1. Social structure and habitat preferences of moose population in Biebrza National Park Mammal Research Institute, Polish Academy of Sciences, Białowieża Project: Biodiversity protection of Red Bog (Czerwone Bagno) - relic of raised bogs in Central Europe Włodzimierz Jędrzejewski Bogumiła Jędrzejewska Tomasz Borowik Kris Hundertmark Marcin Górny

2. Biebrza National Park

3. Biebrza Moose Data Observations by park personnel –group size and composition –location –date 1998-2010 (most from Sept. 2006-February 2010) 1534 observations 2770 total moose Classified as adult male, adult female, juvenile, or calf 2041 adults of known sex

5. Sexual segregation Sexes use different areas/habitats outside of the mating season Common in ungulates with body-size dimorphism –Larger difference in body size is associated with more extreme sexual segregation Some hypotheses (among others) –Predator avoidance by females and calves Females choose less risky environments whereas males choose riskier environments to gain back condition after breeding season –Food selection Females (smaller body size) seek out high-quality forage Males (larger body size) seek out more abundant but more fibrous forage due to their gut morphology –Activity budget Larger-bodied males spend more time ruminating and less time feeding than smaller-bodied females and therefore can’t synchronize movements

6. Moose group dynamics Group size: mean = 1.6 (SD 1.3), range = 1—20 Adult males: 1—11 Adult females: 1—8 67% of all observations are singletons (cows with calves included)

9. Group type for multi-adult groups G = 1.76, p = 0.18

11. Seasonal trends Segregation most pronounced in July and August –Mixed sex groups least likely to be seen together in July/August (2% and 4% of all observations, respectively); most likely to be seen in March and November (17% each) –Cow-calf pairs are not seen with adult males in July and August All-male groups most often seen in June and September (pre-rut)—December (post-rut) All-female groups most often seen in January—May

12. F = 4.68, p < 0.0001

14. OpenClosed SeasonMeanSDMeanSDtP (1-tailed) Spring1.891.331.791.14-0.690.25 Summer1.421.201.451.09 0.180.43 Fall1.681.831.471.07-1.670.048 Winter1.741.051.440.89-2.550.0055 Group size by habitat closure by season ( α’ = 0.0127)

15. Are the sexes distributed differently across the landscape? Create 2041 random points within the park Compare with 2041 observations of adults

16. Tested moose locations vs. random locations –Dispersion significantly different (p < 0.0001) Tested males vs. females seasonally –Šidák correction for multiple comparisons: α’ = 0.0127 –Winter (21 Oct.—30 Apr.) p < 0.0001 –Spring (1 May—30 June) p = 0.011 –Summer (1 July—20 Sept.) p = 0.005 –Rut (21 Sept.—20 Oct.) p = 0.35 Are the sexes distributed differently across the landscape?

17. Social segregation Measured by social segregation coefficient –Varies from 0 (complete aggregation) to 1 (complete segregation)

18. What habitat characteristics are driving the segregation of the sexes? Classify observations by habitat characteristics Account for potential location error

19. Forest Meadow 100 m

20. Classify observations by habitat characteristics Account for potential location error Characterize random points in the same manner to represent available habitat Randomly assign a sex to each random point Use stepwise logistic regression to identify factors influencing space use What habitat characteristics are driving the segregation of the sexes?

21. Possible predictors Landscape metrics Distance to road Distance to village Distance to arable land Distance to railway Distance to forest edge Distance to marsh/water Year Season Habitat metrics Deciduous forest Coniferous forest Other forest Marsh Meadow Arable land/orchard Wetland (yes/no) Edge (yes/no) Temporal metrics

22. How do moose respond to those factors? Selection coefficients = (use – availability)/(use + availability) – 1 (complete avoidance) to +1 (complete preference)

23. * * * * Habitat selection * Males ≠ females

24. * * * * * * *

25. Importance of marsh to calving 16 May-15 June 90% of all female locations in marsh 50% of all male locations in marsh (40% in deciduous forest) Use

26. Selection during calving

27. What patterns are there in space use that could lead to segregation? –Use variables identified by logistic regression –Test sex × location (random vs. observed) –Test sex × season × location

28. Distance to road Location observedrandom FMFM

29. observed random deciduous coniferous F FM M

30. observed random Distance to road FMFM winter calving summer breeding winter calving summer breeding

31. observed random coniferous forest winter calving summer breeding winter calving summer breeding FMFM

32. observed random deciduous forest winter calving summer breeding winter calving summer breeding FMFM

33. winter calving summer breeding winter calving summer breeding MFMF Distance to forest edge observed random

34. Conclusions Group size in Biebrza moose is small Group size in open habitats is greater than in closed habitats, particularly in winter Mixed-sex groups are not common, males and females are equally solitary Moose in Biebrza segregate outside of the breeding season on a very fine scale In general, open habitats were avoided and forested habitats were selected Males have a higher preference for deciduous forest in winter and summer and are farther from roads Females have a higher preference for coniferous forest in winter and are closer to roads Females use coniferous forest more in winter than do males Marsh habitat, although generally avoided, is very important during the calving season

35. Thank you