Abstract
At the treeline in the Central European Alps, adverse climate conditions impair tree growth and cause krummholz formation of Swiss stone pine (Pinus cembra L.). Multi-stemmed trees (tree clusters) are frequently found in the treeline ecotone and are generally thought to originate from seed caches (multiple genets) of the European nutcracker (N. caryocatactes) or due to repeated damage of the leader shoot by browsing or mechanical stress (single genet). Additionally, lack of apical control can lead to upward bending of lateral branches, which may obscure single-genet origin if the lower branching points are overgrown by vegetation and the humus layer. The multi-stemmed growth form may serve as a means of protection against extreme environmental stress during winter, especially at wind-exposed sites, because leeward shoots are protected from, e.g., ice particle abrasion and winter desiccation. The aims of this study therefore were to analyze in an extensive field survey: (i) whether weak apical control may serve as a protection against winter stress; and (ii) to what extent the multi-stemmed growth form of P. cembra in the krummholz zone is originating from a single genet or multiple genets. To accomplish this, the growth habit of P. cembra saplings was determined in areas showing extensive needle damage caused by winter stress. Multi-stemmed saplings were assigned to single and multiple genets based on determination of existing branching points below the soil surface. The findings revealed that upward bending of lateral branches could protect saplings against winter stress factors, and, although multi-stemmed P. cembra trees were primarily found to originate from multiple genets (most likely seed caches), about 38% of tree clusters originated from upward bending of (partially) buried branches. The results suggest that weak apical control of P. cembra in the sapling stage might be an adaptation to increase survival rate under severe climate conditions prevailing above treeline during winter.
Highlights
The alpine treeline is a conspicuous climate-driven ecological boundary, which designates the upper elevational limit of tree growth [1,2,3]
Three growth forms of P. cembra can be distinguished: (1) single-trunk; (2) single genet multi-trunk caused by repeated damage of the leader shoot; and (3) multi-genet tree cluster attributable to multiple germination of seed caches [10]
Weak apical control may exist in P. cembra at the sapling stage, which can be explained by the competitive-sink hypothesis developed in [21]
Summary
The alpine treeline is a conspicuous climate-driven ecological boundary, which designates the upper elevational limit of tree growth [1,2,3]. Three growth forms of P. cembra can be distinguished: (1) single-trunk; (2) single genet multi-trunk caused by repeated damage of the leader shoot; and (3) multi-genet tree cluster attributable to multiple germination of seed caches [10]. The latter two forms are morphologically similar and generally thought to be distinguishable only by genetic analysis [11]. Most pine species have a dominant main stem and distinct lateral branches which grow shorter and more horizontal than the vertical leader shoot [15], i.e., strong apical control is exerted by the leader shoot. Shoot architecture of P. cembra is generally characterized by a vertical main trunk and upward bending of lateral branches
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