. In: Transitando la diversidad, Paisajes naturales y culturales: La Paz. TROPICO-Asociación Boliviana para la conservación/Gobierno Municipal de La Paz/FUNDESNAP- Fundación para el Desarrollo del Sistema Nacional de Áreas Protegidas, La Paz, p. 21-35.) and; (2) the Cloud Forests (or High-Montane Forests) of the Yungas ecoregion, present from 2,500 to 3,300 m, with dense, steep forest vegetation and with a great abundance of epiphytes (Pulgar-Vidal 1987PULGAR-VIDAL, J. 1987. Geografía del Perú: las ocho regiones naturales: la regionalización transversal: la micro regionalización. Peisa, Lima., SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ).

The climate of the PHNS and its buffer zone presents a gradual variation along its altitudinal ranges (Junquas et al. 2018JUNQUAS, C., TAKAHASHI, K., CONDOM, T., ESPINOZA, J.C., CHÁVEZ, S., SICART, J.E. & LEBEL, T. 2018. Understanding the influence of orography on the precipitation diurnal cycle and the associated atmospheric processes in the central Andes. Clim. Dyn. 50(11):3995-4017. https://doi.org/10.1007/s00382-017-3858-8
https://doi.org/10.1007/s00382-017-3858-... , Eghdami & Barros 2019EGHDAMI, M., & BARROS, A.P. 2019. Extreme orographic rainfall in the eastern Andes tied to cold air intrusions. Front. Environ. Sci. 7:101. https://doi.org/10.3389/fenvs.2019.00101
https://doi.org/10.3389/fenvs.2019.00101... ). The highest part of the sanctuary where the Wet Puna and the S.P. Churco village are located corresponds to the climate of High-Montane Dry Tundra (ETH) according to Köppen (1936) (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). The average annual precipitation varies from 2,000 to 3,000 mm, with greater concentration in the summer and with an average annual temperature of 6 to 10ºC (Yarupaitan & Giraldo 2007YARUPAITAN, G. & GIRALDO, U.F. 2007. Expediente Técnico de Categorización de la Zona Reservada Pampa Hermosa. Ministerio de Agricultura. Instituto Nacional de los Recursos Naturales. Intendencia de Áreas Naturales Protegidas, Lima. http://repositorio.unas.edu.pe/handle/UNAS/642 (Last access on 30/04/2021).
http://repositorio.unas.edu.pe/handle/UN... , SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). As the altitude decreases to 2,600 m, there is a zone that comprises, the Continental Humid Boreal (Dwb) climate (Köppen 1936). This range also presents low average annual temperatures, ranging from 10 to 12ºC and high average annual precipitation between 3,000 and 4,000 mm (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... , Eghdami & Barros 2019EGHDAMI, M., & BARROS, A.P. 2019. Extreme orographic rainfall in the eastern Andes tied to cold air intrusions. Front. Environ. Sci. 7:101. https://doi.org/10.3389/fenvs.2019.00101
https://doi.org/10.3389/fenvs.2019.00101... ).

2. Data collection

We distributed 16 sampling points in the study site, eight inside the PHNS and eight in its buffer zone (Figure 1). We used two complementary sampling methods, camera traps and active search through transects, at a sampling unit scale of 1 km². Each sampling point was centered on a camera trap (Bushnell Trophy Cam HD 2018 ©), respecting a minimum distance of 1 km between them, with a maximum deviation of 20% being accepted, due to inaccessibility and difficulty of installation in some areas. We installed the camera traps in places commonly used by mammals through the recognition of signs and animal tracks and no bait was used to avoid the artificial attraction of animals (Srbek-Araújo & Chiarello 2013SRBEK-ARAÚJO, A.C. & CHIARELLO, A.G. 2013. Influence of camera-trap sampling design on mammal species capture rates and community structures in southeastern Brazil. Biota Neotrop. 13(2):51-62. https://doi.org/10.1590/S1676-06032013000200005
https://doi.org/10.1590/S1676-0603201300... ). We configured the equipment to capture three consecutive photos every 10 seconds, whenever the motion sensor was triggered. We carried out the maintenance of the camera traps every six months to change the batteries and collect the stored data. The sampling time was one year and six months (May/2019 - November/2020), a sampling effort of 4,402 traps/day.

For the methodology of active search for transects, we covered 1 km trails (of people and animals) in each of the 16 sample units, in search of mammal tracks (e.g., feces, tracks, food consumed, marks on trees) and direct visualization. The tracks found along each transect were registered and georeferenced. We carried out three visits to each sampling unit (1st: May/2019; 2nd: October/2019; 3rd: November/2020), totaling approximately 48 km of transects covered within the sample units.

3. Data analysis

For the independence of the records obtained by camera traps, we considered an interval of 1 h between the photographic records of all mammals with more than 1 kg (Srbek-Araújo & Chiarello 2005SRBEK-ARAÚJO, A.C. & CHIARELLO, A.G. 2005. Is camera-trapping an efficient method for surveying mammals in Neotropical forests? A case study in south-eastern Brazil. J. Trop. Ecol. 21(1):121-125. https://doi.org/10.1017/S0266467404001956
https://doi.org/10.1017/S026646740400195... ). In order to measure the sampling sufficiency of the survey, we constructed species rarefaction curves using the Estimate S 9.1.0 program (Colwell 2013) with the Chao 2 estimator. For this purpose, we used the independent records obtained through camera traps and transects, using one week as the sampling effort unit (n=76). We also calculated the relative frequency (RF) by camera traps for each taxon using the formula given in percentage: (nº records of the species/nº total records) x 100.

The scientific nomenclature used followed Pacheco et al. (2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ) and Nascimento et al. (2021)NASCIMENTO, F.O., CHENG, J. & FEIJÓ, A. 2021. Taxonomic revision of the pampas cat Leopardus colocola complex (Carnivora: Felidae): an integrative approach. Zool. J. Linn. Soc. 191(2):575-611. https://doi.org/10.1093/zoolinnean/zlaa043
https://doi.org/10.1093/zoolinnean/zlaa0... . For species’ identifications we followed Eisenberg & Redford (1989), Pereira & Aprile (2012)PEREIRA, J. & APRILE, G. 2012. Felinos de Sudamérica: uma guia de identificación integral. Londaiz Laborde Ediciones, Buenos Aires., and Duarte & González (2010)DUARTE, J. M. B., & GONZÁLEZ, S. (eds.). 2010 Neotropical cervidology: biology and medicine of Latin American deer. Funep/IUCN, Jaboticabal.. We also consulted specialists in cases of uncertainty in the identifications. For the identification of tracks, we used the guides of Torres (2011)TORRES, D. 2011. Guía Básica para la identificación de señales de presencia de oso frontino (Tremarctos ornatus) en los Andes Venezolanos. Fundación Andígena, Mérida. https://www.andigena.org/descargas/Torres_D_2006_Guia_Senales_Oso_Andino.pdf (Last access on 27/04/2021).
https://www.andigena.org/descargas/Torre... and Becker & Dalponte (2013)BECKER M. & DALPONTE, J.C. 2013. Rastros de mamíferos silvestres brasileiros - Guia de campo. 3 ed. Technical Books, Rio de Janeiro.. The threat status of native species at the national level followed the Red List of Threatened Species of Peru (SERFOR 2018SERFOR- Servicio Nacional Forestal y de Fauna Silvestre. 2018. Libro Rojo de la Fauna Silvestre Amenazada del Perú. 1 ed. SERFOR, Lima. https://www.serfor.gob.pe/portal/wp-content/uploads/2018/10/Libro-Rojo.pdf (Last access on 07/05/2021).
https://www.serfor.gob.pe/portal/wp-cont... ), and at the global level followed the International Union for Conservation of Nature’s Red List of Threatened Species (IUCN 2021IUCN - International Union for the Conservation of Nature. 2021. The IUCN Red List of Threatened Species. Version 2021-1. http://www.iucnredlist.org (Last access on 28/04/2021).
http://www.iucnredlist.org... ).

Results

We recorded 11 native and three domestic species of medium and large mammals distributed in five orders and nine families (Table 1 and Figures 2, 3). We obtained 525 independent records through camera traps of 10 native species and three domestic mammal species in the study area. By the transect method, we obtained 70 records from four species of native mammals and three domestic species, with Puma concolor being the only species recorded by feces alone.

Figure 2
Medium and large native mammals recorded by camera traps in the high-Andean region of the Pampa Hermosa National Sanctuary and its buffer zone, departament of Junín, Peru: a) Lycalopex culpaeus; b) Leopardus garleppi; c) Leopardus pardalis; d) Tremarctos ornatus; e) Conepatus chinga; f) Pudu mephistopheles; g) Odocoileus virginianus; h) Cuniculus taczanowskii.

Figure 3
Medium and large sized mammals recorded through tracks and direct views in the high-Andean region of the Pampa Hermosa National Sanctuary and its buffer zone, department of Junín, Peru: a) Mustela frenata carcass; b) bromeliad of the genus Puya eaten by Tremarctos ornatus; c) T. ornatus feces; d) Puma concolor feces; e) Creole Bos taurus grazing native vegetation; f) Equus caballus grazing in the high-Andean grasslands.

The richness of native mammals sampled (S obs.= 11) reached 91.7% of the estimated richness (S est.= 11.99 ± 1.85), showing a stabilization of the rarefaction curve from the 40th week onwards (Figure 4). The order Carnivora was the most representative, accounting for 72.7% of the native species recorded, followed by Artiodactyla and Rodentia. The native species with the highest total RF was Odocoileus virginianus (25.5%), followed by Conepatus chinga (7.8%) and Lycalopex culpaeus (7.0%). In addition, 90.9% of native mammals were registered in the PHNS buffer zone, surpassing the proportion found within the sanctuary boundaries, which was 63.6% (Table 1).

Figure 4
Rarefaction curve of medium and large mammals recorded in the high-Andean region of the Pampa Hermosa National Sanctuary and its buffer zone, department of Junín, Peru. Legend: Obs.: total observed richness; S est.: richness estimated by Chao 2 (95% confidence interval - red dashed lines).

The proportion of endemic species of Tropical Andes recorded in the area was 27.2% (Tremarctos ornatus, Pudu mephistopheles and Cuniculus taczanowskii). Only two species of mammals surveyed are categorized as Vulnerable, T. ornatus and P. mephistopheles; nonetheless, only the former species is also listed as Vulnerable globally. On the other hand, P. mephistopheles together with Leopardus garleppi present deficient data for a categorization of their conservation status globally. In addition, two species (P. concolor and C. taczanowskii) are listed as Near Threatened within Peruvian territory. Finally, 63.6% of recorded native mammals have a status of Least Concern as pertains their conservation at a global level, 72.7% of them show a trend towards population reduction according to the IUCN (2021)IUCN - International Union for the Conservation of Nature. 2021. The IUCN Red List of Threatened Species. Version 2021-1. http://www.iucnredlist.org (Last access on 28/04/2021).
http://www.iucnredlist.org... .

The domestic species Bos taurus had the highest number of photographic records among all the surveyed mammals and, consequently, the highest RF (31.04%), in addition to being found both inside the PHNS and in its buffer zone. In total, native species obtained 45.24% of photographic records against 54.75% of records of domestic species (Table 1).

Discussion

The present study is the first and only systematic survey that sought to inventory the medium and large mammals present in the PHNS and its buffer zone. The richness of mammals was higher than most surveys of medium and large mammals carried out between the Wet Puna and Cloud Forests (2,000 - 4,000 m) in Peruvian Eastern Andes, where the reported richness ranged from 4 to 13 species (Shanee & Shanee 2018SHANEE, S. & SHANEE, N. 2018. Diversity of large mammals in the Marañón-Huallaga landscape, Peru: with notes on rare species. Zool. Ecol. 28(4):313-328. https://doi.org/10.1080/21658005.2018.1516277
https://doi.org/10.1080/21658005.2018.15... , López 2020LÓPEZ, N. M. 2020. Fototrampeo de mamíferos medianos y grandes durante el periodo de cuarentena-aislamiento social por la pandemia del COVID-19 en el Sector San Alberto del Parque Nacional Yanachaga Chemillén, Perú. Notas sobre Mamíferos Sudam. 1:1-10. http://doi.org/10.31687/saremNMS.20.0.38
http://doi.org/10.31687/saremNMS.20.0.38... ) (Table 2). Taking into account the altitude range of the surveys, only studies carried out in regions with a difference greater than or equal to 1,000 m obtained a higher number of species than in our study (Medina et al. 2012MEDINA, C.E., ZEBALLOS, H. & LÓPEZ, E. 2012. Diversidad de mamíferos en los bosques montanos del Valle de Kcosñipata, Cusco, Perú. Mastozool. Neotrop. 19(1):85-104. https://www.redalyc.org/pdf/457/45723408008.pdf
https://www.redalyc.org/pdf/457/45723408... , Shanee & Shanee 2018SHANEE, S. & SHANEE, N. 2018. Diversity of large mammals in the Marañón-Huallaga landscape, Peru: with notes on rare species. Zool. Ecol. 28(4):313-328. https://doi.org/10.1080/21658005.2018.1516277
https://doi.org/10.1080/21658005.2018.15... ), which was performed at an altitudinal amplitude of only 500 m (Table 2). Furthermore, in most of aforementioned surveys the study areas are located mostly in the Yungas region, which has a known greater diversity of mammals (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ). Unlike our study area, where the Puna ecoregion is dominant, there is a recognizably lower diversity of mammals compared to the Yungas (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ). In addition, the present study is one of the few surveys that used only primary data in its methodology compared to other studies conducted in similar altitudinal ranges in the Peruvian Eastern Andes (Table 2).

The richness of medium and large mammals found in the high-Andean region of the PHNS and its buffer zone represents about 65% of the native mammal species of this group with known distribution for the study area (17 spp.) (Bernal 2016BERNAL, N. 2016. Lagidium viscacia. The IUCN Red List of Threatened Species 2016: e.T11148A22190789. https://dx.doi.org/10.2305/IUCN.UK.2016-2.RLTS.T11148A22190789.en. (Last access on 23/04/2021).
https://dx.doi.org/10.2305/IUCN.UK.2016-... , Cassola 2016CASSOLA, F. 2016. Sciurus spadiceus (errata version published in 2017). The IUCN Red List of Threatened Species 2016: e.T20022A115155594. https://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T20022A22246240.en. (Last access on 23/04/2021).
https://dx.doi.org/10.2305/IUCN.UK.2016-... , Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ). Thus, six potential species were not recorded for the area: taruca (Hippocamelus antisensis), llama (Lama glama), Andean cat (Leopardus jacobita), southern mountain viscacha (Lagidium viscacia), hairy long-nosed armadillo (Dasypus pilosus), and Andean opossum (Didelphis pernigra). The first four species are typically found in higher rocky areas of the Andes (FAO 2005FAO - Food and Agriculture Organization. 2005. Situación actual de los camélidos sudamericanos del Perú. Proyecto de Cooperación Técnica en apoyo a la crianza y al aprovechamiento. Camélidos Sudamericanos en la Región Andina TCP/RLA/2914. UNICEF/FAO, Lima., Barrio 2010BARRIO, J. 2010. Taruka Hippocamelus antisensis (d’Orbigny 1834). Neotropical cervidology: biology and medicine of Neotropical deer (J. M. B. Duarte & S. González, eds.). FUNEP/IUCN, Jaboticabal., Cossíos et al. 2012aCOSSÍOS, E.D., WALKER, R.S., LUCHERINI, M., RUIZ-GARCÍA, M. & ANGERS, B. 2012a. Population structure and conservation of a high-altitude specialist, the Andean Cat Leopardus jacobita. Endanger. Species Res. 16(3):283-294. https://doi.org/10.3354/esr00402
https://doi.org/10.3354/esr00402... , Bernal 2016BERNAL, N. 2016. Lagidium viscacia. The IUCN Red List of Threatened Species 2016: e.T11148A22190789. https://dx.doi.org/10.2305/IUCN.UK.2016-2.RLTS.T11148A22190789.en. (Last access on 23/04/2021).
https://dx.doi.org/10.2305/IUCN.UK.2016-... ), making it difficult to record these mammals in transition zones between Puna and Yungas (J. Barrio 2021, personal communication), especially taruca, llama and southern mountain viscacha that only occur in the Puna (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ). The hairy long-nosed armadillo and the Andean opossum are found mainly in the Yungas (Pacheco et al. 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ), so possibly the presence of only 18% of our camera traps in the Cloud Forests of the Yungas may have rendering the recording of these two species in the area difficult.

Regarding the species richness of the two ecoregions in which the study area is located, we recorded 41.1% of the community of medium and large mammals known to occur in the Peruvian Puna ecoregion (17 spp.) (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ). While, in relation to the Cloud Forests (>2,500 m) of the Peruvian Yungas, we found 40.7% of medium and large mammals known for this ecosystem (27 spp.) (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ). Based on literature, P. mephistopheles, L. pardalis and E. barbara were only known to occur in the Yungas and other forest ecosystems in Peru (Sanborn 1953, Hurtado et al. 2016, Shanee & Shanee 2018SHANEE, S. & SHANEE, N. 2018. Diversity of large mammals in the Marañón-Huallaga landscape, Peru: with notes on rare species. Zool. Ecol. 28(4):313-328. https://doi.org/10.1080/21658005.2018.1516277
https://doi.org/10.1080/21658005.2018.15... , Pacheco et al. 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ). Nonetheless, we recorded theses three species in areas of Puna (Table 3). These are the first well-documented and georeferenced records within the Peruvian territory that confirm the presence of these three species in this Andean ecoregion. These records demonstrate the ecological plasticity of these typically forest mammals in the use of different types of environments for foraging and obtaining resources (Lyra-Jorge et al. 2008aLYRA-JORGE, M.C., CIOCHETI, G. & PIVELLO, V.R. 2008a. Carnivore mammals in a fragmented landscape in northeast of São Paulo State, Brazil. Biodivers. Conserv. 17(7):1573-1580. https://doi.org/10.1007/s10531-008-9366-8
https://doi.org/10.1007/s10531-008-9366-... , Escamilo et al. 2010ESCAMILO, L.L., BARRIO, J., BENAVIDES, J. & TIRIRA, D. 2010. Northern Pudu, Pudu mephistophiles (De Winton 1896). In Neotropical cervidology: biology and medicine of Neotropical deer (J.M.B. Duarte & S. González, eds.). FUNEP/IUCN, Jaboticabal, p. 133-139., Lima et al. 2020LIMA, K.C.B., PASSAMANI, M. & ROSA, C. 2020. Daily tayra (Eira barbara, Linnaeus 1758) activity patterns and habitat use in high montane tropical forests. Acta Oecol. 108:103624. https://doi.org/10.1016/j.actao.2020.103624
https://doi.org/10.1016/j.actao.2020.103... , Pasa et al. 2021PASA, J.B., ARRAIS, R.C., MASSARA, R.L., PEREIRA, G. & DE AZEVEDO, F.C.C. 2021. Factors influencing the habitat use by ocelots in one of the last large Atlantic Forest remnants in southeastern Brazil. Ecol. Evol. 11(9):4631-4643.).

In addition, we recorded the species P. mephistopheles, L. pardalis and E. barbara at the highest altitudes known to the literature in Peruvian territory (Pacheco et al. 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ) (Table 3). For P. mephistopheles the increment was 56 m (3,506 m) in relation to the previous record of highest altitude of the species in Peru (3,450 m) (Shanee & Shanee 2018SHANEE, S. & SHANEE, N. 2018. Diversity of large mammals in the Marañón-Huallaga landscape, Peru: with notes on rare species. Zool. Ecol. 28(4):313-328. https://doi.org/10.1080/21658005.2018.1516277
https://doi.org/10.1080/21658005.2018.15... ). However, throughout its distribution, this small deer has already been found at 4,500 m in the Ecuadorian Páramos (Escamilo et al. 2010ESCAMILO, L.L., BARRIO, J., BENAVIDES, J. & TIRIRA, D. 2010. Northern Pudu, Pudu mephistophiles (De Winton 1896). In Neotropical cervidology: biology and medicine of Neotropical deer (J.M.B. Duarte & S. González, eds.). FUNEP/IUCN, Jaboticabal, p. 133-139.). As for L. pardalis, the altitudinal increment was 244 m (3,623 m) whereas for E. barbara it was 60 m (3,439 m) compared to previous records of 3,379 m recorded by Jiménez et al. (2010)JIMÉNEZ, C.F., QUINTANA, H., PACHECO, V., MELTON, D., TORREALVA, J. & TELLO, G. 2010. Camera trap survey of medium and large mammals in a montane rainforest of northern Peru. Rev. Peru. Biol. 17(2):191-196. http://sisbib.unmsm.edu.pe/BVRevistas/biologia/biologiaNEW.htm
http://sisbib.unmsm.edu.pe/BVRevistas/bi... for both species in northern Peru (Table 3). These species are rarely reported for such high altitudes, and the altitudinal distribution of E. barbara hardly exceeds 1,200 m (Cuarón et al. 2016CUARÓN, A.D., REID, F., HELGEN, K. & GONZÁLEZ-MAYA, J.F. 2016. Eira barbara. The IUCN Red List of Threatened Species 2016: e.T41644A45212151. https://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T41644A45212151.en. (Last access on 01/05/2021).
https://dx.doi.org/10.2305/IUCN.UK.2016-... ), while L. pardalis is uncommon to be found above 3,000 m (Paviolo et al. 2016PAVIOLO, A., CRAWSHAW, P., CASO, A., DE OLIVEIRA, T., LOPEZ-GONZALEZ, C.A., KELLY, M., DE ANGELO, C. & PAYAN, E. 2016. Leopardus pardalis (errata version published in 2016). The IUCN Red List of Threatened Species 2015: e.T11509A97212355. https://dx.doi.org/10.2305/IUCN.UK.2015-4.RLTS.T11509A50653476.en. (Last access on 01/05/2021).
https://dx.doi.org/10.2305/IUCN.UK.2015-... ). Thus, these records of L. pardalis and E. barbara are also the ones with the highest altitude for the entire area of occurrence of these two Neotropical carnivores.

The order Carnivora was the most representative in the study area, having 72.7% of the species belonging to this taxon, although most of them have a low relative frequency. Our results are similar to other studies with medium and large mammals conducted in transitional Puna-forest regions in the Peruvian Andes (Pacheco et al. 2007PACHECO, V., SALAS, E., CAIRAMPOMA, L., NOBLECILLA, M., QUINTANA, H., ORTIZ, F., PALERMO, P. & LEDESMA, R. 2007. Contribución al conocimiento de la diversidad y conservación de los mamíferos en la cuenca del río Apurímac, Perú. Rev. Peru. Biol. 14(2): 169-180. http://www.scielo.org.pe/scielo.php?script=sci_arttext&pid=S1727-99332007000300001
http://www.scielo.org.pe/scielo.php?scri... , Jiménez et al. 2010JIMÉNEZ, C.F., QUINTANA, H., PACHECO, V., MELTON, D., TORREALVA, J. & TELLO, G. 2010. Camera trap survey of medium and large mammals in a montane rainforest of northern Peru. Rev. Peru. Biol. 17(2):191-196. http://sisbib.unmsm.edu.pe/BVRevistas/biologia/biologiaNEW.htm
http://sisbib.unmsm.edu.pe/BVRevistas/bi... , Medina et al. 2012MEDINA, C.E., ZEBALLOS, H. & LÓPEZ, E. 2012. Diversidad de mamíferos en los bosques montanos del Valle de Kcosñipata, Cusco, Perú. Mastozool. Neotrop. 19(1):85-104. https://www.redalyc.org/pdf/457/45723408008.pdf
https://www.redalyc.org/pdf/457/45723408... , Shanee & Shanee 2018SHANEE, S. & SHANEE, N. 2018. Diversity of large mammals in the Marañón-Huallaga landscape, Peru: with notes on rare species. Zool. Ecol. 28(4):313-328. https://doi.org/10.1080/21658005.2018.1516277
https://doi.org/10.1080/21658005.2018.15... ). Despite occurring at low densities, species of this order have large home ranges and high mobility (Robinson & Redford 1986, Hodge & Arbogast 2016HODGE, A.M.C. & ARBOGAST, B.S. 2016. Carnivore diversity at a montane rainforest site in Ecuador’s Gran Sumaco Biosphere Reserve. Oryx 50(3):474-479. https://doi.org/10.1017/S0030605315000101
https://doi.org/10.1017/S003060531500010... ), increasing the probability of these mammals being detected at least once, certainly depending on the sampling effort. Furthermore, most of the registered carnivores have generalist habits, having the ability to explore resources in different environments (Lyra-Jorge et al. 2008bLYRA-JORGE, M.C., CIOCHETI, G., PIVELLO, V.R. & MEIRELLES, S.T. 2008b. Comparing methods for sampling large-and medium-sized mammals: camera traps and track plots. Eur. J. Wildl. Res. 54(4):739-744. https://doi.org/10.1007/s10344-008-0205-8
https://doi.org/10.1007/s10344-008-0205-... , Cossíos et al. 2012bCOSSÍOS, E.D., ALCÁZAR, P., FAJARDO, U., CHÁVEZ, K., ALFARO-SHIGUETO, J., CÁRDENAS-ALAYZA, S., VALQUI, J., MONTERO, F.G., LESCANO, J., QUEVEDO, M. & VIVAR, E. 2012b. El orden Carnivora (Mammalia) en el Perú: Estado del conocimiento y prioridades de investigación para su conservación. Rev. Peru. Biol. 19(1):17-26. http://www.scielo.org.pe/scielo.php?pid=S1727-99332012000100003&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ), moving from more forested landscapes such as the Cloud Forests, to open areas like the Puna. The presence of large top-chain carnivores, such as Puma concolor and Tremarctos ornatus, also demonstrates that the high-Andean region of the PHNS and its buffer zone has healthy populations of prey and good environmental integrity (Cardillo et al. 2004CARDILLO, M., PURVIS, A., SECHREST, W., GITTLEMAN, J.L., BIELBY, J., MACE, G.M. & MORITZ, C. 2004. Human population density and extinction risk in the world’s carnivores. PLoS Biol. 2(7):e197. https://doi.org/10.1371/journal.pbio.0020197
https://doi.org/10.1371/journal.pbio.002... ).

Among the recorded native species, Odocoileus virginianus had the highest relative frequency (25.5%), and this rate is even higher when compared only among records of native mammals (56%). Such a high relative frequency of this deer species for the Eastern Andes has never been reported in the literature. The highest relative frequency recorded for O. virginianus reported for the Eastern Andes was 15.2% found by Jiménez et al. (2010)JIMÉNEZ, C.F., QUINTANA, H., PACHECO, V., MELTON, D., TORREALVA, J. & TELLO, G. 2010. Camera trap survey of medium and large mammals in a montane rainforest of northern Peru. Rev. Peru. Biol. 17(2):191-196. http://sisbib.unmsm.edu.pe/BVRevistas/biologia/biologiaNEW.htm
http://sisbib.unmsm.edu.pe/BVRevistas/bi... in a survey carried out in the Páramos and Cloud Forests of northern Peru. For other areas of Latin America with occurrence of the species, such as Protected Areas of Mexico, Honduras and Ecuador, this frequency rate varies from 1 to 9% only (Monroy-Vilchis et al. 2011MONROY-VILCHIS, O., ZARCO-GONZÁLEZ, M.M., RODRÍGUEZ-SOTO, C., SORIA-DÍAZ, L. & URIOS, V. 2011. Fototrampeo de mamíferos en la Sierra Nanchititla, México: abundancia relativa y patrón de actividad. Rev. Biol. Trop. 59(1):373-383. https://www.scielo.sa.cr/scielo.php?pid=S0034-77442011000100033&script=sci_arttext&tlng=en
https://www.scielo.sa.cr/scielo.php?pid=... , Gonthier & Castañeda 2013GONTHIER, D.J. & CASTAÑEDA, F.E. 2013. Large-and medium-sized mammal survey using camera traps in the Sikre River in the Río Plátano Biosphere Reserve, Honduras. Trop. Conserv. Sci. 6(4):584-591. https://doi.org/10.1177%2F194008291300600409
https://doi.org/10.1177%2F19400829130060... , Lizcano et al. 2016LIZCANO, D.J., CERVERA, L., ESPINOZA-MOREIRA, S., POAQUIZA-ALVA, D., PARÉS-JIMÉNEZ, V. & RAMÍREZ-BARAJAS, P.J. 2016. Riqueza de mamíferos medianos y grandes del Refugio de Vida Silvestre Marina y Costera Pacoche, Ecuador. Therya 7(1):135-145. https://doi.org/10.12933/therya-16-308
https://doi.org/10.12933/therya-16-308... ). Despite occurring in several ecosystems of the Neotropical region, O. virginianus presents greater abundances in dry tropical forests and regions of grassland-forest ecotone, such as our study area (Delfin 2002, Gallina et al. 2010). Furthermore, in the high-Andean zones there is a partitioning of habitat use between O. virginianus and Hippocamelus antisensis, with O. virginianus tending to avoid such high altitudes (> 3,700 m) and rocky environments (Barrios 2006), but it also does not have a great preference for tropical rainforests, such as the Yungas (Palomino & Ataucusi 2019PALOMINO, M. & ATAUCUSI, Y. 2019. Distribución de mamíferos según rango altitudinal en la Reserva Nacional de Huascarán. Natura@ Economía 4(1):38-52. http://dx.doi.org/10.21704/ne.v4i1.1359
http://dx.doi.org/10.21704/ne.v4i1.1359... ). For these reasons, the high relative abundance of this deer is due to the altitudinal range and the puna-forest ecotone that the study area is located in, generating very favorable environmental conditions for the establishment of the species. Therefore, the high presence of O. virginianus in the area makes the high-Andean zone of PHNS a key location in Peru for potential ecological studies of the species, which still lacks information on its population and conservation status in the Peruvian territory (Gallina et al. 2010).

Among all recorded mammals, Bos taurus was the most frequent. The high presence of this domestic species in the area is due to the extensive livestock by the high-Andean rural communities (Paisley 2001, Goldstein 2002). The absence of more productive livestock management makes the rustic cattle move freely over extensive areas, enabling interaction with native species, generally in a negative way (Goldstein et al. 2006). A study by Barrio (2006)BARRIO, J. 2006. Manejo no intencional de dos especies de cérvidos por exclusión de ganado en la parte alta del Parque Nacional Río Abiseo, Perú. Rev. Electrónica Manejo Faujournalna Silv. Latinoam. 1(2):1-10. in the Peruvian Andes showed that the presence of cattle has a negative effect on the habitat use of native deer O. virginianus and H. antisensis, displacing these species to other areas. On the other hand, this extensive livestock makes the cattle very vulnerable to attacks from native predators such as pumas, foxes and Andean bears, generating retaliation by local communities and, consequently, persecution of these carnivores (Goldstein et al. 2006, Rojas-Vera Pinto et al. 2019ROJAS-VERA PINTO, R., ZEGARRA, R.E, GUTIÉRREZ, R. & BERAÚN, Y. 2019. Conviviendo con el Oso Andino en el Perú. El manejo, diagnóstico y pautas para los conflictos humano-oso. 1 ed., Sociedad Zoológica de Fráncfort Perú (FZS Perú), Cusco. https://cdn.www.gob.pe/uploads/document/file/1940651/Conviviendo%20con%20el%20oso%20andino%20en%20el%20Pe%C3%BA.pdf.pdf (Last access on 09/05/2021).
https://cdn.www.gob.pe/uploads/document/... ). In addition, unmanaged cattle grazing in these high-Andean regions can impact negatively the soil, the water retention of the high-Andean mash, the biodiversity of plants, aquatic macro invertebrates and even the abundance of specialist birds in the upper Andean grasslands (Astudillo et al. 2018ASTUDILLO, P.X., BARROS, S., SIDDONS, D.C. & ZÁRATE, E. 2018. Influence of habitat modification by livestock on páramo bird abundance in southern Andes of Ecuador. Stud. Neotrop. Fauna Environ. 53(1):29-37. https://doi.org/10.1080/01650521.2017.1382122
https://doi.org/10.1080/01650521.2017.13... , Avellaneda-Torres et al. 2018AVELLANEDA-TORRES, L.M., SICARD, T.E.L. & ROJAS, E.T. 2018. Impact of potato cultivation and cattle farming on physicochemical parameters and enzymatic activities of Neotropical high Andean Páramo ecosystem soils. Sci. Total Environ. 631:1600-1610. https://doi.org/10.1016/j.scitotenv.2018.03.137
https://doi.org/10.1016/j.scitotenv.2018... , Machaca et al. 2018MACHACA, N.C., CONDORI, B., PARDO, A.R., ANTHELME, F., MENESES, R.I., WEEDA, C.E. & PEROTTO-BALDIVIESO, H.L. 2018. Effects of grazing pressure on plant species composition and water presence on bofedales in the Andes mountain range of Bolivia. Mires Peat 21:1-15, 2018. http://mires-and-peat.net/media/map21/map_21_15.pdf
http://mires-and-peat.net/media/map21/ma... , Meza-Salazar et al. 2020MEZA-SALAZAR, A.M., GUEVARA, G., GOMES-DIAS, L. & CULTID-MEDINA, C.A. 2020. Density and diversity of macroinvertebrates in Colombian Andean streams impacted by mining, agriculture and cattle production. PeerJ 8:e9619. https://doi.org/10.7717/peerj.9619
https://doi.org/10.7717/peerj.9619... ).

The proportion of endemic species of Tropical Andes as Cuniculus taczanowskii, T. ornatus, and P. mephistopheles was relatively low for the area. This might be explained by the fact that the study was mostly carried out in the Puna ecorregion, which has an endemism rate of only 15.4% in Peruvian territory (Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... ). In turn, the Yungas and the Low Jungle are the ecoregions with the largest number of endemic mammal species in the Tropical Andes (Myers et al. 2000MYERS, N., MITTERMEIER, R.A., MITTERMEIER, C.G., DA FONSECA, G.A. & KENT, J. 2000. Biodiversity hotspots for conservation priorities. Nature 403(6772):853-858. https://doi.org/10.1038/35002501
https://doi.org/10.1038/35002501... , Pacheco et al. 2009PACHECO, V., CADENILLAS, R., SALAS, E., TELLO, C. & ZEBALLOS, H. 2009. Diversidad y endemismo de los mamíferos del Perú. Rev. Peru. Biol. 16(1):5-32. http://www.scielo.org.pe/scielo.php?pid=S1727-99332009000100002&script=sci_arttext&tlng=en
http://www.scielo.org.pe/scielo.php?pid=... , 2020PACHECO, V., GRAHAM-ANGELES, L., DIAZ, S., HURTADO, C.M., RUELAS, D., CERVANTES, K. & SERRANO-VILLAVICENCIO, J. 2020. Diversidad y distribución de los mamíferos del Perú I: Didelphimorphia, Paucituberculata, Sirenia, Cingulata, Pilosa, Primates, Lagomorpha, Eulipotyphla, Carnivora, Perissodactyla y Artiodactyla. Rev. Peru. Biol. 27(3):289-328. http://dx.doi.org/10.15381/rpb.v27i3.18356
http://dx.doi.org/10.15381/rpb.v27i3.183... ).

According to the IUCN, about 73% of the species registered in the study show a trend towards population reduction, indicating the relevance of the area for mammal conservation. However, only two species (T. ornatus and P. mephistopheles) are listed as threatened in Peru (Velez-Liendo & García-Rangel 2018VELEZ-LIENDO, X. & GARCÍA-RANGEL, S. 2018. Tremarctos ornatus (errata version published in 2018). The IUCN Red List of Threatened Species 2017: e.T22066A123792952. https://dx.doi.org/10.2305/IUCN.UK.2017-3.RLTS.T22066A45034047.en. (Last access on 28/04/2021).
https://dx.doi.org/10.2305/IUCN.UK.2017-... , SERFOR 2018SERFOR- Servicio Nacional Forestal y de Fauna Silvestre. 2018. Libro Rojo de la Fauna Silvestre Amenazada del Perú. 1 ed. SERFOR, Lima. https://www.serfor.gob.pe/portal/wp-content/uploads/2018/10/Libro-Rojo.pdf (Last access on 07/05/2021).
https://www.serfor.gob.pe/portal/wp-cont... ), and globally, P. mephistopheles still lacks data to define its current conservation status (Barrio & Tirira 2019BARRIO, J. & TIRIRA, D.G. 2019. Pudu mephistophiles. The IUCN Red List of Threatened Species 2019: e.T18847A22163836. https://dx.doi.org/10.2305/IUCN.UK.2019-1.RLTS.T18847A22163836.en. (Last access on 28/04/2021)
https://dx.doi.org/10.2305/IUCN.UK.2019-... ). Considered as smallest deer in the world, P. mephistopheles is still a poorly known species, especially the populations of central Peru, which are isolated from populations north of the Andes (Ecuador and Colombia), where there is more information about the species (Escamilo et al. 2010ESCAMILO, L.L., BARRIO, J., BENAVIDES, J. & TIRIRA, D. 2010. Northern Pudu, Pudu mephistophiles (De Winton 1896). In Neotropical cervidology: biology and medicine of Neotropical deer (J.M.B. Duarte & S. González, eds.). FUNEP/IUCN, Jaboticabal, p. 133-139.). Due to its low density, the record of three individuals of P. mephistopheles at the study site highlights the importance of the conservation of this area to maintain this species in the region.

Although the area belonging to the PHNS has a good conservation status of its Andean-Amazonian ecosystems, its surrounding areas are being impacted by human activities at different levels (SERNANP 2012SERNANP - Servicio Nacional de Áreas Naturales Protegidas por el Estado. 2012. Diagnóstico - Plan Maestro del Santuario Nacional Pampa Hermosa 2012-2017. Resolución Presidencial Nº 213-2012-SERNANP. http://old.sernanp.gob.pe/sernanp/archivos/biblioteca/planes_maestros_2014/pampa_hermosa/Diagnostico%20Plan%20Maestro%202012-2017%20SN%20Pampa%20Hermosa%20ver%20pub.pdf (Last access on 19/04/2021).
http://old.sernanp.gob.pe/sernanp/archiv... ). Extensive livestock and firewood extraction are the main threats to the ecosystems of the high-Andean region of the sanctuary’s buffer zone, where the S.P. Churco village is located. Despite having a high environmental vulnerability, our results showed that more than 90% of the species recorded were found outside the sanctuary boundaries, showing that this high-altitude buffer zone has an important value for the high-Andean mammals. However, these anthropogenic disturbances added to climate change, could strongly impact the diversity, abundance, and distribution of these mammals in the long term, mainly if natural resources are not rationally used.

As the first systematic survey of medium and large mammals for the PHNS region, we demonstrate that the area has a good representation of this biological community in the high zone of the Peruvian Eastern Andes. In addition, this study is one of the few surveys concentrated on the Wet Puna that only uses primary data and one of the first for the entire Peruvian department of Junín. Biological knowledge gaps in the high-Andean ecosystems are still huge in most Peru. The results we presented here offer new and important information on the local diversity of mammals, as well as updates on the altitudinal distribution of some species. However, further studies are needed on the biodiversity of the high-Andean zone of the PHNS and its surroundings, mainly because the high-Andean ecosystems are the most vulnerable and threatened to climate change throughout the Tropical Andes.

Acknowledgments

We thank the people of S.P. Churco for all support in the field, especially Ms. Cerila, Mr. Amancio Ticse, Mr. Pedro Ticse and Mr. Saturnino. We thank the Servicio Nacional de Áreas Naturales Protegidas por el Estado (SERNANP), through the PHNS rangers, for their support in data collection, especially Julio César Vilcarano. We are also grateful to Letícia Ribeiro, Davi Alves, Laodicéia Pereira and Paolo Ramoni Perazzi for their help in sorting the data. We thank Adriano Chiarello and Adriano Paglia for critically reviewing the manuscript. MMD thanks the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brazil (CAPES) for providing his grant. We thank the anonymous reviewers for comments and improvements to the present manuscript. The data collection for this work was carried out under authorization code Nº. AUT-IFS-2020-045 issued by MINAGRI/SERFOR - Peru.

Ethics

The authors declare compliance with all guidelines established by ethics committees’ universities and government agencies involved. They also state that the manuscript has not been previously published.

Data Availability

The authors declare that the data present in this manuscript may have open access. The data resulting from this research has been archived in the public data repository Biota Neotropica Dataverse, that provides free access and guaranteed preservation. Access URL: https://data.scielo.org/dataset.xhtml?persistentId=doi:10.48331/scielodata.3KOVKP.

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