Global Ecology and Conservation 26 (2021) e01451

Contents lists available at ScienceDirect

Global Ecology and Conservation

journal homepage: http://www.elsevier.com/locate/gecco

Original Research Article

Mammal conservation in Amazonia’s protected areas: A case

ndor National
study of Peru’s Ichigkat Muja - Cordillera del Co
Park
Jason J. Scullion a, *, Jacqueline Fahrenholz b, Victor Huaytalla c, d,
Edgardo M. Rengifo e, f, Elisabeth Lang g
a
Environmental Studies Department. McDaniel College, 2 College Hill, Maryland, USA
b
Duke University. Nicholas School of the Environment, 9 Circuit Drive, North Carolina, USA
c
Parque Nacional Ichigkat Muja-Cordillera Del Co
ndor Mz. 1 Lote 1 AA.HH Pueblo Joven Juan Velasco Alvarado, Amazonas, Peru
d
Faculty of Forest Sciences, Universidad Nacional La Agraria, Lima, Peru
e
Centro de Investigacio
n Biodiversidad Sostenible (BioS), Francisco de Zela 1556, Lima 14, Lima, Peru
f
dua
Universidade de Saeo Paulo, Escola Superior de Agricultura “Luiz de Queiroz”, Centro de Energia Nuclear na Agricultura. Avenida Pa
Dias, 11, Saeo Dimas, CEP 13418-900, Piracicaba, Brazil
g
Johns Hopkins University. Krieger School of Arts and Sciences, 3400 N. Charles St, Maryland, USA

a r t i c l e i n f o a b s t r a c t

Article history: The protected areas of the Amazon Basin provide a core refuge for many threatened forest
Received 31 May 2020 mammals species, but their effectiveness in wildlife conservation is poorly documented. To
Received in revised form 31 December 2020 better understand the impact of protected areas on mammal conservation in the Amazon
Accepted 31 December 2020
Basin, this study evaluates the diversity, abundance, and conservation status of medium
and large forest mammals in a borderland sector of Peru’s Ichigkat Muja - Cordillera del
Keywords:
ndor National Park. Over two field seasons, a total of 26 medium and large forest-
Co
Amazon basin
dwelling mammal species belonging to 19 families were recorded using camera traps
Camera trapping
Forest mammals
and field surveys. In comparison with regional reference sites, 63% of the mammal species
Protected area effectiveness recorded in the regional reference sites were documented in the study area. The drivers of
Mammal conservation species diversity and abundance within the study area were assessed using park guard
Peru reports, published literature, and regression analysis, and the intactness of the local
mammal fauna was evaluated using regional comparison sites. A combination of factors
explain the diversity and abundance of the mammal fauna recorded in the protected study
area, including regional geography and human activities. Given the location of the pro-
tected area in Peru and its adjacency to the border of Ecuador, long-term conservation of
the park’s mammals depends on binational efforts that invest in collaborative institutions
and local communities. This case study provides an example of how borderland protected
areas can present challenging wildlife conservation scenarios that require binational
collaboration. The current condition and at-risk status of mammal fauna in the protected
study area also underlines the importance of consistent field-based monitoring to inform
management and evaluate protected area effectiveness.
© 2021 The Author(s). Published by Elsevier B.V. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

* Corresponding author.
E-mail address: jscullion@mcdaniel.edu (J.J. Scullion).

https://doi.org/10.1016/j.gecco.2021.e01451
2351-9894/© 2021 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/
licenses/by-nc-nd/4.0/).
J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

1. Introduction

Terrestrial mammals are vital to the production of ecosystem goods and services that support human well-being, including
increasing carbon storage and ecosystem diversity (Ceballos and Ehrlich 2002; Estes et al., 2011; Ripple et al., 2014). The
benefits that terrestrial mammals provide humanity are presently at risk, with 21% of all assessed mammals (IUCN 2018) and
61% of all large-bodied carnivores threatened with extinction (Ripple et al., 2014). Mammals facing the highest risks globally
are medium and large in size and those associated with forest ecosystems (Vie
et al., 2009). Core refuges for forest dwelling
mammals include protected areas (Ripple et al., 2016; Pacfici et al., 2020) and intact forests (Betts et al., 2017; Watson et al.,
2018). Research shows that protected areas are generally effective in protecting forest cover (Joppa and Pfaff 2011; Bowker
et al., 2017), but less is known about their efficacy in protecting wildlife (Geldmann et al., 2013; Dudley et al., 2016).
Numerous studies have found that biodiversity inside protected areas is generally higher than in nearby areas under alter-
native land-uses (Geldmann et al., 2013; Coetzee et al., 2014). However, many protected areas have been shown to ineffec-
tively protect their wildlife (Harrison 2011; Laurance et al., 2012; Geldmann et al., 2013; Rija et al., 2020). Causes of wildlife
declines in tropical protected areas include insufficient funding and management (Bruner et al., 2001; Harrison 2011; Rija
et al., 2020) and human hunting (Harrison 2011; Benítez-Lo
pez et al., 2017; Rija et al., 2020). Environmental changes and
threats near reserve boundaries are also affect wildlife declines (Woodroffe and Ginsberg 1998; Laurance et al., 2012). Other
factors influencing wildlife persistence in protected areas can include reserve size (May, 1975; Higgs and Usher 1980), habitat
fragmentation (Saunders et al., 1991; Fahrig 2003), road and human access (Suarez et al., 2009; Espinosa et al., 2014; Benítez-
Lo
pez et al., 2017), and national levels of human development (Rija et al., 2020).
The mammal assemblages of the Amazon Basin are largely intact relative to other regions of the world (Belote et al., 2020)
and the region’s forests are considered a core refuge for large mammalian carnivores such as the jaguar (Panthera onca)
(Je˛ drzejewski et al., 2018). The size and isolation of forests in the Amazon Basin have historically helped to protect the region’s
wildlife (Sollmann et al., 2008; Harrison 2011). However, the future of the Amazon Basin’s mammal species is increasingly
threatened by accelerating infrastructure development, deforestation, and overhunting (Grelle 2005; Killeen 2007; Finer
et al., 2008; Suarez et al., 2009; Bogoni et al., 2020). The core refuge for the Amazon Basin’s mammals is the 46% of the
region that lies within official indigenous territories and protected areas (RAISG 2019). These areas are critical to ensure the
long-term persistence of the region’s mammal species (Peres 2000; Azevedo-Ramos et al., 2006). Indigenous territories and
protected areas in the Amazon Basin have in recent decades been largely effective in reducing deforestation and degradation
(e.g., Nepstad et al., 2006; Scullion et al., 2014; Blackman et al., 2017; Jusys 2018), but less is known about their effectiveness in
protecting wildlife. Existing case studies find that wildlife persistence inside protected areas and indigenous lands located in
the Amazon Basin is affected by a range of biophysical factors, including the existence of wildlife source populations and their
dispersal ability (Novaro et al., 2000; Ohl-Schacherer et al., 2007; Sampaio et al., 2010), the distance to settlements and roads
(Franzen 2006; Suarez et al., 2009; Espinosa et al., 2014) and protected area status (de Carvalho and Morato 2013). To better
understand the effectiveness of protected areas in conserving wildlife in the Amazon Basin and to conduct the first inventory
of mammal fauna in the study area, the diversity and abundance of medium and large forest dwelling mammals was eval-
uated using camera traps and field surveys in a borderland sector of Peru’s Ichigkat Muja - Cordillera del Co
ndor National Park
(CNP) (Fig. 1). The objectives of this research were to (1) assess the diversity and abundance of mammals in the study area (2)
identify the factors that influence the diversity and abundance of mammals in the study area; and (3) inform mammal
conservation efforts in CNP and the beyond.

2. Materials and methods

2.1. Study area

Located in the department of Amazonas, Peru, and along the border with Ecuador, CNP (03
19ʹ S, 078
07ʹ W) was
established in 2007. Portions of CNP are covered by the steep slopes of the Cordillera del Co
ndor Mountains, hereafter
“Co
ndor Mountains.” The Co
ndor Mountains have a year-round tropical wet climate and a variety of ecosystems, including
lowland tropical forests, cloud forests, and high elevation tepui-like ecosystems (RAP 1997). The average annual rainfall of the
CNP ranges from 2400 mm to 4500 mm and the average annual temperature is 24
C (SERNANP 2012). The elevation of the
Co
ndor Mountains within the CNP ranges from 200 m to 2600 m (SERNANP 2018). The Co
ndor Mountains are located be-
tween the Andes Mountains and the Amazon Basin and represent the longest and tallest of the sub-Andean ranges (Neill
2005). Based on their pre-Andean history and geographic complexity, the Co
ndor Mountains are recognized for their high
biological diversity and species endemism (Weigend 2002). The study area is located within CNP and covers 111 km2
(11,059 ha) across an elevation gradient of 223e713 m.a.s.l. The site is covered with tropical lowland rainforest and pre-
montane forests and is located between the steep slopes of the Co
ndor Mountains and the Santiago River (Fig. 1). Within
the study area are small rivers and streams that dissect hill forests, riparian forests, and palm swamps. As of 2016, 86% of the
study area was categorized as “intact forest landscape” (Potapov et al., 2008). There is no history of commercial logging or
agriculture inside CNP. However, the study area is categorized as a ‘Recovery Zone’ due to historical damage from selective
logging, hunting, and three regional conflicts waged between Peru and Ecuador in the 20th century, as recently as 1997 (RAP
1997; SERNANP 2018). The park and its buffer zone are the ancestral home of the Wampis indigenous group, who today live in
communities in the CNP buffer zone near the southern border of the park (Fig. 1). Access to the Wampis’ communities from

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J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

Fig. 1. Regional map showing the study area and camera trap locations. The map shows the study area and camera trap locations inside Peru’s Ichigkat Muja -

ndor National Park (CNP). The study area is bordered to the north by Ecuador, to the east by the Rio Santiago and the Kampankis Mountains, and
Cordillera del Co

ndor.
to the west by the Cordillera del Co

the nearest city in Peru (Nieva) requires approximately a 10 hour, 235 km boat trip. Across the border from the CNP in
Ecuador, the indigenous Shuar communities have road access to nearby villages, including Puerto Minas, (3 km from the
reserve’s border), where they are reported to sell wild meat (SERNANP 2018). The indigenous communities adjacent to CNP in
Peru and Ecuador are located in regions with high rates of poverty (INEC 2014; INEI 2017). Like many indigenous communities
in the Amazon Basin, they hunt forest wildlife as a cultural practice and depend on wild meat as a critical source of dietary
protein (Nasi et al., 2011; Coad et al., 2019). The Peruvian Wampis communities near the CNP live in the buffer zone have
access to large areas of forest to hunt, while the Ecuadorian Shuar communities have limited nearby forest access outside CNP
(SERNANP 2018).

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J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

2.2. Data collection and analysis

Two field seasons of camera trap data were collected by placing camera traps along a regular grid of 24 sampling points at a
density of one camera per 2 km using a modified version of the Tropical Ecology Assessment and Monitoring Network
methodology (Jansen et al., 2014). The cameras were deployed for 104 trap nights entering the dry season of 2018 and 105
trap nights in 2019 (specifically, July 7th - October 20th, 2018 and July 23rd e October 30th, 2019). Browning Strikeforce and
Bushnell Trophy Cam cameras were used and programmed to take three photos at 1-min intervals when activated. Cameras
were placed 30e50 cm from the ground and located within 100 m of each GPS-referenced sampling point along paths, creeks,
and other sites of animal activity. To minimize human disturbance, the cameras were not checked during the study periods.
Camera theft and malfunction resulted in the loss of data from five cameras in 2018 and six cameras in 2019. Species presence
was recorded for each camera by the co-authors based on their local knowledge and regional field guides (Eisenberg and
Redford 2000; Emmons and Feer 1997; Noboa 2017). The relative abundance, or capture frequency, of a given species was
calculated as the number of photos (trap events)/trap nights x 1000 (Tobler et al., 2008). To ensure data independence, image
captures of the same species were excluded if they occurred within 1-h of the first capture (Tobler et al., 2008). An OLS
regression analysis in ArcGIS Pro (version 2.7) was used to evaluate relationships between the camera trap frequency capture
data and local biophysical variables. The taxa evaluated as dependent variables were predators, herbivores, and herbivores
sensitive to hunting. The explanatory variables tested for their relationship with the capture frequency data included distance
to roads, rivers, Ecuadorian communities, Peruvian communities, ranger posts, and military posts as well as point-based data,
including habitat type and park ranger incident records 2017e2019.
This study did not assess arboreal mammals or primate species using camera traps due to the camera’s placement at
ground level. All mammals with an average body mass of less than 0.5 kg were excluded from the study because smaller
mammals are more difficult to consistently detect and identify at the species level using camera traps (Kelly and Holub 2008;
Tobler et al., 2008). Opossums and small rodents were excluded from this study given the difficulty of distinguishing local
Amazonian species based on morphology (Cerqueira and Lemos 2000). Over both field seasons during the camera trap
fieldwork, diurnal primate species in the study area were recorded using sight and sound. CNP park guard patrol reports for
the years 2017e2019 were analyzed for reported mammal fauna and human activity. To evaluate the study area’s mammal
diversity, a list of forest dwelling mammal species that may be recorded in the study area was developed based on regional
case studies with study sites within 400 m of elevation of the study area (Patton et al., 1982; RAP, 1997; Vivar and Rosa, 2004;
Pitman et al., 2012) (See Appendix A for site locations). All four regional studies were field-based species inventories and only
species recorded by the researchers using sight or sound were included in the list of regionally recorded species (Table 1).
Sampling effort of the camera trap study was measured using a species accumulation curve combining both field seasons
using the software SDR 4 (Pisces Conservation). The species accumulation curve generated confirms that total sampling effort
was adequate as the curve reached an asymptote around sample 15 (Appendix B). To better characterize the status of the
mammal fauna in the study area, the mammal assemblage observed in the study area was compared with similar camera-
based inventories of terrestrial mammals using a Jaccard analysis. Specifically, the Jaccard analysis included 15 studies
from Peru and Ecuador that had sufficient sampling effort (see Appendix C). The similarity of mammal fauna was compared
with a presence/absence species matrix using the PAST statistical software (Hammer et al., 2001).

3. Results

In the 2018 survey, 826 images of 18 mammal species were captured in 1976 trap nights. In the 2019 survey, 611 images of
16 mammal species were captured in 1890 trap nights (Appendices D and E). Between the camera trapping and field surveys,
a total of 1437 images and 26 species belonging to 19 families were recorded (Table 1). A total of 63% of the 41 medium and
large forest dwelling mammal species reported in the regional case studies were recorded in the study area (Table 1). The
Jaccard analysis of 15 regional camera trap mammal studies showed that the two regional studies with the highest similarity
scores to the study area were Blake and Loiselle (2018) (Tipuni) and Tobler et al. (2008) (Los Amigos) (Appendices F and G).
Comparing the species-specific capture frequencies from the study area with the Tipuni and Los Amigos studies shows that
the capture frequencies in the study area contrast with the comparison sites. For example, the Amazonian tapir (Tapirus
terrestris) and jaguar (Panthera onca) were recorded less frequently, and the black agouti (Dasyprocta fuliginosa) and lowland
paca (Cuniculus paca) were comparatively more frequent (Table 2). Among others, threatened mammals not recorded in the
study area but recorded regionally include the bush dog (Speothos venaticus), short-eared dog (Atelocynus microtis), and the
brown woolly monkey (Lagothrix lagotricha). Two primate species were observed during the camera trap fieldwork: the
common squirrel monkey (Saimiri sciureus) and the white-tailed titi (Plecturocebus discolor). While primate species were not
assessed with camera traps, the white-fronted capuchin (Cebus albifrons) was recorded once.
The regression analysis shows a significant relationship between decreasing predator occurrence and decreasing distance
to Ecuadorian communities (Table 3). None of the other explanatory variables resulted in a significant finding in any of the
models run using different combinations of explanatory variables. In the 2018 season, at least three jaguar individuals were
recorded, including a rare melanistic jaguar (Rengifo et al., 2019). During the 2019 field season, up to two jaguar individuals
were recorded and during the fieldwork a melanistic jaguar was shot in the study area by a local hunter who reported he was
being attacked. No other hunting incidents were reported during the fieldwork, but evidence of hunting was seen. Ranger
reports for the study area 2017e2019 recorded 186 records of suspected hunting activity, including hunter encounters, shell

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J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

Table 1
Mammal Species Found in this Study Compared to Other Regional Studies.

Family Common Name Scientific Name Status Recorded in Study Area Previous Regional Record
Atelidae Brown woolly monkey Lagothrix lagotricha VU (EN) D
Atelidae Poeppig’s woolly monkey Lagothrix poeppigii VU (VU) D
Atelidae Venezuelan red howler Alouatta seniculus LC (VU) X* A, C, D
Atelidae White-bellied spider monkey Ateles belzebuth EN (EN) X* B, D
Caviidae Capybara Hydrochoerus hydrochaeris LC D
Cebidae Common squirrel monkey Saimiri sciureus LC X* A, C
Cebidae White-fronted capuchin Cebus albifrons LC X A, B
Canidae Bush dog Speothos venaticus NT A, D
Canidae Short eared dog Atelocynus microtis NT (VU) A, B, C, D
Cervidae Red brocket deer Mazama americana VU (VU) X A, B, C, D
Chlamyphoridae Giant armadillo Priodontes maximus LC A, C, D
Chlamyphoridae Southern naked-tailed armadillo Cabassous unicinctus LC A, C
Cuniculidae Lowland paca Cuniculus paca LC X B, D
Cyclopedidae Silky anteater Cyclopes ida LC A, C, D
Dasypodidae Nine-banded armadillo Dasypus novemcinctus LC X A, B, C, D
Dasypodidae Seven banded armadillo Dasypus septemcinctus LC D
Dasypodidae Greater-long nosed armadillo Dasypus kappleri LC D
Dasyproctidae Black agouti Dasyprocta fuliginosa LC X A, B, C, D
Dasyproctidae Green acouchi Myoprocta pratti LC X A, D
Dinomyidae Pacarana Dinomys branickii LC (VU) A, C, D
Erethizontidae Bicolored porcupine Coendou bicolor LC X A, C, D
Erethizontidae Ecuadorian dwarf porcupine Coendou ichillus DD D
Felidae Jagarundi Herpailurus yagouaroundi LC X A, D
Felidae Margay Leopardus wiedii DD (NT) X A, B, C, D
Felidae Ocelot Leopardus pardalis LC X A, B, C, D
Felidae Puma Puma concolor LC (NT) X B, D
Felidae Jaguar Panthera onca NT (NT) X A, B, C, D
Felidae Oncilla Leopardus tigrinus VU (DD) D
Mustelidae Tayra Eira barbara LC X A, B, C, D
Mustelidae Greater grison Galictis vittata LC A, B, C, D
Mustelidae Neotropical otter Lontra longicaudis NT X* A, D
Mustelidae Amazon weasel Mustela africana LC X D
Myrmecophagidae Southern tamandua Tamandua tetradactyla LC X A, B, D
Myrmecophagidae Giant anteater Myrmecophaga tridactyla VU (VU) X A, B, C, D
Pitheciidae White-tailed titi Plecturocebus discolor (LC) X* A
Procyonidae South American coati Nasua nasua LC X A, B, D
Procyondiae Kinkajou Potos flavus LC C, D
Procyondiae Crab eating raccoon Procyon cancrivorous LC X A, B, C, D
Tapiridae Amazonian tapir Tapirus terrestris VU (NT) X A, B, C, D
Tayassuidae White-lipped peccary Tayassu pecari VU (NT) X* A, B, C, D
Tayassuidae Collared peccary Pecari tajacu LC X A, C

The table shows medium and large forest dwelling mammals recorded in the CNP study area and comparable regional sites using camera traps and visual
and auditory signs. Species recorded in the study area marked with “X*” were recorded by park rangers during their 2017e2019 field patrols and/or the
2018e2019 camera trap fieldwork and not on the camera traps. Only diurnal primates were recorded given the daytime patrols and fieldwork. The species
conservation “Status” reported above is from the IUCN Red List (2019) and the secondary status in parentheses is from the Peruvian Red Book (SERFOR
2018): LC, least concern; NT, near threatened; VU, vulnerable; EN, endangered; CR, critically endangered. Regional studies that reported the presence of
medium to large terrestrial mammals are shown as A ¼ Patton et al., 1982; B ¼ RAP 1997; C ¼ Vivar and Rosa 2004; D ¼ Pitman et al., 2012.

casings, hunter trails, and sleeping shelters. The ranger reports also recorded single visual observations of four additional
mammal species: the Venezuelan red howler monkey (Alouatta seniculus) the white-bellied spider monkey (Ateles belzebuth),
the neotropical otter (Lontra longicaudis), and the white-lipped peccary (Tayassu pecari). Three other mammal species were
recorded in the ranger reports based on footprints, feces, or burrow structure: the capybara (Hydrochoerus hydrochaeris), the
giant armadillo (Priodontes maximus), and the kinkajou (Potos flavus).

4. Discussion

4.1. Drivers of diversity and abundance in the CNP study area

This study found that the CNP study area contains a diversity of medium and large forest mammals, including at least 26
species belonging to 19 families. Apex predators were infrequent and selected medium-sized herbivores and meso-predators
were common (Table 1). A large percentage of regionally known species were found in the study area (63%), but a number of
vulnerable species were recorded at low densities or not at all. As explored in the section that follows, the diversity and
abundance of forest mammal species recorded in the study area and the level of species turnover documented between the
study area and regional reference sites is explained by a combination of factors, including ecology, geology, and human
activities.

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J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

Table 2
Selected CNP species captures rates relative to comparison studies.

Average Capture Frequency

Species Common Name This Study Tipuni Los Amigos

Puma 0.82 15.16 6.51
Jaguar 1.64 5.70 13.28
Predator Species Jaguarundi 1.64 0.42 0.52
Ocelot 9.84 10.85 11.97
Margay 0.82 0.70 3.90
Black Agouti 194.15 46.88 25.52
Red Brocket Deer 81.21 76.78 10.32
Lowland Paca 65.35 34.08 12.23
Prey Species
Collared Peccary 14.77 104.19 10.93
White-Lipped Peccary 0.00 45.90 54.68
Amazonian Tapir 9.57 34.78 26.56

The table shows species captures rates for the CNP study site and two similar sites identified using a Jaccard analysis Blake and Loiselle (2018) - Tipuni and
Tobler et al., (2008) - Los Amigos (see Appendices F and G). For each comparison study in the table, the original species capture data was combined and
reanalyzed to derive total capture frequencies based on the formula of number of photos (trap events)/trap nights x 1000 (Tobler et al., 2008). Total sampling
effort for each study was reported in trap nights as 3840 for Los Amigos, 3866 in this study, and 7189 for Tipuni.

Ecological and geographic factors are likely to play a large role in the mammal species recorded in the study area as well as
the regional species distribution as these factors often determine the distribution of wildlife, including via physical barriers
(Harcourt and Wood 2012) and landscape heterogeneity (Kozakiewicz 1993). Mountainous environments, such as the Condor
Mountains, present unique geophysical conditions that influence the distribution and diversity of wildlife, including pro-
nounced barriers to dispersal and broad climate diversity over small regions (Rahbek et al., 2019a; Rahbek et al., 2019b).
surrounding the study area is known to generate a regional biodiversity gradient affecting forest dwelling mammals. Between
the Rio Santiago, which sits on the eastern boundary of the study area, and the Rio Cenepa, Patton et al. (1982) identified
distinct east-west species diversity gradients as well as noticeable elevational shifts in mammal fauna. Differential habitat
complexity between the river basins is hypothesized to play a role, and the location of the study area between the Condor
Mountains and within the flood plains and foothills of the Rio Santiago likely affects species dispersal and recolonization
(Fig. 1). The sampling intensity and dry season focus may also have influenced the camera trapping results, with limited
seasonality (Wearn and Glover-Kapfer 2017) and the low abundance of rare species known to reduce observed capture rates
(Tobler et al., 2008).
Another set of factors impacting the diversity and abundance of local mammal species is human activity. Patton et al.
(1982) thought that centuries of indigenous hunting may explain missing primate fauna and that the region’s biogeog-
raphy helped to explain the distribution of regional mammal species. Beginning with the study area’s history of human
warfare in the 20th century, including as recently as 1997 (RAP, 1997; SERNANP 2018), war undoubtedly impacted the di-
versity and abundance of mammals found in the study area. The long-term impacts of earlier conflicts on local forest
mammals is unknown due to a lack of historical data, but the modern link between armed conflict and reduced mammal
fauna is well documented (Dudley et al., 2002; Daskin and Pringle 2018). More recent human impacts on the study area were
documented in this research and center on threats due to habitat loss and human hunting. Habitat loss and degradation are
leading threats to mammal fauna worldwide (Ripple et al., 2014; IUCN 2018), but their impact on the study area’s mammal
fauna has not be studied. The forest cover around the study area in Peru is intact beyond local community sites, but in nearby
Ecuador, land-cover change is more pronounced, including the presence of small subsistence farms near the reserve boundary
(SERNANP 2018). While more research is needed, negative impacts on the mammal fauna in CNP from regional land use
change may continue to grow with local communities.

Table 3
Regression results of predator capture frequency.

Explanatory Variable Coefficient Std. Error Probability Variance Inflation Factor

Distance to Military Posts 0.000323 0.0002 0.12462 5.040769
Distance to Ranger Posts 0.000276 0.000175 0.133941 6.74916
Distance to Ecuadorian Communities 0.000539 0.000193 0.012963* 6.362209
Distance to Peruvian Communities 0.000146 0.000134 0.293588 5.069056

Statistic Probability
Joint F-Statistic 0.014974*
Koenker Statistic 0.075957
Jarque-Bera Statistic 0.549362

The table above shows the results of OLS regression analysis. *Indicates significant result at 0.01. Large variance inflation factor (>7.5) indicates redundancy
between explanatory variables. Joint F-statistic indicates overall model significance. A significant Koenker statistic indicates non-stationarity or hetero-
skedasticity. A significant Jarque-Bera statistic indicates that the residuals are not normally distributed and that model predictions are biased.

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J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

Hunting is also a leading worldwide threat facing mammals (Ripple et al., 2016; IUCN 2018) and a key threat in the study
area. Hunting activity was documented in the study area through the park guard reports and the camera traps and field
surveys. The results of the camera trapping and surveys found that species known to be vulnerable to hunting were rarely
detected or not recorded, including the brown woolly monkey (Lagothrix lagotricha), the Amazonian tapir (Tapirus terrestris),
and the collared peccary (Pecari tajacu). Such species, along with others recorded at low densities, are often at risk of
overhunting due to their large home ranges, relatively large bodies, and slow rates of reproduction (McKinney 1997; Purvis
et al., 2000, Benítez-Lo
pez et al., 2019). Evidence of recent human impacts on the study area’s mammal fauna is also shown by
the regression results of the camera trap data showing that the frequency of felid predators declined closer to the Ecuadorian
communities. Interestingly, no other biophysical factors were significantly related to the capture frequencies recorded. This
may be due to low abundance of selected taxa, the consistency of biophysical conditions across the study area, and the study’s
sampling design. Other evidence of human impacts on the study area’s mammal fauna recorded include relatively low capture
frequencies for large predators and vulnerable herbivores and higher frequencies of less sensitive herbivores for the CNP
study area compared to two sites identified in the Jaccard analysis with similar mammal fauna, Tipuni and Los Amigos (Tobler
et al., 2008; Blake and Loiselle 2018). Likewise, a comparison of the mammal species recorded in the study area with regional
mammal surveys found a lower diversity of mammal fauna recorded in this study than nearby sites, particularly the adjacent
Kampankis Mountains, which contains all but one mammal “missing” from this study, likely due to the area’s low hunting
activity (Pitman et al., 2012).
Given the lack of historical wildlife data for the study area, this study could not determine the effectiveness of park
management and enforcement efforts in wildlife conservation; only follow-up wildlife monitoring at the site can answer this
question. The cumulative impacts of hunting on the reserve are also unknown and will remain so without further research,
but the risk for local people and the protected area is high as long-term hunting impacts include forest ecosystem changes and
reduced food resources for local communities. Studies have consistently found that the loss of apex predator species alters the
composition and function of the forest vegetation via trophic cascades that release local herbivores and meso-predators
(Wright et al., 2000; Ripple and Beschta 2006; Prugh et al., 2009). More research is needed to identify potential trophic
cascades in the study area, but the abundance of medium mammal herbivores and mesopredators recorded may indicate
ecosystem changes are underway. A corollary example may come from Ecuador’s Machalilla National Park, where high
abundance of brocket deer was detected, and the extirpation of jaguars and pumas was presumed to be the cause of the high
deer population (Cervera et al., 2016). Overall, the findings of this research echo research worldwide (e.g., Redford 1992;
Sreekar et al., 2015; Benítez-Lo
pez et al., 2019) in finding that the mammal community of the study area is being negatively
affected by human hunting despite its protected status.

4.2. Mammal conservation in borderlands and the CNP study area

The transboundary CNP study area examined here provides a useful example of how mammal conservation in protected
areas can be affected by their transboundary context. Previous research has shown that transboundary areas face increased
threats in remote borderland regions due to weak government enforcement and complex political conditions that enable
poaching, illegal hunting, and the wildlife trade (Liu et al., 2020). Borderland case studies also find that poverty motivates
many of those involved in wildlife hunting and trafficking (e.g., van Vliet et al., 2014; Krishnasamy et al., 2018; Uprety et al.,
2020). To address the threats facing wildlife in the CNP, the most recent park management plan (SERNANP 2018) prioritizes
species monitoring, surveillance and control of unauthorized entry into the reserve, and creating conservation agreements
with nearby indigenous communities in Peru. However, engaging the local communities in Ecuador is not within the
jurisdiction of SERNANP, thus requiring support from institutions in Ecuador.
New regulations, binational collaborative agencies, and development projects to improve border governance and improve
local livelihoods in this region were agreed to as part of the 1998 peace agreement signed between Peru and Ecuador: the
Peru-Ecuador Binational Plan (PEBP). This agreement ended their most recent war, but the governance and investment
initiatives envisioned have lacked sufficient funding to be fully implemented (Ali 2019). The PEBP seeks to improve the
livelihoods of populations living on the border of Peru and Ecuador through border integration and public investment
projects (PEBP, 2018). Community-based projects have already been carried out in the region through the PEBP mechanism,
and that may indirectly contribute to the conservation of ecosystems and wildlife. Also, in recent years in the Presidential
Meetings and Binational Cabinets of Peru and Ecuador, the “Roadmap for the Conservation of CNP” was proposed in which
binational actions were established with the participation of institutions from both countries. The implementation of this
Roadmap would help improve border governance and control the entry of hunters into the CNP.
Taken together, the CNP study area is a refuge for a diversity of forest dwelling mammals native to the Amazon Basin. A
combination of factors explains the contemporary mammal fauna recorded in the CNP study area, including regional ge-
ography and human activities. Additional wildlife monitoring is needed to evaluate the effectiveness of the CNP in conserving
its wildlife. Given the local economic conditions and the regional demand for wild meat, reducing hunting in the reserve will
require a combination of proven wildlife conservation strategies that include poverty alleviation for local communities (Rija
et al., 2020) and the use of participatory wildlife management mechanisms that involve all stakeholders (Campos-Silva et al.,
2017). Such efforts depend on binational efforts between Peru and Ecuador to further develop collaborative cross-border
institutions and provide adequate funding as envisioned in the 1998 peace agreement ending the war between the two
countries. This case study finds that protected areas in borderlands can present unique wildlife conservation challenges that

7
J.J. Scullion, J. Fahrenholz, V. Huaytalla et al. Global Ecology and Conservation 26 (2021) e01451

require binational cooperation and targeted community investment. It is also shown that ongoing wildlife monitoring is
needed to inform protected area management and assess reserve effectiveness. This monitoring effort will fill a crucial gap in
our knowledge of protected area efficacy (Geldmann et al., 2019) and increase the accuracy of global analyses of ecological
integrity and faunal intactness (Plumptre et al., 2019).

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have
appeared to influence the work reported in this paper.

Acknowledgments

This study was carried out with research permit 08-2019-SERNANP-JEF (PNIM-CC) and would not have been possible
without the support of Jessica Tsamajain, manager of the Ichigkat Muja National Park - Cordillera del Co
dor (SERNANP), and
her park rangers and technicians, who helped deploy and collect the camera traps, especially Diego Tataje, Tercero Lirio and
Yamir Tenorio. This study also had the support and enthusiasm of students from the McDaniel College Conservation Biology
course (ENV 3106) who conducted the initial wildlife image analysis and contributed to the ideas and results presented
(Spring 2018 and 2019) A special thanks to Luis Cueto for his invaluable support with the fieldwork and project planning.
Thanks to Provost Julia Jasken of McDaniel College for her critical support of this project. Many thanks to the two anonymous
reviewers whose thoughtful feedback and expertise greatly improved the manuscript. McDaniel College provided interna-
tional travel support to the first author.

Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.gecco.2021.e01451.

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