European Journal of Forest Research

, Volume 138, Issue 4, pp 653–672 | Cite as

Scale dependence shapes how plant traits differentially affect levels of pre- and post-dispersal seed predation in Scots pine

  • Fiona R. WorthyEmail author
  • Philip E. Hulme
Original Paper


To understand the demographic consequences of seed predation, it is essential to scale-up losses over space and time. We tested how individual tree characteristics, forest attributes, and the local environment affected rates of both pre- and post-dispersal seed predation in Scots pine Pinus sylvestris, in Scotland’s ancient native pinewoods. The sampling strategy was hierarchical, allowing assessment of how the influence of these factors differed with spatial scale. This approach revealed that the fate of a tree’s seeds depended on many factors, at multiple scales. Levels of seed predation were highly variable over space and time. Seed predation by local post-dispersal seed predators (invertebrates and small mammals) varied at the finest spatial scales, whereas mobile avian pre-dispersal seed predators discriminated among seeds at every spatial scale. Tree crown shape affected removal of seed by all seed predators; both crossbills Loxia spp. and generalist avian granivores selected trees with broad crowns. However, trees with narrow crowns were selected by red squirrels Sciurus vulgaris, and the associated tall understory vegetation supported the highest levels of post-dispersal seed predation. Seed chemistry was an effective seed defence that reduced seed loss to all seed predators. In contrast, increases in physical cone defence were ineffective at reducing seed loss and showed a potential trade-off with investment in seed viability. Large seeds carried a survival cost, being preferentially consumed by squirrels and post-dispersal seed predators. All post-dispersal seed predators consumed a higher proportion of seeds at the times and locations where seed fall was greatest.


Conifer Granivory Pinus sylvestris Native pinewood Seed fate Spatio-temporal variation 



Mick Marquiss and Richard Law made valuable contributions throughout this study. The Royal Society for the Protection of Birds permitted fieldwork at Abernethy, provided accommodation, and many staff gave useful advice. Scottish Natural Heritage permitted fieldwork at Mar Lodge and provided accommodation. Ron Summers and Robert Proctor provided additional data on cone counts for Abernethy Forest. Statistical advice was provided by David Elston. This research was funded by a Natural Environment Research Council studentship to FRW (Grant No. NER/S/A/2002/11035) and while writing by a Yunnan Provincial Post-doctoral Grant to FRW.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

10342_2019_1198_MOESM1_ESM.pdf (451 kb)
Online Resource 1 Map of study sites (PDF 450 kb)
10342_2019_1198_MOESM2_ESM.pdf (178 kb)
Online Resource 2 Forest structure (PDF 178 kb)
10342_2019_1198_MOESM3_ESM.pdf (119 kb)
Online Resource 3 Timeline of seed development and seed predation (PDF 119 kb)
10342_2019_1198_MOESM4_ESM.xlsx (21 kb)
Online Resource 4 Sectors marked for cone collection (XLSX 20 kb)
10342_2019_1198_MOESM5_ESM.pdf (122 kb)
Online Resource 5 Supplementary methods (PDF 122 kb)
10342_2019_1198_MOESM6_ESM.pdf (204 kb)
Online Resource 6 Common factor analysis (PDF 204 kb)
10342_2019_1198_MOESM7_ESM.pdf (211 kb)
Online Resource 7 Supplementary results (PDF 210 kb)


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Authors and Affiliations

  1. 1.Centre for Mountain Ecosystem StudiesKunming Institute of BotanyKunmingChina
  2. 2.The Bio-Protection CentreLincoln UniversityCanterburyNew Zealand

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