Translator Disclaimer
1 June 2000 Population trends and harvest management of pine marten Martes martes in Scandinavia
Author Affiliations +
Abstract

In this paper, I describe historical and present harvesting and population trends of pine marten Martes martes in Scandinavia, based on a literature review and analyses of harvest statistics and population indices. The pine marten population has experienced two periods of over-harvesting with subsequent large-scale declines in population density and local extinctions; in the 1500–1600s and in the early 1900s. The principal incentive for harvesting appears to have been economic (valuable pelt), but eradication efforts may have compounded the effect on the population. In the last decades, the pine marten population density has increased. At present, it is receding but pine martens are still harvested intensively. I discuss implications for management, and caution about over-harvesting of the Scandinavian pine marten population in a near future.

Most species of the genus Martes are forest associates, sensitive to the conversion of natural habitats following human forest exploitation (e.g. Buskirk 1994). In addition, their fur is extraordinarily luxurious and valuable, and the history of marten harvesting is both long and intensive. Because of their generally low reproductive potentials (late maturation and small litters; Mead 1994), marten populations are particularly susceptible to increased mortality. As a result, marten populations world-wide have undergone long-term declines caused by over-harvesting and habitat loss. The sable M. zibellina was hunted to near-extinction in Siberia in the 1600s (Bakeyev & Sinitsyn 1994), and in China in the 1800s (Ma & Xu 1994). In North America, populations of American marten M. americana and fisher M. pennanti decreased in the early 1900s, and were locally extirpated, because of excessive harvesting in combination with extensive habitat loss (Strickland 1994). Predator control severely reduced numbers and distribution of the pine marten M. martes in Great Britain in the 1800s (Langley & Yalden 1977), and Grakov (1978) found that populations of pine marten in Russia decreased during periods of heavy harvest, and increased during periods of partial or total protection. Krott & Lampio (1983) described a marked decrease in the pine marten population in Finland during the first half of the 20th century, probably due to over-harvesting.

In this paper, I summarise data on harvesting and population trends of pine marten on the Scandinavian Peninsula, including Norway and Sweden. These two countries constitute a relatively homogenous and well defined geographic area, show several similarities in history and hunting traditions, and probably share the same continuous pine marten population. I review historic accounts, compile harvesting statistics, and present some new data on the recent situation. Where possible, I separate data from northern and southern Sweden, because the hunting traditions differ between these areas (e.g. Ekman 1910), and because Lindström, Brainerd, Helldin & Overskaug (1995) noticed dissimilarities in recent population trends. I discuss implications for the management of the Scandinavian pine marten population.

Methods

Sources for historical review

The historical review was based primarily on: (i) magazines on hunting and natural history published prior to 1960 by the Swedish Association for Hunting and Wildlife Management, the Swedish Museum of Natural History, and the Norwegian Zoological Society (Tidskrift för Jägare och Naturforskare 1834–1836, Svenska Jägareförbundets Nya Tidskrift 1863–1907, Svenska Jägareförbundets Tidskrift 1908–1939, Svensk Jakt 1940–1960, Fauna och Flora 1906–1960, Fauna 1948–1960), and (ii) essays on traditional Saami and peasant hunting and trapping (Olaus Magnus 1555, Ekman 1910, Lundmark 1982, Fjellström 1985, Kjellström 1995).

Harvest statistics

The number of bounties paid in Norway during 1896–1930 were obtained from Statistics Norway (1978a), and Norwegian bag records from the period 1973–1995 were estimated in Statistics Norway (1978b, 1980, 1983, 1987, 1991, 1996). Swedish bag records for 1941–1995 were estimated by the Swedish Association for Hunting and Wildlife Management (1960–1995). During 1960–1963, Swedish bag records were separated between the six northernmost counties and the remaining 16, and from 1964 onwards they were reported by county (Fig. 1). Detailed descriptions of the methods for the bag record estimates are given in Statistics Norway (1996) and on the home page of the Swedish Association for Hunting and Wildlife Management (in Swedish;  www.jagareforbundet.se/forsk/viltovervakning). I present bag records summarised for Sweden and Norway, but Swedish bag records from 1960 onwards are also presented as means per land area, separately for northern Sweden (the six northernmost counties, situated entirely within the boreal zone) and southern Sweden (the remaining 16 counties, at least partly including the boreo-nemoral or nemoral zone). Throughout the paper, winters (harvesting seasons) are denoted by the calendar year after new-year, so that the winter and harvesting season of for instance 1996/97 is denoted 1997.

Figure 1.

The Scandinavian Peninsula (Norway and Sweden), with indication of the southern border of the boreal region (bold line), the location of the counties in northern (grey) and southern (white) Sweden, and the Grimsö Wildlife Research Area (black square) in the northern part of the county of Örebro.

Marten population trends

The Swedish Association for Hunting and Wildlife Management (1960–1995) reported changes in the Swedish game populations by county in the harvesting seasons of 1960–1995, as perceived by county game managers. Based on these reports, I calculated indices of marten population change for each year by giving ‘increase’ a score of +1, ‘stable’ 0, and ‘decrease’ -1. I then obtained one average for the northern counties, and one for the southern (the county of Örebro excluded). Hunters (N = 150–355) in the county of Örebro (see Fig. 1) reported perceived changes in the game populations on their hunting-grounds in the harvesting seasons of 1974–1997. I calculated an index of marten population change for the entire county for each year with the same procedure as above.

I also used changes in trapping success and scat counts to estimate marten population trends. I recorded marten trapping success (martens captured/100 trap-nights) in the harvest seasons of 1990–1997 for 18 marten trappers in or near the northern part of the county of Örebro with a total trapping effort of 8,000–10,000 trap-nights/season. I counted marten scats left in the winters of 1991–1997 on four forest trails (each measuring 1.1–2.4 km) situated in a 55 km2 untrapped wildlife reserve within the Grimsö Wildlife Research Area in the county of Örebro (see Fig. 1). Trails were cleared of scats in mid-October, and searched once or twice when snow conditions were suitable, plus once in mid-April, after the snow had melted.

Results

Historical harvest and population trends before 1900

Records of pine marten harvesting in Scandinavia date back to the Mesolithic Period (Trolle-Lassen 1986). The harvesting has probably been of great economic importance to many people, in Scandinavia most notably the Saamis, the ethnic minority in the north, for thousands of years (Fjellström 1985). The rapid human population growth starting in the 500s A. D. in Sweden, with increasing settlement in remote forest areas and conversion of forest to farmland (Berglund, Helmfrid & Hyenstrand 1994), probably resulted in pine marten densities that gradually declined over several centuries. The Swedish national law code from 1347 regulated the harvest by season; to secure pelt primeness martens should not be killed between mid-March and early November (Ekman 1910). In the 1500s, the Saamis used marten furs to pay taxes to the Swedish and Norwegian states (Olaus Magnus 1555, Lundmark 1982), and the Swedish State and Court took an interest in the marten fur trade (Anonymous 1833). At this time, marten furs were highly valued all over Europe, and were exported from both Sweden and Norway in large quantities (Olaus Magnus 1555, Fjellström 1985). In the 1600s, the pine marten was increasingly recognised as a vermin on game and poultry (Kjellström 1995), and hunting regulations stated that martens could be killed year-round, no matter where or how (Kjellström 1995). In the 1500–1600s over-harvesting likely occurred; the number of marten furs acquired by the Swedish State from the Saamis declined (Lundmark 1982; Fig. 2), and in the late 1600s, pine martens in northern Scandinavia were rare, if not extinct (Dass ca 1670 in Selås 1990). Scattered reports (e.g. Thunberg 1798, Wergeland 1834, Modin 1917) indicate that populations remained low until the late 1800s, when numbers apparently started building up, at least locally, in boreal Scandinavia (Gustafson 1888, Modin 1917, Fjellström 1985, Kjellström 1995). Pelt prices were temporarily low at this time (Modin 1917).

Figure 2.

Number of pine marten furs acquired by the Swedish state from the Saamis in Lule Lappmark, a region of northern Sweden during 1561–1610 and 1614–1615 (redrawn after Lundmark 1982).

Population decline during 1900–1930

In the early 1900s, a country-wide predator extermination programme was initiated in Norway (Steen, Yoccoz & Ims 1990), and also Sweden was embraced by extensive anti-predator propaganda. As part of the predator extermination programme, Norwegian authorities started paying bounties for killed pine martens, and after a few years' delay, the number of martens bountied reached a maximum (Fig. 3). Concurrently, pelt prices increased rapidly, because marten fur wraps and collars became fashionable (Modin 1917, Mackrell 1986). At that time, the value of a marten pelt corresponded to as much as 30 days work for a peasant or a lumberjack (Kjellström 1995). As a result, harvesting became excessive for a period of 2–3 decades (Modin 1915, Anonymous 1916, Eliasson 1943, Selås 1990, Kjellström 1995). The low numbers of several other furbearers, e.g. red fox Vulpes vulpes and lynx Lynx lynx (Hjeljord 1980), and an increased use of modem shotguns (Ekman 1910), may have promoted marten hunting even further. At this time, martens were mainly hunted by snow tracking, and shot after being forced out of their daybed (Ekman 1910). It was told that when a marten trail had been found, people left their occupation and went marten hunting (Modin 1915, Selås 1990). Contemporary accounts describe the hunting for pine marten as “severe persecution” (Modin 1915, author's translation) and “a ruthless war of extermination” (Anonymous 1916, author's translation).

Figure 3.

Number of marten bounties paid in Norway during 1896–1930 based on data from Statistics Norway (1978a).

Already in the 1910s, pine marten numbers had declined dramatically, and the species was probably exterminated from large areas in the north (Ekman 1910, Modin 1915, von Post 1925, Eliasson 1943, Selås 1990, Kjellström 1995). The decline may have been amplified by the lack of hole trees caused by the developing modem forestry (Modin 1917). From 1915, marten harvesting was regionally banned in Sweden for 5-year periods, but this apparently did not halt the trend, as pelt prices were still high, and poaching occurred (Modin 1925, 1926). The number of bounties paid in Norway declined until 1930 (see Fig. 3), at which time the bounty payment was discontinued and the species received full protection in both Norway and Sweden.

General population increase during 1930–1990

Soon after the protection, pine marten numbers started to increase, and continued to do so through the 1960s (von Essen 1941, Dahl 1943, Eliasson 1943, SelJs 1990; Fig. 4a). In 1939 (Sweden) and 1946 (Norway) harvesting seasons were reinstated, however to a limited extent; the open season was set to one month, and the species was still protected regionally (Selås 1990). During the 1940s and 1950s, the reported yearly bag in Sweden was stable at 600–1,000 martens (Fig. 5). In the 1960s and 1970s, the open season was gradually prolonged, and harvesting was extended to include all Scandinavia (Swedish Association for Hunting and Wildlife Management 19601995, Selås 1990). Bag records increased slowly in northern Sweden (Fig. 6a), but remained stable at a low level in southern Sweden (see Fig. 6b). The latter was probably due to a low interest in marten hunting in southern Sweden, as the marten population was generally reported to increase in the 1960s (see Fig. 4a). In Norway the situation was probably similar to that in northern Sweden (see Fig. 5). In the 1940s pelt prices were still relatively high (Bergkvist 1944), but subsequently decreased.

Figure 4.

Perceived changes in the population density of pine marten in Sweden during 1960–1997, based on annual reports from hunters; A) for southern Sweden (the county of Örebro excluded; ) and northern Sweden () , reports by county were obtained from the Swedish Association for Hunting and Wildlife Management (1960–1995); B) for the county of Örebro, reports were supplied by local hunters during 1974–1997. Reports of ‘increase’ were given a score of +1, ‘stable’ 0, and ‘decrease’ -1, and the index gives the average value for all counties or hunters.

Figure 5.

Bag records of pine marten in Sweden during 1941–1995 () and in Norway during 1973–1995 () based on data from Swedish Association for Hunting and Wildlife Management (1960–1995) and Statistics Norway (1978b, 1980, 1983, 1987, 1991, 1996).

Figure 6.

Pine marten bag/1,000 km2 (±SE of the mean) in the Swedish counties during 1960–1995 for: A) six counties in northern Sweden, and B) 16 counties in southern Sweden based on data from the Swedish Association for Hunting and Wildlife Management (1960–1995).

In the 1970s, the increase in marten numbers ceased in northern Sweden; the hunter's index of population change fluctuated around zero (see Fig. 4a). Also in southern Sweden the increase was temporarily slowed down (see Fig. 4), although harvest was low (see Fig. 6b). From the early 1980s, the increase in pine marten numbers was amplified, at least in southern Sweden (Lindström et al. 1995; see Fig. 4). In northern Sweden indices of population change still fluctuated around zero (see Fig. 4a). During this time, bag records also increased, most pronouncedly in southern Sweden (see Figs 5 and 6). In Norway and northern Sweden harvesting reached a maximum in the late 1980s (see Figs 5 and 6a). Trapping gained increased attention (Ødegård, Pedersen & Oppegård 1994, Helldin 1994), and as pelt prices were low, predator control was the main objective for the trapping (Ødegård et al. 1994).

Figure 7.

Trapping success expressed as martens captured/100 trap-nights (±SE of the mean; /) of 18 marten trappers in or near the northern part of the county of Örebro, Sweden 1990–1997, and number of scats found/km trail (±SE of the mean; /) along four forest trails in the Grimsö Wildlife Research Area, the county of Örebro, Sweden, in winters 1991–1997.

Harvest and population trend in the 1990s

Indices of population change imply that marten numbers have decreased in northern Sweden after 1990, and that the increase has ceased in southern Sweden (see Fig. 4). Between 1990 and 1997, pine marten trapping success in the county of Örebro declined (Friedman's test, X2 = 25.1, df = 7, P = 0.007; Fig. 7), and scat counts also showed a declining trend, however not significant (Friedman's test, X2 = 9.84, df = 6, P = 0.13; see Fig. 7). This suggests that marten density decreased also in southern Sweden. The bag records decreased in Norway and northern Sweden, but were still high relative to before 1980 (see Figs 5 and 6a). Bag records remained high in southern Sweden; harvest per unit area in the south was 3–4 times as high as in the north (see Fig. 6b).

Discussion

The historic accounts and data presented suggest that the population density of pine marten in Scandinavia has been largely governed by harvesting intensity in most historic time. The population has experienced two periods of over-harvesting with subsequent large-scale declines in population density and local extinctions; in the 1500–1600s and in the early 1900s. The main incentive for harvesting appears to have been economic, as marten numbers have recovered in times of low pelt prices. However, eradication efforts may have compounded the effect on the marten population.

Recent research has suggested that Scandinavian pine marten population density since the 1970s has been limited by red fox predation (Storch, Lindström & de Jounge 1990, Lindström et al. 1995), possibly in combination with modem forestry practices (Brainerd 1997). Structurally diverse forests supply martens with escape cover from enemies such as the red fox, as well as shelter at harsh weather (Overskaug 1992, Brainerd, Helldin, Lindström, Rolstad, Rolstad & Storch 1995, Eide 1995, Gundersen 1995). Several authors (Grakov 1972, Overskaug 1992, Thompson & Harestad 1994, Brainerd 1997) have also hypothesised that marten foraging success increases with structural diversity. However, Clevenger (1994) suggested that the pine marten, due to its generalised diet, should not be food limited, and Helldin (1999) supported this with data indicating that pine marten density was independent of prey abundance. Forest fragmentation could adversely affect pine marten density (Grakov 1972, Bjärvall, Nilsson & Norling 1977, Brainerd 1997), but may favour red foxes through increased small mammal abundance (Christiansen 1979, Christensen 1985, Henttonen 1989, Gundersen 1995). Accordingly, the red fox populations increased along with the development of the clear-cutting practices in the 1950s and 1960s (Christiansen 1979, Hjeljord 1980). This may explain the interrupted increase in the pine marten population in the 1970s. In the 1980s, the red fox population declined dramatically due to an epizootic of sarcoptic mange, and the increase in the marten population was linked to this decline in both time and amplitude (Lindström et al. 1995).

Lindström et al. (1995) predicted that the pine marten population density should decrease with the recovery of the red fox population in the 1990s. Although the data presented here support this prediction, I cannot exclude the possibility that the intensive harvesting has contributed to the depression of the pine marten population density during the 1990s. Based on a combination of radio-tracking data, harvest records, and age structure in the harvest, I estimated the harvest rate in the county of Örebro and surroundings to be about 6% of the autumn population during 1989–1993 (Helldin 1994), which could probably be considered low, despite the marten's low reproductive capacity. Since then (i.e. between the median year of that study, 1991, and 1997), trapping success and the number of scats found in the area have declined to about one-third or one-fourth of the previous level (bearing in mind that the trend for the scat count was not significant; see Fig. 7). If these measurements represent the actual population change, and the number of martens harvested is constant (as reported from southern Sweden; see Fig. 6b), then the present harvest rate may be 3–4 times higher than that reported for 1989–1993, i.e. around 20%. Although Bakeyev & Sinitsyn (1994) suggested that populations of sable can sustain harvest rates as high as 25–30%, Strickland (1994) pre sented data showing that harvest rates of >20% may affect sex and age ratios of a fisher population. Furthermore, if harvesting is additive to other mortality, it may cause an extended low in a population that is initially decreasing for other reasons (Thompson & Colgan 1987).

Implications for management

Caughley & Sinclair (1994) identified three alternative goals for wildlife population management: conservation, control, or harvesting for a sustained yield. They further stressed the importance of monitoring populations as a tool for achieving the desired goal. Several methods of monitoring marten population changes are available, including track counts, questionnaires, assessing harvest per effort or sex and age ratio in harvest, and others (e.g. Raphael 1994, Strickland 1994). Marten harvesting may be regulated by seasons, refugia, quotas and licensing (Strickland 1994). Juveniles generally dominate the catch early in the season, whereas the proportion of adults increases in late winter (Strickland & Douglas 1987, Helldin 1994). In late winter there is also a higher risk that harvesting is additive to other winter mortality. Hence, a concentrated catch soon after pelts are prime (late October) may affect the population less than if the catch was distributed over the winter. Martens are territorial, and a system of untrapped refugia larger than twice the size of a mean marten home range may secure a population ‘reservoir’, and allow a fast recolonisation of trapped areas (Strickland 1994).

Today the need for careful management of populations of all marten species is widely recognised (Bakeyev & Sinitsyn 1994, Fortin & Cantin 1994, Strickland 1994). Populations of American marten and fisher in North America, and of sable in Russia, are harvested for a sustained yield (Strickland 1994, Bakeyev & Sinitsyn 1994). In populations where numbers are low, efforts are put into reintroductions and habitat protection (Strickland 1994, Bakeyev & Sinitsyn 1994, Ma & Xu 1994, Tatara 1994).

In Scandinavia from the 1600s through the 1800s (in Norway until 1930), the effective policy of governments was to allow an unregulated pine marten harvest, or even to promote eradication efforts. This policy evidently was detrimental to the pine marten population. Present legislation restricts harvesting only by season, primarily for ethical reasons; martens should not be regularly trapped or hunted in the period in which young may depend on their mother. Furthermore, there is practically no monitoring of population changes. With the current low pelt prices, the economic incentives for marten harvesting is largely eliminated, which probably leads to a self-regulating harvest rate. On the other hand, predator control is put forward as an important objective for present marten harvesting, which implies that trappers actually intend to depress the marten population.

Intensive harvesting of a population with a low rate of increase, in a fluctuating environment, may eventually lead to its extinction or severe depletion, irrespectively of the density of the harvested population (Lande, Engen & Sæther 1995). To secure maintained viable populations of pine marten, future management should include carefully considered goals, reliable continuous monitoring of population changes, and tools efficient in regulating the harvest.

Acknowledgements

I thank E. Lindström for numerous discussions on the subject, and H. Andrén, L. Hansson, K. Kauhala, V. Selås, and J. Swenson for valuable comments of earlier drafts of this manuscript. The study was financed by the Swedish Environmental Protection Agency and the private foundations ‘Oskar och Lili Lamms Minne’ and ‘Olle och Signhild Engkvists Stiftelser’.

References

1.

Anonymous 1833: Konung Gustaf I:s bref. - Tidskrift för Jägare och Naturforskare 2: 443. (In Swedish). Google Scholar

2.

Anonymous 1916: Fridlysning af mård. - Svenska Jägare-förbundets Tidskrift 54: 318. (In Swedish). Google Scholar

3.

Bakeyev, N.N. & Sinitsyn, A.A. 1994: Status and conservation of sables in the Commonwealth of Independent States. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 246–254. Google Scholar

4.

Bergkvist, R. 1944: Om m&rden. - Svensk Jakt 82: 399–402. (In Swedish). Google Scholar

5.

Berglund, B., Helmfrid, S. & Hyenstrand, Å. 1994: Tiotusen år i Sverige. - In: Helmfrid, S. (Ed.); Kulturlandskapet och bebyggelsen. SNA, Bra Böcker, Höganäs, Sweden, pp. 12–17. (In Swedish). Google Scholar

6.

Bjärvall, A., Nilsson, E. & Norling, L. 1977: Urskogens betydelse för tjäder och mård. - Fauna och Flora 72: 31–38. (In Swedish). Google Scholar

7.

Brainerd, S.M. 1997. Habitat selection and range use by the Eurasian pine marten (Martes martes) in relation to commercial forestry practices in southern boreal Scandinavia. - PhD thesis, Agricultural University of Norway, Ås, Norway, 155 pp. Google Scholar

8.

Brainerd, S.M., Helldin, J-O., Lindström, E.R., Rolstad, E., Rolstad, J. & Storch, I. 1995: Pine marten (Martes martes) selection of resting and denning sites in Scandinavian managed forest. - Annales Zoologici Fennici 32: 151–157. Google Scholar

9.

Buskirk, S.W. 1994: Introduction to the genus Martes. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 1–10. Google Scholar

10.

Caughley, G. & Sinclair, A.R.E. 1994: Wildlife ecology and management. - Blackwell Science, Cambridge, Massachusetts, 334 pp. Google Scholar

11.

Christensen, H. 1985: Biotopvalg, atferd og næring hos rev (Vulpes vulpes) vinterstid i et barskogsområde på Østlandet. - M.Sc. thesis, University of Oslo, Norway, 76 pp. (In Norwegian). Google Scholar

12.

Christiansen, E. 1979: Skog- og jordbruk, smågnagare og rev. - Tidskrift for Skogbruk 87: 115–119. (In Norwegian). Google Scholar

13.

Clevenger, A.P. 1994: Feeding ecology of Eurasian pine martens and stone martens in Europe. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and Conservation. Cornell University Press, Ithaca, New York, pp. 326–340. Google Scholar

14.

Dahl, E. 1943: Dalamas ryggradsdjur. - Fauna och Flora 38: 220–236. (In Swedish). Google Scholar

15.

Eide, N. 1995: Ecological factors affecting the spatial distribution of mountain hare, red fox, pine marten and roe deer in a southern boreal forest during winter. - M.Sc. thesis, Agricultural University of Norway, Ås, Norway, 41 pp. Google Scholar

16.

Ekman, S. 1910: Norrlands jakt och fiske. - Norrländskt Handbibliotek 4, Uppsala, Sweden (in facsimile print 1983, Två Förläggare Bokförlag, Umeå, Sweden), 481 pp. (In Swedish). Google Scholar

17.

Eliasson, N.A. 1943: Kommer mArden tillbaka? - Svensk Jakt 81: 64–66. (In Swedish). Google Scholar

18.

Fjellström, P.M. 1985: Samemas samhälle i tradition och nutid. (In Swedish with English summary: Lappish society in tradition and the present day). - P.A. Norstedt & Söners Förlag, Stockholm, Sweden, 640 pp. Google Scholar

19.

Fortin, C. & Cantin, M. 1994: The effects of trapping on a newly exploited American marten population. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 179–191. Google Scholar

20.

Grakov, N.N. 1972: Vliyanie kontsentrirovannykh rubok lesa na chislennost lesnoy kunitsy. (In Russian with English summary: Effect of concentrated clear fellings on the abundance of the pine marten [Martes martes]). - Biulleten Moskovskoe Obshchestva Ispytatelei Prirody, Otdel Biologij 77: 14–23. Google Scholar

21.

Grakov, N.N. 1978: Izmenenie chislennosti lesnoy kunitsy i nekotorie zakonomemosti etogo protsessa. (In Russian with English summary: Long-term changes in the abundance of the pine marten [Martes martes] and some pat-tems of this process). - Biulleten Moskovskoe Obshchestva Ispytatelei Prirody, Otdel Biologij 83: 46–56. Google Scholar

22.

Gundersen, V.S. 1995: Habitatbruk hos mår vinterstid. - M.Sc. thesis, Agricultural University of Norway, Ås, Norway, 40 pp. (In Norwegian). Google Scholar

23.

Gustafson, W. 1888: Notiser. - Svenska Jägareförbundets Nya Tidskrift 26: 102. (In Swedish). Google Scholar

24.

Helldin, J-O. 1994: Mårdjakten i Mellansverige 1989–1993. - Viltforum 1994(1), Swedish Association for Hunting and Wildlife Management, Uppsala, Sweden, 8 pp. (In Swedish). Google Scholar

25.

Helldin, J-O. 1999: Diet, body condition, and reproduction of Eurasian pine martens (Martes martes) during cycles in microtine density. - Ecography 22: 324–336. Google Scholar

26.

Henttonen, H. 1989: Metsien rakenteen muutoksen vaikutuksesta myyräkantoihin ja sitä kautta pikkupetoihin ja kanalintuihin - hypoteesi. (In Finnish with English summary: Does an increase in the rodent and predator densities, resulting from modem forestry, contribute to the long-term decline in Finnish tetraonids?). - Suomen Riista 35: 83–90. Google Scholar

27.

Hjeljord, O. 1980: Viltbiologi. - Landbruksforlaget, Olso, Norway, 318 pp. (In Norwegian). Google Scholar

28.

Kjellström, R. 1995: Jakt och fångst i södra Lappland i äldre tid. - Nordiska museets förlag, Stockholm, Sweden, 326 pp. (In Swedish). Google Scholar

29.

Krott, P. & Lampio, T. 1983: Näädän esiintymisestä Soumessa 1900-luvulla. (In Finnish with English summary: Occurrence of the pine marten (Martes martes) in Finland in the 1900s). - Suomen Riista 30: 60–63. Google Scholar

30.

Lande, R., Engen, S. & Sæther, B-E. 1995: Optimal harvesting of fluctuating populations with a risk of extinction. - American Naturalist 145: 728–745. Google Scholar

31.

Langley, P.J.W. & Yalden, D.W. 1977: The decline of the rarer carnivores in Great Britain during the nineteenth century. - Mammal Review 7: 95–116. Google Scholar

32.

Lindström, E.R., Brainerd, S.M., Helldin, J-O. & Overskaug, K. 1995: Pine marten - red fox interactions: a case of intraguild predation? - Annales Zoologici Fennici 32: 123–130. Google Scholar

33.

Lundmark, L. 1982: Uppbörd, utarmning, utveckling: det samiska fågstsamhällets övergång till rennomadism i Lule lappmark. (In Swedish with English summary: Taxation, expropriation and development: the transition of Saami (Lapps) hunting society to reindeer nomadism in Lule Lappmark). - PhD thesis, Umeå University, Umeå, Sweden, 230 pp. Google Scholar

34.

Ma, Y. & Xu, L. 1994: Distribution and conservation of sables in China. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York: 255–261. Google Scholar

35.

Mackrell, A. 1986: Shawls, stoles and scarves. - Batsford, London, UK, 96 pp. Google Scholar

36.

Mead, R.A. 1994: Reproduction in Martes. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 404– 422. Google Scholar

37.

Modin. E. 1915: Rörande mårdens fridlysning. - Svenska Jägareförbundets Tidskrift 53: 377. (In Swedish). Google Scholar

38.

Modin, E. 1917: Något om mården i Norrland. - Svenska Jägareförbundets Tidskrift 55: 206–208. (In Swedish). Google Scholar

39.

Modin, E. 1925: M&rden i Ångermanland. - Svenska Jägareförbundets Tidskrift 63: 125. (In Swedish). Google Scholar

40.

Modin, E. 1926: De smärre pälsdjuren. - Svenska Jägareförbundets Tidskrift 64: 62. (In Swedish). Google Scholar

41.

Olaus Magnus, 1555: Historia de Gentibus Septentrionalibus, Rome, Italy. - (Swedish translation in 1909–1925, Michaelsgillet, Stockholm, Sweden; in facsimile print 1976, Gidlunds, Stockholm, Sweden, 1804 pp. (In Swedish). Google Scholar

42.

Overskaug, K. 1992: Studier av m˚r (M. martes) og rødrev (V. vulpes) i et midt-norsk barskogsområde gjennom vinteren. - M.Sc. thesis, University of Trondheim, Norway, 61 pp. (In Norwegian). Google Scholar

43.

Raphael, M.G. 1994: Techniques for monitoring populations of fishers and American martens. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 224–240. Google Scholar

44.

Selås, V. 1990: Maren. Norges dyr I. - Cappelens Forlag, Oslo, Norway, pp. 142–151. (In Norwegian). Google Scholar

45.

Statistics Norway 1978a: Hunting Statistics 1846–1977. - Official Statistics of Norway, report A 955. Statistics Norway, Oslo, Norway, 195 pp. Google Scholar

46.

Statistics Norway 1978b: Hunting Statistics 1977–1995. - Official Statistics of Norway, report A 974. Statistics Norway Oslo, Norway, 70 pp. Google Scholar

47.

Statistics Norway 1980: Hunting Statistics 1979. - Official Statistics of Norway, report B 144. Statistics Norway, Oslo, Norway, 60 pp. Google Scholar

48.

Statistics Norway 1983: Hunting Statistics 1982. - Official Statistics of Norway, report B 417, Statistics Norway, Oslo, Norway, 60 pp. Google Scholar

49.

Statistics Norway 1987: Hunting Statistics 1986. - Official Statistics of Norway, report B 721, Statistics Norway, Oslo, Norway, 58 pp. Google Scholar

50.

Statistics Norway 1991: Hunting Statistics 1990. - Official Statistics of Norway, report B 993, Statistics Norway, Oslo, Norway, 56 pp. Google Scholar

51.

Statistics Norway 1996: Hunting Statistics 1995. - Official Statistics of Norway, report C 331, Statistics Norway, Oslo, Norway, 58 pp. Google Scholar

52.

Steen, H., Yoccoz, N.G. & Ims, R.A. 1990: Predators and small rodent cycles: an analysis of a 79-year time series of small rodent population fluctuations. - Oikos 59: 115–120. Google Scholar

53.

Storch, I., Lindström, E. & de Jounge, J. 1990: Diet and habitat selection of the pine marten in relation to competition with the red fox. - Acta Theriologica 35: 311–320. Google Scholar

54.

Strickland, M.A. 1994: Harvest management of fishers and American martens. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 149–164. Google Scholar

55.

Strickland, M.A. & Douglas, C.W. 1987: Marten. - In: Novak, M., Baker, J.A., Hobbard, M.E. & Malloch, B. (Eds.); Wild furbearer management and conservation in North America. Ministry of Natural Resources, Ontario, Canada, pp. 531–546. Google Scholar

56.

Swedish Association for Hunting and Wildlife Management 1960: Svenska jagareförbundets årsredogörelser 59/60. - Svenska Jägareförbundet, Spånga, Sweden, 286 pp. (In Swedish). Google Scholar

57.

Swedish Association for Hunting and Wildlife Management 1961: Svenska jagareförbundets årsredogörelse 60/60. - Svenska Jägareförbundet, Spånga, Sweden, 233 pp. (In Swedish). Google Scholar

58.

Swedish Association for Hunting and Wildlife Management 1962: Svenska jagareförbundets årsredogörelse 61/62. - Svenska Jägareförbundet, Spånga, Sweden, 240 pp. (In Swedish). Google Scholar

59.

Swedish Association for Hunting and Wildlife Management 1963: Svenska jagareförbundets årsredogörelse 62/63. - Svenska Jägareförbundet, Spånga, Sweden, 267 pp. (In Swedish). Google Scholar

60.

Swedish Association for Hunting and Wildlife Management 1964: Svenska jagareförbundets årsredogörelse 63/64. - Svenska Jägareförbundet, Spånga, Sweden, 93 pp. (In Swedish). Google Scholar

61.

Swedish Association for Hunting and Wildlife Management 1965: Svenska jagareförbundets årsredogörelse 64/65. - Svenska Jägareförbundet, Spånga, Sweden, 104 pp. (In Swedish). Google Scholar

62.

Swedish Association for Hunting and Wildlife Management 1966: Svenska jagareförbundets årsredogörelse 65/66. - Svenska Jägareförbundet, Spånga, Sweden, 111 pp. (In Swedish). Google Scholar

63.

Swedish Association for Hunting and Wildlife Management 1967: Svenska jagareförbundets årsredogörelse 66/67. - Svenska Jägareförbundet, Spånga, Sweden, 164 pp. (In Swedish). Google Scholar

64.

Swedish Association for Hunting and Wildlife Management 1968: Svenska jagareförbundets årsredogörelse 67/68. - Svenska Jägareförbundet, Spånga, Sweden, 174 pp. (In Swedish). Google Scholar

65.

Swedish Association for Hunting and Wildlife Management 1969: Svenska jagareförbundets årsredogörelse 68/69. - Svenska Jägareförbundet, Spånga, Sweden, 174 pp. (In Swedish). Google Scholar

66.

Swedish Association for Hunting and Wildlife Management 1970: Svenska jagareförbundets årsredogörelse 69/70. - Svenska Jägareförbundet, Spånga, Sweden, 175 pp. (In Swedish). Google Scholar

67.

Swedish Association for Hunting and Wildlife Management 1971: Svenska jagareförbundets årsredogörelse 70/71. - Svenska Jägareförbundet, Spånga, Sweden, 190 pp. (In Swedish). Google Scholar

68.

Swedish Association for Hunting and Wildlife Management 1972: Svenska jagareförbundets årsredogörelse 71/72. - Svenska Jägareförbundet, Spånga, Sweden, 185 pp. (In Swedish). Google Scholar

69.

Swedish Association for Hunting and Wildlife Management 1973: Svenska jagareförbundets årsredogörelse 72/73. - Svenska Jägareförbundet, Spånga, Sweden, 165 pp. (In Swedish). Google Scholar

70.

Swedish Association for Hunting and Wildlife Management 1974: Svenska jagareförbundets årsredogörelse 73/74. - Svenska Jägareförbundet, Spånga, Sweden, 148 pp. (In Swedish). Google Scholar

71.

Swedish Association for Hunting and Wildlife Management 1975: Svenska jagareförbundets årsredogörelse 74/75. - Svenska Jägareförbundet, Spånga, Sweden, 173 pp. (In Swedish). Google Scholar

72.

Swedish Association for Hunting and Wildlife Management 1976: Svenska jagareförbundets årsredogörelse 75/76. - Svenska Jägareförbundet, Spånga, Sweden, 192 pp. (In Swedish). Google Scholar

73.

Swedish Association for Hunting and Wildlife Management 1977: Svenska jagareförbundets årsredogörelse 76/77. - Svenska Jägareförbundet, Spånga, Sweden, 138 pp. (In Swedish). Google Scholar

74.

Swedish Association for Hunting and Wildlife Management 1978: Svenska jagareförbundets årsredogörelse 77/78. - Svenska Jägareförbundet, Spånga, Sweden, 148 pp. (In Swedish). Google Scholar

75.

Swedish Association for Hunting and Wildlife Management 1979: Svenska jagareförbundets årsredogörelse 78/79. - Svenska Jägareförbundet, Spånga, Sweden, 145 pp. (In Swedish). Google Scholar

76.

Swedish Association for Hunting and Wildlife Management 1980: Svenska jagareförbundets årsredogörelse 79/80. - Svenska Jägareförbundet, Spånga, Sweden, 168 pp. (In Swedish). Google Scholar

77.

Swedish Association for Hunting and Wildlife Management 1981: Svenska jagareförbundets årsredogörelse 80/81. - Svenska Jägareförbundet, Spånga, Sweden, 159 pp. (In Swedish). Google Scholar

78.

Swedish Association for Hunting and Wildlife Management 1982: Svenska jagareförbundets årsredogörelse 81/82. - Svenska Jägareförbundet, Spånga, Sweden, 192 pp. (In Swedish). Google Scholar

79.

Swedish Association for Hunting and Wildlife Management 1983: Svenska jagareförbundets årsredogörelse 82/83. - Svenska Jägareförbundet, Spånga, Sweden, 151 pp. (In Swedish). Google Scholar

80.

Swedish Association for Hunting and Wildlife Management 1984: Svenska jagareförbundets årsredogörelse 83/84. - Svenska Jägareförbundet, Spånga, Sweden, 127 pp. (In Swedish). Google Scholar

81.

Swedish Association for Hunting and Wildlife Management 1985: Svenska jagareförbundets årsredogörelse 84/85. - Svenska Jägareförbundet, Spånga, Sweden, 156 pp. (In Swedish). Google Scholar

82.

Swedish Association for Hunting and Wildlife Management 1986: Svenska jagareförbundets årsredogörelse 85/86. - Svenska Jägareförbundet, Spånga, Sweden, 118 pp. (In Swedish). Google Scholar

83.

Swedish Association for Hunting and Wildlife Management 1987: Svenska jagareförbundets årsredogörelse 86/87. - Svenska Jägareförbundet, Spånga, Sweden, 122 pp. (In Swedish). Google Scholar

84.

Swedish Association for Hunting and Wildlife Management 1988: Svenska jagareförbundets årsredogörelse 87/88. - Svenska Jägareförbundet, Spånga, Sweden, 160 pp. (In Swedish). Google Scholar

85.

Swedish Association for Hunting and Wildlife Management 1989: Svenska jagareförbundets årsredogörelse 88/89. - Svenska Jägareförbundet, Spånga, Sweden, 120 pp. (In Swedish). Google Scholar

86.

Swedish Association for Hunting and Wildlife Management 1990: Svenska jagareförbundets årsredogörelse 89/90. - Svenska Jägareförbundet, Spånga, Sweden, 117 pp. (In Swedish). Google Scholar

87.

Swedish Association for Hunting and Wildlife Management 1991: Svenska jagareförbundets Srsredogorelse 90/91. - Svenska Jägareförbundet, Spånga, Sweden, 130 pp. (In Swedish). Google Scholar

88.

Swedish Association for Hunting and Wildlife Management 1992: Vilttillgång och avskjutning för jaktåret 1991/92. - Svenska Jägareförbundet Spånga, Sweden, 17 pp. (In Swedish). Google Scholar

89.

Swedish Association for Hunting and Wildlife Management 1993: Vilttillgång och avskjutning för jaktåret 1992/93. - Svenska Jägareförbundet Spånga, Sweden, 11 pp. (In Swedish). Google Scholar

90.

Swedish Association for Hunting and Wildlife Management 1994: Vilttillgång och avskjutning för jaktåret 1993/94. - Svenska Jägareförbundet, Spånga, Sweden, 11pp. (In Swedish). Google Scholar

91.

Swedish Association for Hunting and Wildlife Management 1995: Vilttillgång och avskjutning för jaktiret 1994/95. - Svenska Jägareförbundet, Spånga, Sweden, 11 pp. (In Swedish). Google Scholar

92.

Tatara, M. 1994: Ecology and conservation status of the Tsushima marten. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 272–279. Google Scholar

93.

Thompson, I.D. & Colgan; P.W. 1987: Numerical responses of martens to a food shortage in northcentral Ontario. - Journal of Wildlife Management 51: 824–835. Google Scholar

94.

Thompson, I.D. & Harestad, A.S. 1994: Effects of logging on American martens, and models for habitat management. - In: Buskirk, S.W., Harestad, A.S., Raphael, M.G. & Powell, R.A. (Eds.); Martens, sables, and fishers: biology and conservation. Cornell University Press, Ithaca, New York, pp. 355–367. Google Scholar

95.

Thunberg, C.P. 1798: Beskrivning pö svenske djur. - J.F. Edmans Kungliga Academiska Boktryckeriet, Uppsala, Sweden, 112 pp. (In Swedish). Google Scholar

96.

Trolle-Lassen, T. 1986: Human exploitation of the pine marten (Martes martes) at the late mesolithic settlement of Tybrind Vig in Western Funen. - In: Konigsson, L-K. (Ed.); Nordic Late Quartemary Biology and Ecology. Striae 24, Uppsala, Sweden, pp. 119–124. Google Scholar

97.

Wergeland, H. 1834: Strödda underrättelser. - Tidskrift för Jägare och Naturforskare 3: 856. (In Swedish). Google Scholar

98.

von Essen, K. 1941: Fågelobservationer från Strömsberg vid Jönköping 1940. - Fauna och Flora 36: 184. (In Swedish). Google Scholar

99.

von Post, A. 1925: Om förändringar i faunan och floran i och omkring östra delen av Hjälmaren (Sörmlandsdelen) under det senaste halvseklet. - Fauna och Flora 20: 209. (In Swedish). Google Scholar

100.

Ødegård, F.E., Pedersen, H.B. & Oppegård, B. 1994: Mår i Akershus: undersökelse av jakt og fangst på mår i Akershus. - Akershus Jeger- og Fiskerforbund, Oslo, Norway, 11 pp. (In Norwegian). Google Scholar
© WILDLIFE BIOLOGY
J-O. Helldin "Population trends and harvest management of pine marten Martes martes in Scandinavia," Wildlife Biology 6(4), 111-120, (1 June 2000). https://doi.org/10.2981/wlb.2000.006
Received: 11 December 1998; Accepted: 31 March 2000; Published: 1 June 2000
JOURNAL ARTICLE
10 PAGES


SHARE
ARTICLE IMPACT
Back to Top