Content uploaded by H. Okarma
Author content
All content in this area was uploaded by H. Okarma on Oct 12, 2017
Content may be subject to copyright.
Status and management of the wolf in Poland
Abstract
The history, past status and distribution of the wolf Canis lupus in Poland is presented. After being persecuted as a pest, the wolf was given game species status in 1975, with a close season from 1 April to 31 July. The wolf inhabits about half of Poland with two core areas in the southeast and northeast of the country. Two main methods of hunting wolves (from a tower and using fladry) are described. Management attitudes and threats to the wolf population are also discussed.
... Persistently persecuted, they recovered during periods of wars (Jędrzejewska et al. 1996). The last Polish eradication campaign lasted from 1955 to 1975 and resulted in the near extinction of wolves (Okarma 1989, 1993, Jędrzejewska et al. 1996. The government ceased persecution and upgraded the wolf status from pest to game species in 1975 (Okarma 1993). ...
... The last Polish eradication campaign lasted from 1955 to 1975 and resulted in the near extinction of wolves (Okarma 1989, 1993, Jędrzejewska et al. 1996. The government ceased persecution and upgraded the wolf status from pest to game species in 1975 (Okarma 1993). At this time, the estimated population of wolves in Poland was <100 individuals, and their range was mainly restricted to north-eastern and south-eastern parts of the country (Okarma 1993). ...
... The government ceased persecution and upgraded the wolf status from pest to game species in 1975 (Okarma 1993). At this time, the estimated population of wolves in Poland was <100 individuals, and their range was mainly restricted to north-eastern and south-eastern parts of the country (Okarma 1993). In 1995, wolves became protected in most regions of Poland, and in 1998, the strict protection was extended to the entire country (Gula 2008). ...
The wolf Canis lupus population occupying the lowlands of central Europe is divided into two management units: the Baltic population east of the Vistula river and the Central European population to the west. We re-evaluated arguments for this division in the context of the ongoing wolf recovery and its usefulness for wolf management in Poland. To do so, we 1) compared the recovery stage on each side of the Vistula, 2) investigated the history of wolf occurrence in western Poland after the eradication campaign of 1955–1975, 3) evaluated dispersal corridors, dispersal distances and genetic datafor evidence of a possible isolation of the two alleged populations and 4) compared habitat characteristics in Poland on each side of the Vistula.
The total area of forest occupied by wolves was 56 600 km2 in 2015 and grew by 5340 km2 until June 2017. Wolves in eastern Poland occurred in more areas than predicted by a habitat model, whereas wolves in the west have not yet recolonized all suitable habitats. Wolves have never been extinct west of the Vistula after the eradication campaign, but their recovery started only in the 1980s. Areas currently occupied by wolves on both sides of the Vistula are interconnected by dispersal corridors less than 100 km long, and population genetic studies show that wolves inhabiting the Polish lowlands constitute one genetic cluster. The wolf habitats west of the Vistula have a higher proportion of forests are less fragmented. We conclude that wolves inhabiting the lowlands on both sides of the Vistula river belong to the same population, have similar conservation status, and should be treated as the same management unit.
... Hunters were interested in the highest possible reduction of its populations to limit losses caused by wolf depredation on wild ungulates and to remove alleged threats to humans [54]. Following accounts of the increasing predation of wolves on farm animals and wildlife, in 1955, the national government excluded the wolf from the list of game species and proclaimed a programme for the extirpation of the 'pest' [55]. "Akcja wilcza" [eng. ...
... At that time, there were less than 60 wolves in Poland, mainly in the eastern-most parts of the country [16]. This relisting of the wolf as a game species was spurred by the warnings of scientists that the species might be exterminated and by the growing awareness of the potential positive impact of wolves on ecosystems [54,55]. However, the lobbying of groups supporting wolf eradication contributed to the reintroduction of high bounties in 1984 (without cancelling of the game status). ...
... Already in 1989, wolf hunting was prohibited in the Białowieża Forest (East Poland) due to ecological research concerning wolves carried out by wildlife biologists [55,63]. One of the biologists involved explained in an interview: "In 1989, after political changes in Poland, our academic tutor [name] became vice-minister of environmental protection (. . ...
Poland was one of the first countries of Central and Eastern Europe with stable wolf populations to effectively introduce year-round protection of the species. This paper traces the process of policy change using institutional theory as an organizational perspective. Based on the analysis of data from desk research and semi-structured interviews, we propose a model of institutional change and argue that in the 1990s, environmental activists and wildlife biologists successfully used a political window of opportunity connected with socio-economic transformation after 1989 and managed to induce the government to move the species from the domain of hunting to the domain of nature conservation. The new policy, informed by an ecological paradigm, diverged from the historical path dominated by hunters and the vision of the wolf as a pest and a hunting target. The improved protection led to the numerical growth of Poland's wolves and ultimately to their westward expansion.
... In Poland, wolves have a history of persecution and recovery, providing the chance to explore how historical wolf presence can influence local attitudes toward the carnivore. By the early 1900s state-sanctioned eradication campaigns had eliminated wolves from most of Poland except a few forested regions in the east of the country (Wolsan et al. 1992, Okarma 1993. During the world wars, wolf numbers began to recover (Wolsan et al. 1992), but renewed eradication efforts after WWII had reduced wolves in Poland to <100 individuals by the early 1970s (Okarma 1993). ...
... By the early 1900s state-sanctioned eradication campaigns had eliminated wolves from most of Poland except a few forested regions in the east of the country (Wolsan et al. 1992, Okarma 1993. During the world wars, wolf numbers began to recover (Wolsan et al. 1992), but renewed eradication efforts after WWII had reduced wolves in Poland to <100 individuals by the early 1970s (Okarma 1993). In 1975, the status of the wolf was upgraded from pest to game species, which allowed hunting with firearms only (no poisoning or trapping), with a nation-wide 4-month closed season introduced in 1981 (Okarma 1993). ...
... During the world wars, wolf numbers began to recover (Wolsan et al. 1992), but renewed eradication efforts after WWII had reduced wolves in Poland to <100 individuals by the early 1970s (Okarma 1993). In 1975, the status of the wolf was upgraded from pest to game species, which allowed hunting with firearms only (no poisoning or trapping), with a nation-wide 4-month closed season introduced in 1981 (Okarma 1993). The wolf population subsequently recovered to an estimated 700-900 individuals by the mid-1990s, but wolves were still rare in western Poland (Okarma 1993, Gula 2008a, Nowak and Mysłajek 2017. ...
Human dimensions are a critical part of large carnivore conservation. We investigated how the historical presence of wolves (Canis lupus) influences public attitudes toward the carnivore and its management in rural areas of Poland. From March 2016 to March 2017, we used a self-administered questionnaire to assess attitudes of rural residents (n = 292) and foresters (n = 325) in 6 regions where wolves have either been continuously present, or where they have recently recovered after a period of absence. While we found that attitudes toward wolves were generally neutral or positive, differences in attitudes did exist across target groups and regions with long and short histories of wolf presence. Foresters tended to have more negative attitudes toward wolves than did rural residents, and their attitudes remained stable across regions. In contrast, rural residents from the regions with an uninterrupted history of wolf presence, which also suffer greater rates of livestock depredation, tended to have less positive attitudes than did residents from regions where wolves have recovered more recently. Knowledge of wolves and wildlife value orientations were also positive predictors of attitudes. Negative attitudes among local residents and lower support for wolf conservation in response to wolf attacks on livestock could be a major obstacle for the continued recovery of wolves in Europe. Our results reinforce the need for proactive approaches, involving both information campaigns and actions to mitigate wolf predation on livestock, to maintain local support for wolf conservation.
... In Poland, after World War II, the government launched an eradication campaign that nearly eliminated the wolves from the country, leaving only small populations remaining in the easternmost part of the Carpathian Mountains and in the lowlands in the northeast of the country (Okarma, 1993). After listing the wolf as a game species in 1975 and then strictly protected species in 1998, wolves started to recolonize forested areas in Poland, Germany, and other parts of western Europe (Gula et al., 2020;Linnell et al., 2008;Szewczyk et al., 2019). ...
... 65% of the region. Wolf densities in Carpathians are the highest in the country, especially in the eastern part of the region (Chief Inspectorate of Environmental Protection, 2019; Gula, 2008b;Okarma, 1993). ...
As the wolf Canis lupus populations continue to recover across Europe, livestock depredation becomes increasingly challenging for their effective conservation. We aim to (1) analyze the spatiotemporal variation in wolf attacks on livestock in relation to the landscape structure, livestock species, and the phase of wolf expansion in Poland and (2) discuss the implications for conservation and management in an expanding, protected wolf population. From 2008 to 2018, farmers reported 5499 attacks on livestock with 13,164 killed individuals, and the number of attacks increased 2.7-fold at the country scale. Sheep were among the most frequently killed livestock, and surplus killing (>2 killed individuals) was relatively common in captive deer and sheep depredation. The attacks were patchily distributed; 59% of all kills occurred in municipalities constituting 1% of the country surface. The probability and number of attacks were positively influenced by forest and pasture cover, and by the occurrence of depredation in previous years. Spatial variation and long-term dynamics in livestock depredation by wolves can be attributed to different husbandry practices and phases of wolf recovery in three regions of the country. Our results indicate that accelerating increase in depredation rates during the initial phases of wolf recovery is likely to be followed by stabilization or decrease in attacks. A detailed spatiotemporal analysis of wolf–livestock conflict can help in management decisions in areas with ongoing wolf population recovery.
... Potwierdza to fakt, że myśliwi często wiążą niższą skuteczność polowań z obecnością tych drapieżników (37). Co więcej, wielu myśliwych chętnie widziałoby wilka z powrotem na liście gatunków łownych (9,36 Analiza macierzy korelacji pomiędzy liczebnością wilka szarego a liczebnością bydła (r 5 = 0,96) i owiec (r 6 = 0,97) wykazała wysoką dodatnią korelację pomiędzy zmiennymi, co świadczy o braku wpływu zwiększającej się populacji wilka szarego na liczebność tych gatunków hodowlanych. W przypadku pogłowia kóz wysoka ujemna korelacja (r 7 = -0,85) sugeruje istotną zależność, niemniej jednak spadek liczebności tego gatunku w Polsce nie jest spowodowany preferencjami żerowymi wilka, tylko zmniejszonym zainteresowaniem hodowlą tego gatunku (tab. ...
The aim of this study was to determine the predation pressure of the grey wolf (Canis lupus) on wild ungulate populations and the relationship between the size of the grey wolf population and the number of confirmed kills of livestock and companion animals in Poland over eight hunting seasons from 2011/2012 to 2018/2019. There is a positive and complete relationship between the increasing wolf population and the number of confirmed deer kills. The size of the wolf population in Poland was compared to the size of domestic and companion animal population and the number of specimens that fell prey to the grey wolf. It was demonstrated that as the wolf population in Poland increases, the number of confirmed kills of farm and companion animals rises. It seems rational to take steps aimed at mitigating this problem as soon as possible. It is necessary to consider implementation of an integrated protection system that involves culling and employing available methods to deter wolves from attacking grazing herds.
... Recent studies have shown that the Central European population originates from animals inhabiting the western edges of the Baltic population (Szewczyk et al., 2019). Baltic population had never been eradicated, yet it experienced a bottleneck lasting at least 20 years (Flousek et al., 2014;Wolsan et al., 1992), and between 1960 and 1980 its size dropped below 50 individuals (Okarma, 1989(Okarma, , 1993Sumiński, 1975;Wolsan et al., 1992). In the highlands, the Carpathian population is also spreading westwards, and past bottlenecks that happened in the 20th century are visible in contemporary genetic diversity (Hulva et al., 2018). ...
Aim
The aim of the study was to assess the effect of changes in population size and migration on variation in functional immunity genes in the previously bottlenecked population of the grey wolf, Canis lupus.
Location
Eastern/Central Europe: Poland, Czechia, Slovakia.
Methods
We genotyped 7 immunity genes: three MHC-DLA genes (dog leukocyte antigen) and four Toll-like receptor (TLR) genes among 130 wolves originating from three populations: two lowland (Baltic and Central European), and highland Carpathian. We contrasted the population structure in immunity genes with a neutral structure based on 13 microsatellites, and we analysed signatures of selection in the immunity loci.
Results
We found high overall genetic variance in immunity genes and no evidence for decreased diversity in the recently established populations. The population structure in immunity loci was weak, with pairwise FST lower than for neutral markers. Although the results of neutrality tests were not significant, we identified codons under selection, both positive and negative.
Main Conclusions
We demonstrated that despite recent population expansion which is expected to result in decreased genetic diversity, the diversity of immunity genes in the newly established wolf population is similar to those in the source population. This suggests that migrations do not cause allele loss in grey wolf. Signatures of selection on codon level, but not in tests using allele frequencies, suggest the contrasting effects of demography and selection.
... Wolves tend to prefer wild prey and seem to cause harm to livestock when starving and when livestock is easy to take i.e not well protected with infrastructures such as fencing (Pulliainen, 1993;Bibikov, 1985). In Poland it was observed that a four fold increase in wild ungulate populations led to wolf attacks on livestock falling fourfold in correlation with the ungulate population boom (Okarma, 1993). Despite the evidence, wolf prejudices remain, see Figure. ...
The following study tested the viability of managing wolf populations via the presence of managed "wolf restaurants". Wolf restaurants in this context refers to supplementary feeding of wolves in close proximity to their den or areas where wolves frequent within their territory. An individual based model was built using the Netlogo software. The model was used to computationally test this theory as it has not yet been attempted in the field. This form of management was tested in two simulated wilderness areas; a large area (3000km 2 ) and a smaller area (312km 2 ). This study found that in a large wilderness area with a stable wolf population and medium densities of wild prey, nutrition management was effective in reducing dispersion by 33% and livestock depredation by 55%. The model was then tested for a smaller wilderness area. It was discovered that when prey density was high and wolf restaurants were present, livestock depredation was reduced by 450% and movement ecology was reduced by as much as 300% compared to unmanaged wolves in the same environment. The results of this model suggest that wolf nutrition management is possible to mitigate instances of wolf-human conflicts in large wild communities, but that smaller ecological islands of wilderness with high prey density is where it has the potential to work best. The smaller simulated wilderness area emulates that which wolves are moving into throughout Europe. Wolf nutrition management can be used as a tool to enhance the success of future wolf reintroductions and re-establishments.
... Wolves tend to prefer wild prey and seem to cause harm to livestock when starving and when livestock is easy to take i.e not well protected with infrastructures such as fencing (Pulliainen, 1993;Bibikov, 1985). In Poland it was observed that a four fold increase in wild ungulate populations led to wolf attacks on livestock falling fourfold in correlation with the ungulate population boom (Okarma, 1993). Despite the evidence, wolf prejudices remain, see Figure. ...
The following study tested the viability of managing wolf populations via the presence of managed "wolf restaurants". Wolf restaurants in this context refers to supplementary feeding of wolves in close proximity to their den or areas where wolves frequent within their territory. An individual based model was built using the Netlogo software. The model was used to computationally test this theory as it has not yet been attempted in the field. This form of management was tested in two simulated wilderness areas; a large area (3000km 2 ) and a smaller area (312km 2 ). This study found that in a large wilderness area with a stable wolf population and medium densities of wild prey, nutrition management was effective in reducing dispersion by 33% and livestock depredation by 55%. The model was then tested for a smaller wilderness area. It was discovered that when prey density was high and wolf restaurants were present, livestock depredation was reduced by 450% and movement ecology was reduced by as much as 300% compared to unmanaged wolves in the same environment. The results of this model suggest that wolf nutrition management is possible to mitigate instances of wolf-human conflicts in large wild communities, but that smaller ecological islands of wilderness with high prey density is where it has the potential to work best. The smaller simulated wilderness area emulates that which wolves are moving into throughout Europe. Wolf nutrition management can be used as a tool to enhance the success of future wolf reintroductions and re-establishments.
... Another potential control measure is fladry, which is a fence with ropes marked by hanging colored flags that sway in the wind and provide a visual warning signal . For centuries, fladry had been used to corner and hunt down wolves in Eastern Europe (Okarma, 1993), but in recent years it has been adopted as a measure to fence off livestock and thus to prevent wolf attacks (Musiani et al., 2003;Iliopoulos et al., 2019). Additionally, new technological measures are proposed such as electric fences, shock collars put on wolves, or deterrents such as the radio-activated guard (RAG) box, a device which keeps wolves away by emitting strobe lights and sounds when triggered by a signal from radio collars (Breck et al., 2002;Schultz et al., 2005;Salvatori and Mertens, 2012). ...
Wolves (Canis lupus) can kill domestic livestock resulting in intense conflicts with humans. Damage to livestock should be reduced to facilitate human-wolf coexistence and ensure positive outcomes of conservation efforts. Current knowledge on the effectiveness of livestock protection measures from wolves is limited and scattered in the literature. In this study, we compiled a dataset of 30 cases describing the application of 11 measures of protecting cattle and smaller livestock against wolves, estimated their effectiveness as a relative risk of damage, and identified the best measures for damage reduction. We found that: (1) lethal control and translocation were less effective than other measures, (2) deterrents, especially fladry which is a fence with ropes marked by hanging colored flags that sway in the wind and provide a visual warning signal, were more effective than guarding dogs; (3) deterrents, fencing, calving control and herding were very effective, but the last two measures included only one case each; and (4) protection of cattle was more effective than that of small stock (sheep and goats, or sheep only) and mixed cattle and small stock. In all of these cases, the relative risk of damage was reduced by 50-100%. Considering Germany as an example of a country with a recovering wolf population and escalating human-wolf conflicts, we suggest electric fences and electrified fladry as the most promising measures, which under suitable conditions can be accompanied by well-trained livestock guarding dogs, and the temporary use of deterrents during critical periods such as calving and lambing seasons. Further research in this field is of paramount importance to efficiently mitigate human-wolf conflicts.
p>The present study is an effort towards the international and multidisciplinary approach to conservation of European biodiversity. The main aim was to map the distribution of suitable areas for the conservation of bears, lynx and wolves in the Carpathian Mountains. It was done applying a distance classifier, the Mahalanobis distance, over a set of environmental variables representing the region.
The results suggested that 41, 58 and 65% of the Carpathian Ecoregion is highly suitable for bear, lynx and wolf, respectively. Considering the three carnivores at once, 20% of the area is highly suitable.
Suitable areas are fragmented, but interspersed with areas of less suitability value, without being isolated, and spatially distributed all along the Mountain range.
The results were validated with an independent data set and results suggest that the model produced an acceptable estimate of the areas effectively occupied by the carnivores. The comparison between suitability maps obtained with the two independent data sets showed that they were consistent, always reaching values of K-statistics > 0.5.
The development of human activities over the land poses problems of how to integrate land exploitation and biodiversity conservation. The outputs of the environmental modelling exercise were used for estimating the distribution of potential conflicts between the presence of carnivores and livestock husbandry practices. Results suggested an effective management would avoid the summer grazing of livestock in carnivore areas and the use of damage prevention measures. The actual effect of currently protected areas in the region was assessed and the need of an increased portion of protected land, particularly in Romania and Ukraine emerged after analysing the proportion of highly suitable areas for large carnivores under any kind of legal protection.</p
Patterns of the changes in distribution and numerical status of the wolf Canis lupus L. in Poland during the Quaternary are presented based on both published and unpublished palaeontological, archaeological and historical records. The wolf inhabited the territory of Poland as far back as the Upper Pleistocene. It was widely distributed in Poland during much of the Holocene, particularly from the Neolithic through Middle Ages. Its populations west of the Vistula were exterminated by the end of the 1860s. The species was very rare on the turn of the 19th century when its range was restricted to the east-central, easternmost and southeastern Poland. Early in the 1900s it became more numerous and subsequently reoccupied some portions of its former range. During the 1920s and 1930s it again declined in number and distribution. The species abruptly increased in number during the 1940s, with a peak of above 800 individuals at the mid-century. Early in the 1950s it was dispersed almost throughout Poland. A drastic decrease in the Polish wolf population began about the mid-1950s and continued until about the mid-1960s when it was estimated to be above 100 specimens only. During the late 1960s the wolf population was established between 100 and 200, but already in the early 1970s the species was nearly extinct in Poland (under 60 individuals in 1972). At that time its range was already diminished to the Sobibór Woods, Roztocze, and northeasternmost and southeasternmost Poland. Since about the mid-1970s it has generally been becoming more and more abundant up to about 900 individuals between 1984 and 1986. In consequence, its range has expanded westward once again, including most of the forested areas of Poland. — In: B. Bobek, K. Perzanowski and W.L. Regelin (eds.), Global trends in wildlife management (vol. 2, pp. 375–380). Świat Press, Cracow.
The wolf Canis lupus Linnaeus, 1758 has had "game species" status since the late 1970's. Hunting is permitted only by licensed hunters from August 1 to March 31. Official reports show that recently over 160 wolves were killed during every hunting season in Poland. Numbers and distribution of kills showed that the wolf has expanded its range since the late 1970's and occupied two core areas: northeastern and southeastern Poland.
Presently as many as 29 species and subspecies of deer alive today are threatened with extinction (recorded in the Red Data Book, IUCN). In some cases the number of individuals of certain species and subspecies is very limited. Breeding such small groups of animals has negative inbreeding effects and can even be responsible for the extinction of some species in the near future. One such example is the Schomburgk's deer (Ce.rvus schomburgki Blyth, 1963). The last Schomburgk's deer in captivity died in 1911 while the last Schomburgk's deer in the wild was shot in 1938 (Fradrich, 1980). Significant progress in domestic animal reproduction suggests that both semen freezing and embryo transfer can have practical application in protecting animals threatened with extinction. At the moment these methods are used in preventing the total loss of rare breeds of domestic animals. Not only have semen banks been established but also frozen embryo banks (Betteridge, 1977; Seidel, 1981). It would seem that to protect rare deer species these zootechnical methods could be used. The importance of freezing semen and embryo transfer was widely discussed at the 1st International Symposium on Artificial Breeding of Non-Domestic Animals in 1977 in London (Polge, 1978). The fact, that experiments with wild animals create many additional problems was taken into consideration. la deer, the usefulness of opening semen banks was stressed at deer workers meetings in Longview, USA — 1977 (Whitehead, 1978) and in Frcnt Royal, USA — 1982.
Age, sex and legbone marrow fat content of 90 red deer Cervus elaphus killed by wolves Canis lupus in winter in southeastern Poland were examined 1984-1988. The majority of kills were calves (44%) and hinds (40%); stags formed only 16%. The average age of hinds was 7.2 yr compared to 5.3 yr in stags. Animals older than 10 yr comprised only 13% of prey. Adults showed high femoral fat content (> 80%) throughout the winter (76% in early winter and 52% in late winter). Among calves femur marrow fat varied through the winter. In early winter 70% of calves had a high fat content (> 80%), though by late winter only 20% had such a high fat content, whilst 43% had low fat content (< 20%).
Wolf-ungulate interactions were studied in the pristine deciduous and mixed forests of the Bialowiea National Park in 1985–1989. The study period included two severe and two mild winters. The community of ungulates inhabiting Bialowiea National Park consisted of red deer Cervus elaphus, 55% of all ungulates; wild boar Sus scrofa, 42%; and roe deer Capreolus capreolus, moose Alces alces, and European bison Bison bonasus, about 1% each. The average size of red deer groups increased from 2.7 (SD 2.35) in spring and summer to 6.9 (SD 6.84) in autumn and winter. In winter the group size of red deer was positively correlated with the depth of snow cover and negatively correlated with the mean daily temperature. Average group size of wild boar did not change significantly between seasons; it was 6.8 (SD 5.16) in spring and summer and 5.7 (SD 4.67) in autumn and winter. Analysis of 144 wolf scats showed that wolves preyed selectively on red deer. In October–April, Cervidae (mostly red deer) constituted 91% of biomass consumed by wolves, while wild boar made up only 8%. In May–September deer formed 77% of prey biomass, and the share of wild boar increased to 22%. In all seasons of the year wolves selected juveniles from deer and boar populations: 61% of red deer and 94% of wild boar of determined age recovered from wolves' scats were young a National Park showed that predation was the predominant mortality factor for red deer (40 killed, 10 dead from causes other than predation) and roe deer (4 killed, none dead). Wild boar suffered most from severe winter conditions (8 killed, 56 dead). The percentage of ungulates that had died from undernutrition and starvation in the total mortality was proportional to the severity of winter.
Thirty-one stomach of voles shot in Bieszczady Mountains were collected during two consecutive hunting seasons. Deer tissues amounted to over 55% and wild boar tissues to almost 11% of stomachs content. It appears, that the carcasses of domestic livestock used as a bait consist a significant component of voles diet in winter (over 30% of stomachs content). The authors estimated, that the food requirements of an average wolf in winter, could be covered by consuming two red deer.
Zusammenfassung Durch Vergleich von 145 Wolfsschädeln mit 165 Schädeln großer Hunde fand der Verfasser sieben Merkmale am Schädel und ein Merkmal am Unterkiefer, welche die Unterscheidung der beiden Arten erlauben. Es folgt die Beschreibung der einzelnen Merkmale an Hand von Zeichnungen und einer Tabelle, die die prozentuale Häufigkeit und die Signifikanzdaten enthält. Die hohen Ziffern dieser Berechnungen berechtigen zu der Schlußfolgerung, daß das gleichzeitige Vorkommen von sechs der erwähnten Merkmale an einem Schädel ein ausreichender Beweis für seine Zugehörigkeit bedeutet. Wegen der hohen Polymorphie der beiden Arten kann die angewandte Methode zoogeographisch beschränkt sein und muß an außereuropäischem Material überprüft werden. Die vorliegende Abhandlung enthält die “subjektive” Merkmalserfassung, ihre Objektivierung ist in Bearbeitung.