The European pine marten is distributed through most portions of continental Eurasia from western Europe in the west to western Siberia in the east, from the northern edge of coniferous forest in the north to Asia Minor in the south. The species also inhabits the Caucasus and many Mediterranean islands including the Balearic Islands, Corsica, Sardinia, Elba and Sicily. The degree to which island populations are due to human introduction is not entirely known. Formerly widespread in Great Britain, it is now restricted to Ireland and northern portions of mainland Britain (Corbet and Southern 1977, Nowak 1999).
Martes martes prefer forest habitats, including deciduous, mixed, and coniferous forest. Old-growth forest is often preferred over young forest (Overskaug et al 1994). The species is considered to be a habitat specialist. Having a closed treetop as cover from predation is thought to be an important habitat criterion for pine martens. They are found outside of forest, however. On the island of Minorca, Martes martes showed no habitat preference, living in shrubland and seemingly indifferent to tree cover (Clevenger 1994). It is thought that the absence of predators on the island has allowed the martens to become habitat generalists. In Scotland, pine martens frequent many habitat types. They are seen in young forest plantations, coarse grassland, heather and grass moorland, and borders. Stone dykes are used as runways to get from area to another (Gurnell et al 1994).
Martes martes is a medium-sized carnivore, about the size and proportions of a large domestic cat. Head and body length is 450-580 mm, tail length is 160-280 mm. Substantial size variation is found geographically. Sexual dimorphism is also seen in size, with males outweighing females by 12-30%. The fur is a rich brown coat that is thick and silky in the winter and short and coarse in the summer. Pads on the soles are completely covered with fur in the winter. Juveniles acquire their adult pelage in their first winter, and a complete molt occurs only once a year, in the spring. The winter fur grows in September. The coloration includes an irregular, creamy-orange throat patch, a grayish tint on the belly, and darkening on the paws. The tail is long and bushy and the ears are relatively large and triangular. An abdominal scent gland is present, as well as anal scent glands.
The pine marten is an adept tree climber, with many adaptations including bone and muscle structure for powerful forelimbs, long tail to aid in balancing, and well-developed claws (Grzimek 1990, Corbet and Southern 1977, Nowak 1999).
Reproduction in European pine martens is tied closely to the seasonality of their temperate habitats, as it is in many members of the family Mustelidae. Mating and fertilization in July and August is followed by a period of delayed implantation that lasts about seven months. Implantation occurs in late February and March. The timing of implantation responds to photoperiod, specifically to the spring increase in duration of daylight. Postimplantation development lasts 30-35 days, and parturition occurs in late March through April. Each litter produces an average of three young (range, 2-5). Adult females only have four functional mammae.
Adult male pine martens exhibit a distinct seasonal testicular cycle. Males complete testicular development a full month before estrous, and begin regression about the time females enter estrous. The first visible sign of estrous in females is an enlarged vulva. Most matings occur within a 30- to 45-day period, during which females may exhibit one to four periods of sexual receptivity. These periods last for 1-4 days and have an interval of 6-17 days. Copulation is prolonged, lasting 30-50 minutes, and may occur on the ground or in trees. In captivity, multiple copulations with one or more males can occur during each period of receptivity. A "false heat" occurs in February and March, corresponding to implantation and the beginnings of pregnancy. The increased social activity and intrasexual aggression may facilitate late dispersal of yearlings (Helldin and Lindstrom 1995).
At birth, young of Martes martes weigh about 30 g. They are blind, deaf and toothless, and have thick, short fur. The eyes open at 34-38 days. Young martens begin consuming solid food at 36-45 days, and weaning occurs about six weeks after parturition. At 7-8 weeks, young emerge from the den and may begin dispersing at 12-16 weeks, during the breeding season. Some young may overwinter in the natal territory and disperse in the following spring.
In the wild, male and female Martes martes may mate in their first summer, at 14 months of age. The first mating season, however, may typically be deferred until the second or third year. In captivity, most males do not breed until 27 months old. This could be due to stress incurred under captive conditions or inaccuracy in aging wild animals. In captivity, Martes martes lives about 15 years, and males can be sexually vigorous throughout their lives. (Mead 1994, Helldin and Lindstrom 1995).
Martes martes is mostly active during the night and at dusk. Hollow trees are the preferred nesting site, and an individual has several nests within the home range. Squirrel nests, abandoned bird nests and rock crevices are also used as hideaways. At colder ambient temperatures in the winter, martens more frequently choose to rest underground. Animals are solitary except when young are in the nest, and male-female bonds are transitory. Field studies indicate that males may successfully guard a mated female through territory defense if his range encompasses that of the female (Schroepfer et al 1997). Home range size estimates vary widely between studies. It is clear that male ranges are larger than female ranges and that they overlap those of one or more females. Homes ranges within both sexes, however, are usually non-overlapping. There is either high individual and geographical variation in home range size, or difficulty in measurement by researchers. Some sources give an average of 23 sq km for males and 6.5 sq km for females (Nowak 1999), others estimate only 2.2 sq km for males and 1.5 sq km for females (Zalewski et al 1995). On the island of Minorca, ranges were measured as 0.5 sq km for females and 6.9 sq km for males (Clevenger 1993). In the majority of the species' range, where the habitat is highly seasonal, range size changes seasonally as well. The autumn/winter ranges can be 8-54% smaller than the spring/summer ranges (Zalewski et al 1995). This range difference is also reflected in the distances covered by males at night, with further travel in the summer than the winter (Schroepfer et al 1997). Density of martens is about 0.3-0.8 sq km. At high densities intrasexual ranges can overlap. Independent subadults are tolerated within the range of adult animals; these may represent offspring that did not disperse in their first fall.
Communication is known to occur through scent-marking of home ranges. Abdominal and anal scent glands are used to mark areas throughout the territory. In the nest, young communicate to their mother by twittering.
As discussed earlier, social behavior intensifies in late winter, as evidenced by increased scent marking frequency, intersexual tolerance, intrasexual aggression, and hormonal levels. This was initially though to be a mating period, but it is clear that mating does not occur until summer. In addition, males have not undergone spermatogenesis yet. The term "false heat" has been applied to this behavioral enigma (Helldin and Lindstrom 1995). Due to the unexplained occurrence of extra, subadult individuals only in the early winter months, it has been suggested that the aggressive behavior serves to force dispersal of these remaining young from the previous summer before the new litter is born. (Helldin and Lindstrom 1995, Schroepfer et al 1997, Clevenger 1993, Zalewski et al 1995, Nowak 1999)
The pine marten is omnivorous. It favors animal food, relying on small mammals for most of the year. The diet composition and proportion often change according to season and local conditions. Populations respond to the unpredictable cycles of rodents, such as voles, by drastically increasing their consumption of these prey items (Zalewski et al 1995). The reproductive characteristics of Martes martes prevent it from closely tracking the rodent cycles: a population increase is seen a full year after a rodent boom. When fruits and berries become abundant in the autumn, martens may fill 30% of their diet with these resources (in Scotland and on the island of Minorca). In other regions, such as Poland, fruits may never be eaten (Zalewski et al 1995, Clevenger 1993, Gurnell et al 1994). Aside from the effects of seasonally available fruits and unpredictable rodent booms, diet is otherwise reasonably constant. Favored foods include voles, squirrels, other small mammals, birds, insects, carrion, and frogs, reptiles, and snails. Diets of pine martens that forage along a loch in Scotland have been recorded to include crabs, echinoderms, and barnacles (Gurnell et al 1994).
Food is stored in the summer and autumn to compensate for low winter resources (Helldin and Lindstrom 1995). Martens are skillful treetop hunters, racing on thin, swinging branches and leaping from one treetop to another in pursuit of a squirrel. Their arboreal adaptations discussed earlier allow this acrobatic ability. Foraging also occurs extensively on the forest floor. In habitats other than forest, all foraging is completed on the ground. Several skull attributes allow martens to be remarkable predatators: elongated braincase allows for insertion of an elarged temporalis muscle; a large flange on the mandibular fossae prevent dislocation of the lower jaw; and, well-developed shearing and crushing cheekteeth. These characteristics aid martens in capture, restraint and processing of prey (Zalewski et al 1995, Clevenger 1993, Gurnell et al 1994, Grzimek 1990, Helldin and Lindstrom 1995).
The winter coat of the European pine marten has always been much in demand. The species has been successfully kept on fur farms. Life history characteristics, however, prevent trade of pine marten fur from being feasible on a large commercial scale (Grzimek 1990).
No negative effects. Martes martes avoids human settlements, and has never been known as a pest. (pp. 410-411 in Grzimek 1990)
Lisa Schwanz (author), University of California, Berkeley, James Patton (editor), University of California, Berkeley.
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
uses smells or other chemicals to communicate
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
having the capacity to move from one place to another.
the area in which the animal is naturally found, the region in which it is endemic.
reproduction that includes combining the genetic contribution of two individuals, a male and a female
uses touch to communicate
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
A terrestrial biome found in temperate latitudes (>23.5? N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
Clevenger, A. 1994. Habitat characteristics of Eurasian pine martens Martes martes in an insular Mediterranean environment.. Ecography, 17: 257-263.
Clevenger, A. 1993. Pine marten (/Martes martes/) home ranges and activity patterns on the island of Minorca, Spain.. Zeitschrift fuer Saeugetierkunde, 58: 137-143.
Corbet, G., H. Southern. 1977. The Handbook of British Mammals. London: Blackwell Scientific Publications.
Grzimek, 1990. Grzimek's Encyclopedia of Mammals. San Francisco: McGraw-Hill Publishing Company.
Gurnell, G., T. Vennig, B. Maccaskill, D. Maccaskill. 1994. The food of pine martens (/Martes martes/) in West Scotland. Journal of Zoology (London), 234: 680-683.
Mead, R. 1994. Reproduction in /Martes/. Pp. 404-422 in S Buskirk, A Harestad, M Raphael, R Powell, eds. Martens, Sables, and Fishers: Biology and Conservation. Ithaca: Cornell University Press.
Nowak, R. 1999. Walker's Mammals of the World, 6th Edition, Volume 1. Baltimore: Johns Hopkins University Press.
Overskaug, K., H. Broseth, B. Knutsen. 1994. Area and habitat use of pine martens, /Martes martes/, in mid-Norway.. Lutra, 37: 81-88.
Schroepfer, R., P. Wiegand, H. Hogrefe. 1997. The implications of territoriality for the social system of the European pine marten, /Martes martes (L., 1758).. Zeitschrift fuer Saeugetierkunde, 62: 209-218.
Trees for Life, 2004. "Species profile: pine marten" (On-line). Trees for life. Accessed May 05, 2004 at http://www.treesforlife.org.uk/tfl.pinemarten.html.
Zalewski, A., W. Jedrzejewski, B. Jedrzejewska. 1995. Pine marten home ranges, numbers and predation on vertebrates in a deciduous forest (Bialowieza National Park, Poland).. Annales Zoologici Fennici, 32: 131-144.