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Martesmartens

By Lauren Dubberley

Diversity

The Genus Martes contains a total 7 species of martens and the sable. Members of the Family Mustelidae, they typically have large paws with partially retractile claws, long slender bodies with short legs, and luxurious coats. Distributed throughout North America, Europe and Asia these animals are forest-dwelling and call the taiga, deciduous, and coniferous forest home. Martes are omnivores with diets including small mammals, fish, birds, fruit, and seeds. Like many other Mustelidae martens live solitary only to meet during breeding season in the late spring/early summer. Young are born altricial and stay with the mother for 3-4 months before going off on their own. The most distinctive feature of this genus is the typically yellow/brown fur coat, a popular choice for fur hunters for hundreds of years. ("Marten", 2019; Proulx, 2000)

Geographic Range

Martes species are only found within the Northern Hemisphere and no species of martes has successfully established an introduced population in another non-native area. The pine marten (Martes martes) is indigenous over most of Europe, from Mediterranean to Fennoscandian taiga, and to western Siberia and Iran. The stone/beech marten (M. foina) occurs from Mongolia and the northern Himalayas to most of Europe. The American marten (M. americana) occurs in large contiguous populations in forested habitats of North America north of 35° latitude. The sable M. zibellina occurs in Russia, Mongolia, China, North Korea, and Japan. The yellow-throated marten (M. flavigula) and the Nilgiri marten, (M. gwatkinsi) occur from the Himalayas to eastern Russia, south to the Malay Peninsula and Sunda Shelf to Taiwan. The Japanese marten (M. melampus) occurs in forests of the main Japanese archipelago and the Korean peninsula. (Clark, et al., 1987; "Marten", 2019; MONAKHOV, 2011; Nowak, 2005; Proulx, 2000)

Habitat

The pine marten (Martes martes) is found in wooded areas, shrublands, and coniferous forests. Large open areas are actively avoided by this species, while not restricted to the forest a dependence on trees exists. In urban areas (M. martes) use hedgerows and small wood plots as habitat. The Beech marten (M. foina) frequents forests, woodlands and pastures, and is expanding in suburban and urban areas. In urban areas, beech martens den almost entirely in buildings, particularly during winter. The beech marten does not dig burrows or occupy another species abandoned burrow. Instead, it nests in naturally occurring fissures and clefts in rocks, spaces between stones in rock slides and inhabited or uninhabited stone structures. It may live in tree holes at a height of up to 9 metres. The sable (M. zibellina) over most of its distribution inhabits coniferous taiga forests dominated by spruce, pine, larch, cedar, and birch in both lowland and mountainous terrain. Sables live in burrows near riverbanks and in the thickest parts of woods. These burrows are commonly made in between the roots of trees to make it more secure. The yellow-throated marten (M. flavigula); including the Nilgiri marten, (M. gwatkinsi) occurs in sub-tropical and tropical forests. These two species are almost completely arboreal and do not often come down to the ground. The American marten (M. americana) is associated with coniferous and mixed forests with overhead cover and a dense underbrush. Species occupy exceptionally large territories, the males larger than the females, ideal territories are highly productive and have high canopy cover. (Clark, et al., 1987; "Marten", 2019; MONAKHOV, 2011; Pereboom, et al., 2008; Proulx, 2000; Smith and Schaefer, 2002)

Systematic and Taxonomic History

Martes is located inside the subfamily Guloninae which also contains 'Wolverines Gulo', the Fisher 'Pekania', and the 'Tayra Eira'. The fisher used to be a member of Martes before genetic data was used to place it in its own genus. Martes species are in the same family,<Mustelidae>, and more distantly related to 'Weasels Mustela' and 'Otters Lurinae'. Within Martes data suggested that the lineages of Martes differentiated with five radiation stages from the generic divergences (stage I) to the intraspecific divergences (stage V). The first offshoots are of M. flavigula, M. pennanti, and G. gulo (stage II), the second is M. foina (stage III), and the third are M. americana, M. martes, M. melampus, and M. zibellina (stage IV). (Hosoda, et al., 2000; Marmi and Francisco López‐Giráldez, 2004; Wilson and Reeder, 2005)

  • Synonyms
    • Charronia
    • Foiana
    • Lamprogale
    • Mustela
    • Pekania
    • Zibellina

Physical Description

All 7 species in Martes have a long, slender-body with relatively large rounded ears, triangular head, short limbs, and typically a bushy tail. Males of this genera are typically 18 to 24 in and 4 to 12 lbs and females are 12 to 16 in and 1 to 4 lbs. The American Marten (M. americana) has coloration from pale yellowish buff to tawny brown to almost black. Their head is usually lighter than the rest of their body, while the tail and legs are darker. American marten usually have a characteristic throat and chest bib ranging in color from pale straw to vivid orange. In the Yellow-throated marten (M. flavigula) the top of the head is black brown with shiny brown highlights and a mixture of white hair tips. The fur is a shiny brownish-yellow color with a gold shine along the surface of the back.The legs and belly are bright yellow, the chest and lower part of the throat are a brighter, orange-golden color than the back and belly. The front paws and lower forelimbs are pure black, while the upper parts of the limbs are the same color as the front of the back. The tail is of a shiny pure black color. The Beech marten has a dark-brown tail and a white throat patch. The patch is large and generally has two projections extending backwards to the base of the forelegs and upward on the legs. The Nilgiri marten (M. gwatkinsii) has the most striking markings with deep brown color from head to rump, with the forequarters being almost reddish, with a bright throat ranging in colour from yellow to orange. The European Marten is a solid light to dark brown with a creme/yellow colored bib. The Japanese Marten varies in color from dark brown to dull yellow with a cream-colored throat. The Sable ranges from dark brown to light brown, geographic location has created local color variations with individual amounts of grey, white, and yellow. Seasonal variation does exist in their coats. Summer fur is shorter, sparser, less compact and lustrous when compared to winter coats. Young marten have their adult patterning and coat from a young age (Allen and Heptner, 2019; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; Gimranov and Kosintsev, 2015; "Marten", 2019; MONAKHOV, 2011; "Nilgiri Marten", 2020; Reig, 1992)

  • Sexual Dimorphism
  • male larger

Reproduction

Martes species are usually solitary organisms, only coming together to mate. Males usually fight violently with each other for the right to breed with a female, the largest and strongest male getting to mate with the most females. Courtship behavior can also include chasing and other "playful" behaviors as seen in the Sable (M.zibellina). (Allen and Heptner, 2019; Barja, et al., 2011; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; MONAKHOV, 2011; "Nilgiri Marten", 2020; Ruggiero and Henry, 1993)

Martes species are all seasonal breeders, June to August for the majority of species. Additionally, the Yellow-throated marten (M. flavigula) has another breeding season from mid-February to March while the Japanese marten's (M. melampus) is from March to mid-may. Embryonic implantation is delayed for about 8 months. The Beech and American Marten (M. foina and M. americana) have a gestation period of 236 to 237 days. The Sable (M. zibellina) takes longer with 245 to 298 days.The European Pine Marten (M. martes) has the shortest gestation period at only a month, in other species the development period is around a month but they have a longer gestation period. Litter size ranges significantly within this genera with the Yellow-throated marten and Sable having 3 kits tops to the Beech marten's 7. Weaning typically occurs around 1 to 1.5 months old and sexual maturity at 1 to 2 years old. (Allen and Heptner, 2019; "European Pine Marten", 2020; Frederickson, 2020; Helldin, 2000; "Marten", 2019; "Nilgiri Marten", 2020; Nowak, 2005)

In Martes species the parental investment comes only from the female in every species except for the Sable (M.zibellina). Pre-fertilization maternal investment includes constructing a suitable dening space, typically off the ground and away from predators. Pre-independence the female martin is responsible for feeding, protecting, and teaching her kits the hunting skills they need. In the Sable the male assists the female by protecting her territory and providing her food. Martin generally are weaned around 1 to 1.5 months and ready to leave the mothers den around 3 to 4 months old. (Allen and Heptner, 2019; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; MONAKHOV, 2011; "Nilgiri Marten", 2020)

  • Parental Investment
  • altricial
  • female parental care
  • pre-hatching/birth
    • provisioning
      • male
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

In general the lifespan of Martes species in the wild is unknown and only a few species are kept commonly enough in captivity to generate significant data. European pine marten (M. martes) has been recorded to have an average lifespan of 3-4 years in the wild and 18 years in captivity. Japanese marten have an unknown lifespan in the wild, but in captivity specimens live up to 12 years. Threats most likely to limit the lifespan of these species are conflict with humans and predators. (Allen and Heptner, 2019; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; "Marten", 2019; MONAKHOV, 2011; "Nilgiri Marten", 2020)

Behavior

Martes species are solitary, males and females live in separate territories that may overlap. In the European pine marten (M.martes) both sexes use fecal markings to mark the boundaries of a territory and display reproductive status. Yellow-throated martens (M. flavigula) are an unique exception, pairs hunt and live in a territory together. Martes species are varied in their lifestyles; Nilgiri martens (M. gwatkinsii) are diurnal and arboreal. Beech martens (M. foina) are crepuscular and terrestrial. Competition among Martes species occurs often and has resulted in dietary niches in the Pine and Beech marten. Martes are active year-round avoiding heavy snow by traveling through the trees. Rainfall is also actively avoided by the Sable (M. zibellina) and the American pine marten (M. americana) who seek out shelter as a rule due to an increased rate of mortality when wet. Marten den in hollow trees, rock fissures, or scrub covered fields. (Allen and Heptner, 2019; Barja, et al., 2011; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; Gazzola and Balestrieri, 2019; MONAKHOV, 2011; "Nilgiri Marten", 2020; PARR and DUCKWORTH, 2007)

Communication and Perception

Martes species are athletic species with well developed senses to take down prey and survive in harsh climates. Sensitive ears pick up a variety of vocal calls, 7 specifically distinguished in the American pine marten (M. americana) alone. A well developed sense of smell allows marten species to pick up on pheromones and fecal markings. Fecal markings communicate territory boundaries, stress levels, and reproductive status to other members of that species. Yellow-throated martens (M. flavigula) use sharp eyesight, ear position, vocalizations, and physical contact to communicate with its hunting partner. (Allen and Heptner, 2019; Barja, et al., 2011; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; Gazzola and Balestrieri, 2019; MONAKHOV, 2011; "Nilgiri Marten", 2020; PARR and DUCKWORTH, 2007)

Food Habits

Martes species are all omnivorous with varying main diets based on where the species is found and lifestyle. American martens (M. americana) primarily prey on 'Snowshoe hares (Lepus americanus)' with Glaucomys sp., Tamias sp., birds, reptiles, and in season fruits and berries (Vaccinium sp.). Sables (M. zibellina) primarily prey on mice (Peromyscus sp.) and voles. Winter stores have contained least weasels (Mustela nivalis), up to 7 voles (Microtus and Myodes), and pine cones of Pinus sibirica and Pinus pumila. The main diet of European pine martens (M. martes) is 'voles microtines' with hares as the most common alternative. Japanese pine martens (M. melampus) are piscivores eating fish, frogs, and small mammals like the Japanese hare (Lepus brachyurus). Insects, fruits, and seeds are eaten when necessary. The diet of yellow-throated martens is less well known, but probably includes small mammals, poultry, and muntjac fawns (Muntiacus). Beech martens (M. foina) are unusual in that their diet is the highest in plant content of all Martes species. Winter diets consisting predominantly of fruits. Summer diets are mainly sparrow-like birds, small mammals, and eggs. Beech, Sable, and American martens all eat carrion as well. (Clark, et al., 1987; Frederickson, 2020; Helldin, 2006; MONAKHOV, 2011; Nowak, 2005; PARR and DUCKWORTH, 2007)

Predation

All Martes species except for the yellow-throated martens (M. flavigula) are vulnerable to 'birds of prey Accipitridae', 'foxes Vulpes', 'wild cats Feildae', and 'canids Canidae'. No specific anti-predator adaptations have been identified in this group. Martes will hide in trees and rock fissures when pursued by predators, they will use their flexible body and sharp claws to defend themselves and avoid capture. Yellow-throated martens have few predators outside of Siberian tigers and Asiatic black bears. (Allen and Heptner, 2019; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; MONAKHOV, 2011)

Ecosystem Roles

Martes species are secondary level consumers in their ecosystem, being both predator and prey. Japanese martens (M. melampus) play an important role in seed dispersing; their large size and large home ranges mean seeds can be carried farther away from the parent plant. American pine martens (M. americana) and European pine martens (M. martes) both have evolutionary relationships with tree squirrel Tamiasciurus sp. and European red squirrels (Sciurus vulgaris) respectively. They consume squirrels and then live in their dens. Martes species are home to a host of parasitic infections. In the sable (M. zibellina) 14 species of parasitic worms and 21 species of fleas have been identified. (Clark, et al., 1987; "European Pine Marten", 2020; MONAKHOV, 2011)

  • Ecosystem Impact
  • disperses seeds
Commensal/Parasitic Species

Economic Importance for Humans: Positive

Martes species are valued furbearers, Sable (M. zibellina) are harvested at rates of 300,000-400,000 pelts from the wild and 10,000-20,000 pelts from fur farms. Beech martens (M. foina) have a lower quality fur and are less valuable than Sables and Pine martens, Beech marten pelts make up 10-12% of the market. Beech martens and Yellow-throated martens (M. flavigula) are infamously tamable, zoologist George Rolleston theorized that the "domestic cat" of the Ancient Greeks and Romans was in fact the beech marten. Marine biologist Jeanne Villepreux-Power kept two tame beech martens. The Japanese marten (M. melampus) primarily preys upon the Japanese hare (Lepus brachyurus) which is considered a pest as they reduce the quality of trees with their browsing. (Allen and Heptner, 2019; Frederickson, 2020; Herr, et al., 2009; MONAKHOV, 2011)

  • Positive Impacts
  • pet trade
  • body parts are source of valuable material
  • controls pest population

Economic Importance for Humans: Negative

Martes species are not a threat to people but cause damage to agriculture, cars, and livestock. Beech martes (M. foina) bite through the cables and wires of cars, the reasoning for this is not fully understood but typically peaks in the spring when young martens begin to explore. On 29 April and 21 November 2016, two beech martens shut down the Large Hadron Collider, the world's most powerful particle accelerator, by climbing on 18–66 kV electrical transformers. Japanese martens (M. melampus) consume insects important to agriculture decreasing crop yields and long-term field health. The Yellow-throated marten (M. flavigula) kill and consume cats and various types of poultry. Old reports say that it was able to attack an unarmed man in groups of three or four but scavenging on human corpses is more common. (Frederickson, 2020; Nowak, 2005)

  • Negative Impacts
  • crop pest

Conservation Status

All martes species are listed as Least Concern by the IUCN except for the Nilgiri marten (M. gwatkinsii) which is listed as vulnerable. Sable (M. zibellina) populations are increasing, populations of Japanese martens (M. melampus), Pine martens (M. martes), Beech martens (M. foina), Nilgiri martens are stable, and American martens (M. americana) and Yellow-throated martens (M. flavigula) populations are in decline. Threats to conservation include the fur trade, hunting, and habitat loss. To conserve the American marten den boxes were placed in the forest to enhance the habitat and allow for proper population monitoring. (Croose, et al., 2016; "European Pine Marten", 2020; Frederickson, 2020; Herr, et al., 2009; "Nilgiri Marten", 2020; Nowak, 2005; Sato, et al., 2009)

  • IUCN Red List [Link]
    Not Evaluated

Other Comments

Martes species have cultural and historical significance in their native region. People in the Iga region, Mie Prefecture, have a saying about the Japanese marten (M. Melampus), "the fox has seven disguises, the tanuki has eight, and the marten has nine," and a legend relates how the marten has greater ability in shapeshifting than the fox or tanuki. In the Akita Prefecture and the Ishikawa Prefecture, if a marten crosses in front of someone, it is said to be an omen for bad luck, and in the Hiroshima Prefecture, if one kills a marten, one is said to soon encounter a fire. In the Fukushima Prefecture, they are also called heko, fuchikari, komono, and haya, and they are said to be those who have died in avalanches in disguise. In Croatia M. Martes pelts were highly valued goods used as a form of payment. The banovac, a coin struck and used between 1235 and 1384, included the image of a marten. This is one of the reasons why the Croatian word for marten, kuna, is the name of the modern Croatian currency. A running marten is shown on the coat of arms of Slavonia and subsequently on the modern design of the coat of arms of Croatia. The official seal of the Croatian Sabor (parliament) from 1497 until the late 18th century had a similar design. The etymology of Martes comes from the Middle English 'Mearth' or martryn in turn borrowed from the Anglo-French martrine and Old French martre (Latin martes). (Allen and Heptner, 2019; Clark, et al., 1987; "European Pine Marten", 2020; Frederickson, 2020; "Marten", 2019; "Nilgiri Marten", 2020)

Contributors

Lauren Dubberley (author), Colorado State University, Tanya Dewey (editor), University of Michigan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

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.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

delayed implantation

in mammals, a condition in which a fertilized egg reaches the uterus but delays its implantation in the uterine lining, sometimes for several months.

diurnal
  1. active during the day, 2. lasting for one day.
endothermic

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.

female parental care

parental care is carried out by females

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

piscivore

an animal that mainly eats fish

polygynous

having more than one female as a mate at one time

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

stores or caches food

places a food item in a special place to be eaten later. Also called "hoarding"

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

taiga

Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

2020. "European Pine Marten" (On-line). Accessed February 16, 2020 at en.wikipedia.org/wiki/European_pine_marten.

2019. "Marten" (On-line). Accessed February 09, 2020 at https://en.wikipedia.org/wiki/Marten.

2020. "Nilgiri Marten" (On-line). Accessed February 16, 2020 at en.wikipedia.org/wiki/Nilgiri_marten.

Allen, G., V. Heptner. 2019. "Yellow-throated marten" (On-line). Accessed February 16, 2020 at en.wikipedia.org/wiki/Yellow-throated_marten.

Barja, I., G. Silván,, L. Martínez-Fernández, J. Carlos Illera. 2011. Physiological Stress Responses, Fecal Marking Behavior, and Reproduction in Wild European Pine Martens (Martes martes). Journal of Chemical Ecology, 37: 253–259.

Clark, T., E. Anderson, C. Douglas, M. Strickland. 1987. Martes americana. The American Society of Mammalogists, 289: 1-4.

Cook, J. 2002. Molecular evolution of Holarctic martens (genus Martes, Mammalia: Carnivora: Mustelidae). Molecular Phylogenetics and Evolution, 24(2): 169-179.

Croose, E., J. Birks, J. Martin. 2016. Den boxes as a tool for pine marten Martes martes conservation and population monitoring in a commercial forest in Scotland. Conservation Evidence, 13: 57-61.

Frederickson, B. 2020. "Beech Marten" (On-line). Accessed February 16, 2020 at en.wikipedia.org/wiki/Beech_marten.

Gazzola, A., A. Balestrieri. 2019. Nutritional ecology provides insights into competitive interactions between closely related Martes species. Mammal Review, 50(1): 82-90.

Gimranov, D., P. Kosintsev. 2015. Differentiation of three Martes species (M. martes, M. zibellina, M. foina) by tooth morphotypes. Vertebrate Palaeontology, 14(8): 647-656.

Helldin, J. 2000. Population trends and harvest management of pine marten Martes martes in Scandinavia. Wildlife Biology, 6(4): 11-120.

Helldin, J. 2006. Diet, body condition, and reproduction of Eurasian pine martens Mantes martes during cycles in microtine density. Ecography, 22(3): 324-336.

Herr, J., L. Schley, T. Roper. 2009. Stone martens (Martes foina) and cars: investigation of a common human–wildlife conflict. European Journal of Wildlife Research volume, 55: 471–477.

Hosoda, T., H. Suzuki, M. Harada. 2000. Evolutionary trends of the mitochondrial lineage differentiation in species of genera Martes and Mustela. Genes & Genetic Systems, 75(5): 259-267.

MONAKHOV, V. 2011. Martes zibellina (Carnivora: Mustelidae). The American Society of Mammalogists, 43: 75-86.

Marmi, J., J. Francisco López‐Giráldez. 2004. Phylogeny, evolutionary history and taxonomy of the Mustelidae based on sequences of the cytochrome b gene and a complex repetitive flanking region. Zoologica Scripta, 33(6): 481-499.

Nowak, R. 2005. Walker's Carnivores of the World. Baltimore: Johns Hopkins Press.

PARR, J., J. DUCKWORTH. 2007. Notes on diet, habituation and sociality of Yellow-throated Marten Martes fl avigula. Small Carnivore Conservation, 36: 27-29.

Pereboom, V., M. Mergey, N. Villerette, R. Helder, J. Gerard, T. Lodé. 2008. Movement patterns, habitat selection, and corridor use of a typical woodland-dweller species, the European pine marten (Martes martes), in fragmented landscape. Canadian Journal of Zoology, 86: 983-991.

Proulx, G. 2000. World Distribution and Status of the Genus Martes in 2000. Martens and Fishers (Martes) in Human-Altered Environments, 1: 21-76.

Reig, S. 1992. Geographic Variation in Pine Marten (Martes martes) and Beech Marten (M. foina) in Europe. Journal of Mammalogy, 73(4): 744–769.

Ruggiero, L., S. Henry. 1993. Courtship and Copulatory Behavior of Martes americana. Northwestern Naturalist, 74(1): 18-22.

Sato, J., S. Yasuda, T. Hosoda. 2009. Genetic Diversity of the Japanese Marten (Martes melampus) and Its Implications for the Conservation Unit. Zoological Science, 26(7): 457-466.

Schwartz, M., A. Ruiz-Gonzalez, R. Masuda, C. Pertoldi. Conservation genetics of the genus Martes: Assessing within-species movements, units to conserve, and connectivity across ecological and evolutionary time. Conservation Genetics: 398-428.

Smith, A., J. Schaefer. 2002. Home-range size and habitat selection by American marten (Martes americana) in Labrador. Canadian Journal of Zoology, 80(9): 1602-1609.

Wilson, D., D. Reeder. 2005. "Martes" (On-line). Mammal Species of the World: A Taxonomic and Geographic Reference. Accessed March 25, 2020 at https://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?s=y&id=14001203.

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