Pinus nigra Arn. ssp. salzmannii seedling recruitment is affected by stand basal area, shrub cover and climate interactions

Published Date

September 2016, Volume 73, Issue 3, pp 649-656

First online: 01 April 2016

Title 

Pinus nigra Arn. ssp. salzmannii seedling recruitment is affected by stand basal area, shrub cover and climate interactions

• Author 

• Manuel E. Lucas-Borja 
• , David Candel-Pérez
• , Francisco A. García Morote
• , Thierry Onkelinx
• , Pedro A. Tíscar
• , Philippe Balandier

Abstract

Key message

Shrub cover has visible effects on Pinus nigra Arn ssp.salzmannii seedling emergence, but only in drier years under moderate basal area (25–30 m 2   ha −1 ). In the wetter year, shrub cover favours seedling survival without basal area influence.

Context

Shrubs are known to favour tree seedling recruitment in difficult climate environments, but facilitation may prove optimal in intermediate-level rather than harsh conditions, although such an effect remains to be specified.

Aims

The main aim of this work was to evaluate the influence of stand basal area (15–20, 25–30 and 35–40 m2 ha−1) × with/without shrub facilitation on seedling recruitment in Spanish black pine (P. nigra Arn. ssp. salzmannii), which has been struggling to regenerate natural forests since the late nineteenth century, sometimes jeopardizing stand persistence.

Methods

In spring 2012 and 2013, 72 subplots of 25 × 25 cm were established in Cuenca Mountains, central-eastern Spain (four replicates, each counting three subplots × 2 shrub conditions, both under and outside shrub cover, in three different stand basal areas). All plots were randomly distributed at least 500 m apart within each stand basal area. Each subplot comprised 20 seeds previously collected at the experimental forest site. Seedling emergence and early survival were monitored every month in 2012 and 2013.

Results

Seedling recruitment was strongly influenced by year. The year 2012 was drier and warmer than 2013. In 2012, seedling emergence was higher under the 25–30-m2 ha−1stand basal area and favoured by shrub cover, whereas in 2013, neither basal area nor shrub cover had significant effects. No seedling survived the summer in 2012. Higher seedling survival was found outside shrub cover at every basal area interval in 2013 with living seedlings until the end of the year.

Conclusion

Climate is the main factor controlling seedling emergence and survival. Shrub facilitation has visible effects on seedling emergence, but only in drier years under moderate light (moderate basal area). Facilitation collapses at extreme stress levels. In wetter years, shrub cover does not promote seedling survival and basal area is not an influential factor.

References

1. Adili B, El Aouni MH, Garchi S, Balandier P (2009). Natural regeneration of Pinus pinea L. in Tunisia as influenced by canopy cover, litter biomass and understorey vegetation. In: Final COST E47 conference. Forest vegetation management towards environmental sustainability. Vejle, Denmark. University of Copenhagen, Forest and Landscape Denmark, pp 66–68
2. Adili B, El Aouni MH, Balandier P (2013) Unraveling the influence of light, litter and understory vegetation on Pinus pinea natural regeneration. Forestry 86:297–304CrossRef
3. Agestam E, Ekö PM, Nilsson U, Welander NT (2003) The effects of shelterwood density and site preparation on natural regeneration of Fagus sylvatica in southern Sweden. For Ecol Manag 176:61–73CrossRef
4. Allué JL (1990) Atlas fitoclimático de España. Taxonomías. Ministerio de Agricultura, Pesca y Alimentación, Instituto Nacional de Investigaciones Agrarias, Madrid
5. Balandier P, Collet C, Miller JH, Reynolds PE, Zedacker SM (2006a) Designing forest vegetation management strategies based on the mechanisms and dynamics of crop tree competition by neighbouring vegetation. Forestry 79:3–27CrossRef
6. Balandier P, Sonohat G, Sinoquet H, Varlet-Grancher C, Dumas Y (2006b) Characterisation, prediction and relationships between different wavebands of solar radiation transmitted in the understorey of even-aged oak (Quercus petraea, Q. robur) stands. Trees 20:363–370CrossRef
7. Barbeito I, Fortin MJ, Montes F, Cañellas I (2009) Response of pine natural regeneration to small-scale variation on a managed Mediterranean mountain forest. Appl Veg Sci 12:488–503CrossRef
8. Barbier S, Gosselin F, Balandier P (2008) Influence of tree species on understory vegetation diversity and mechanisms involved—a critical review for temperate and boreal forests. For Ecol Manag 254:1–15CrossRef
9. Bertness MD, Callaway R (1994) The role of positive forces in natural communities. Trends Ecol Evol 9:191–193CrossRefPubMed
10. Callaway RM (1995) Positive interactions among plants. Bot Rev 61:306–349CrossRef
11. Callaway RM, Walker LR (1997) Competition and facilitation: a synthetic approach to interactions in plant communities. Ecology 78:1958–1965CrossRef
12. Callaway RM, Brooker RW, Choler P, Kikvidze Z, Lortie CJ, Michalet R et al (2002) Positive interactions among alpine plants increase with stress. Nature 417:844–848CrossRefPubMed
13. Candel-Pérez D, Linares JC, Viñegla B, Lucas-Borja ME (2012) Assessing climate-growth relationships under contrasting stands of co-occurring Iberian pines along an altitudinal gradient. For Ecol Manag 274:48–57CrossRef
14. Castro J, Zamora R, Hódar JA, Gómez JM (2005) Ecology of seed germination of Pinus sylvestris L. at its southern, Mediterranean distribution range. Investig Agrar: Sist Recur For 14:143–152
15. Catovsky S, Bazzaz FA (2000) The role of resource interactions and seedling regeneration in maintaining a positive feedback in hemlock stands. J Ecol 88:100–112CrossRef
16. Del Cerro Barja A, Lucas Borja ME, Martínez García E, Navarro López R, Andrés Abellán M, López Serrano FA (2009) The influence of stand density and soil treatment on the Spanish black pine (Pinus nigra Arn. ssp. salzmannii) regeneration in Spain. Investig Agrar: Sist Recur For 18:167–180
17. Emborg J (1998) Understorey light conditions and regeneration with respect to the structural dynamics of a near-natural temperate deciduous forest in Denmark. For Ecol Manag 106:83–95CrossRef
18. Fyllas NM, Dimitrakopoulos PG, Troumbis AY (2008) Regeneration dynamics of a mixed Mediterranean pine forest in the absence of fire. For Ecol Manag 256:1552–1559CrossRef
19. Germino MJ, Smith WK, Resor C (2002) Conifer seedling distribution and survival in an alpine-treeline ecotone. Plant Ecol 162:157–168CrossRef
20. Gómez-Aparicio L, Zamora R, Gómez JM, Hódar JA, Castro J, Baraza E (2004) Applying plant facilitation to forest restoration in Mediterranean ecosystems: a meta-analysis of the use of shrubs as nurse plants. Ecol Appl 14:1128–1138CrossRef
21. Gómez-Aparicio L, Valladares F, Zamora R, Quero JL (2005) Response of tree seedlings to the abiotic heterogeneity generated by nurse shrubs: an experimental approach at different scales. Ecography 28:757–768CrossRef
22. Granda E, Escudero A, De La Cruz M, Valladares F (2012) Juvenile-adult tree associations in a continental Mediterranean ecosystem: no evidence for sustained and general facilitation at increased aridity. J Veg Sci 2:164–175CrossRef
23. Holmgren M, Scheffer M, Huston MA (1997) The interplay of facilitation and competition in plant communities. Ecology 78:1966–1975CrossRef
24. Howard KSC, Eldridge DJ, Soliveres S (2012) Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland. Basic Appl Ecol 13:159–168CrossRef
25. Kerr G (2000) Natural regeneration of Corsican pine (Pinus nigra subsp. laricio) in Great Britain. Forestry 73:479–488CrossRef
26. Kerr G (2008) Managing continuous cover forests. Forestry commission operational guidance booklet 7. Forestry Commission, Edinburgh
27. Kitzberger T, Steinaker DF, Veblen TT (2000) Effects of climatic variability on facilitation of tree establishment in Northern Patagonia. Ecology 81:1914–1924CrossRef
28. Kunstler G, Curt T, Bouchaud M, Lepart J (2006) Indirect facilitation and competition in tree species colonization of sub-Mediterranean grasslands. J Veg Sci 17:379–388CrossRef
29. Le Bagousse-Pinguet Y, Maalouf JP, Touzard B, Michalet R (2014) Importance, but not intensity of plant interactions relates to species diversity under the interplay of stress and disturbance. Oikos 123:777–785CrossRef
30. Lee CS, Kim JH, Yi H, You YH (2004) Seedling establishment and regeneration of Korean red pine (Pinus densifloraS. et Z.) forests in Korea in relation to soil moisture. For Ecol Manag 199:423–432CrossRef
31. Legras EC, Vander Wall SB, Board DI (2010) The role of germination microsite in the establishment of sugar pine and Jeffrey pine seedlings. For Ecol Manag 260:806–813CrossRef
32. López-Serrano FR, Rubio Caballero E, Andrés Abellán M, Del Cerro Barja A, García Morote FA, De las Heras Ibañez J, Lucas-Borja ME, Moya Navarro D, Odi Lara M (2009) Efectos del Cambio climático en los bosques castellano-manchegos. In: Rodríguez Torres A, Fernández Castro H, Rojano Saura I (eds) Impactos del Cambio climático en Castilla La Mancha. Fundación General de Medio Ambiente, Castilla-La Mancha, Spain
33. Lucas-Borja ME, Silva-Santos P, Fonseca T, Tíscar PA, López Serrano FR, Andrés Abellán M, Martínez García E, Del Cerro Barja A (2010) Modelling Spanish black pine postdispersal seed predation in Central-eastern Spain. Forest Syst 19:393–403CrossRef
34. Lucas-Borja ME, Fonseca T, Parresol B, Silva-Santos P, García-Morote FA, Tíscar PA (2011) Modelling Spanish black pine seedling emergence: establishing management strategies for endangered forest areas. For Ecol Manag 262:195–202CrossRef
35. Lucas-Borja ME, Fonseca Fidalgo T, Linares JC, García-Morote FA, López-Serrano FR (2012) Does the recruitment pattern of Spanish black pine (Pinus nigra Arn ssp. salzmannii) change the regeneration niche over the early life cycle of individuals? For Ecol Manag 284:93–99CrossRef
36. Madrigal J, Martínez E, Hernando C, Guijarro M, Díez C, Vega JA, Pérez-Gorostiaga P, Fonturbel MT, Cuiñas P, Alonso M, Beloso MC (2004) Respuesta a corto plazo del regenerado post-incendio de Pinus pinaster Ait. Aclareos mecanizados intensos. Silva Lusitana 12:1–14
37. Maestre FT, Cortina J (2004) Are Pinus halepensis plantations useful as a restoration tool in semiarid Mediterranean areas? For Ecol Manag 198:303–317CrossRef
38. Maestre FT, Valladares F, Reynolds JF (2005) Is the change of plant-plant interactions with abiotic stress predictable? A meta-analysis of field results in arid environments. J Ecol 93:748–757CrossRef
39. Maher EL, Germino MJ, Hasselquist NJ (2005) Interactive effects of tree and herb cover on survivorship, physiology, and microclimate of conifer seedlings at the alpine tree-line ecotone. Can J For Res 35:567–574CrossRef
40. Michalet R, Brooker RW, Cavieres LA, Kikvidze Z, Lortie CJ, Pugnaire FI, Valiente-Banuet A, Callaway RM (2006) Do biotic interactions shape both sides of the humped-back model of species richness in plant communities? Ecol Lett 9:767–773CrossRefPubMed
41. Michalet R, Le Bagousse-Pinguet Y, Maalouf JP, Lortie CJ (2014) Two alternatives to the stress-gradient hypothesis at the edge of life: the collapse of facilitation and the switch from facilitation to competition. J Veg Sci 25:609–613CrossRef
42. Padilla FM, Pugnaire FI (2007) Rooting depth and soil moisture control Mediterranean woody seedling survival during drought. Funct Ecol 21:449–489CrossRef
43. Page LM, Cameron AD, Clarke GC (2001) Influence of overstorey basal area on density and growth of advance regeneration of Sitka spruce in variably thinned stands. For Ecol Manag 151:25–35CrossRef
44. Pagès JP, Pache G, Joud D, Magnan N, Michalet R (2003) Direct and indirect effects of shade on four forest tree seedlings in the French Alps. Ecology 84:2741–2750CrossRef
45. Prévosto B, Amandier L, Quesney T, De Boisgelin G, Ripert C (2012) Regenerating mature Aleppo pine stands in fire-free conditions: site preparation treatments matter. For Ecol Manag 282:70–77CrossRef
46. Rodríguez-García E, Ordóñez C, Bravo F (2011) Effects of shrub and canopy cover on the relative growth rate of Pinus pinaster Ait. seedlings of different sizes. Ann For Sci 68:337–346CrossRef
47. Simard MJ, Bergeron Y, Sirois L (1998) Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate. J Veg Sci 9:575–582CrossRef
48. Soil Atlas of Europe (2005) European soil bureau network. European Commission, Luxembourg
49. Sthultz CM, Gehring CA, Whitham TG (2007) Shifts from competition to facilitation between a foundation tree and a pioneer shrub across spatial and temporal scales in a semiarid woodland. New Phytol 173:135–145. doi:10.​1111/​j.​1469-8137.​2006.​01915.​x CrossRefPubMed
50. Tíscar PA (2007) Dinámica de regeneración de Pinus nigra subsp. salzmannii al sur de su área de distribución: etapas, procesos y factores implicados. Investig Agrar: Sist Recur For 16:124–135
51. Tíscar PA, Linares JC (2014) Large-scale regeneration patterns of Pinus nigra Subsp. salzmannii: poor evidence of increasing facilitation across a drought gradient. Forests 5:1–20. doi:10.​3390/​f5010001 CrossRef
52. Tíscar PA, Lucas-Borja ME, Candel Pérez D (2011) Changes in the structure and composition of two Pinus nigrasubsp. salzmannii forests over a century of different silvicultural treatments. Forest Syst 20:525–535CrossRef
53. Valladares F (2004) Ecología del bosque mediterráneo en un mundo cambiante. Ministerio de Medio Ambiente. EGRAF, Madrid
54. Vicente-Serrano SM, Beguería S, López-Moreno JI (2010) A multiscalar drought index sensitive to global warming: the Standardized Precipitation Evapotranspiration Index. J Clim 23:1696–1718CrossRef

For further details log on website :
http://link.springer.com/article/10.1007/s13595-016-0550-9