Habitat selection of European pine marten in Central Italy: from a tree dependent to a generalist species

Studies at small spatial scale are often fundamental to highlight the behavioural plasticity of a species and thus have important implications for conservation planning, in particular for species usually considered as habitat specialists. We investigated second-order habitat selection of the European pine marten in an area dominated by deciduous oak forest and open fields in central Italy, by radio-tracking 16 pine martens (eight males, eight females). Pine martens placed home ranges in areas with more open field than in the study area, whereas woodland (oak and conifer forests) comprised a smaller portion of the home range than predominant forest character of the studied area. Although the presence of the species in the open habitats has been documented, to our knowledge, our results provide the first evidence of home range establishment in this cover type by pine marten at population level. The combination of low predation risk and high availability of resources could allow pine martens to occupy open fields in our study area. We highlighted different individual strategies of habitat selection, with some individuals placing home ranges in areas with high forest coverage while others occupying open areas. We found no effects of sex and body condition on habitat selection, and this could indicate that in the study area, both forested and non-forested cover types, such as open fields, shrub and anthropic areas, can provide adequate food, overhead cover and resting sites for all individuals. Pine marten ability to occupy open fields seems thus more related to the behavioural flexibility of the species, rather than to the need to supplement dens and forage from complementary lower quality habitat. The high quality of the Mediterranean continental area studied could also explain the selection of open areas by the pine marten. Our results offer useful information on pine marten ecology and may be helpful for conservation management of this species in southern Europe.     

Spatial Niche Segregation of Sympatric Stone Marten and Pine Marten – Avoidance of Competition or Selection of Optimal Habitat?

Coexistence of ecologically similar species relies on differences in one or more dimensions of their ecological niches, such as space, time and resources in diel and/or seasonal scales. However, niche differentiation may result from other mechanisms such as avoidance of high predation pressure, different adaptations or requirements of ecologically similar species. Stone marten (Martes foina) and pine marten (Martes martes) occur sympatrically over a large area in Central Europe and utilize similar habitats and food, therefore it is expected that their coexistence requires differentiation in at least one of their niche dimensions or the mechanisms through which these dimensions are used. To test this hypothesis, we used differences in the species activity patterns and habitat selection, estimated with a resource selection function (RSF), to predict the relative probability of occurrence of the two species within a large forest complex in the northern geographic range of the stone marten. Stone martens were significantly heavier, have a longer body and a better body condition than pine martens. We found weak evidence for temporal niche segregation between the species. Stone and pine martens were both primarily nocturnal, but pine martens were active more frequently during the day and significantly reduced the duration of activity during autumn-winter. Stone and pine martens utilized different habitats and almost completely separated their habitat niches. Stone marten strongly preferred developed areas and avoided meadows and coniferous or deciduous forests. Pine marten preferred deciduous forest and small patches covered by trees, and avoided developed areas and meadows. We conclude that complete habitat segregation of the two marten species facilitates sympatric coexistence in this area. However, spatial niche segregation between these species was more likely due to differences in adaptation to cold climate, avoidance of high predator pressure and/or food preferences by both species than competitive interaction between them.     

The influence of forest fragmentation and landscape pattern on American martens

1. We investigated the effects of forest fragmentation on American martens ( Martes americana Rhoads) by evaluating differences in marten capture rates (excluding recaptures) in 18 study sites with different levels of fragmentation resulting from timber harvest clearcuts and natural openings. We focused on low levels of fragmentation, where forest connectivity was maintained and non-forest cover ranged from 2% to 42%.
2. Martens appeared to respond negatively to low levels of habitat fragmentation, based on the significant decrease in capture rates within the series of increasingly fragmented landscapes. Martens were nearly absent from landscapes having > 25% non-forest cover, even though forest connectivity was still present.
3. Marten capture rates were negatively correlated with increasing proximity of open areas and increasing extent of high-contrast edges. Forested landscapes appeared unsuitable for martens when the average nearest-neighbour distance between open (non-forested) patches was <100 m. In these landscapes, the proximity of open areas created strips of forest edge and eliminated nearly all forest interior.
4. Small mammal densities were significantly higher in clearcuts than in forests, but marten captures were not correlated with prey abundance or biomass associated with clearcuts.
5. Conservation efforts for the marten must consider not only the structural aspects of mature forests, but the landscape pattern in which the forest occurs. We recommend that the combination of timber harvests and natural openings comprise <25% of landscapes ≥9 km² in size.
6. The spatial pattern of open areas is important as well, because small, dispersed openings result in less forest interior habitat than one large opening at the same percentage of fragmentation. Progressive cutting from a single patch would retain the largest amount of interior forest habitat.     

The European Pine Marten Martes martes in the Balearic Islands, Spain

Between March and December 1990, a survey of Pine Marten Martes martes L. on the Balearic Islands of Minorca and Majorca was carried out. Pine Marten were found distributed over 55% of Minorca and occupied Aleppo Pine and Holm Oak forests, mediterranean shrublands and riparian/ cliff habitats. In Majorca, Pine Marten were found on 48% of the island and three broad physiographic areas, the Sierra de Tramuntana, Sierra de Levant and the Massif de Randa. Droppings were collected along four designated routes during five bimonthly intervals in Minorca to describe the Pine Marten's differential use of habitat types. The upland/open pine forest had the highest use index while the Holm Oak forest had the lowest. The number of Pine Marten faeces collected was greatest during the months of July-August. The Balearic Island Pine Marten populations were once on the verge of extinction; however, protection during the last 20 years has enabled the species to recover throughout most of the forested habitats on both islands. Resource managers must carefully monitor the Pine Marten populations to provide the species and its habitat with adequate protection and management in the future.     

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.     

Estimation of population density of European pine marten in central Italy using camera trapping

Evaluating presence and abundance of small carnivores is essential for their conservation. In Italy, there is scarce information on European pine marten distribution, and no data are published on its abundance. Camera traps have been widely used to estimate population density applying capture–recapture models for species in which individual recognition is possible. Here we estimate the abundance of European pine martens in central Italy using camera trapping and a model that allows the estimation of population density without the need for individual recognition Rowcliffe et al. (Anim Conserv 11:185–186, 2008). Camera trapping was also used to evaluate habitat use patterns by martens. Fifteen camera traps were deployed in 90 placements for 15 days each, for a total of 1,334 camera days. Pine martens were captured in 24% of camera trap placements with a mean trap success rate of 0.33 photographs per camera placement. Estimated pine marten population density in the study area was 0.34 individuals km⁻². Marten trap rate was not strongly associated with any habitat type, although there were trends towards lower probability of records at locations with high coverage of cultivated fields and higher probability of records at locations with high coverage of human-made woodland. The results suggest that pine martens in this area are not confined to wooded habitat. To our knowledge, this study is the first application of the Rowcliffe et al. (Anim Conserv 11:185–186, 2008) method to a wild carnivore population and, furthermore, the first estimation of population density of pine martens in Italy.     

The NATURA 2000 database as a tool in the analysis of habitat selection at large scales: factors affecting the occurrence of pine and stone martens in Southern Europe

Most studies on habitat selection among animals are conducted at local scales, whereas reliable determination of species requirements at larger spatial scales can be problematic. We used data available for NATURA 2000 sites to determine the habitat requirements of two relatively widespread and common species—pine marten Martes martes and stone marten Martes foina—in Southern Europe. Using presence-absence data, we applied statistical models at two spatial scales. At the local scale (within the dispersal distances of the species), sites occupied by martens were compared with unoccupied sites using buffers of different sizes, whereas at the regional scale, unoccupied sites were selected randomly. To adjust for spatial autocorrelation of data, penalised quasi-likelihood approximations were used. Both species of martens demonstrated preferences for areas with higher proportions of forest cover and steeper terrain. At the local, but not at the regional, scale, pine martens occurred at lower elevations, whilst stone martens occurred at higher elevations. We found that climatic variables (mean temperature, precipitation) had no significant effect on the presence of the martens. The results of our analyses generally confirmed findings of previous studies on habitat selection of both marten species in Southern Europe. This demonstrates the utility of data collected for NATURA 2000 sites for use in various analyses such as conservation planning and evaluating the impact of climate change on the distribution of animal species.     

Decline in American marten occupancy rates at Sagehen Experimental Forest,California

We compared the distribution and frequency of American marten (Martes americana) detections during historic surveys and a recent survey on the Sagehen Experimental Forest (SEF) in the Sierra Nevada Mountains, California. This area has been the location of 9 previous marten surveys during 1980–1993, each involving a systematic detection/non-detection survey on the same grid. These data are a time series of information on the occupancy of martens that can be related to habitat change in the study area. Our objectives were to 1) resurvey martens in SEF using methodology similar to previous studies to assess current marten occupancy; 2) evaluate changes in marten occupancy during the period 1980–2008; and 3) examine associations between marten occurence and changes in habitat and landscape metrics. Current marten occupancy was estimated using surveys conducted in summer 2007, winter 2007–2008, and summer 2008. From 1978 to 2007 there was a decrease in predicted habitat patch size, core area, and total amount of marten habitat in the study area, as well as an increase in distance between patches. Marten detections in 2007–2008 were approximately 60% lower than in surveys in the 1980s. We detected no martens in the summers of 2007 and 2008, and 10 detections in winter 2007–2008 were limited to higher elevations in the southwestern portion of SEF. No martens were detected in the lower elevations where most of the recent forest management activity occurred. We suggest that the marten population at SEF has been negatively affected by the loss and fragmentation of habitat. We recommend that future management of forests in the Sagehen basin focus on restoring and connecting residual marten habitat to improve habitat quality for martens. © 2011 The Wildlife Society.     

Far from the madding crowd: Tolerance toward human disturbance shapes distribution and connectivity patterns of closely related Martes spp.

To assess niche overlap between the most similar European sympatric carnivores, the pine marten Martes martes and stone marten Martes foina, and outline their potential distributions and connectivity corridors in Central Italy, we applied a multivariate kernel density procedure which allowed to assess both species' ecological hypervolumes based on a set of 16 environmental predictors and used the resulting probability of occurrence map as a resistance surface in electrical circuit theory-based models. Distance to watercourses and percent cover of deciduous forest and shrubland were the most relevant factors shaping pine marten ecological niche, while stone marten distribution was mainly shaped by human population density and cover of both human settlements and deciduous forest. Overlap between the hypervolumes of the two martens was low-to-moderate, while, on average, landscape connectivity was higher for the stone marten. The inclusion in the models of human disturbance-related variables enabled to define a possible mechanism driving habitat partitioning in human-altered landscapes. Based on our results, increasing human density and urbanization of European lowland and hilly landscapes are expected to represent a greater threat to the pine marten than the stone marten.     

Snow tracking reveals different foraging patterns of red foxes and pine martens

Red fox (Vulpes vulpes) shares similar prey preferences and co-occurs with several other carnivores, and is together with pine marten (Martes martes), the most common mesocarnivore in the northern boreal forest. Voles are important prey for both species, but it is unclear to what extent they compete for the same food resources in winter. Here, we use 2139 km and 533 km of meticulous snow tracking of red foxes and pine martens to evaluate their food niches. We measured hunting and digging behaviour, whether successful or not, and the effect of snow depth and temperature. Pine martens were restricted to forested habitats, whereas red foxes used a wide range of habitats. Red foxes were found to dig more often than pine martens, 0.67 vs. 0.39 digging events per kilometre. Hunting was less common and similar in both species, about 0.1 hunting event per kilometre. Pine martens were more efficient in hunting and finding food remains compared to red foxes. Increasing snow depth reduced hunting success and also reduced dig success of red foxes. Food niche overlap was small. Red foxes used mostly voles and carrion remains of ungulates, whereas pine martens used cached eggs and small birds. We suggest that caching eggs is an important strategy for pine martens to survive winter in northern latitudes. Snow depth was important for capturing voles, and thick snow cover appeared to mask the effect of vole peaks. Intensified land use, as clear-cutting and leaving slaughter remains from harvest, will benefit red foxes on the expense of pine martens. The ongoing climate change with warmer winters and less snow will likely further benefit the red fox.     

Where to sleep in a rural landscape? A comparative study of resting sites pattern in two syntopic Martes species

The niche‐complementarity hypothesis predicts that two sympatric species must differ in their requirements for one of the three main ecological dimensions (i.e. habitat use, diet, and activity time) to coexist. European pine marten Martes martes and stone marten M. foina are syntopic medium‐sized mustelids with very similar morphology and ecology for which resting sites are a key resource. To better understand how these species coexist, we investigated whether key features of their resting site pattern (number of resting sites, area over which they are distributed, main habitat type used for resting) differed. We used diurnal telemetry to identify resident individuals (e.g. spatially stable individuals over time) and to locate them during resting periods in a fragmented forested area in France. Stone marten used fewer resting sites distributed over a smaller surface area than pine marten. Most stone marten resting sites were located in open habitat (83%) in the proximity of human habitations, whereas pine martens rested almost exclusively in forest (98%). Sex, age, and season explained some variability in both the number of resting sites and the probability of resting within forested habitat for stone marten but not pine marten. The area covered by resting sites was larger in males than in females, but age modulated this difference in an opposite way for the two species. Such a pattern was expected given the intra‐sexual territoriality and the reproductive phenology of these species. Overall, stone marten showed higher inter‐individual variability in resting site pattern than pine marten. The particular pattern observed in subadult male stone martens during summer (increase in resting site surface area and in the probability to rest in forest) may reflect an attempt to settle in forests, and we discuss these implications in the context of interspecific competition.     

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.     

Biological invasions provide a framework for reintroductions: selecting areas in England for pine marten releases

We suggest that reintroductions, like biological invasions, have two phases: establishment when a new population becomes self-sustaining, and spread when a population increases its distribution. Stochastic effects on mortality and sex ratios are most likely to determine whether a population becomes established, while factors influencing birth rates will probably most influence spread. Using this establishment-spread structure, we evaluate the autecological suitability of regions in England for pine marten Martes martes reintroductions. Risks of mortality from predator control, traffic accidents and predation by foxes were used to evaluate suitability for establishment. Mortality risk was higher in all potential release regions in England (selected as having 25% or more woodland cover) than in regions of current pine marten distribution in Scotland; risk of predation was higher in the latter. Indices of prey abundance were used to evaluate suitability for populations to spread. Prey indices in potential release regions were generally higher than in regions of current distribution. A relation between prey, woodland cover, and known pine marten densities suggested that potential release regions are capable of supporting relatively high densities of pine martens, though these might be reduced by higher mortality. We concluded that all potential release regions are suitable for pine marten populations to spread. However, reintroductions should first be to regions with lower risk of mortality, in case higher levels of the latter prevent establishment. The suitability of relict regions of distribution in northern England was low until post-war afforestation, suggesting that habitat suitability constrained recovery of relict populations. These regions remain less suitable for reintroductions.     

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.     

Widespread distribution of Pine martens (<Emphasis Type="Italic">Martes martes</Emphasis>) in a fragmented suburban landscape

Pine martens (Martes martes) are forest specialists that depend either on large patches of forest or on the absence of barriers between smaller forest patches. It thus seems likely that they would disappear from landscapes with heavily fragmented forests and a high density of presumptive barriers that are typical of Central Europe. We studied the distribution of pine martens in a suburban landscape characterised by fragmented forests and a high density of presumptive barriers (e.g. housing areas, roads). We assessed the presence of pine martens using camera traps in a sample of 170 1?×?1-km quadrats and used occupancy modelling to account for imperfect detection. We found an almost complete and countrywide distribution of the pine marten. Larger roads and housing areas were not negatively related to its current distribution as we predicted. We conclude that pine martens are capable to survive in some heavily fragmented landscapes.     

Fate of translocated wild-caught and captive-reared stone martens (<Emphasis Type="Italic">Martes foina</Emphasis>)

We radio-tracked one urban wild-caught and four orphaned captive-reared stone martens (Martes foina) after translocation to novel habitats. The wild-caught marten rapidly returned to urban habitat but died 7 days after release. The captive-reared martens survived >130 days after release. An initial period of extensive movements was followed by decreased activity. Captive-reared martens apparently adapted well to living in the wild. However, we recommend that exclusion of martens from den sites should be used to avoid having to resort to the capture and translocation of problem martens.     

Fate of translocated wild-caught and captive-reared stone martens (<Emphasis Type="Italic">Martes foina</Emphasis>)

We radio-tracked one urban wild-caught and four orphaned captive-reared stone martens (Martes foina) after translocation to novel habitats. The wild-caught marten rapidly returned to urban habitat but died 7 days after release. The captive-reared martens survived >130 days after release. An initial period of extensive movements was followed by decreased activity. Captive-reared martens apparently adapted well to living in the wild. However, we recommend that exclusion of martens from den sites should be used to avoid having to resort to the capture and translocation of problem martens.     

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.

     

Fur snagging as a method of evaluating the presence and abundance of a small carnivore, the pine marten (Martes martes)

The pine marten Martes martes was once considered one of Ireland's rarest carnivores, but is now recolonizing many of the areas where it was once extirpated. It is important to monitor its population expansion so that conservation management decisions can be made regarding the status of this species. In this study, we investigate the use of fur snagging devices as a method of estimating pine marten presence and abundance in lowland and upland broadleaf woodland. We investigated whether these devices reliably detected presence, and how quickly. It was found that hair traps functioned very well in detecting pine martens on all survey transects and was essential in their detection in the most isolated site. A survey length of 6 days was sufficient to detect pine marten in all survey sites, with most detections occurring within the first 4 days. The hair traps functioned equally well as scat location for detecting pine marten and had the added advantage of there being no confusion as to the origin of the sample. Live trapping was used to verify the estimates of relative abundance from hair traps. There was a trend for the number of hair samples per site and the number of times animals were trapped per site to be positively correlated, although this was not significant. Hair traps do provide a real alternative for the monitoring of pine martens in both lowland and upland broadleaf woodlands, overcoming many of the disadvantages associated with scat location.