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Nutritional ecology of martens (Martes foina and Martes martes) in the

western Italian Alps

Article  in  Ecological Research · November 2021

DOI: 10.1111/1440-1703.12277

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DOI: 10.1111/1440-1703.12277

ORIGINAL ARTICLE

Nutritional ecology of martens (Martes foina and Martes

martes) in the western Italian Alps

Marco Granata1 | Andrea Mosini2 | Manuel Piana2 | Francesco Zambuto3 |

Enrica Capelli3 | Alessandro Balestrieri3,4

1
Department of Life Sciences and Systems
Biology, University of Turin, Turin, Italy
2
Valgrande Società Cooperativa-Studi,
Verbania Possaccio, Italy
3
Department of Earth and Environmental
Sciences, University of Pavia, Pavia, Italy
4
Department of Environmental Sciences
and Policy, University of Milan,
Milan, Italy
Correspondence
Marco Granata, Department of Life
Sciences and Systems Biology, University
of Turin, Via Accademia Albertina
13, 10123 Turin, Italy.
Email: marcocgranata@gmail.com
Funding information
Val Grande National Park
Abstract
The macronutrient requirements of the strictly related pine marten (Martes
martes) and stone marten (Martes foina) are almost identical, but, at range
scale, in areas of putative sympatry (overlapping European ranges) the stone
marten tends to be more frugivorous, which makes the contribution of carbo-
hydrate energy to be higher than the target. In contrast, the macronutrient
intake of the pine marten would be unaffected by the occurrence of the stone
marten, supporting the putative dominance of the first in interspecific interac-
tions. Most available studies examined marten diets separately, highlighting
the need for further studies in areas of actual co-occurrence. With this aim, we
compared the two martens' diets in the Italian Alps both in sympatric and allo-
patric conditions, as assessed by the genetic identification of scats. Although
fruit and rodents formed the bulk of both species' diets, as predicted the stone
marten consumed twice as many fruit species as the pine marten and ate fruit
more often in areas of sympatry, thus consuming less protein and more carbo-
hydrates respect to its intake target. This competition-driven, nutritionally
imbalanced diet may affect the fitness of stone marten populations and play an
important and still underreported role in regulating the relative abundance of
marten communities.

KEYWORDS
allopatry, competition, diet, macronutrient ratio, sympatry

1 | INTRODUCTION very similar in terms of morphology, habitat preferences,

For closely related, morphologically similar species, inter-
specific relationships are a major biological factor that can
restrict the potential distribution of the less competitive spe-
cies when resource availability is limited (MacArthur, 1972;
Violle et al., 2011). Pine marten Martes martes and stone
marten Martes foina are medium-sized mustelids which are
and foraging behavior (Larroque et al., 2015). Although
stone martens are usually more flexible in their habitat
requirements, both species prefer forested habitats and tend
to avoid open areas (Virgo
s et al., 2012). They co-exist over
a large area, including most of central and southern Europe
(Mitchell-Jones et al., 1999), setting the conditions for com-
petition to occur.

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© 2021 The Authors. Ecological Research published by John Wiley & Sons Australia, Ltd on behalf of The Ecological Society of Japan.

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2 GRANATA ET AL.
Although both the pine marten and stone marten can
consume a large variety of food items, their trophic
niches overlap largely (Remonti et al., 2012; Zalewski,
2004). Local and seasonal variations in food availability
make it difficult to compare the diets of generalist spe-
cies, an impediment that can be overcome by integrating
the traditional approach of evaluating dietary niches in
terms of foods alone with their macronutrient composi-
tion (Machovsky-Capuska et al., 2016). Both laboratory
and field studies have demonstrated that animals can
self-select species-specific ratios of macronutrients—pro-
tein, lipids, and carbohydrates (Balestrieri et al., 2019;
Raubenheimer & Simpson, 1997), and macronutrient
intakes can be used to effectively compare the diets of
generalist predators (Gazzola & Balestrieri, 2020).
Recently, by reviewing available data on the foods
eaten by both pine and stone martens throughout Europe
in the framework of nutritional ecology, it has been dem-
onstrated that although the macronutrient requirements
of the two species are almost identical, in areas of puta-
tive sympatry (areas of overlap of their respective
European ranges) the stone marten tends to be more fru-
givorous, entailing a lower contribution of protein energy
respect to the intake target (Gazzola & Balestrieri, 2020).
This result would bring new evidence in support of the
dominant role played by the pine marten in the two spe-
cies' interaction (Delibes, 1983; Virgo
s et al., 2012). None-
theless, with a few exceptions (Monterroso et al., 2016;
Posłuszny et al., 2007), most of the studies reviewed by
Gazzola and Balestrieri (2020) (93.1%) examined the diet
of only one marten species, preventing the investigation
of the effects of interspecific competition on their food
niches and highlighting the need for further studies in
areas of co-occurrence of the two martens.
The outcome of competition depends largely on the com-
plexity of habitats (MacArthur, 1972). In mountainous areas,
extensive seasonality (Vanham et al., 2008) and pronounced
altitudinal gradients for climatic factors such as temperature,
rainfall, and solar radiation (Legates & Willmott, 1990;
Rolland, 2003) are expected to affect the distribution and
availability of food resources. The two martens co-occur
throughout the Italian Alps (Fonda et al., 2021), offering the
opportunity to investigate their competitive interactions at
the altitudinal limits of their overlapping ranges.
Nonetheless, due to their elusive behavior and need
for genetic analysis to identify scats (Davison et al., 2002;
Statham et al., 2005), marten ecology has been poorly
studied in the Alps so far. Scat-based surveys were used
to assess both the diet (Cantini, 1991; Prigioni
et al., 2008) and habitat preferences (Pedrini et al., 1995)
of genus Martes as a whole, while roadkills provided the
opportunity to gather some information on the stomach
content of pine martens (Agnelli & De Marinis, 1995).
With the aim of comparing the diets of pine and stone
martens in mountain areas of either sympatry or allopa-
try, we (1) carried out an extensive genetic survey for
marten scats in the western Italian Alps in 2019
(Figure 1c) and (2) analyzed genotyped fecal samples col-
lected in NE Piedmont in 2015–2017 (Figure 1a,b), as part
of a study aimed at assessing the distribution of mustelids
in the Val Grande National Park and neighboring areas
(Lepontine Alps; Balestrieri et al., 2018). In the first area
both martens are known to occur (based on museum
archives and records collected in the period 1990–2010;
Sindaco & Carpegna, 2010), while the pine marten is
suspected to have recolonised the latter area only in the
last decade (Balestrieri et al., 2021). Following Remonti
et al. (2016) the estimated volumes of food items used by
both martens were then transformed in macronutrient
energy ratios to allow the sound comparison of their feed-
ing requirements (Gazzola & Balestrieri, 2020).
We expected the stone marten to be more frugivorous
when sympatric with the pine marten than in allopatry.
However, the supposed reduced availability of food
resources along the altitudinal gradient was expected to
narrow the trophic niches of both martens and increase
food niche overlap.
2 | METHODS
2.1 | Study area
The study was conducted in the western Italian Alps
(Piedmont region, NW Italy; Figure 1), including the
Maritime and Cottian Alps in the southern sector, and
the Graian, Pennine and Lepontine Alps in the northern
sector (International Standardized Mountain Subdivision
of the Alps; Marazzi, 2005). The whole study area was
included within the biogeographical Alpine region
(Figure 1). In such a wide area (11,275 km2), ranging
between 188 and 4565 m a.s.l., climate varies greatly:
southern areas are generally characterized by a more
Mediterranean climate, with warm and dry summers and
mild and wet winters, while northern areas show a typi-
cal Alpine climate, with cool summers and cold winters.
Anyway, air temperature (yearly means range between
6.5 and 8.0
C) strictly depends on altitude and, as a gen-
eral scheme, three main vegetation types can be
described along the altitudinal gradient:
• 500–1500 m a.s.l.: broad leaves and mixed forests, gen-
erally dominated by beech Fagus sylvatica, chestnut
Castanea sativa, common ash Fraxinus excelsior, and
green alder Alnus viridis; fruit trees and shrubs include
whitebeam Sorbus aucuparia and S. aria, wild cherry
GRANATA ET AL. 3

F I G U R E 1 Study areas: Graian and SW Alps (a) and Lepontine Alps (b). The first area was sampled by superimposing a 10  10 km2
square grid coinciding with the UTM kilometric grid (c) [Color figure can be viewed at wileyonlinelibrary.com]

Prunus sp., blackberries and raspberries Rubus spp.
and rose berries Rosa spp.
• 1500–2000 m a.s.l.: coniferous forests, mainly larches
Larix decidua, followed by Scot's pines Pinus sylvestris,
Norway spruces Picea abies, silver fir Abies alba, and
Arolla pines Pinus cembra.
• 2000–2500 m a.s.l.: Alpine prairies, sometimes delimited
by a discontinuous shrub belt, generally with alpenrose
Rhododendron ferrugineum, juniper Juniperus sp., rasp-
berries and bilberries Vaccinium spp.
The limit of perennial snows is about 3000 m a.s.l.
4 GRANATA ET AL.
2.2 | Sampling and species identification
Fresh scats were searched for by two surveyors along linear
transects coinciding with paths (N = 53; mean
length = 7.2 km, min-max: 2.5–14.8 km), ranging in alti-
tude between 500 and 2000 m a.s.l. The southern sector
and Graian Alps (provinces of Turin and Cuneo; Figure 1a)
were sampled between February 2019 and December 2019
(N = 30). To survey this large area as uniformly as possible,
we superimposed a 10  10 km2 square grid coinciding
with the UTM kilometric grid, obtaining 116, 100 km2
squares (Figure 1c). Those including only a small portion of
the study area or lacking suitable paths were excluded. Of
the resulting 52 squares, a subset of 30 squares (57.7%) was
selected. The Lepontine Alps (23 transects; Figure 1b) were
surveyed between July 2015 and August 2017 (Balestrieri
et al., 2018). Only apparently “fresh” scats suspected to
belong to martens based on their size and morphology were
collected; surveys were delayed by at least 1 week after
heavy rain. All collected scats were geo-referenced with a
portable GPS. A small portion (about 10%) of each scat was
stored in an autoclave tube containing 96% ethanol and
then frozen at 20
C until processed (Balestrieri
et al., 2016; Ruiz-Gonz
alez et al., 2008).
Fecal DNA was extracted using the QIAamp DNA
Stool Mini Kit (Quiagen) according to the instructions pro-
vided by the manufacturer. The specific identification of
scats was accomplished by the PCR-RFLP method devel-
oped by Ruiz-Gonz
alez et al. (2008). A 276 bp fragment of
mDNA (D-loop) was amplified by two primers (Mm_L1
and Mm_H1), specific for the genera Martes and Mustela.
Then, PCR products were digested by two restriction
enzymes (RsaI and HaeIII), which generate different
restriction patterns for each species, allowing their unam-
biguous identification (Ruiz-Gonz
alez et al., 2008).
2.3 | Diet analysis
Each scat was washed over a sieve with a 1-mm mesh
size and undigested remains were inspected to identify
food items based on the occurrence of diagnostic ele-
ments (e.g., seeds for fruit, wings, legs, and cuticle for
insects, guard hairs and teeth for mammals) and their
comparison with reference collections and identification
keys (Debrot et al., 1982; Teerink, 2003). According to the
method by Kruuk and Parish (1981), the relative volume
of each food item “as ingested” was assessed by eye; this
method has been widely used for assessing carnivore
diets and provides volume estimates as accurate as those
obtained by the analysis of stomach contents (Balestrieri
et al., 2011). Results were expressed as: percent frequency
(F% = number of scats containing a specific food
item  100/total number of fecal samples), percent vol-
ume (V% = total estimated volume of each food item as
ingested/number of scats containing that item), and per-
cent mean volume (mV% = F%  V%/100), which out-
lines the proportional contribution of each food item to
the overall diet (Kruuk & Parish, 1981). With the aim of
assessing as precisely as possible the relative volume of
each prey in each scat, we inferred the minimum number
of items of each prey type based on the number of diag-
nostic hard parts following Balestrieri et al. (2019).
Trophic niche overlap was assessed by Pianka's O
index (Pianka, 1973),
P
n
pij pik
i¼0
Ojk ¼ sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
ffi,
P n P n
p2ij p2ik
i¼0 i¼0
where pij and pik being the mean volumes (mV) of each
food item in the diets of species j and k. According to
Krebs (1989), to assess niche overlap food items were cat-
egorized to the lowest possible level (i = 34; Table 1). For
exploratory analyses, food items were grouped in 10 main
food categories (fruit, Rosaceae, cultivated fruit, insects,
birds, rodents, Gliridae, Arvicolidae, Muridae, ungulates),
and raw frequency and volume data were compared
between species by the chi-square test (χ 2) and Mann–
Whitney's U test, respectively. To account for multiple
tests on related data, the level of significance was calcu-
lated by Bonferroni's sequential technique (Rice, 1989).
Available literature on the nutritional composition of
food items was checked to obtain, on a wet-weight basis,
the mean percentage of protein, lipids, and carbohydrates
of those used by martens (Remonti et al., 2016;
USDA, 2014). For the calculation of the nutrient profiles,
percent mean volumes were then converted in macronutri-
ent energy contents (expressed as percentage of the total
metabolizable energy per food item; Raubenheimer, 2011)
using the following coefficients: protein = 14.64 kJ/g;
fat = 35.56 kJ/g; nonstructural carbohydrates = 14.64 kJ/g
(Hewson-Hughes et al., 2011; Remonti et al., 2016). Right-
angled mixture triangles, which allow representation of the
three components (i.e., protein, lipids, and carbohydrates)
in a two-dimensional nutrient space (Raubenheimer, 2011),
were used to visually compare the macronutrient composi-
tion of the diet of different marten populations.
3 | RESULTS
Out of 311 fecal DNA samples collected, 253 (81.3%)
could be assigned to one of the target species. As
GRANATA ET AL. 5

T A B L E 1 Diet of pine- (PM) and stone martens (SM) in the western Italian Alps, in areas of either allopatry or sympatry; F% is the
percent frequency of occurrence, V% is the estimated percent of volume of the food items

Allopatric SM Sympatric SM Allopatric PM Sympatric PM

Food items F% V% F% V% F% V% F% V%
Fruit 26.0 63.9 48.6 85.6 26.3 60.0 36.2 50.3
Rosaceae 22.0 65.8 35.1 80.0 21.1 50.0 27.7 42.7
Rosa sp. 0.7 100.0 8.1 73.3 8.5 41.3
Rubus sp. 12.7 65.3 5.4 55.0 10.5 25.0 10.6 29.0
Prunus sp. 2.7 51.3 13.5 100.0 5.3 100.0 6.4 48.3
Sorbus aucuparia 4.0 48.3 2.7 100.0 5.3 100.0 2.1 100.0
Fragaria 2.7 100.0
Plum 1.3 17.5
Cherry 2.7 60.0
Apple 2.7 50.0
Pear 2.0 100.0 5.3 50.0
Vaccinium sp. 5.4 100.0 2.1 50.0
Juniperus communis 0.7 10.0
Castanea sativa 0.7 10.0 2.1 100.0
Grape 3.3 46.0 2.1 50.0
Persimmon 0.7 50.0
Undetermined 0.7 20.0 5.4 100.0 2.1 100.0
Insects 10.7 51.9 10.8 63.8 5.3 100.0 6.4 38.3
Coleoptera larvae 2.0 45.0 8.1 50.0
Coleoptera 0.7 5.0 2.7 5.0 5.3 100.0
Orthoptera 1.3 7.5
Hymenoptera 6.0 61.7 6.4 38.3
Dermaptera 0.7 100.0
Undetermined 1.3 10.0 2.7 100.0
Passeriformes 8.0 66.7 5.4 52.5 4.3 75.0
Mammals 74.7 94.9 54.1 90.0 84.2 95.0 83.0 89.2
Rodents 74.0 94.9 51.4 89.5 84.2 91.3 63.8 83.3
Sciuridae 1.3 75.0
Gliridae 16.0 94.8 2.7 100.0 31.6 85.0 10.6 89.0
Glis glis 16.0 94.8 2.7 100.0 31.6 85.0 8.5 98.8
E. quercinus 2.1 50.0
Arvicolidae 32.7 93.9 40.5 93.3 21.1 100.0 23.4 88.2
C. glareolus 19.3 96.9 40.5 93.3 15.8 100.0 23.4 88.2
Chionomys nivalis 5.3 80.0 5.3 100.0
Undetermined 8.0 95.8
Muridae 21.3 94.8 10.8 75.0 31.6 75.0 38.3 82.2
Apodemus sp. 21.3 94.8 10.8 75.0 31.6 75.0 38.3 82.2
Undetermined 4.0 86.7 10.5 70.0
Ungulates 1.3 100.0 10.6 97.0
Carnivores 0.7 100.0 2.1 100.0
Garbage 2.1 100.0
6 GRANATA ET AL.

expected, in the Lepontine Alps (Figure 1a) the stone
marten outnumbered the pine marten (150 vs. 19), while
in the second study area (Figure 1b,c) the two species
were confirmed to occur in sympatry, surveys yielding
almost equal numbers of genetically identified scats
(37 vs. 47, respectively).
In both areas, fruit and rodents formed the bulk of
both martens' diet, with other food resources (insects,
birds, and carrion) usually totaling less than 5% in mV%
(Table 1 and Figure 2). In the diet of both martens fruit
mainly consisted of Rubus berries, wild cherries, and rose
berries (Table 1). Wood mice (Apodemus sp.) and bank
voles (Clethrionomys glareolus), together with the edible
dormouse (Glis glis) in the Lepontine Alps, were the most
preyed rodents by both species (Table 1). Overall, pine
martens ate ungulate carrions (χ 2 = 7.7, 1 df, p < 0.05)
and mice (χ 2 = 7.9, 1 df, p < 0.05) more frequently.
In sympatry, food niche overlap was 0.66. Mice were
used the most by pine martens in terms of both frequency
(χ 2 = 8.1, 1 df, p < 0.05) and volume (U = 625.5,
p < 0.05). In sympatry, pine and stone martens used
almost the same number of fruits (7 vs. 8, respectively),
while in the Lepontine Alps the stone marten used twice
as much fruit species than the pine marten (10 vs. 5).
The diet of the pine marten in sympatry and allopatry
did not differ (no significant difference was found for
both the frequency of use and volume of the 10 main
food categories), while the stone marten preyed less on
rodents and ate fruit more often (χ 2 = 7.2, 1 df, p < 0.05,
for both) in areas of sympatry with the pine marten.
Juniper berries, chestnuts, and rose berries were the
most carbohydrate-rich fruit (98.0%–81.0% energy), although

F I G U R E 2 Percent frequency of occurrence (F%) versus the estimated percent volume (V%) of 10 main food categories used stone
martens (a) and pine martens (b) in allopatry and by stone martens (c) and pine martens (d) in sympatry; isopleths connect points of equal
percent mean volume (Vm%) in the diet (see section 2)
GRANATA ET AL.
F I G U R E 3 Right-angled mixture triangle showing the
estimated macronutrient intake of allopatric (All) and sympatric
(Sym) Martes martes (MM) and Martes foina (MF). Data are
expressed as percentages of metabolizable energy derived from each
macronutrient
only the latter appreciably contributed to the diet of the pine
marten (mV% = 2.5). Considering foods of animal origin,
insects and, particularly, Coleoptera larvae were the lipid-
richest resource (65.0%), while small mammals were the
main source of protein energy (≈50%, on average).
Pine marten diets in the two study areas provided
similar ratios of protein to nonprotein energy, averaging
46% protein, 44% lipid, and 10% carbohydrate energy
(Figure 3). Stone marten macronutrient ratios were con-
sistent with those of the pine marten when allopatric
(46%, 46%, and 8%, respectively), while in sympatry stone
martens consumed less protein (39%) and more carbohy-
drates (20%; Figure 3).
4 | DISCUSSION
Both martens are generalist feeders, with small rodents
and fruits often forming the bulk of their diets
(Clevenger, 1994; Papakosta et al., 2014; Zalewski, 2004;
Zhou et al., 2011). By reviewing available diet data at
range scale and comparing the macronutrient ratios of
pine marten and stone marten, Gazzola and
Balestrieri (2020) demonstrated that where the two species'
ranges overlap the stone marten tends to eat more carbo-
hydrates and less protein than the pine marten, while the
macronutrient intake of the latter seems to be unaffected
by the occurrence of its congener. Nonetheless, most of
the studies reviewed by these authors did not compare the
7
feeding requirements of the two martens in the same envi-
ronmental conditions. To overcome this gap, we applied
mDNA-based identification of scats to assess and compare
the macronutrient composition of both martens' diets in
the sector of the biogeographical Alpine area coinciding
with the western Italian Alps.
In this study, contrary to our expectations, food avail-
ability allowed some differential use of food resources by
sympatric martens; trophic niche overlap was lower than
that reported for the Swiss Alps (0.94; Marchesi
et al., 1989) and similar to that recorded by Posłuszny
et al. (2007) in central Poland (0.67). In accordance with
the range-scale trend (Gazzola & Balestrieri, 2020), in
areas of sympatry variation in the frequency of use of
fruit and rodents made stone marten's protein intake
lower than that of the pine marten. Results are consistent
with those reported for genetically identified sympatric
martens in Poland, where fruit was the major food
resource for stone martens throughout the year
(Posłuszny et al., 2007). In the western Italian Alps, the
diversity of fruit species eaten by stone martens was
almost twice that of the pine marten; consistent with
available data (Gazzola & Balestrieri, 2020), this differ-
ence mostly depended on the largest use of cultivated
fruits by the more synanthropic stone marten.
In the Lepontine Alps, where pine marten occurrence
was negligible (see also Balestrieri et al., 2021), the pro-
portional contribution of protein energy to stone marten's
diet did not differ from pine martens'. The only other
study which, to our knowledge, investigated the diets of
both martens in an area where the stone marten was
dominant in terms of relative abundance was carried out
in northern Portugal, at the southwestern limit of pine
marten distribution (Monterroso et al., 2016). In the
breeding period (April–May), fruits formed the bulk of
pine marten diet (51.6% of ingested biomass), while
accounted for only 10% of all the biomass ingested by
stone martens. While sample size did not allow us to pre-
cisely assess the composition of pine marten diet in the
Lepontine Alps, both the results of Monterroso
et al.'s (2016) study and ours suggest that competition
plays a major role in enhancing frugivory in the stone
marten, unless it largely outnumbers its competitor.
Carnivores are highly adapted to high-protein diets
compared with herbivores and several omnivores (Kohl
et al., 2015), their protein metabolizable energy intake
target being around one-half of the total intake (Hewson-
Hughes et al., 2011; Wilder et al., 2016). Carnivores with
protein-rich diets tend to have higher body conditions
and aerobic capacity, lower erythrocyte turnover, and
increasing immune function (Santos et al., 2020;
Whiteman et al., 2019). In contrast, many predators can-
not down-regulate rates of protein catabolism, hindering
8 GRANATA ET AL.
their capabilities of using low-protein food resources
(Eisert, 2011; Walton, 1986). The balancing of protein to
nonprotein may thus be expected to be the main driver of
foraging in mustelids (Jensen et al., 2014; Remonti
et al., 2011). Conversely, carnivores' diet usually includes
little carbohydrate energy, as predators are not able to
effectively regulate glucose homeostasis (Myers &
Klasing, 1999; Schermerhorn, 2013). Excess of carbohy-
drates hinders the digestion of protein and absorption of
minerals (Schunemann et al., 1989), and strictly carnivo-
rous species, such as cats and salmonids, have a thresh-
old level for carbohydrate intake beyond which food
intake is reduced (Ruohonen et al., 2007).
Unbalanced diets have been reported to have sharp
effects on life span and reproduction (crickets: Maklakov
et al., 2008; mice: Solon-Biet et al., 2014). Although mus-
telids may be more tolerant toward carbohydrates than
obligate carnivores (Kohl et al., 2015; Remonti
et al., 2016), the consistency of martens' macronutrient
ratios in allopatry suggests that frugivory, as a makeshift
strategy to cope with interspecific competition, may
lower the fitness of the less competitive species.
Fluctuations in prey numbers are considered to medi-
ate the competitive interactions between strictly related
mustelids in nonequilibrium communities, with coexis-
tence being enhanced by large prey populations
(Powell & Zielinski, 1983). The nutritional ecology of
martens suggests that prey availability may regulate the
relative abundance of sympatric populations of pine and
stone martens in a more subtle way, by driving the less
competitive species to eat nutritionally imbalanced food
(i.e., that do not optimally satisfy the animals' nutrient
requirements; Raubenheimer & Simpson, 2003). To sub-
stantiate this hypothesis, future studies should aim to
investigate the relative abundance of sympatric martens
in relation to fluctuations in small mammal abundance
and fruit consumption.
A C K N O WL E D G M E N T S
Research in northern Piedmont was supported by the Val
Grande National Park (2014–2017), as part of the project
“Monitoraggio della biodiversità animale in ambiente
alpino” (Monitoring of animal biodiversity on the Alps).
We are grateful to M. Gilardi, L. Ricci, and P. Tirozzi for
their help with field work. Research on the western Alps
was part of the degree thesis of M. Granata, who was
tutored by S. Bertolino. J. Lovejoy kindly revised the
English language.
CONFLICT OF INTEREST
The authors declare that there is no conflict of interest
that could be perceived as prejudicing the impartiality of
the research reported.
ORCID
Marco Granata https://orcid.org/0000-0002-9445-7137
Alessandro Balestrieri https://orcid.org/0000-0001-
5444-2806
RE FER EN CES
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How to cite this article: Granata, M., Mosini, A.,
Piana, M., Zambuto, F., Capelli, E., & Balestrieri,
A. (2021). Nutritional ecology of martens (Martes
foina and Martes martes) in the western Italian
Alps. Ecological Research, 1–10. https://doi.org/10.
1111/1440-1703.12277