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Diet of badgers living in a deciduous forest in Hungary
Mammalian Biology Zeitschrift fuÈr SaÈugetierkunde www.elsevier.de/mammbiol
Short communication
Diet of badgers living in a deciduous forest in Hungary By J. LANSZKI Ecological Research Group, University of KaposvaÂr, KaposvaÂr, Hungary Receipt of Ms. 13. 10. 2003 Acceptance of Ms. 07. 01. 2004
Key words: Meles meles, feeding habits, amphibians, earthworms The Eurasian badger Meles meles (L.) is a widely occurring predator. Its population density has shown a wide range and a general increase in Europe since the 1980s (Griffiths 1993; Neal and Cheeseman 1991; Mitchell-Jones et al. 1999; Kowalczyk et al. 2003). The increasing badger population (in Hungary from 1987: 2.0 ind./ 1 000 ha to 2001: 3.2 ind./1 000 ha; Heltai 2002) can cause problems in agriculture (e. g. Roper et al. 1995; Heltai 2002). The feeding habits of the badger have been studied in areas of agricultural land and natural habitats with different environmental conditions (Goszczynski et al. 2000). In northern latitudes of Europe the most important food taxa for the badger are earthworms and vertebrates, but plant matter and insects become more significant in its diet the further south it lives. In forests in temperate zones of Europe the main food for badgers are earthworms. This biogeographical pattern of feeding habits can be explained by the variation in abundance and availability of earthworms (Goszczynski et al. 2000). The main food (earthworms) and the mean annual temperature are crucial factors, influencing the density of badgers in the European temperate and 1616-5047/04/69/05-354 $ 30.00/0.
boreal zones (Kowalczyk et al. 2003). Little is known about the feeding habits of badgers living in deciduous forest in the Pannonia region; a seasonal investigation has been performed only in an agricultural environment (Lanszki et al. 1999). The aim of this study was to investigate the seasonal feeding habits of badgers living in deciduous forest surrounding ponds. The study area is situated in south-western Hungary (46°24¢ N, 17°27¢ E) in the Boronka Nature Conservation Area (or BNCA, 78.3 km2). Determining factors in the BNCA district are areas of associated forest, which surround six dammed valley fish ponds established a hundred years ago. The ponds situated in the designated study area are surrounded by willow marshes and alder swamps, and also extensive forests of hornbeam and oak. (More information on the area is given by Lanszki et al. 2001). The climate is continental: during the study (December 1996 to February 2001) the mean (± s. e.) winter temperature is 1.7 ± 0.6 °C, duration of snow covering 33Ô12 days, and snow depth 65 ± 11 mm; summer temperature 20.7 ± 0.5 °C, mean annual precipitation 711 ± 104 mm. Mamm. biol. 69 (2004) 5´ 354±358
Diet of badgers living in a deciduous forest in Hungary The feeding habits of the badger were investigated by analysis of scats. Scats were collected twice a month, mainly at two badger setts from latrines and partially on a standard route (approx. 5 000 m long), where badger footprints occurred. A total of 156 badger scats were analysed by a standard procedure (Jedrzejewska and Jedrzejewski 1998). Prey determination was performed by microscope on the basis of feather, bone, dentition and hair characteristics (e. g. MaÈrz 1972; Teerink 1991; Brown et al. 1993; and own collection). Diet composition was expressed in two ways: as the relative frequency of occurrence and as an estimate of the percentage fresh weight (biomass) of food consumed. The presence of earthworms in the scats was determined by microscopic identification of chaetae, and the biomass of earthworms was calculated following Kruuk (1989) and Jedrzejewska and Jedrzejewski (1998). For the quantitative estimation, the scats were washed through filters (0.5 mm pore diameter) and dried, after which the remains of items were separated. All dry prey or plant remains were weighed and multiplied by coefficients of digestibility described for the badger (Jedrzejewska and Jedrzejewski 1998; Goszczynski et al. 2000). Trophic niche breadth was calculated in acP cordance with Levins: B = 1/ pi2, where pi is the relative frequency of a given taxon (Krebs 1989). The following food taxa were used in these calculations: voles, mice, other rodents and insectivores, medium-sized mammals and carcasses of ungulates, birds, other vertebrates, invertebrates and plant matter. Distribution of prey body weight was determined in accordance with Clevenger (1993), to create categories (< 15 g; 15±50 g; 51±100 g; 101±300 g; > 300 g). Prey classification on the basis of characteristic zonation was performed according to published data from Gittleman (1985) on predators (1: terrestrial; 2: arboreal and 3: aquatic). The SPSS 7.5 statistics program (Green et al. 1997) was used for the processing of the data obtained. The proportion according to biomass of small mammals consumed by the badger
355
(Tab. 1, Fig. 1) characteristically varied around a low value; only in autumn did it show a small rise. Remains of pine marten Martes martes (in summer) and dog Canis familiaris (in autumn) occurring in the diet were, as with the Cervidae (Cervus elaphus and Capreolus capreolus) in winter and spring, probably derived from carrion consumption, the significance of which was found to be insignificantly small. The badger consumed birds' eggs in spring and small Passeriformes throughout the year except in winter, but total bird biomass in its diet proved to be low. From spring to autumn, reptiles and amphibians were the dominant food resource. Amphibians (mainly frogs Rana sp.) occurring in the diet were found to account for a much more significant percentage than reptiles. Fish occurred (probably as carrion) occasionally in the diet only in winter and autumn. Invertebrates (mainly earthworms Lumbricidae) played the major role in the diet only in winter; after this their significance decreased gradually until autumn. The proportion of plant matter (mainly maize Zea mays, originated from feed distributors installed for ungulates) proved low in winter and spring. This subsequently rose substantially in summer, and in autumn consumption of forest fruits actually became of secondary significance to the badger. The minimun known numbers of food taxa in the diet were 47 prey and 11 plant taxa. The trophic niche was narrow (Tab. 1). The diet composition of the badger showed significant differences among seasons (v2 = 5 027.5, df = 24, P < 0.001). Prey species weighing less than 15 g (mainly invertebrates) occurred the most frequently (69%) in the diet of the badger, 25% being classifiable in the 15±50 g weight category (consisting mostly of amphibians and small mammals). The heavier prey taxa (above 50 g, e. g., carrions, reptiles, fish) played no substantial role (6%). The determinant majority (83%) of the diet was accounted for terrestrial species. The proportion accounted for by species associated with wetlands proved considerable (13%), while the role of arboreal species was found to be insignificant (4%).
356
J. LANSZKI
Table 1. Diet composition of badgers living in deciduous forests surrounding ponds in the BNCA. %O = percentage of occurrence; %B = percentage of biomass consumed; + = under 0.05%. Winter Item Bank vole Clethrionomys glareolus Common vole Microtus arvalis Field mouse Apodemus sp. Other Rodentia Shrews Soricidae Carrion Small bird Passeriformes sp. Bird egg Swamp turtle Emys orbicularis egg Grass snake Natrix natrix Undetermined Reptilia Common toad Bufo bufo Edible frog Rana esculenta Common spadefoot Pelobates fuscus Tree frog Hyla arborea Undetermined Bombina sp. Undetermined Rana sp. Fish Carabidae beetles Dung beetle Geotrupes sp. Other Coleoptera Wasp Vespidae sp. Other invertebrates Earthworm Lumbricidae Wild raspberry Rubus sp. Strawberry Fragaria sp. Pear Pyrus sp. Other fruits Maize Zea mays Other plant matter No. of scats analysed Items per scat Niche breadth (B index)
%O
Spring
%B
%O
Summer
%B
1.0
0.5
1.1
1.7
1.0 1.0
0.5 0.3
8.3
0.7
0.2 0.2 0.7 0.4 0.9 0.5
0.1 + 0.8 + 0.4 +
0.2 1.3 5.3 11.7 3.0 3.2 0.5 1.1
0.1 0.5 16.1 38.8 10.6 6.7 0.4 2.9
7.7 24.4 6.6 0.4 9.2 16.4
+ 2.2 + + 0.1 16.1
0.2
0.6
0.2 2.1 2.5
0.1 1.3 0.3
1.0
+
3.1 9.3
7.9 26.3
1.0
+
4.1 1.0 2.1 14.5
8.8 0.1 + 1.2
3.1 17.5
+ 47.9
13.4 19.2
5.1 0.6
25 3.9 3.50 2.32
On the basis of this investigation it can be established that the badger consumed a wide spectrum of food; there were large differences in the diet composition among seasons, similar to other studies in Europe (Neal and Cheeseman 1991; Goszczynski et al. 2000). In temperate zones numerous food resources are available during each season; this seems to influences prey choice greatly. Badgers have adapted well to the food availability, as is indicated by the abundance of food taxa in their diet. In contrast to its varied food, it utilized only a nar-
96 4.6 2.04 1.63
%O
%B
0.8 0.8
1.0 0.3
0.8 0.8 0.8
0.5 + 0.2
3.8
11.0
1.5 1.5 6.8 1.5 2.3 0.8 3.0
2.0 13.3 33.2 7.6 5.6 1.3 1.3
15.9 18.0 6.8 4.5 5.2 11.3 4.5 2.3 3.0
0.5 0.9 0.1 0.2 + 3.5 2.3 8.5 5.6
3.8
0.6
19 7.0 2.26 1.67
Autumn %O
%B
1.6
7.4
3.2 4.8
1.5 3.3
1.6 1.6
0.1 +
6.5 6.5
35.6 14.3
3.2
7.9
1.6 11.3 9.7
0.1 2.2
1.6 3.2 19.4 1.6
+ + 1.4 4.6
6.5 3.2 4.8 8.0
6.6 9.8 4.2 0.7
16 3.9 3.34 2.43
row spectrum of the food resources available. Depending on habitat opportunities the badger was able to switch over to specialize on earthworms or amphibians. Due to this, the trophic niche was narrow (contrary to experience in the Mediterranean area Ciampalini and Lovari 1985). Thus, two thirds of the prey consumed was very small in size. These results are in accordance with the food specialist hypothesis (Kruuk 1989; Roper and LuÈps 1995). However, badger living in the forest utilized seasonally dependent food resources; hence,
Diet of badgers living in a deciduous forest in Hungary
357
References
Fig. 1. Seasonal changes in the diet of badgers (percentage of biomass consumed) in the BNCA.
they were characteristically opportunistic food generalists (Roper and LuÈps 1995). The badger succeeded in exploiting the limited range of food resources available. Studies have shown resource exploitation to differ in forest and agricultural habitats (Kruuk 1989; Neal and Cheeseman 1991; Goszczynski et al. 2000). The badger possesses a great capacity for adapting, which is also indicated by its prevalence of occurrence, i. e., the fact that it is becoming increasingly frequent (in Hungary: Heltai 2002). In winter, as results from observations performed in northern Europe have shown, the most significant species occurring in its diet proved to be earthworms; subsequently, in contrast to the diet of the badger in northern and central European forests and agricultural land and in Mediterranean areas (Neal and Cheeseman 1991; Goszczynski et al. 2000), amphibians (partially reptiles) played the major role in its diet from spring to autumn. In agricultural environments in Hungary the consumption of plants was more frequent (Lanszki et al. 1999), compared with the badgers living in the forest. The nearest agricultural areas are at least 1.5±2.0 km away and the closest villages 4±5 km distant from the badger setts examined in this study. The consumption of food items associated with agricultural and inhabited areas, e. g., common vole and domestic animals (dog), was found to occur only very rarely.
Acknowledgements The author is grateful to D. L. Moss for her help with the English manuscript. This work was supported by the OTKA (F 23057) and the Bolyai scholarship.
Brown, R.; Ferguson, J.; Lawrence, M.; Lees, D. (1993): Federn, Spuren und Zeichen der VoÈgel Europas: Ein FeldfuÈhrer. Wiesbaden: Aula-Verlag. Ciampalini, B.; Lovari, S. (1985): Food habits and trophic niche overlap of the badger (Meles meles L.) and the red fox (Vulpes vulpes L.) in a Mediterranean coastal area. Z. SaÈugetierkunde 50, 226±234. Clevenger, A. P. (1993): Pine marten (Martes martes LinneÂ, 1758) comparative feeding ecology in an island and mainland population of Spain. Z. SaÈugetierkunde 58, 212±224. Gittleman, J. L. (1985): Carnivore body size: ecological and taxonomic correlates. Oecologia 67, 540±554. Goszczynski, J.; Jedzejewska, B.; Jedrzejewski, W. (2000): Diet composition of badger (Meles meles) in a pristine forest and rural habitats of Poland compared to other European populations. J. Zool. (London) 250, 495±505. Green, S. B.; Salkind, N. J.; Akey, T. M. (1997): Using SPSS for Windows: analyzing and understanding data. New Jersey: Prentice Hall. Griffiths, H. I. (1993): Badger game-bag data estimates of badger (Meles meles) population sizes in Europe. Small Carnivore Conservation 9, 9±10. Heltai, M. (2002): The status and distribution of mammal predators in Hungary. Diss. thesis, St. Stephen University, GoÈdoÈllo², Hungary. Jedrzejewska, B.; Jedrzejewski, W. (1998): Predation in Vertebrate Communities. The Bialowieza Primeval Forest as a case study. Berlin Heidelberg, New York: Springer-Verlag. Kowalczyk, R.; Zalewski, A.; Jedrzejewska, B.; Jedrzejewski, W. (2003): Spatial organization and demography of badgers (Meles meles) in Bialowieza Primeval Forest, Poland, and the influence of earthworms on badger densities in Europe. Can. J. Zool. 81, 74±87. Krebs, C. J. (1989): Ecological Methodology. New York: Harper Collins Publ. Pp. 372±374. Kruuk, H. (1989): The Social Badger. Oxford: Oxford Univ. Press. Lanszki, J.; KoÈrmendi, S.; Hancz, C.; Zalewski, A. (1999): Feeding habits and trophic niche overlap in a Carnivora community of Hungary. Acta Theriol. 44, 429±442. Lanszki, J.; KoÈrmendi, S.; Hancz, C.; Martin, T. G. (2001): Examination of some factors affecting selection of fish prey by otters (Lutra lutra) living by eutrophic fish ponds. J. Zool. (London) 255, 97±103.
358
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MaÈrz, R. (1972): GewoÈll- und Rupfungskunde. Berlin: Akademie Verlag. Mitchell-Jones, A. J.; Amori, G.; Bogdanowicz, W.; Krystufek, B.; Reijnders, P. J. H.; Stubbe, M.; Thissen, J. B. M.; VohralõÂk, V.; Zima, J. (1999): The Atlas of European Mammals. London: Academic Press. Neal, E. G.; Cheeseman, C. L. (1991): Badger. In: The Handbook of British Mammals. Ed. by G. B. Corbet and S. Harris. Oxford: Blackwell Scientific Publ. Pp. 415±423. Roper, T. J.; Findlay, S. R.; LuÈps, P.; Shepherdson, D. J. (1995): Damage by badgers Meles meles to wheat Triticum vulgare and barley
Hordeum sativum crops. J. Appl. Ecol. 32, 720±726. Roper, T. J.; LuÈps, P. (1995): Diet of badgers (Meles meles) in central Switzerland: an analysis of stomach contents. Z. SaÈugetierkunde 60, 9±19. Teerink, B. J. (1991): Hair of West-European mammals. Cambridge: Cambridge Univ. Press. Author's address: JoÂzsef Lanszki, Ecological Research Group, University of KaposvaÂr, P. O. Box 16, 7401 KaposvaÂr, Hungary (e-mail: [email protected])
Short communication
Diet of badgers living in a deciduous forest in Hungary By J. LANSZKI Ecological Research Group, University of KaposvaÂr, KaposvaÂr, Hungary Receipt of Ms. 13. 10. 2003 Acceptance of Ms. 07. 01. 2004
Key words: Meles meles, feeding habits, amphibians, earthworms The Eurasian badger Meles meles (L.) is a widely occurring predator. Its population density has shown a wide range and a general increase in Europe since the 1980s (Griffiths 1993; Neal and Cheeseman 1991; Mitchell-Jones et al. 1999; Kowalczyk et al. 2003). The increasing badger population (in Hungary from 1987: 2.0 ind./ 1 000 ha to 2001: 3.2 ind./1 000 ha; Heltai 2002) can cause problems in agriculture (e. g. Roper et al. 1995; Heltai 2002). The feeding habits of the badger have been studied in areas of agricultural land and natural habitats with different environmental conditions (Goszczynski et al. 2000). In northern latitudes of Europe the most important food taxa for the badger are earthworms and vertebrates, but plant matter and insects become more significant in its diet the further south it lives. In forests in temperate zones of Europe the main food for badgers are earthworms. This biogeographical pattern of feeding habits can be explained by the variation in abundance and availability of earthworms (Goszczynski et al. 2000). The main food (earthworms) and the mean annual temperature are crucial factors, influencing the density of badgers in the European temperate and 1616-5047/04/69/05-354 $ 30.00/0.
boreal zones (Kowalczyk et al. 2003). Little is known about the feeding habits of badgers living in deciduous forest in the Pannonia region; a seasonal investigation has been performed only in an agricultural environment (Lanszki et al. 1999). The aim of this study was to investigate the seasonal feeding habits of badgers living in deciduous forest surrounding ponds. The study area is situated in south-western Hungary (46°24¢ N, 17°27¢ E) in the Boronka Nature Conservation Area (or BNCA, 78.3 km2). Determining factors in the BNCA district are areas of associated forest, which surround six dammed valley fish ponds established a hundred years ago. The ponds situated in the designated study area are surrounded by willow marshes and alder swamps, and also extensive forests of hornbeam and oak. (More information on the area is given by Lanszki et al. 2001). The climate is continental: during the study (December 1996 to February 2001) the mean (± s. e.) winter temperature is 1.7 ± 0.6 °C, duration of snow covering 33Ô12 days, and snow depth 65 ± 11 mm; summer temperature 20.7 ± 0.5 °C, mean annual precipitation 711 ± 104 mm. Mamm. biol. 69 (2004) 5´ 354±358
Diet of badgers living in a deciduous forest in Hungary The feeding habits of the badger were investigated by analysis of scats. Scats were collected twice a month, mainly at two badger setts from latrines and partially on a standard route (approx. 5 000 m long), where badger footprints occurred. A total of 156 badger scats were analysed by a standard procedure (Jedrzejewska and Jedrzejewski 1998). Prey determination was performed by microscope on the basis of feather, bone, dentition and hair characteristics (e. g. MaÈrz 1972; Teerink 1991; Brown et al. 1993; and own collection). Diet composition was expressed in two ways: as the relative frequency of occurrence and as an estimate of the percentage fresh weight (biomass) of food consumed. The presence of earthworms in the scats was determined by microscopic identification of chaetae, and the biomass of earthworms was calculated following Kruuk (1989) and Jedrzejewska and Jedrzejewski (1998). For the quantitative estimation, the scats were washed through filters (0.5 mm pore diameter) and dried, after which the remains of items were separated. All dry prey or plant remains were weighed and multiplied by coefficients of digestibility described for the badger (Jedrzejewska and Jedrzejewski 1998; Goszczynski et al. 2000). Trophic niche breadth was calculated in acP cordance with Levins: B = 1/ pi2, where pi is the relative frequency of a given taxon (Krebs 1989). The following food taxa were used in these calculations: voles, mice, other rodents and insectivores, medium-sized mammals and carcasses of ungulates, birds, other vertebrates, invertebrates and plant matter. Distribution of prey body weight was determined in accordance with Clevenger (1993), to create categories (< 15 g; 15±50 g; 51±100 g; 101±300 g; > 300 g). Prey classification on the basis of characteristic zonation was performed according to published data from Gittleman (1985) on predators (1: terrestrial; 2: arboreal and 3: aquatic). The SPSS 7.5 statistics program (Green et al. 1997) was used for the processing of the data obtained. The proportion according to biomass of small mammals consumed by the badger
355
(Tab. 1, Fig. 1) characteristically varied around a low value; only in autumn did it show a small rise. Remains of pine marten Martes martes (in summer) and dog Canis familiaris (in autumn) occurring in the diet were, as with the Cervidae (Cervus elaphus and Capreolus capreolus) in winter and spring, probably derived from carrion consumption, the significance of which was found to be insignificantly small. The badger consumed birds' eggs in spring and small Passeriformes throughout the year except in winter, but total bird biomass in its diet proved to be low. From spring to autumn, reptiles and amphibians were the dominant food resource. Amphibians (mainly frogs Rana sp.) occurring in the diet were found to account for a much more significant percentage than reptiles. Fish occurred (probably as carrion) occasionally in the diet only in winter and autumn. Invertebrates (mainly earthworms Lumbricidae) played the major role in the diet only in winter; after this their significance decreased gradually until autumn. The proportion of plant matter (mainly maize Zea mays, originated from feed distributors installed for ungulates) proved low in winter and spring. This subsequently rose substantially in summer, and in autumn consumption of forest fruits actually became of secondary significance to the badger. The minimun known numbers of food taxa in the diet were 47 prey and 11 plant taxa. The trophic niche was narrow (Tab. 1). The diet composition of the badger showed significant differences among seasons (v2 = 5 027.5, df = 24, P < 0.001). Prey species weighing less than 15 g (mainly invertebrates) occurred the most frequently (69%) in the diet of the badger, 25% being classifiable in the 15±50 g weight category (consisting mostly of amphibians and small mammals). The heavier prey taxa (above 50 g, e. g., carrions, reptiles, fish) played no substantial role (6%). The determinant majority (83%) of the diet was accounted for terrestrial species. The proportion accounted for by species associated with wetlands proved considerable (13%), while the role of arboreal species was found to be insignificant (4%).
356
J. LANSZKI
Table 1. Diet composition of badgers living in deciduous forests surrounding ponds in the BNCA. %O = percentage of occurrence; %B = percentage of biomass consumed; + = under 0.05%. Winter Item Bank vole Clethrionomys glareolus Common vole Microtus arvalis Field mouse Apodemus sp. Other Rodentia Shrews Soricidae Carrion Small bird Passeriformes sp. Bird egg Swamp turtle Emys orbicularis egg Grass snake Natrix natrix Undetermined Reptilia Common toad Bufo bufo Edible frog Rana esculenta Common spadefoot Pelobates fuscus Tree frog Hyla arborea Undetermined Bombina sp. Undetermined Rana sp. Fish Carabidae beetles Dung beetle Geotrupes sp. Other Coleoptera Wasp Vespidae sp. Other invertebrates Earthworm Lumbricidae Wild raspberry Rubus sp. Strawberry Fragaria sp. Pear Pyrus sp. Other fruits Maize Zea mays Other plant matter No. of scats analysed Items per scat Niche breadth (B index)
%O
Spring
%B
%O
Summer
%B
1.0
0.5
1.1
1.7
1.0 1.0
0.5 0.3
8.3
0.7
0.2 0.2 0.7 0.4 0.9 0.5
0.1 + 0.8 + 0.4 +
0.2 1.3 5.3 11.7 3.0 3.2 0.5 1.1
0.1 0.5 16.1 38.8 10.6 6.7 0.4 2.9
7.7 24.4 6.6 0.4 9.2 16.4
+ 2.2 + + 0.1 16.1
0.2
0.6
0.2 2.1 2.5
0.1 1.3 0.3
1.0
+
3.1 9.3
7.9 26.3
1.0
+
4.1 1.0 2.1 14.5
8.8 0.1 + 1.2
3.1 17.5
+ 47.9
13.4 19.2
5.1 0.6
25 3.9 3.50 2.32
On the basis of this investigation it can be established that the badger consumed a wide spectrum of food; there were large differences in the diet composition among seasons, similar to other studies in Europe (Neal and Cheeseman 1991; Goszczynski et al. 2000). In temperate zones numerous food resources are available during each season; this seems to influences prey choice greatly. Badgers have adapted well to the food availability, as is indicated by the abundance of food taxa in their diet. In contrast to its varied food, it utilized only a nar-
96 4.6 2.04 1.63
%O
%B
0.8 0.8
1.0 0.3
0.8 0.8 0.8
0.5 + 0.2
3.8
11.0
1.5 1.5 6.8 1.5 2.3 0.8 3.0
2.0 13.3 33.2 7.6 5.6 1.3 1.3
15.9 18.0 6.8 4.5 5.2 11.3 4.5 2.3 3.0
0.5 0.9 0.1 0.2 + 3.5 2.3 8.5 5.6
3.8
0.6
19 7.0 2.26 1.67
Autumn %O
%B
1.6
7.4
3.2 4.8
1.5 3.3
1.6 1.6
0.1 +
6.5 6.5
35.6 14.3
3.2
7.9
1.6 11.3 9.7
0.1 2.2
1.6 3.2 19.4 1.6
+ + 1.4 4.6
6.5 3.2 4.8 8.0
6.6 9.8 4.2 0.7
16 3.9 3.34 2.43
row spectrum of the food resources available. Depending on habitat opportunities the badger was able to switch over to specialize on earthworms or amphibians. Due to this, the trophic niche was narrow (contrary to experience in the Mediterranean area Ciampalini and Lovari 1985). Thus, two thirds of the prey consumed was very small in size. These results are in accordance with the food specialist hypothesis (Kruuk 1989; Roper and LuÈps 1995). However, badger living in the forest utilized seasonally dependent food resources; hence,
Diet of badgers living in a deciduous forest in Hungary
357
References
Fig. 1. Seasonal changes in the diet of badgers (percentage of biomass consumed) in the BNCA.
they were characteristically opportunistic food generalists (Roper and LuÈps 1995). The badger succeeded in exploiting the limited range of food resources available. Studies have shown resource exploitation to differ in forest and agricultural habitats (Kruuk 1989; Neal and Cheeseman 1991; Goszczynski et al. 2000). The badger possesses a great capacity for adapting, which is also indicated by its prevalence of occurrence, i. e., the fact that it is becoming increasingly frequent (in Hungary: Heltai 2002). In winter, as results from observations performed in northern Europe have shown, the most significant species occurring in its diet proved to be earthworms; subsequently, in contrast to the diet of the badger in northern and central European forests and agricultural land and in Mediterranean areas (Neal and Cheeseman 1991; Goszczynski et al. 2000), amphibians (partially reptiles) played the major role in its diet from spring to autumn. In agricultural environments in Hungary the consumption of plants was more frequent (Lanszki et al. 1999), compared with the badgers living in the forest. The nearest agricultural areas are at least 1.5±2.0 km away and the closest villages 4±5 km distant from the badger setts examined in this study. The consumption of food items associated with agricultural and inhabited areas, e. g., common vole and domestic animals (dog), was found to occur only very rarely.
Acknowledgements The author is grateful to D. L. Moss for her help with the English manuscript. This work was supported by the OTKA (F 23057) and the Bolyai scholarship.
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