# Long-term scatterhoarding by Eurasian red squirrels (Sciurus vulgaris) > How many seeds does a red squirrel bury in a year? In two Belgian woodlands, researchers counted the small dig holes squirrels leave when they recover their buried food. Each squirrel hid about 2,300 to 2,800 pine cones, acorns and beechnuts a year, and found most of them again, carrying it through the lean months of late winter and spring. Summary by Kersey Lawrence, Original Wisdom. Updated 5 November 2026. Page: https://www.originalwisdom.com/tracking-literature/1996-wauters-casale-long-term-scatterhoarding-eurasian/ ## Citation and access - **Authors:** L.A. Wauters, P. Casale - **Year published:** 1996 - **Journal or source:** Journal of Zoology, London 238(2): 195–207 - **Full citation:** Wauters, L.A., and Casale, P. 1996. Long-term scatterhoarding by Eurasian red squirrels (Sciurus vulgaris). Journal of Zoology, London 238(2): 195–207. - **DOI:** https://doi.org/10.1111/j.1469-7998.1996.tb05389.x - **Access status:** Publisher link only - **Where to find it:** https://doi.org/10.1111/j.1469-7998.1996.tb05389.x ## Tags - **Species:** Mammals > Eurasian red squirrel - **Region:** Europe > Belgium - **Tracking:** Caches; Digs; Tracking; Tracks and signs identification - **Behaviour and ecology:** Behaviour; Energy budget; Feeding; Food caching; Interspecific competition; Scatterhoarding; Seed dispersal - **Methods and technology:** Direct observation; Plot searches; Radio telemetry - **Data type:** New field data - **Observer qualification:** Not reported - **Discipline:** Behaviour and ecology; Wildlife tracking - **Year published:** 1996 ## Detailed keywords - **Species:** Studied: Eurasian red squirrel (Sciurus vulgaris). Foods hoarded: pine cones of Scots pine (Pinus sylvestris) and Corsican pine (Pinus nigra), acorns of pedunculate oak (Quercus robur), beechnuts of beech (Fagus sylvatica), and more rarely hazelnuts and sweet chestnuts (Castanea sativa). In the study areas but not studied: larch (Larix decidua) and spruce (Picea abies). Mentioned for comparison: grey squirrel (Sciurus carolinensis), fox squirrel (Sciurus niger), Japanese squirrel (Sciurus lis), American red squirrel (Tamiasciurus hudsonicus), Douglas squirrel (Tamiasciurus douglasii), great spotted woodpecker (Dendrocopos major, the only other animal in western Europe that can extract seeds from closed pine cones). - **Location and habitat:** Two 30 ha study areas in the Province of Antwerpen, northern Belgium. The coniferous woodland is the Merodese Bossen at Herenthout (212 ha), dominated by Scots and Corsican pine, with some larch and spruce, and oak and beech only along paths or in small patches. The mixed deciduous woodland is the Peerdsbos at Schoten (150 ha), dominated by mature oak and beech with some chestnut, a few larch and spruce, and a small 2 ha patch of pine on the edge. - **Topics:** food hoarding, scatterhoarding versus larderhoarding, cache contents and density, hoarding rate, cache recovery and its timing through the year, energy value of caches, how woodland type shapes hoarding, seed dispersal and the squirrel–tree relationship, competition between red and introduced grey squirrels. - **Statistics or models:** Descriptive statistics (means, standard deviations, ranges, percentages). Energy budget calculations combining time spent hoarding and eating, hoarding and feeding rates, daily energy intake and the energy content of each seed type. A sign test comparing the two woodlands. - **Data type:** New field data. Cache contents: 100 hoards dug up on 10 plots of 5 × 5 m in the coniferous woodland. Recovered caches: counted on 2 plots of 10 × 75 m in the coniferous woodland, from early September to the end of June in 1984 and 1985. Hoarding behaviour: watched in radio-tracked squirrels from August to December, 1985–1987: 413 minutes in the coniferous area and 223 minutes in the deciduous area. Earlier data: activity budgets, feeding rates and energy figures from the authors’ earlier studies of the same populations. ## Synopsis ### 1. What they did Many tree squirrels bury food to eat later. Eurasian red squirrels are “scatterhoarders”: they bury one or a few seeds in each spot, spread across hundreds or thousands of spots in their home range. Until this study, no one had measured how much food Sciurus squirrels store this way, or how much of a squirrel’s yearly energy needs it supplies. The authors studied red squirrels in two woodlands in northern Belgium: one mainly pine, the other mainly oak and beech. They dug up 100 caches to see what was in them, and counted recovered caches on the forest floor through the year. They also followed squirrels carrying radio transmitters and watched them burying food. They combined these observations with their earlier measurements of how long squirrels are active, how fast they eat each seed type and how much energy they need each day. From this they worked out how many seeds a squirrel buries in a year, how many it recovers, and how many days of food that represents. ### 2. What they found What is in a cache - The 100 caches held 155 food items: 65 had a single item, and the most in one cache was 5. - Of caches with more than one item, 43% mixed seeds from different tree species. - Overall, 63% of the items were pine cones, 21% acorns and 16% beechnuts. - Seeds were buried about 2–6 cm deep in the soil or leaf litter. How much is hoarded - Recovered caches showed as small round holes about 5 cm deep. About 5,100 to 6,700 cached seeds were dug up per hectare each year in the pine wood. With about 2.3 squirrels per hectare, that is about 2,200 to 2,900 seeds recovered per squirrel per year. - Squirrels hoarded mostly in September and October, tailing off through to January. - Each seed took about 44–50 seconds to hide. Acorns and beechnuts were hidden faster than pine cones, because they were often picked up from the ground. - Almost all food was buried in the ground. Only twice did a squirrel hide food in a tree. - A squirrel spent about 32 hours a year hoarding in the pine wood and 34 hours in the oak–beech wood, burying about 2,300 and 2,800 seeds a year respectively. - What they hoarded depended on what was abundant: mostly pine cones in the pine wood, mostly acorns and beechnuts in the oak–beech wood. How much is recovered - Squirrels recovered about 83% of their caches in the pine wood and 79% in the oak–beech wood. - They recovered nearly all buried pine cones (99–100%) and most beechnuts (83–92%), but only about half of the acorns (54–62%). - The recovered food was enough for about 43 days of a squirrel’s energy needs in the pine wood and 68 days in the oak–beech wood. When caches are used - Oak–beech wood: caches were used mostly from January to April, with acorns eaten first. - Pine wood: caches were used mostly from March to May, with pine cones eaten from April to June, when no more seeds were left on the trees. ### 3. What they concluded - Red squirrels hoard less than American red squirrels in similar Canadian forests (about one-quarter in pine woods). Even so, their caches matter during the hardest part of the year, January to May, when energy-rich food is scarce. - Woodland type shapes the behaviour. In oak–beech woods, seeds are available in the canopy for only a short time and fallen seeds are soon eaten by other animals. So squirrels hoard more there and use their caches earlier. - Red squirrels digest acorns poorly and never eat more than 4–6 a day, which may explain why so many cached acorns are left. Unrecovered beechnuts and acorns may grow into seedlings, so squirrels may help disperse these trees. - Hoarding may be more important in regions with colder winters or less varied food. - Introduced grey squirrels forage more on the ground, and they may raid red squirrels’ caches in early spring, when reds depend on them most. This competition could help explain why grey squirrels have replaced red squirrels in much of Britain. ### 4. How tracking was or was not used No trailing was reported. The study relied on reading signs on the forest floor: - Recovered caches: recognised as small round holes about 5 cm deep, and counted on fixed plots through the year to estimate how many caches were dug up per hectare. - Buried caches: found by digging through the litter and soil to about 10 cm. They were distinguished from seeds that had simply fallen because they were buried 2–6 cm deep. - Who made them: the authors state that in Belgium squirrels are the only animals that store pine cones and beechnuts this way. Hoarding behaviour itself was watched directly, by finding radio-tagged squirrels and observing them. ### 5. Tracker or observer certification Not reported. The cache recovery estimates rest on recognising squirrel dig holes and buried caches, but no qualification or experience in tracks and signs identification is described. ### 6. Other technologies, skills or innovations - Radio transmitters, used to find squirrels so their behaviour could be watched for 30 to 60 minutes at a time - Energy budget calculations linking behaviour to the energy value of each seed type ### 7. Implications for conservation (from the paper) Competition with introduced grey squirrels for buried seeds, especially in late winter and early spring, may lower red squirrel survival or breeding. This could be a critical part of understanding why red squirrels are being replaced by grey squirrels in much of Britain. ### 8. Implications for tracking (Original Wisdom's observations) _These are Original Wisdom's observations, not the authors'._ Small round dig holes where caches were recovered are a sign trackers can read and count, and this study shows that such counts can be turned into estimates of how much food animals store and use. The paper also shows when to look: hoarding in autumn, and cache recovery in late winter and spring. ## Cite this summary Lawrence, K. 2026. Summary of: Long-term scatterhoarding by Eurasian red squirrels (Sciurus vulgaris) (Wauters and Casale, 1996). Original Wisdom, Tracking Literature. https://www.originalwisdom.com/tracking-literature/1996-wauters-casale-long-term-scatterhoarding-eurasian/