, Volume 153, Issue 3, pp 625–632 | Cite as

Effects of herbivory and patch size on tree seedling survivorship in a fog-dependent coastal rainforest in semiarid Chile

  • Ek del-ValEmail author
  • Juan J. Armesto
  • Olga Barbosa
  • Pablo A. Marquet
Plant Animal Interactios


The landscape (matrix) surrounding habitat fragments critically affects the biodiversity of those fragments due to biotic interchange and physical effects. However, to date, there have been only a limited number of studies on plant–animal interactions in fragmented landscapes, particularly on how tree seedling herbivory is affected by fragmentation. We have examined this question in a fog-dependent mosaic of rainforest fragments located on coastal mountaintops of semiarid Chile (30°S), where the effects of the surrounding semiarid matrix and forest patch size (0.1–22 ha) on tree seedling survival were simultaneously addressed. The rainforest is strongly dominated by the endemic evergreen tree species Aextoxicon punctatum (Olivillo, approx. 80% of basal area). To assess the magnitudes and causes of Olivillo seedling mortality, we set up a field experiment where 512 tree seedlings of known age were transplanted into four forest fragments of different sizes in four 1.5 × 3-m plots per patch; one-half of each plot was fenced off with chicken wire to exclude small mammals. The plots were monitored for 22 months. Overall, 50% of the plants died during the experiment. The exclusion of small mammals from the plots increased seedling survival by 25%, with the effect being greater in smaller patches where matrix-dwelling herbivores are more abundant. This experiment highlights the important role of the surrounding matrix in affecting the persistence of trees in forest fragments. Because herbivores from the matrix cause greater tree seedling mortality in small patches, their effects must be taken into account in forest conservation–restoration plans.


Habitat fragmentation Plant–animal interactions Tree regeneration Water limitation 



The Corporación Nacional Forestal (CONAF, IV Región, Chile) kindly gave permission to work at Fray Jorge National Park. We thank J. Monardes, N. Davies, V. Matus, B. González, A. Gutiérrez and P. Chacón for assistance in the field and F. Matus, J. Larco and C. Tejo for the laboratory work. We especially thank C. Garín and Y. Hussein for small mammal trapping. Pablo Necochea kindly drew the Fray Jorge map. We thank K. Boege, J. Benítez-Malvido and two anonymous reviewers for comments on earlier versions. This work was supported by FONDAP–FONDECYT 1501-0001 (CASEB), Biocores Project funded by EC under INCO IV programme (Contract ICA 4-CT-2001–10095), CMEB (PO2-051-F), ICM (P05-002) and Project P02-051-F ICM (IEB). O. Barbosa thanks the A. Mellon foundation for a doctoral fellowship. Part of this work was conducted while PAM was a Sabbatical Fellow at the National Center for Ecological Analysis and Synthesis, a Center funded by NSF (Grant no. DEB-0072909), the University of California and the Santa Barbara campus. The experiments performed for this investigation comply with the current laws of Chile.


  1. Alverson WS, Waller DM, Solheim SL (1988) Forest too deer: edge effects in Northern Wisconsin. Conserv Biol 2:348–358CrossRefGoogle Scholar
  2. Arnold EA, Asquith NM (2002) Herbivory in a fragmented tropical forest: patterns from islands at Lago Gatún, Panama. Biodivers Conserv 11:1663–1680CrossRefGoogle Scholar
  3. Asquith NM, Mejia-Chang M (2005) Mammals, edge effects, and the loss of tropical forest diversity. Ecology 86:379–390CrossRefGoogle Scholar
  4. Asquith NM, Wright SJ, Clauss MJ (1997) Does mammal community composition control recruitment in neotropical forests? Ecology 78:941–946CrossRefGoogle Scholar
  5. Barbosa O, Marquet PA (2001) Effects of forest fragmentation on the beetle assemblage at the relic forest of Fray Jorge, Chile. Oecologia 132:296–306CrossRefGoogle Scholar
  6. Benítez-Malvido J (1998) Impact of forest fragmentation on seedling abundance in a tropical rain forest. Conserv Biol 12:380–389CrossRefGoogle Scholar
  7. Benítez-Malvido J, Lemus-Albor A (2005) The seedling community of tropical rain forest edged and its interaction with herbivores and pathogens. Biotropica 37:301–313CrossRefGoogle Scholar
  8. Cadenasso M, Pickett STA (2000) Linking forest edge structure to edge function: mediation of herbivore damage. J Ecol 88:31–44CrossRefGoogle Scholar
  9. Cornelius C, Cofré H, Marquet P (2000) Effects of habitat fragmentation on bird species in a relict temperate forest in semiarid Chile. Conserv Biol 14:534–543CrossRefGoogle Scholar
  10. Crawley MJ (1990) Rabbit grazing, plant competition and seedling recruitment in acid grassland. J Appl Ecol 27:803–820CrossRefGoogle Scholar
  11. Curran LM et al (1999) Impact of El Niño and logging on canopy tree recruitment in Borneo. Science 286:2184–2188PubMedCrossRefGoogle Scholar
  12. del-Val E, Armesto JJ, Barbosa O, Christie DA, Gutierrezet AG et al. (2006) Rainforest islands in the Chilean semiarid region: a fog-dependent ecosystem under adverse climate. Ecosystems 9:598–608Google Scholar
  13. Di Castri F, Hajek E (1976) Bioclimatología de Chile. Universidad Católica de Chile, SantiagoGoogle Scholar
  14. Dirzo R (2001) Plant-mammal interactions: lessons for our understanding of nature and implications for biodiversity conservation. In: Press MC, Huntly N, Levin S (eds) Ecology: achievement and challenge. Blackwell, Oxford pp 319–336Google Scholar
  15. Gutiérrez JR, Meserve PL (2000) Density and biomass responses on ephemeral plants to experimental exclusions of small mammals and their vertebrate predators in the Chilean semiarid zone. J Arid Environ 45:173–181CrossRefGoogle Scholar
  16. Holmgren M, Scheffer M, Ezcurra E, Gutiérrez JR, Mohren GMJ (2001) El Niño effects on the dynamics of terrestrial ecosystems. Trends Ecol Evol 16:89–94PubMedCrossRefGoogle Scholar
  17. Jaksic F (2001) Ecological effects of El Niño in terrestrial ecosystems of western South America. Ecography 24:241–250CrossRefGoogle Scholar
  18. Jaksic F, Silva-Aranguiz E, Silva SI (2004) Fauna del Parque Nacional Bosque Fray Jorge: una revisión bibliográfica. In: Squeo FA, Gutiérrez JR, Hernández IR (eds) Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena, La Serena, pp 93–114Google Scholar
  19. Janzen DH (1983) No park is an island: increase in interference from outside as park size increases. Oikos 41:402–410CrossRefGoogle Scholar
  20. Janzen DH (1986) The eternal external threat. In: Soulé ME (eds) Conservation biology. The science of scarcity and diversity. Sinauer Assoc, Suderland, pp 286–303Google Scholar
  21. Kummerow J (1966) Aporte al conocimiento de las condiciones climáticas del bosque de Fray Jorge. Boletín Técnico, Universidad de Chile, Facultad de Agronomía 24:21–24Google Scholar
  22. Laurance WF (2000) Do edge effects occur over large spatial scales? Trends Ecol Evol 15:134–135PubMedCrossRefGoogle Scholar
  23. Laurance WF, Vasconcelos HL, Bruna EM, Didham RK, Stouffer PC et al. (2002) Ecosystem decay of Amazonian forest fragments: a 22-year investigation. Conserv Biol 16:605–618Google Scholar
  24. Lienert J, Fischer M (2003) Habitat fragmentation affects the commom wetland specialist Primula farinosa in north-east Switzerland. J Ecol 91:587–599CrossRefGoogle Scholar
  25. López-Cortés F, López D (2004) Antecedentes bioclimáticos del Parque Nacional Bosque Fray Jorge. In: Squeo FA, Gutiérrez JR, Hernández IR (eds) Historia Natural del Parque Nacional Bosque Fray Jorge. Universidad de La Serena, La Serena, pp 45–60Google Scholar
  26. Lusk CH (2002) Leaf area accumulation helps juvenile evergreen trees tolerate shade in a temperate rainforest. Oecologia 132:188–196CrossRefGoogle Scholar
  27. McEuen AB, Curran LM (2006) Plant recruitment bottlenecks in temperate forest fragments: seed limitation and insect herbivory. Plant Ecol 184:297–309CrossRefGoogle Scholar
  28. Meserve PL (1981a) Resource partitioning in a Chilean semiarid small mammal community. J Anim Ecol 50:745–757CrossRefGoogle Scholar
  29. Meserve PL (1981b) Trophic relationships among small mammals in a Chilean semiarid thorn scrub community. J Mammal 62:304–314CrossRefGoogle Scholar
  30. Montecinos A, Aceituno P (2003) Seasonality of the ENSO-related rainfall variability in central Chile and associated circulation anomalies. J Climate 16:281–296CrossRefGoogle Scholar
  31. Muñoz C, Pisano E (1947) Estudio de la vegetación y flora de los Parques Nacionales de Fray Jorge y Talinay. Agric Tech 7:71–190Google Scholar
  32. Murúa R (1996) Comunidades de mamíferos del bosque templado de Chile. In: Armesto JJ, Villagrán C, Arroyo MD (eds) Ecología de los bosques nativos de Chile. Editoria Univesitaria, Santiago, pp 113–134Google Scholar
  33. Novoa-Jerez J, Viada-Ovalle JM, López D, Squeo FA (2004) Localización espacial del bosque Fray Jorge en los Altos del Talinay, IV Región de Coquimbo. In: Squeo FA, Gutiérrez JR, Hernández IR (eds) Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena, La Serena, pp 161–171Google Scholar
  34. Núñez-Avila M, Armesto JJ (2006) Relict islands of the temperate rainforest tree Aextoxicon punctatum (Aextoxicaceae) in semi-arid Chile: genetic diversity and biogeographic history. Aust J Bot 54:733–743CrossRefGoogle Scholar
  35. Paton PWC (1994) The effect of edge on avian nest success: how strong is the evidence? Conserv Biol 8:17–26CrossRefGoogle Scholar
  36. Pérez C, Villagrán C (1994) Influencia del clima en el cambio florístico, vegetacional y edáfico de los bosques de “olivillo” (Aextoxicon punctatum R. et Pav.) de la Cordillera de la Costa de Chile: implicaciones biogeográficas. Rev Chil Hist Nat 67:77–90Google Scholar
  37. Rao M, Terborgh J, Nuñez P (2001) Increased herbivory in forest isolates: implications for plant community structure and composition. Conserv Biol 15:624–632CrossRefGoogle Scholar
  38. Saunders DA, Hobbs RJ, Margules CR (1991) Biological consequences of ecosystem fragmentation: a review. Conserv Biol 5:18–32CrossRefGoogle Scholar
  39. Silva SI (2005) Posiciones tróficas de pequeños mamíferos en Chile: una revisión. Rev Chil Hist Nat 78:589–599Google Scholar
  40. Squeo FA, Arancio G, Novoa-Jerez J (2004) Heterogeneidad y diversidad florística del Bosque de Fray Jorge. In: Squeo FA, Gutiérrez JR, Hernández IR (eds) Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena, La Serena, pp 173–185Google Scholar
  41. Villagrán C, Armesto JJ (1980) Relaciones florísticas entre las comunidades relictuales del Norte Chico y la Zona Central con el bosque del Sur de Chile. Bol Mus Nac Hist Nat 37:85–99Google Scholar
  42. Villagrán C, Armesto JJ, Hinojosa F, Cuvertino J, Pérez C, Medina C (2004) El enigmático origen del bosque relicto de Fray Jorge. In: Squeo FA, Gutiérrez JR, Hernández IR (eds) Historia Natural del Parque Nacional Bosque Fray Jorge. Ediciones Universidad de La Serena, La Serena, pp 173–185Google Scholar
  43. Weltzin JF, Archer SR, Heitschmidt RK (1998) Defoliation and woody plant (Prosopis glandulosa) seedling regeneration: potential versus realized herbivory tolerance. Plant Ecol 138:127–135CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Ek del-Val
    • 1
    • 4
    Email author
  • Juan J. Armesto
    • 1
    • 2
  • Olga Barbosa
    • 1
    • 5
  • Pablo A. Marquet
    • 1
    • 2
    • 3
  1. 1.Center for Advanced Studies in Ecology and Biodiversity (CASEB), Departamento de Ecología, Facultad Ciencias BiológicasPontificia Universidad Católica de ChileSantiagoChile
  2. 2.Instituto de Ecología y Biodiversidad (IEB)Facultad de Ciencias Universidad de ChileSantiagoChile
  3. 3.National Center for Ecological Analysis and SynthesisSanta BarbaraUSA
  4. 4.Centro de Investigaciones en EcosistemasUniversidad Nacional Autónoma de MéxicoMoreliaMexico
  5. 5.BIOMEUniversity of SheffieldSheffieldUK

Personalised recommendations