Trophic and spatial relationships of the pine marten (Martes martes) and American mink (Neovison vison) on mountain rivers of the Southern Urals

The American mink (Neovison vison) and pine marten (Martes martes) are the most common mustelid species in mountain forests of the Southern Urals. The bulk of the diet (85.3%) in the American mink consists of small mammals (35.8%), insects (22.7%), gastropods (14.1%), and fish (12.7%). In the pine marten, the main components comprising 79.6% of the diet are vegetable foods (29.8%), small mammals (27.0%), and insects (22.8%). The low diversity of their food spectra is accounted for by landscape and ecological features of the study region. The abundance of rodents is low, their distribution has a mosaic pattern, and other kinds of food are more abundant near rivers and streams, which explains the preference of both mustelid species for riparian biotopes. The breadth of trophic niches in the pine marten and American mink are 1.67 and 1.62, respectively, and the coefficient of their overlap is 0.75.     

Biogeographical patterns in the diet of an opportunistic predator: the red fox Vulpes vulpes in the Iberian Peninsula

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Effect of felling on red fox (Vulpes vulpes) and pine marten (Martes martes) diets in transitional mixed forest in Belarus

In transitional mixed forests in northern and central Belarus the influence of intensified felling on the diets of red foxes Vulpes vulpes L. and pine martens Martes martes L. was investigated in two model forested terrains with sandy and clay top-grounds. A total of 1904 scats of red foxes and 1624 scats of pine martens were analysed over two periods differed by logging rate. When logging rate was conservative, red fox and pine marten diets were found to be similar, but under heavy logging feeding of the predator species shifted. In both model woodlands we found the same pronounced dietary trend of higher consumption of rodents, first of all, Microtus voles. The dietary changes were well related to the registered increase in Microtus vole numbers and total number of rodents in felling areas. The increased preying on rodents caused lower consumption of other food items, particularly medium-sized mammals (year-round) or/and birds or/and fruits (in the warm season) or/and mammalian carrion (in the cold season). In the conditions of intensified felling the food niches of the red fox and pine marten diverged mostly because of the great difference in the species structure of rodents consumed. Red foxes turned to preying on Microtus voles more frequently, but less on bank voles Myodes glareolus; while pine martens increased their taking of Microtus voles, continued foraging for bank voles and began taking slightly more of Apodemus mice. Before heavy logging dietary similarity between the red fox and pine marten was high and did not vary considerably through seasons and study areas, whereas after felling was intensified their diet overlap became lower.     

Spring and summer food habits and habitat use of the Europena pine marten (Martes martes) on the island of Micorca, Spain

The food habits and habitat use of pine marten ( Martes martes L.) on the Balearic Island of Mincorca were studied from March-August 1990. In a 28.2 km² area, a series of hiking trails/forest roads were used to collect pine marten faeces bi-monthly and study te species' differential use of habitats. A total of 28 different food items were identified in 723 faeces. Small mammals were the most important food overall, constituting 34% of the volume During March-April, small mammals were the principal food consumed (63% of volume), followed by birds (19%). In May-June, birds were the main food (40%), then small mammals. Plant material and insects were the most important foods in July-August, both made up 68% of the pine marten diet. The abundance of certain foods and the degree of difficulty in obtaining them appear to be important factors that govern pine marten food selection in Minorca. Habitat use was studied along 12 routes totalling 32.6 km. Pine marten showed preference in March-April for Coastal Shrubland habitat, while during May-June they were found most in the Upland Pine type. During July-August, the Upland Pine and Open Pine habitats were most frequented by pine marten. According to bimonthly food habits data, habitat use seems largely determined by the abundance and availability of their prey items in the available habitat types. Martens may concentrate hunting efforts in areas different from when they are not hunting. The Minorcan pine marten differs from continental populations in its wide use of shrublands, in addition to forests.     

Diet of stone martens: an example of ecological flexibility

The feeding ecology of the stone marten in a rural area of Central Italy was assessed by faecal analysis. Habitat analysis, performed through a Geographical Information System, and intensive radio-tracking allowed us to investigate intra-population variations of diet in relation to habitat. The species'feeding habits were opportunistic: fruit and berries were the staple diet but mammals and birds were also important. Diet varied seasonally in relation to resource availability, with a predominance of fruit in autumn and frequent presence of insects in summer. Individuals living in adjacent areas but in different habitats (wooded and rural) showed significant variations of diet, adapting their feeding habits to local availability of foods. The great adaptability of the stone marten to very different resource conditions can represent a key to the species'success through its wide range.     

Role of fruits in the diet of small mustelids (<Emphasis Type="Italic">Mustela</Emphasis> sp.) from the western Italian Alps

The food habits of both weasels and stoats (Mustela nivalis and M. erminea) and the occurrence of fruit in their diet were studied by scat analysis in the Gran Paradiso National Park (western Italian Alps). Being impossible to distinguish between stoats and weasels scats, they were grouped together as the genus Mustela. Despite the high consumption of small rodents, which represented the main food item in the diet of small mustelids (41.5% in mean volume), wild fruits (mainly Rosaceae and Ericaceae) were also relatively common (23.3% in mean volume). Fruits were exploited throughout the year, with an intensive use in seasons of high availability. Their consumption did not significantly differ between habitat types, despite a significant decrease of fruit availability from deciduous mixed forest to alpine prairie. This evidence suggests that this trophic behaviour could represent a consolidated feeding strategy of both weasels and stoats, which seem to adopt an opportunistic behaviour as shown by larger size mustelids. We argue that fruits, owing to their good availability and high profitability, could play an important role in the diet of small mustelids in alpine habitats, slackening their well-documented dependence on rodents.     

Comparative analyses on food habits of Japanese marten,Siberian weasel and leopard cat in the Tsushima islands,Japan

Food habits of three sympatric carnivore mammals in the Tsushima islands of Japan were studied during 1986–91. Scats of the Tsushima marten (n=1236), the Siberian weasel (n=218) and the Tsushima leopard cat (n=350) were collected monthly and the food items were determined by scat contents analysis. Marten was omnivorous showing a high level of diversity of food throughout the year. The important foods for marten were fruits and berries from spring to autumn, insects in summer and autumn and small mammals all year round. Leopard cat preferred to hunt wood/mice and birds, and remained a flesh meat specialist throughout the year. Weasel was intermediate between marten and leopard cat, but was slightly biased towards the flesh meat eater. The three carnivores do not compete against one another for food, except for small rodents. A conflict for food between leopard cat and weasel was suggested to be more intense than that of other combinations based on diet overlapping. Marten may be characterized as an opportunistic generalist. When interspecific competitors existed, or human disturbance to the habitat occurred, the preferential flexibility of the marten to alternative food resources might become more advantageous than the other two species.     

A non-invasive genetic method to identify the sympatric mustelids pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) and stone marten (<Emphasis Type="Italic">Martes foina</Emphasis>): preliminary distribution survey on the northern Iberian Peninsula

The closely related mustelids European pine marten (Martes martes) and stone marten (Martes foina) sympatrically inhabit a large area of Europe. However, given our limited knowledge of their bioecological relationships, their extremely elusive behaviour and the fact that their faeces cannot be distinguished on the basis of morphology alone, it is very difficult to monitor their populations. In this study, we describe a reliable non-invasive polymerase chain reaction (PCR)–restriction fragment length polymorphism (PCR-RFLP) method for distinguishing between M. martes and M. foina based on the analysis of deoxyribonucleic acid extracted from faeces samples. The method was specifically designed to avoid possible interference from potential prey mammals and other sympatric carnivores. The procedure consists of PCR amplifying a mitochondrial D-loop region followed by digesting the resulting 276-bp-long amplicons with the restriction enzymes HaeIII and RsaI. To assess the efficiency of this technique, we conducted a preliminary field study across the potential sympatric distribution areas of both marten species in the northern Iberian Peninsula. Out of 359 faeces samples collected, we identified 80 as specimens from the stone marten and 235 from the pine marten. Unequivocal species identification was thus possible in 88% of the faeces samples collected. These findings reveal the combined use of non-invasive genetic sampling and GIS technology to be a reliable and cost-effective procedure for improving our knowledge of the spatial distributions of sympatric marten species. This protocol could also be used to identify and improve information gaps, to develop effective research and management programmes and in population and landscape genetics studies of marten species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Antipredatory Response and Food Intake in Wood Mice (Apodemus sylvaticus) under Simulated Predation Risk by Resident and Novel Carnivorous Predators

Chemical signals left by predators are a potential source of information about the risk of predation, and small mammals are known to take them into account when making decisions. We investigated whether wood mice (Apodemus sylvaticus) are more likely to avoid the faeces of resident predators (red fox Vulpes vulpes and common genet Genetta genetta) vs. a novel predator (European pine marten Martes martes). Odour recognition would increase perceived predation risk and reduce food intake by individual mice. Wood mice response to predators was analysed by live‐trapping using two untreated controls (baited/non‐baited) and traps experimentally manipulated with three predator treatments (faeces of red fox, common genet or pine marten). Traps were baited with 4 g of toasted corn, and food intake by wood mice was determined as the amount of bait remaining in each trap. We found that traps treated with faeces of resident predators were the most avoided, and the number of captures in traps treated with pine marten faeces was similar to the control‐baited traps. The variation found in food intake was explained by the interaction between the types of treatment and breeding condition. Food intake was similar in control‐baited traps and in traps with faeces of pine marten, but when predation risk by resident predators (red fox and common genet) was simulated, breeders reduced food intake significantly as compared to non‐breeders. These results indicate that predator recognition and feeding behaviour under predation risk depend on individual factors and the balance of costs‐benefits in each particular predation risk situation at a given place and time.     

Spatial organisation and dynamics of the pine marten Martes martes population in Białowieza Forest (E Poland) compared with other European woodlands

We studied factors affecting density and spacing patterns in the pine marten Martes martes population inhabiting temperate forests of Bia?owieza National Park, eastern Poland. From 1985/1986 to 1995/1996 marten densities ranged from 3.63 to 7.57 individuals 10 km^?2 (mean 5.4) and were positively correlated with abundance of forest rodents in the previous year. The rate of marten population growth was inversely density‐dependent and positively related to rodent density. Annual mortality rate averaged 0.384 and tended to be negatively related to marten densities. Mean annual home range of males (2.58 km², SE=0.24) was larger than that of females (1.41 km², SE=0.20). Seasonal home ranges also differed significantly between males and females. Both sexes held the smallest ranges in December–January. Female ranges increased in April–May, whereas those of males increased in June–September when they were mating. Fidelity of pine martens to their home ranges was very high. The mean shift between arithmetic centres of seasonal ranges was 0.25 km, and the ranges recorded in two consecutive seasons overlapped, on average, by 87–90%. We observed very little home range overlap between neighbouring male (mean 4–6%) or female (mean 6%) marten. Year round the neighbouring individuals of the same sex neither avoided nor attracted each other. Females attracted males only during the spring‐summer mating season. A review of other studies has documented that winter severity and seasonal variation in ecosystem productivity were essential factors shaping the biogeographic variation in pine marten densities between 41^o and 68^oN. The density of marten populations increased in areas with mild winters and lower seasonality. Maximum population densities (indicative of habitat carrying capacity) were correlated with mean winter temperature. In Europe, male home ranges increased with decreasing forest cover in a study area, whereas female ranges varied positively with rodent abundance.     

Efficient species identification of pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>) and red fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) scats using a 5′ nuclease real-time PCR assay

Monitoring wildlife species by DNA identification of samples collected non-invasively is an important tool in conservation management. DNA identification of species from faecal (scat) samples is problematic due to the small quantities and poor quality of the DNA isolated from such samples. This study demonstrates the use of real-time PCR technology in the identification of red fox (Vulpes vulpes) and pine marten (Martes martes). It is shown that real-time PCR can be used to identify fox and pine marten by either melting curve analysis (Tm determination) with SYBR Green 1 detection or by the use of species specific fluorogenic probes. The technique is shown to work efficiently with scat DNA.     

Does size dimorphism reduce competition between sexes? The diet of male and female pine martens at local and wider geographical scales

Sex-specific niche segregation is often used to explain sexual size dimorphism (SSD). However, whether food niche partitioning between sexes occurs as a case of sexual size dimorphism or by other mechanisms, such as behavioural dimorphism or habitat segregation, remains poorly understood. To evaluate the nature and extent of food-niche differentiation between sexes in a solitary predator I examined variation in the diet of male and female pine martensMartes martes Linnaeus, 1758 in years of high and low rodent abundance. Small mammals were the most important prey for pine martens in years of both low and high rodent abundance (occurring in more than 49% of scats). Birds, invertebrates and plant material were relatively common food items in summer diet, whereas ungulate carcasses were often consumed in autumn—winter. In general, males consumed more ungulate carcasses, plant material, amphibians and reptiles than did females, whereas females preyed more on squirrels and birds than males. There was significant seasonally dependent, between-sex variation in the occurrence of shrews, small rodents, other mammals, birds and invertebrates in marten diet. Whereas the occurrence of bank vole, birds, carcasses and plant material changed between sexes, seasons and years with various rodent abundances, both sexes consumed larger prey and had increased food niche breadth in years of low compared with high rodent abundance. Neither prey size nor food niche breadth were significantly different between males and females. The food-niche overlap between sexes was consistently lower in spring and in years of low rodent abundance. A wider geographical comparison of different marten populations showed that the diet of males and females varied significantly between locations. Females consistently preyed on squirrels and birds, whereas males fed more often on ungulate carcasses and plant material. Local and geographical comparison of male and female diets suggest that food-niche partitioning between male and female pine martens changes across different habitat and food conditions, and is not related to sexual size dimorphism, but rather to behavioural differences between sexes.     

A PCR-RFLP-based method to distinguish sable (<Emphasis Type="Italic">Martes zibellina</Emphasis>) and pine marten (<Emphasis Type="Italic">Martes martes</Emphasis>)

Species identification is an important issue in conservation and a particular focus for wildlife forensics. Molecular biological methods retain a unique power to differentiate between difficult samples that lack other identifiable characteristics. The pine marten (Martes martes) and sable (Martes zibellina) are closely related species with very similar pelage characteristics and are often difficult to distinguish from each other. The sable, however, in contrast to the pine marten, remains an endangered and protected animal in China with both hunting and fur trade strictly prohibited for this species. Here, we present a polymerase chain reaction-based restriction fragment length polymorphism method for distinguishing the two species. We sequenced a 638-bp fragment of cytochrome b gene in 39 sables, 68 pine martens, and 10 stone martens and identified all variable nucleotides. A new primer pair was subsequently designed to amplify a 316-bp fragment containing restriction sites of enzyme BseG I and BamH I that are different among martens. When the fragment was cut using BseG I, the resulting restriction pattern was identical in the sable and pine marten, but differed from all other martens. When cut using BamH I, the fragment generated two diagnostic fragments in the sable which could distinguish them from pine martens. This method was valid for all haplotypes of sable and pine marten thus far identified and has high potentially applicability for the identification of the two species.     

Nutritional ecology provides insights into competitive interactions between closely related Martes species

1. Closely related predator species often share several prey items, making it hard to differentiate the effects on their feeding habits of variation in food availability and of competition. We hypothesised that we could overcome this obstacle by quantifying and comparing nutritional niches.
2. We reviewed dietary studies that assessed the relative bulk of each food item, as either per cent biomass or per cent mean volume, in the diet of two closely related species, pine marten Martes martes and stone marten Martes foina, and calculated the nutrient profiles (intakes of protein, lipids and carbohydrates) of each diet.
3. Both martens’ diets were tightly clustered (mean values: 47% of energy from protein, 39% from lipid, and 14% from carbohydrate). In allopatry, the nutritional niches of the two species did not differ, but in sympatry, the stone marten ate more carbohydrates and less protein than the pine marten. In allopatry, the protein intake of the stone marten remained high (45–52%) in very different habitats, from cultivated lowland to Alpine forests.
4. Our data suggest that stone marten frugivory may, at least partially, be the result of interspecific competition. By analysing dietary data in the framework of nutritional ecology, we could compare the feeding requirements of pine martens and stone martens more effectively than by using classical estimates of trophic niche overlap at the food item level. This approach may help to shed light on the trophic relationships of other competing species.

     

Intraguild dietary overlap and its possible relationship to the coexistence of mesocarnivores in intensive agricultural habitats

The stone marten (Martes foina) and the pine marten (M. martes) are closely related mammalian carnivores potentially subject to exploitative competition. The recent expansion of the pine marten into the intensively cultivated plain of the River Po (NW Italy), where previously only stone marten occurred, offered an interesting opportunity to analyse their relationships. We studied the distribution and diet of Martes species and trophic niche overlap between martens and red foxes (Vulpes vulpes) in two study areas, each with two pseudoreplicates, by analysing genotyped faeces. Our results seem to confirm the displacement of the stone marten from one study area, the pine marten being the only Martes species occurring where previously the stone marten had been reported. We found a large food niche overlap between red fox and both stone and pine martens, but with evidence of size-related differences in the consumption of some food items. We hypothesised that, due to the poor prey-base of the environment, highly altered by intensive crop cultivation, intense interspecific competition originally occurred between the red fox and stone marten. The heightening of interspecific competition caused by the entry of the pine marten in the predator guild may have caused the displacement of the stone marten, at least temporarily. The mechanism of such displacement needs to be clarified through further surveys in areas where the three species occur sympatrically.     

Annual and seasonal changes in diets of martens: evidence from stable isotope analysis

Theory predicts that generalist predators will switch to alternative prey when preferred foods are not readily available. Studies on the feeding ecology of the American marten (Martes americana) throughout North America suggest that this mustelid is a generalist predator feeding largely on voles (Microtus sp.; Clethrionomys sp.). We investigated seasonal and annual changes in diets of martens in response to the changing abundance of small rodents (Peromyscus keeni, and Microtus longicaudus) on Chichagof Island, Southeast Alaska, using stable isotope analysis. We hypothesized that martens would feed primarily on small rodents during years with high abundance of these prey species, whereas during years of low abundance of prey, martens would switch to feed primarily on the seasonally available carcasses of salmon. We also hypothesized that home-range location on the landscape (i.e., access to salmon streams) would determine the type of food consumed by martens, and martens feeding on preferred prey would exhibit better body condition than those feeding on other foods. We live-captured 75 martens repeatedly, from mid-February to mid-December 1992–1994. We also obtained marten carcasses from trappers during late autumn 1991 and 1992, from which we randomly sub-sampled 165 individuals. Using stable isotope ratios and a multiple-source mixing model, we inferred that salmon carcasses composed a large portion of the diet of martens in autumn during years of low abundance of rodents (1991 and 1992). When small rodents were available in high numbers (1993 and 1994), they composed the bulk of the diet of martens in autumn, despite salmon carcasses being equally available in all years. Selection for small rodents occurred only in seasons in which abundance of small rodents was low. Logistic regression revealed that individuals with access to salmon streams were more likely to incorporate salmon carcasses in their diet during years of low abundance of small rodents. Using stable isotope analysis on repeated samples from the same individuals, we explored some of the factors underlying feeding habits of individuals under variable ecological conditions. We were unable to demonstrate that body weights of live-captured male and female martens differed significantly between individuals feeding on marine-derived or terrestrial diets. Therefore, martens, as true generalist predators, switched to alternative prey when their principal food was not readily available on a seasonal or annual basis. Although salmon carcasses were not a preferred food for martens, they provided a suitable alternative to maintain body condition during years when small rodents were not readily available. Received: 1 May 1996 / Accepted: 24 February 1997     

Genetic analysis of sable (<Emphasis Type="Italic">Martes zibellina</Emphasis>) and pine marten (<Emphasis Type="Italic">M. martes</Emphasis>) populations in sympatric part of distribution area in the northern Urals

Analysis of nucleotide sequences of the mitochondrial DNA (mtDNA) control region (495 bp) of sables (Martes zibellina) and pine martens (M. martes) from allopatric parts of the species ranges has shown a considerable interspecific genetic distance (>3%). In sympatric populations of these species in the northern Urals, differences between two species-specific mtDNA lineages are still large; however, classification of each individual nucleotide sequence with one of the two lineages is not correlated with whether the given animal is phenotypically a sable, a pine marten, or a potential hybrid (the so-called “kidas”). This indicates a high degree of reciprocal introgression of the sable and pine marten mtDNA in the northern Urals and suggests that their interspecific hybridization is common in the sympatric zone.     

A non-invasive approach to determining pine marten abundance and predation

A non-invasive approach was used to investigate variation in pine marten (Martes martes) abundance between the midlands and east of Ireland, and to determine the frequency of occurrence of squirrels and other small mammals in the diet. Remotely plucked hair samples were genotyped to differentiate between individual animals, and real-time polymerase chain reaction (PCR) was used to identify predator and prey DNA in scats. Macro analysis of prey remains was carried out on a sub sample of scats and the results from both methods are compared. Non-invasive techniques were successful in determining the presence and relative abundance of the pine marten at woodland level. As expected, abundance was found to be higher in the core population of the midlands than in the east. Pine martens were found to reach higher numbers per km² of forested habitat in Ireland than their British or European counterparts. Both traditional hard part analysis and molecular dietary analysis of mammalian prey yielded similar results. We provide the first evidence of the European pine marten predating upon the North American grey squirrel (Sciurus carolinensis) in its invasive range. While the grey squirrel was not available as a prey item in any of the midlands sites, it was available in the east, where it featured significantly more frequently in the diet than the native red squirrel. In both the midlands and the east the woodmouse is the most frequently occurring mammal in the diet.     

Range expansion of the pine marten (Martes martes) in an agricultural landscape matrix (NW Italy)

Habitat segregation is considered to favour the coexistence of sympatric pine martens Martes martes and stone martens M. foina, the latter being displaced to agricultural and urbanised areas. Subsequent to the report of pine martens in cultivated areas of the western River Po plain (NW Italy), we reviewed all available information on the presence of this species in plain areas of northern Italy and, for two study areas, applied a non-invasive PCR-RFLP method for the identification of Martes species from faecal mtDNA.A total of 24 pine marten records were collected, grouped in the western part of the River Po plain. The number of records showed an exponential increase from 1988 to 2007, the percentage of woods in a 10 km² wide circular plot surrounding the location of records being inversely correlated to its distance from the 300 m a.s.l. contour line. In the two study areas, 36 out of 119 “marten-like” faeces were assigned to the pine marten, whilst none belonged to the stone marten. In the best monitored area, the pine marten was present almost constantly.Our results suggest that the pine marten is expanding its range to include cultivated areas which were previously considered a prerogative of the more synanthropic stone marten.     

Fruit consumption by carnivores in Mediterranean Europe

• 1 We reviewed patterns of fruit consumption amongst 10 species of mesocarnivores: red fox Vulpes vulpes, weasel Mustela nivalis, stoat Mustela erminea, polecat Mustela putorius, stone marten Martes foina, pine marten Martes martes, Eurasian badger Meles meles, common genet Genetta genetta, Egyptian mongoose Herpestes ichneumon and wildcat Felis silvestris in Mediterranean Europe.
• 2 The 65 reviewed studies recorded 79 different fruits eaten by carnivores, 58 of which were identified to species. Most records (63%) were of fleshy fruits with high pulp content. The frequency of occurrence of fruit items varied widely amongst species and regions. Four of the carnivore species (red fox, stone marten, badger and common genet) included more than 30 fruit species in their diet.
• 3 A longitudinal pattern was detected in the consumption of fruit in the Mediterranean region, with the frequency of occurrence of fruit consumption increasing towards the east.