1. Engineers in the forest. How do foxes and badgers modify forest vegetation? Joanna Czarnecka 1, Justyna Sokal 1, Przemysław Kurek 2, Jan Holeksa 3 1 Maria Curie-Skłodowska University, Lublin, Poland 2 Institute of Botany, Polish Academy of Sciences, Kraków, Poland 3 A. Mickiewicz University, Poznań, Poland The entrance to burrow of the fox (Vulpes vulpes)

2. Introduction or… …what was the starting point of our study  Ecosystem engineers modify the availability of resources to other species  They cause physical changes in biotic and abiotic matter, but do not influence trophic interactions directly Jones CG., Lawton J.H., Shachak M. 1994. Oikos

3. Introduction Ecosystem engineers create, modify or maintain existing habitats

4. Introduction Animals building nests and burrows are good example of engineering species The primary function of the nests: hazard mitigation 1. Nests are habitats theirselves 2. Nests affect habitats in which they are built But also:

5. Introduction Girlatovce, eastern Slovakia, 2009 Nest as habitats Nests of the white stork (Ciconia ciconia) are habitats for many plant and animal species. Seedlings of 97 plant species (Czarnecka, Kitowski 2013) Total N content (g kg -1 ) Nest number Average content in Polish soils

6. Study aim Burrows affect habitats in which they are built Fox (Vulpes vulpes) and badger (Meles meles) disperse seeds and deposit them with faeces D + D Disturbance Sett of badger Dispersal Faeces on ”the dormant”

7. Study aim How do the activity of animals influence qualitative and quantitative structure of understorey trees and shrubs in the vicinity of burrows? Local patches versus matrix The nest closest vicinity versus forest floor vegetation

8. Source: commons.wikimedia.org Study site and methods Landscape parks National parks Lublin Poznań Kraków The Kampinos National Park Warsaw

9. Study site and methods The Vistula river Warsaw The Kampinos National Park (KNP) Source: Matuszkiewicz 2003 Pine and mixed oak-pine forest Ash-alder forests Potential landscape fito-compexes of KNP Oak-hornbeam and mixed forests Oak-hornbeam and mixed forests Oak-hornbeam and mixed forests Oak-hornbeam and mixed forests Studied burrows (N = 36)

10. Study site and methods Matrix Patches Burrow plot 100 m 2 Control plot 100 m 2 50 m Badger N = 19 burrows  exploited intensively all over the year  more complex structure Fox N = 17 burrows  exploited less intensively mostly in mating season and during winter  smaller and simpler structure All plant individuals were counted

11. Matrix differences Burrowing places – preferences of studied animal species Badger versus fox Higher clay and silt content Higher soil fertility Higher content of:  total carbon (C)  total nitrogen (N)  magnesium (Mg)  calcium (Ca)  potassium (K) Plant communities with higher tree species richness and greater broadleaved trees admixture Source: Kurek et al. 2014

12. Matrix differences – control plots Burrowing places – preferences of studied animal species Badger N = 19 Fox N = 17 Total number of species (trees and shrubs) 1812 Frequency (%) Frangula alnus8976 Sorbus aucuparia7935 Pyrus communis4218 Padus avium266 Juniperus communis3253 Mean diversity measures BadgerFox Species number P <0.05 (U-test) * Shannon diversity NS 23 tree and shrub species were found in total

13. Influence of the animal activity PATCHES (burrow plots) versus MATRIX (control plots) Mean number of individuals Badger All speciesFrangula alnus NS Mean diversity measures Species numberShannon diversity P <0.01 (Wilcoxon test) ** P <0.01 (paired samples t-test)

14. Influence of the animal activity PATCHES (burrow plots) versus MATRIX (control plots) Badger Control plot Burrow plot Matrix Patches Axis 1 and diversity index: S r = +0.84; p < 0.0001 Axis 1 and number of species: S r = +0.45; p < 0.01 Axis 1 and abundance of F. alnus: S r = -0.65; p < 0.0001 50 m DCA

15. Influence of the animal activity PATCHES (burrow plots) versus MATRIX (control plots) Fox www.wikimediawww.wikimedia (Malene) Mean number of individuals All speciesFrangula alnus NS Mean diversity measures Species numberShannon diversity P <0.05 (Wilcoxon test) * P <0.05 (paired samples t-test) *

16. Influence of the animal activity PATCHES (burrow plots) versus MATRIX (control plots) Fox www.wikimediawww.wikimedia (Malene) Control plot Burrow plot No obvious pattern Axis 1 and number of species: S r = +0.53; p < 0.01 Axis 1 and abundance of F. alnus: S r = +0.82; p < 0.0001 DCA

17. What is more important? D or D? DISTURBANCE or DISPERSAL? Sample – frequencies (%) of particular species, N = 23 Species Badger Fox MatrixPatchesFaecesMatrixPatchesFaeces 1Frangula alnus 89 0.776820.7 2Rhamnus catharticus 521--12- 3Cerasus avium 112610.16125.1 Low impact Disturbance Dispersal Is any general pattern there?

18. What is more important? D or D? DISTURBANCE or DISPERSAL? Sample – frequencies (%) of particular species, N = 23 Spearman rank correlation for frequencies Badger Fox In patches and in matrix In patches and in faeces Disturbance and seed source Dispersal S r = 0.42; p < 0.05S r = 0.57; p < 0.01 S r = 0.47; p < 0.05S r = 0.54; p < 0.01 Both are important!

19. Conclusions 1.Burrowing animals create patches which are different from forest vegetation 2.They cause evident increase of local plant species diversity. The result depends on animal species 3.There are two main aspects of animal activity: disturbance and internal seed dispersal and their importance differs for different plant species