Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile
Habitat fragmentation is one of the main factors threatening biodiversity, especially in regions where natural ecosystems are under anthropogenic pressure, such as the Araucanía region in southern Chile. This study explores the presence of nestedness patterns in the composition of carnivorous mammal...
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| author | Sanhueza, R. Moya, W. Rau, J. R. |
| author_facet | Sanhueza, R. Moya, W. Rau, J. R. |
| author_institution_txt_mv | [
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"author": "R. Sanhueza",
"institution": "Laboratorio de Ecología, Departamento de Ciencias Biológicas and Biodiversidad, Universidad de Los Lagos, Osorno, Chile",
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},
{
"author": "W. Moya",
"institution": "Laboratorio de Ecología, Departamento de Ciencias Biológicas and Biodiversidad, Universidad de Los Lagos, Campus Osorno. P.O. Box 933, Osorno, Chile",
"orcid": ""
},
{
"author": "J. R. Rau",
"institution": "Laboratorio de Ecología, Departamento de Ciencias Biológicas and Biodiversidad, Universidad de Los Lagos, Osorno, Chile",
"orcid": ""
}
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| author_sort | Sanhueza, R. |
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| datestamp_date | 2026-08-20T12:37:09Z |
| description | Habitat fragmentation is one of the main factors threatening biodiversity, especially in regions where natural ecosystems are under anthropogenic pressure, such as the Araucanía region in southern Chile. This study explores the presence of nestedness patterns in the composition of carnivorous mammal species in six fragments along the southern part of the Coastal Range in the Araucanía region. Using scent stations and actively searched five species were recorder over a nine-month period: hog-nosed skunk, kodkod, cougar, gray fox, and culpeo fox. The analysis revealed that the nested pattern, with a temperature of 13.661 °C, suggests a risk of local extinction in smaller fragments (less than 14 ha). The hog-nosed skunk was detected in four fragments, including one of the smaller ones (3.1 ha), while the cougar and kodkod were more restricted, being recorded in three and two fragments, respectively. The culpeo fox and gray fox were present in only one or two fragments. These results support the island biogeography theory, highlighting that larger fragments sustain greater species diversity. This study highlights the importance of conserving larger forest fragments to maintain carnivore diversity, as specialized species rely on denser and more connected habitats. It is recommended to prioritize the preservation of continuous forest areas and restore connectivity between fragments to mitigate the negative effects of fragmentation. |
| doi_str_mv | 10.15407/zoo2024.06.501 |
| first_indexed | 2025-07-17T12:36:59Z |
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© Publisher Publishing House "Akademperiodyka" of the NAS of Ukraine, 2024. The article is published under
an open access license CC BY-NC-ND (https://creativecommons.org/licenses/by-nc-nd/4.0/)
UDC UDC 599.742(1-13:23.071:835.32)
NESTEDNESS IN A CARNIVOROUS MAMMAL ASSEMBLAGE IN
FOREST FRAGMENTS OF THE ARAUCANÍA REGION,
SOUTHERN CHILE
R. Sanhueza, W. Moya & J. R. Rau*
1Laboratorio de Ecología, Departamento de Ciencias Biológicas and Biodiversidad, Universidad de Los Lagos,
Campus Osorno. P.O. Box 933, Osorno, Chile
*Corresponding author
E-mail: jrau@ulagos.cl
R. Sanhueza (https://orcid.org/0009-0008-4208-7577)
W. Moya (https://orcid.org/0000-0002-5118-8985)
J. R. Rau (https://orcid.org/0000-0003-0444-578X)
urn:lsid:zoobank.org:pub:B6F20359-8074-4BBE-88DC-6D35F827C7A3
Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region,
Southern Chile. Sanhueza, R., Moya, W., Rau, J. R. — Habitat fragmentation is one of the main fac-
tors threatening biodiversity, especially in regions where natural ecosystems are under anthropogenic
pressure, such as the Araucanía region in southern Chile. This study explores the presence of nestedness
patterns in the composition of carnivorous mammal species in six fragments along the southern part of
the Coastal Range in the Araucanía region. Using scent stations and actively searched five species were re-
corder over a nine-month period: hog-nosed skunk, kodkod, cougar, gray fox, and culpeo fox. The analy-
sis revealed that the nested pattern, with a temperature of 13.661 °C, suggests a risk of local extinction in
smaller fragments (less than 14 ha). The hog-nosed skunk was detected in four fragments, including one
of the smaller ones (3.1 ha), while the cougar and kodkod were more restricted, being recorded in three
and two fragments, respectively. The culpeo fox and gray fox were present in only one or two fragments.
These results support the island biogeography theory, highlighting that larger fragments sustain greater
species diversity. This study highlights the importance of conserving larger forest fragments to maintain
carnivore diversity, as specialized species rely on denser and more connected habitats. It is recommended
to prioritize the preservation of continuous forest areas and restore connectivity between fragments to
mitigate the negative effects of fragmentation.
Key words : habitat fragmentation, carnivore density, nestedness, Araucanía Forest, species composition.
Zoodiversity, 58(6):501–512, 2024
DOI 10.15407/zoo2024.06.501
Ecology
502 R. Sanhueza, W. Moya & J. R. Rau
Introduction
Habitat loss and fragmentation are among the most critical threats to biodiversity (Teckentrup et al., 2019;
Ntshanga et al., 2021), significantly impacting ecosystem functions through alterations in species richness and
abundance (Saura, 2021; Palmeirim et al., 2020; Liu et al., 2018). Fragmentation often leads to local extinctions
and distinctive patterns in species richness and composition (Rybicki et al., 2020).
Over the past 30 years, central-southern Chile has experienced ongoing and increasing fragmentation
of forested areas, with some effects also observed in the surrounding mountainous regions (Rau et al., 2001;
Echeverría et al., 2006). In less than a century, the landscape of the La Araucanía region has transformed from a
dense, impenetrable forest (Verniory, 2017) to one marked by novel landscapes and clear signs of environmental
degradation. The severity of forest fragmentation primarily impacts mammal populations, depending on their
habitat requirements (Zúñiga et al., 2014; Palmeirim et al., 2019), diet (Zúñiga et al., 2016), and behavioral
adaptations, such as hunting and territoriality (Bista et al., 2022).
Since species exhibit varying levels of sensitivity to habitat fragmentation, species loss within the
remaining habitat fragments does not occur randomly. Instead, it frequently follows a nested pattern, where
species composition becomes progressively simplified as more sensitive species are lost (Patterson and Atmar,
1986; Atmar and Patterson, 1993, 1995; Deane & He, 2018; Hovestadt et al., 2024). This pattern is driven by
factors promoting compositional disaggregation (García-Quintas and Parada-Isada, 2017; Flores et al., 2018).
Nestedness is especially prevalent in fragmented systems shaped by local extinctions (Patterson and Atmar,
1986; Luther et al., 2020), playing a key role in landscape ecology. It provides a metric for assessing non-random
species presence-absence across different assemblages and communities, enabling the quantification of species
vulnerability to habitat changes (Luther et al., 2020).
Nestedness patterns have been studied in various taxa, including raptors (Rau et al., 2015), bird
assemblages (Dardanelli and Bellis, 2021), ants (Flores et al., 2018), and amphibians (Rodríguez et al., 2023).
However, little is known about the nestedness patterns of carnivorous mammals in fragmented landscapes.
Carnivorous mammals are a valuable focus for this study as they represent the second most researched group in
relation to the role of the matrix in fragmented landscapes (Ferreira et al., 2018). This study aims to investigate
the nestedness patterns of five carnivorous mammal species across six forest habitat fragments to determine the
likelihood of their local extinction. Given the ongoing and increasing habitat fragmentation in the study area,
we expect to observe nestedness at the species composition level, considering the specific habitat requirements
of each species.
Material and methods
Study area
The study area is located in the southern part of the Coastal Range in the Araucanía Region, southern
Chile (fig. 1), extending from the southern side of the Toltén River to the border with the Los Ríos Region,
covering a total area of 209,430 hectares. This region is characterized by Valdivian laurel forest vegetation,
including species such as Laurelia sempervirens, Persea lingue, and Lophozonia obliqua. Historically, the area
has been heavily modified by human activity and is now primarily composed of remnants of mixed vegetation
and secondary growth forests (Gutiérrez and Becerra, 2018).
Sampling method
We selected 20 native forest fragments larger than 1 hectare based on land-use and land cover mapping
derived from the 2003 Landsat ETM+ satellite image (Regional Plan for Urban Development and Territory —
PRDUyT, LPT-UCT 2003). Each fragment was visited to verify its land-use category, level of intervention,
and the feasibility of conducting fieldwork (e. g., accessibility and permission). Following this verification, the
sample was reduced to six fragments, with one sample collected from each, except for the “Boroa Norte 1” and
“Nueva Etruria” fragments, which were sampled twice.
Table 1 presents the characteristics of the six sampling sites within the fragmented forest landscape of the
Coastal Mountains in the Araucanía Region. The fragment sizes ranged from 3.1 to 239.1 hectares and were in
the communes of Pitrufquén, Toltén, Gorbea, and Loncoche. These fragments consisted of old-growth native
forest, secondary native forest, and mixed forest (native species interspersed with exotics). In each of the six
fragments, we identified the presence of carnivorous mammals using indirect detection techniques (Valente
503Nestedness In A Carnivorous Mammal Assemblage In Forest fragmentsof the Araucanía Region, southern Chile
et al., 2018), such as scent stations (Webster & Beasley, 2019) and active searches for signs (feces and tracks) (da
Silva Chaves et al., 2019) along roads and paths.
Mammal survey
We placed scent stations approximately every 200 meters along existing roads and trails within the native
forest fragments and surrounding areas, such as forestry or agroforestry matrices, to avoid repeated visits to
the same stations (Muñoz-Pedreros et al., 1995). To attract carnivores, we used American wildcat (Lynx rufus)
urine as bait (Muñoz-Pedreros et al., 1995). Additionally, we actively searched for tracks and feces along roads
Fig. 1. Location map of the seven sampling sites in the Coastal Mountains of the Araucanía Region, southern
Chile: 1 — Boroa Norte 1; 2— Licancullín; 3 — El Socorro; 4 — Afquintué; 5 — Nueva Etruria; and 6 — Boroa
Norte 3.
T a b l e 1 . Characterization of the seven sampling sites within a fragmented forest landscape in the
Coastal Mountains of the Araucanía Region, southern Chile
ID
Fragment Site Sector Commune Coord. Ref (UTM) Area
(ha)
Category
of use Matrix
1 Boroa Norte 1 Boroa Norte Toltén 665984N/5652334 E 14.0 BNA AF
2 Licancullín Colonia San Jorge Toltén 678416N/5654733 E 46.8 RN AF
3 El Socorro Colonia San Jorge Toltén 672161N/5654558 E 239.1 BNA F
4 Afquintué Afquintué Loncoche 707838N/5644950 E 6.8 BNA AF
5 Nueva Etruria Comuy y Nueva
Etruria/Mantahue
Pitrufquén/
Gorbea
685227N/5665714 E 146.5 RN F
6 Boroa Norte 3 Boroa Norte Toltén 663638N/5653327 E 3.1 RN AF
Note: BNA — Adult native forest (old-growth forest); RN — Native secondary forest; BM — Mixed
forest; AF — Agroforestry; F — Forestry.4
504 R. Sanhueza, W. Moya & J. R. Rau
and paths throughout the entire sampling period in each fragment. Track identification was based on both field
experience and Acosta and Simonetti’s guide (1999).
Data collection took place over a period of nine months, from July 2006 to May 2007, with four
effective sampling weeks per fragment. Scent stations were active for two consecutive nights in each of the
six fragments. The “Licancullín” and “El Socorro” fragments were sampled in February 2007, “Afquintué”
in March 2007, and “Boroa Norte 3” in May 2007. The “Boroa Norte 1” and “Nueva Etruria” fragments
were sampled twice, in July 2006 and January 2007, and in September 2006 and March 2007, respectively.
These fragments were sampled twice due to suboptimal climatic conditions during the initial sampling,
which hindered carnivore detection. Therefore, sampling was repeated at a different time to ensure the
validity of the data.
Data analys is
To identify the presence of nestedness patterns in the composition of carnivorous mammal species,
we utilized the Binary Matrix Nestedness Temperature Calculator (BINMATNEST), which has gained
prominence for detecting and quantifying nestedness in recent years (Rodríguez-Gironés and Santamaría,
2006; Zungu et al., 2019; Ferrarini et al., 2023). This program also incorporates the earlier Nestedness
Temperature Calculator (NTC) developed by Atmar and Patterson (1993, 1995), as well as a second model
developed by Fischer and Lindenmayer (2002) in which has been used in ecological networks studies (Lin
et al., 2018).
BINMATNEST offers several advantages: (1) it implements a perfectly ordered line that is uniquely
defined; (2) it uses genetic algorithms to reorder rows and columns in a way that minimizes matrix temperature;
and (3) it provides three alternative null models; model 1 (Atmar and Patterson, 1993, 1995), model 2 (Fischer
and Lindenmayer, 2002) and model 3 (Rodríguez-Gironés and Santamaría, 2006), to calculate the statistical
significance of matrix temperature. The program also compares the observed adjacency matrix with a “perfectly
ordered” matrix.
The results generated by this program indicate that higher temperatures (approximately 100 °C) reflect a
lower degree of nestedness, suggesting a reduced probability of local extinctions. Conversely, lower tempera-
tures (around 0 °C) indicate a higher degree of nestedness, with an increased likelihood of local extinctions.
Results
Among the carnivore species potentially present in our study area, we recorded the
following five: the hog-nosed skunk (Conepatus chinga Molina, 1782), kodkod (Leopardus
guigna Molina, 1782), cougar (Puma concolor Linnaeus, 1771), gray fox (Lycalopex griseus
Gray, 1837), and culpeo fox (Lycalopex culpaeus Molina, 1837). The hog-nosed skunk was
recorded at four sites: the two largest fragments, El Socorro and Nueva Etruria, as well as
two smaller fragments, Boroa Norte 1 and Boroa Norte 3. The cougar was observed in three
fragments, including the two largest (El Socorro and Nueva Etruria) and one smaller frag-
ment (Boroa Norte 3). The kodkod was detected in two fragments: one large (Nueva Etru-
T a b l e 2 . Number of records of five species of carnivorous mammals in six fragments in the Coastal
Mountains of the Araucanía Region, southern Chile
Fragments/Species Culpeo fox Gray fox Hog-nosed skunk Kodkod Cougar Total per fragment
Nueva Estruria 1 2 1 1 2 7
El Socorro 0 0 1 0 3 4
Boroa Norte 3 0 0 1 0 1 2
Boroa Norte 1 0 0 1 0 0 1
Licancullín 0 0 0 1 0 1
Afquintúe 0 2 0 0 0 2
Total per species 1 4 4 2 6 17
505Nestedness In A Carnivorous Mammal Assemblage In Forest fragmentsof the Araucanía Region, southern Chile
ria) and one small (Licancullín). The gray fox was specifically recorded in Nueva Etruria
and Afquintúe, while the culpeo fox was found only in Nueva Etruria (table 2).
The nesting temperature calculated by the BINMATNEST software was 13.661°C, in-
dicating a nested pattern and the potential for local extinctions. The mean and variance
(mean ± var.) for the three models were as follows: 27.516 ± 92.051 (p = 0.04) for model
1, 23.981 ± 107.290 (p = 0.14) for model 2, and 21.624 ± 102.047 (p = 0.24) for model 3.
Model 1, developed by Atmar and Patterson (1993, 1995), demonstrated greater robust-
ness, as indicated by a significant p-value and the lowest variance.
By other hand, table 3 presents the presence and absence of the five identified carni-
vore species across six forest fragments in the Coastal Mountains of the Araucanía Region.
Discussion
The studies of species composition emphasize life history traits and interactions among
species (i. e., a biological approach), allowing for the identification of those most sensitive
to habitat fragmentation. Our research focused on species composition to better explain
potential nested patterns, as opposed to solely studying species richness, which tends to
adopt a more statistical or mathematical approach (see Rau, 2020).
Our results revealed a nested pattern in the species composition of the carnivorous
mammal assemblage within forest fragments in the Coastal Mountains of the Arau-
canía Region. This pattern persisted despite the high mobility of carnivorous mam-
mals, which is associated with their large home ranges and territorial behavior, as well
as their generalist habitat use. It is important to clarify that this study focused exclu-
sively on forest fragments, meaning that other biotopes, such as open fields, agricul-
tural lands, or wetlands, were not included in the analysis. This limitation is significant
because many carnivorous mammals are known to use a variety of habitats beyond
forests, particularly for foraging or movement between fragmented areas. For exam-
ple, the hog-nosed skunk, with an estimated average home range of 1.9 km² (Iriarte
and Jaksic, 2017), occupies a variety of environments across its distribution, including
shrublands, savannas, forests, mountains, high plateaus, parks, and steppes (Donadio
et al., 2001; Riojas-López et al., 2019)
Another carnivore, the kodkod, with an estimated home range of 2.4 to 2.9 km² (Iriarte
and Jaksic, 2017), is typically a forest specialist (Correa and Roa, 2005; Marín et al., 2022;
Marín, et al., 2022). In contrast, the cougar, with home ranges estimated between 65 and
90 km² for males and 40 to 80 km² for females (Franklin et al., 1999; Iriarte and Jaksic,
2017), prefers more open habitats or exotic plantations, such as Pinus radiata, in areas with
T a b l e 3 . Presence and absences of five species of carnivorous mammals in six fragments in the Coastal
Mountains of the Araucanía Region, southern Chile
Fragments/Species Culpeo fox Gray fox Hog-nosed skunk Kodkod Cougar
Nueva Estruria 1 1 1 1 1
El Socorro 0 0 1 0 1
Boroa Norte 3 0 0 1 0 1
Boroa Norte 1 0 0 1 0 0
Licancullín 0 0 0 1 0
Afquintúe 0 1 0 0 0
506 R. Sanhueza, W. Moya & J. R. Rau
open canopies that facilitate hunting (Lira and Naranjo, 2003; Charre-Medellín et al., 2012;
Dumont et al., 2022). The culpeo fox and gray fox, with home ranges ranging from 4.5 to
1,140 km² and 3 to 4 km², respectively, occasionally inhabit forested areas but generally
favor more open habitats or shrublands, where small mammals are more abundant and
accessible (Quintana et al., 2009; Medel et al., 2022).
In the fragments of Boroa Norte 1, Licancullín, El Socorro, and Afquintúe, the canopy
is denser, accompanied by a thicker and more extensive layer of leaf litter. In contrast, the
fragments of Nueva Etruria and Boroa Norte 3 have a more open canopy, with a greater
prevalence of herbaceous plants and shrubs. Our findings align with previous studies on
the culpeo fox conducted in central Chile (Acosta-Jamett and Simonetti, 2004), central Ar-
gentina (Pia et al., 2013), Colombia (Noguera-Urbano et al., 2016), and the high-Andean
ecosystems of Ecuador (Guntiñas et al., 2019), which found that this species is more abun-
dant in high-altitude areas characterized by high humidity, extreme temperatures, and ho-
mogeneous vegetation typical of open, treeless habitats.
In a separate studies, Lantschner (2012) and Haley-Marie (2023) have found that the
culpeo fox and gray fox primarily use continuous native vegetation but also inhabit dense
conifer plantations. These species tend to be more abundant in fire-affected zones and
sparse plantations than in dense ones. In contrast, the cougar is present across all habi-
tat types, showing no preference (Winkel et al., 2023). The kodkod, on the other hand, is
closely associated with native Mediterranean forests and temperate rainforests, favoring ar-
eas with significant vegetation cover (Schüttler et al., 2017). This species has been sparsely
studied, and its ecology and habitat use remain poorly understood.
However, studies such as Schüttler et al. (2017) have found that the kodkod is for-
est-dependent and makes intensive use of forest edges, particularly near water sources
in Chilean temperate rainforests. The kodkod compensates for the lack of continuous
forest by maintaining a larger habitat range and efficiently utilizing forest edges, likely
due to higher prey availability in these areas. Additionally, this species is adaptable to
fragmented landscapes dominated by human activity, using small forest fragments and
vegetation corridors within agricultural matrices to navigate the landscape (Gálvez,
2013). There are also reports of kodkods inhabiting dense pine plantations (Acosta-
Jamett and Simonetti, 2004).
This species (kodkod) has a restricted distribution, primarily found in central and
southern Chile (30°–48° S), including the large island of Chiloé off the southern coast, and
marginally in adjacent areas of southwestern Argentina (39°–46° S, west of 70° W) (Napoli-
tano et al., 2015). However, more recently, Napolitano et al. (2020) reported new records
from the southern Coquimbo and northern Valparaíso regions, marking the northernmost
confirmed occurrences of this species to date. In contrast, little is known about the habitat
use of the gray fox. A recent study by Osorio (2021) found that this species is tolerant to
landscape changes, showing no significant response to burns, and is often found near hu-
man settlements and rivers.
Although our study attributed the presence of highly mobile carnivores to the for-
est fragments, it is likely that these fragments serve primarily as corridors or stepping
stones within their larger home ranges. For instance, the presence of cougars, which
have large home ranges, is likely more attributable to the overall landscape rather than
the specific fragments. Carnivores, in general, are highly vulnerable to local extinc-
tion due to habitat fragmentation, primarily because of their low population densities
and large territorial requirements (Rocha et al., 2018). However, not all carnivores are
equally sensitive to landscape and local-level variables; their vulnerability is largely de-
507Nestedness In A Carnivorous Mammal Assemblage In Forest fragmentsof the Araucanía Region, southern Chile
termined by body size. Larger, more generalist carnivores, such as the culpeo fox, have
larger home ranges and are therefore more sensitive to habitat fragmentation (Crooks,
2002). In contrast, specialist carnivores, such as the kodkod, are likely the most affected
(Swihart et al., 2003).
In Chile, the effects of native forest fragmentation on the kodkod and culpeo fox
have been studied in Los Queules National Reserve and the surrounding pine frag-
ments and plantations (Acosta and Simonetti, 2004). For the kodkod, it has been sug-
gested that fragmentation and habitat loss may severely impact the species, particularly
females, which are more sedentary and have smaller home ranges (Sanderson et al.,
2002). In contrast, the culpeo fox, a habitat generalist (Medel and Jaksic, 1988), may
be less affected by forest fragmentation. This species appears to prefer large areas of
native forest over fragments and plantations, likely due to the protection they offer.
It is generally expected that smaller fragments will support a less diverse set of species
compared to larger areas (Rau et al., 2015).
On the other hand, the nesting temperature of 13.661 °C obtained in our study indi-
cates a high degree of order in the carnivorous mammal assemblage in the forest fragments
studied. In terms of nestedness theory, a low temperature suggests that communities are
hierarchically organized, with some species being more frequent and widely distributed,
while others are restricted to specific habitats or fragments (Mittelbach and McGill, 2019).
This level of order may reflect local extinction processes or the preferential selection of
habitats by certain species.
Previous studies using BINMATNEST, such as those by Atmar and Patterson (1993,
1995), Lin et al. (2018), Mariani et al. (2024) consider that the rank of each species invari-
ably depends on the interactions stipulated with other species through the adjacency ma-
trix of the network. A common ranking method in economic and ecological networks is
to sort the nodes such that the layout of the reordered adjacency matrix looks maximally
nested with all nonzero entries packed in the upper left corner, called Nestedness Maximi-
zation Problem (NMP) have shown that lower temperatures are typically associated with
highly structured systems and significant levels of nestedness, often linked to habitat frag-
mentation and the differential ability of species to persist in fragmented environments. In
contrast, higher temperatures imply greater randomness in species distribution, indicating
less hierarchical structuring. In our case, the relatively low temperature supports the hy-
pothesis that forest fragments are undergoing local extinction processes, leading to a nested
pattern in the species present.
The presence/absence and the number of species records in each fragment could
indeed be influenced by the specifics of our data collection. Although we conducted
fieldwork over a period of nine months, using scent stations for two consecutive
nights per fragment, it is possible that the number of days dedicated to data collec-
tion was insufficient to capture the full diversity of carnivore species in each frag-
ment. For example, two nights might not be enough time to detect species with large
home ranges, like the cougar, which may not pass through the fragments during such
short timeframes.
Moreover, climatic conditions and the season during which the sampling took
place could have impacted the detection of carnivores. Some species may exhibit sea-
sonal behavior changes, such as reduced activity during unfavorable weather condi-
tions or breeding seasons, which could have reduced the likelihood of detection (Cara-
vaggi et al., 2018; Levy et al., 2019; Guiden and Orrock, 2020; Abrahms et al., 2022).
This is particularly relevant for fragments like “Nueva Etruria” and “Boroa Norte 1,”
508 R. Sanhueza, W. Moya & J. R. Rau
which were sampled twice due to suboptimal weather conditions during the initial
sampling. Therefore, the variation in the number of species recorded across fragments
may reflect not only habitat characteristics but also temporal limitations in the data
collection process. To address these potential biases in future studies, extending the
sampling period, increasing the number of active days per fragment, and considering
seasonal variations in carnivore activity could provide a more comprehensive under-
standing of species presence and distribution within the forest fragments.
We pointed out that to planning an effective biodiversity conservation measure for
fragmented environments, it is essential to safeguard the probability of recolonization
of carnivorous mammals (e. g., to plan the management of the fragmented landscape
so that it is a suitable (permeable) substrate for the dispersal of individuals). This prob-
ability of recolonization depends on the spatial relationships between landscape ele-
ments used by populations, including habitat patches and matrix elements, dispersal
characteristics of the organisms, and the temporal changes in the structure of the land-
scape (Fahrig and Merriam, 1994). Therefore, to minimize the effects of fragmentation
and to protect the associate carnivore mammals its essential to have good knowledge
of species biology and its habitat use.
Conclusion
This study provides important insights into the composition and distribution of
carnivorous mammals across fragmented forest habitats in the Coastal Mountains of
the Araucanía Region, Chile. Five carnivore species were recorded in the study area:
the hog-nosed skunk, kodkod, cougar, gray fox, and culpeo fox. These species exhib-
ited varying degrees of sensitivity to habitat fragmentation, which may reflect their
different ecological requirements and life history traits. Also, the study reveals a nested
pattern in the species composition of carnivores, as indicated by the low nesting tem-
perature. This suggests that local extinctions are likely, particularly in smaller frag-
ments where species such as the hog-nosed skunk and kodkod, which have smaller
home ranges, may be more vulnerable.
The findings suggest that species with larger home ranges and generalist habits,
such as the cougar and culpeo fox, are less sensitive to fragmentation and may use
fragmented habitats as corridors or stepping stones. In contrast, species more closely
associated with dense forest cover, such as the kodkod, are likely more affected by
fragmentation, as they depend on specific habitat features for survival. However, the
species distribution and nested pattern observed in this study may not fully represent
the wider landscape use of these predators, as it was restricted to the forest fragments
and not other biotypes.
Finally, the study highlights the importance of larger forest fragments in maintaining
carnivorous mammal diversity. Smaller fragments are likely to support a less diverse as-
semblage of species, as predicted by the theory of island biogeography. For conservation
efforts, it is essential to prioritize the preservation of larger continuous forest areas and the
restoration of connectivity between fragments to mitigate the negative effects of fragmenta-
tion on sensitive species, particularly the kodkod and hog-nosed skunk. Further research
is needed to monitor the long-term impacts of fragmentation and to explore how differ-
ent species adapt to changing landscapes, particularly in regions like the Araucanía, where
human-driven land-use changes are accelerating.
509Nestedness In A Carnivorous Mammal Assemblage In Forest fragmentsof the Araucanía Region, southern Chile
Acknowledgments
Authors wish to thank the Dirección de Investigación Universidad de Los Lagos for support this study.
As well as to Soraya Sade for her assistance with the final editing of the text, to Gonzalo Rebolledo for a previ-
ous editing of the map and Laboratorio de Planificación Territorial, Universidad Católica de Temuco (LPT-
UCT), for providing cartographic databases. RS wish to thank the Magister en Ciencias mención Producción,
Conservación y Manejo de Recursos Naturales, Universidad de Lagos, Chile. Finally, we thank and anonimous
reviewer and the Invited editor (Dr. Mikhail Rusing) for cogent corrections and suggestions that greatly im-
prove the final version of our manuscript.
Disclosure statement
No potential conflict of interest was reported by the authors
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Received 28 February 2024
Accepted 10 December 2024
|
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| institution | Zoodiversity |
| issn | 2707-7268 |
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| language | English |
| last_indexed | 2026-08-21T01:01:42Z |
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| spelling | oai:ojs.akademperiodyka.org.ua:article-6022026-08-20T12:37:09Z Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile Sanhueza, R. Moya, W. Rau, J. R. habitat fragmentation carnivore density nestedness Araucanía Forest species composition Habitat fragmentation is one of the main factors threatening biodiversity, especially in regions where natural ecosystems are under anthropogenic pressure, such as the Araucanía region in southern Chile. This study explores the presence of nestedness patterns in the composition of carnivorous mammal species in six fragments along the southern part of the Coastal Range in the Araucanía region. Using scent stations and actively searched five species were recorder over a nine-month period: hog-nosed skunk, kodkod, cougar, gray fox, and culpeo fox. The analysis revealed that the nested pattern, with a temperature of 13.661 °C, suggests a risk of local extinction in smaller fragments (less than 14 ha). The hog-nosed skunk was detected in four fragments, including one of the smaller ones (3.1 ha), while the cougar and kodkod were more restricted, being recorded in three and two fragments, respectively. The culpeo fox and gray fox were present in only one or two fragments. These results support the island biogeography theory, highlighting that larger fragments sustain greater species diversity. This study highlights the importance of conserving larger forest fragments to maintain carnivore diversity, as specialized species rely on denser and more connected habitats. It is recommended to prioritize the preservation of continuous forest areas and restore connectivity between fragments to mitigate the negative effects of fragmentation. Publishing House "Akademperiodyka" of the National Academy of Sciences of Ukraine 2024-10-16 Article Article application/pdf https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/602 10.15407/zoo2024.06.501 Zoodiversity; Vol. 58 No. 6 (2024): Zoodiversity Zoodiversity (Vestnik Zoologii); Том 58 № 6 (2024): Zoodiversity 2707-7268 2707-725X 10.15407/zoo2024.06 en https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/602/295 Copyright (c) 2024 R. Sanhueza, W. Moya, J. R. Rau |
| spellingShingle | Sanhueza, R. Moya, W. Rau, J. R. Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title | Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title_full | Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title_fullStr | Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title_full_unstemmed | Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title_short | Nestedness in a Carnivorous Mammal Assemblage in Forest Fragments of the Araucanía Region, Southern Chile |
| title_sort | nestedness in a carnivorous mammal assemblage in forest fragments of the araucanía region, southern chile |
| topic_facet | habitat fragmentation carnivore density nestedness Araucanía Forest species composition |
| url | https://ojs.akademperiodyka.org.ua/index.php/Zoodiversity/article/view/602 |
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