NORTH-WESTERN JOURNAL OF ZOOLOGY 12 (1): 184-187 ©NwjZ, Oradea, Romania, 2016 Article No.: e152701 http://biozoojournals.ro/nwjz/index.html

First observations of occasional carnivory in Tatra , Slovakia

Zuzana BALLOVÁ1,*, Milan BALLO2 and Ladislav HOLKO3

1. Department of Geobotany, Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 9, 845 23, Bratislava, Slovakia. 2. Správa TANAP, Hodžova 11, 031 01, Liptovský Mikuláš, Slovakia. 3. Institute of Hydrology, Slovak Academy of Sciences, Ondrašovská 16, 031 05, Liptovský Mikuláš, Slovakia. *Corresponding author, Z. Ballová, E-mail: [email protected]

Received: 09. January 2015 / Accepted: 29. May 2015 / Available online: 30. May 2016 / Printed: June 2016

Abstract. Marmots are primarily herbivores, a wide variety of plants that grow in alpine and montane meadows which represent their habitats. Upon emerging from in late spring, their diet consists largely of grasses, herbs, and roots. Occasional carnivorous behaviour has been confirmed in the Alpine subspecies of the Tatra Mountains (Marmota marmota latirostris), Slovakia for the first time in 2014. Our experiment shows that under unfavourable conditions (prolonged snow cover and temperatures below freezing) marmots accept meat that they do not eat when plants are an available food resource. The Tatra marmots consumed provided hare carcasses and fly larvae, however, plant food is clearly preferred over foods when available.

Key words: environmental factors, animal food, Carpathians, endemic species, Marmota marmota latirostris, Tatra Mts.

Marmots (Marmota spp.) are primarily herbivo- small birds, and small by marmots has rous . Upon emerging from hibernation in been associated with harsh environmental condi- late spring, their diet consists largely of grasses, tions (Bibikov 1996). However, sometimes the rea- herbs, and roots. Several studies report that mar- son of this behaviour is not clear. In the Görlitz mots eat a wide variety of plants (Bibikov 1996, zoo where alpine marmots and vultures were kept Barrio et al. 2013, Armitage 2014). All species of together, the marmots fed on the carcasses given marmots are characterized by the selective con- to vultures and, even defended them (Artl & Ge- sumption of various plants in a particular region, bauer 1996). as well as season, rather than by preference for The Tatra marmot (M. marmota latirostris) is an certain species of plants. Marmots readily digest endemic subspecies of the alpine marmot that oc- only lush herbs and those parts (shoots, leaves, curs in the Tatra Mountains (the Western Carpa- and flower heads) that contain assimilable protein thian Mountains, boundary between northern cen- and carbohydrates (Bibikov 1996). Their food tral Slovakia and Poland) and inhabits open areas choice may be based on the protein (Frase & Ar- in the subalpine and alpine zones. Within its na- mitage 1989) or the essential fatty acid content tive range, it mainly forages in tall-stem grass- (Florant 1998, Bruns et al. 1999). Wood marmots lands and tall-herb plant communities from the al- sometimes utilize trees (bark, branches, buds, and liances Calamagrostion villosae and Trisetion fusci leaves), like yellow-bellied marmots (M. flaviven- (Ballová & Šibík 2015). Tatra marmots are consid- tris) use trees as an alternative source of forage ered generalist herbivores that selectively feed on during years with high spring snow pack (Olson grasses and herbs (Chovancová & Šoltésová, 1988, et al. 2003). Woodchucks (M. monax) may climb Karč, 2006). Until now, meat consumption had not trees and feed on their leaves (Swihart & Picone been reported for them. Our previous observa- 1991). High mountain and tundra marmot species' tions of small mammals’ hair in marmot excre- habitats do not contain trees, thus they are de- ments suggested that Tatra marmots may occa- prived of this food resource. sionally consume meat (Ballová 2013). This behav- As has already been mentioned from different iour might be expected based on observations on mountain ranges in Asia (Kapitonov 1960), Europe the nominal subspecies in the (Ferrari et al. (Ferrari et al. 2012) and North America (Barash 2012). Since there were doubts about such behav- 1973, Armitage et al. 1979), alpine, subalpine and iour, we decided to experimentally test marmot mountain tundra marmot species may be occa- food preferences in spring, after their emergence sionally carnivorous. Consumption of animal from hibernation. We hypothesized that patterns food, including locusts, molluscs, earthworms, of food choice are based on the climatic and eco- First observations of occasional carnivory in Tatra marmots 185 logical conditions of the area and expected that mots rejected the meat, even though the air tem- marmots will consume meat under harsh envi- perature was below zero (–6°C on April 15). The ronmental conditions. The experiment was con- meat was not consumed by marmots, neither on ducted within the monitoring of protected species days with negative air temperature (April 15 – in cooperation with the Tatra National Park. April 17) nor on warm days during the last week of April with the mean daily temperatures about In spring 2014, after marmots had emerged from hiberna- +4°C. On May 8th, we placed the second hare on tion (April 15 and April 29), we placed photo traps the . The photo trap after snowfalls in the (Bushnell Trophy Cam HD Essential with high-quality middle of May (May 12 – May 16, about 0.4 m – full colour resolution and day/night autosensor) above the entrances of two hibernation in the Jalovecká 1.5 m snow cover) recorded marmots consuming dolina Valley (the Western Tatra Mts., Western Carpathi- meat at the burrow on May 18. The air tempera- ans, Slovakia, coordinates from 49°13'08.43" N and ture was above zero. During snowfall (May 12 – 19°40'01.12" E to 49°10'10.82" N and 20°02'12.86" E). The May 16) and rainfall (May 17) the marmots were cameras were motion triggered with sensor reaches out to in the burrow. We also observed that marmots ate 60 feet or beyond and automatically captured images at fly larvae at the carcass in preference to meat. one second intervals. Study localities were chosen to rep- The marmot settlement in the Hlboká dolina resent two marmot sites with different environmental conditions (Blahout 1960, 1971) – a marmot colony with Valley (the valley site) is located in a glacial gorge the main burrow system situated in a valley (Hlboká with a long duration of snow cover. Marmots in dolina Valley, hibernaculum at 1886 m a.s.l.) and a mar- the glacial gorge hibernated longer than the mar- mot colony with the main burrow system on a slope (Mt. mots on Mt. Grapy. Warmer period with rainfall Grapy, hibernaculum at 1915 m a.s.l.). On hibernation between April 22nd and April 28th was followed burrow mounds in each colony we placed a fresh com- by marmot emergence on the 29th of April. De- mon hare (Lepus europaeus) roadkill, on two different days spite the warming, snow cover did not melt in the (April 15 on the Mt. Grapy and April 29 in the Hlboká dolina Valley, because marmots had not emerged from Hlboká dolina Valley which is surrounded by the hibernation on the same day) and recorded whether the sides of glacial gorge and rocky walls. On April meat was taken by any animal. The distance between 30, the meat was consumed by marmots at this lo- cameras and hares was about 0.7 m. The hares were fixed cality, which was still snow-covered (Fig. 1). The with steel wire to prevent them from being carried away air temperature at the time was about +1°C. by golden eagles (Aquila chrysaetos) or any other preda- tors. A simple test for food preference was conducted by placing rock salt and fresh dandelion leaves from lower elevations. The items were placed under each hare carcass to prevent their removal by the wind or chamois (Rupi- capra rupicapra tatrica). Precipitation and air temperature data from two permanent meteorological stations in the Jalovecká dolina Valley were used to determine meteorological conditions during the experiment (the stations are located at alti- tudes 1500 and 1900 m a.s.l.). Snowfall was assumed on the days with precipitation when mean daily air tempera- ture was below the freezing point. The weather was rela- tively warm and wet from the last week of April. Extreme precipitation occurred on May 15 and May 16. It resulted in floods with the recurrence interval of approximately Figure 1. Adult marmot feeding on the body of dead hare 10–20 years (Danko, 2014). Meteorological data indicate in the Hlboká dolina Valley. that snow pack accumulated at both study sites in the middle of May. Snow accumulation was confirmed by the photographs and also our measurements on selected Despite the fact that other (Aquila dates. chrysaetos, Chionomys nivalis, Martes martes) visited Marmots at Mt. Grapy (the slope site) emerged the hares, cameras recorded that marmots ate al- from hibernation on April 15. Snow cover melted most the entire hare at both sites. Only bones and earlier and more rapidly on the Mt. Grapy than in fur scraps were left. Eagles unsuccessfully at- the Hlboká dolina Valley due to southern orienta- tempted to take the carcasses. Pine martens tion of the slopes. The slopes were almost without (Martes martes) and Alpine voles (Chionomys snow cover when we brought the first hare. Mar- nivalis) were very shy and after short inspection 186 Z. Ballová et al. they left the carcasses. vourable weather conditions resulting in a short- Marmots in both settlements ate the dandelion age of vegetable food. Development of snow cover leaves first and then they accepted the meat. This during the warmer part of the year (after marmot simple preference test showed that meat was not a emergence from hibernation) and long snow sea- preferred food item, but it was taken when the sons seem to be the main reasons causing the oc- vegetable food was in short supply. The rock salt casional carnivory and eventually cannibalism. was licked by the marmots at both sites simulta- Our experiment has shown that meat is not a neously with the meat consumption. Salt licking preferred food item for Tatra marmots, but it will frequently occurs in marmots, woodchucks (M. be eaten in the absence of plants. The observed monax) use small mineral licks, and lick road sur- marmots did not eat meat if they had dandelion faces for residues of winter-applied NaCl (Weeks leaves available. On the basis of our research we & Kirkpatrick, 1978), and Armitage (2000) ob- suggest that similar behaviours are generally served yellow-bellied marmots (M. flaviventris) prevalent in most marmot species and depend on licking the mud surface at salt licks. food accessibility. Our observations confirm this The Tatra marmots in our test behaved very behaviour for the first time in the Carpathian Mts., similarly to Olympic marmots (M. olympus) which for an endemic marmot subspecies. Barash (1973) observed in North America. The writer placed beef hearts at colonies in the spring- time, which were eaten by the marmots when Acknowledgments. This study was supported by the snow covered the ground, but ignored them after Tatra National Park and the grant agency VEGA under the snow melt. Grant 2/0090/2012. The experiment was conducted in Hibernating mammals, like the Tatra and Al- cooperation with the Tatra National Park within the pine marmots, may consume most or all their monitoring of protected species authorized by the reserves during particularly harsh winter so that Ministry of Environment (No. 3847/2014–2.3). Collection at the vernal emergence they rely completely on of meteorological data was supported by the grant agency new food intake. But, especially at high altitudes, a VEGA under Grant 2/0042/11. For English proofreading and for their suggestions for this study, we are indebted long lasting snow cover can delay the availability to Dr. Jozef Šibík and Dr. Július Rusko. of food resources (Ferrari et al. 2012). Under unfa- vourable conditions (mainly in the springtime), animal food was found in stomachs of some mar- mot species. According to Kapitonov (1960), 28% References of the spring diet of M. camtchatica, hibernating in Arlt, D., Gebauer, A. 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