Regeneration and stand dynamics of Fitzroya cupressoides (Cupressaceae) forests of southern Chile’s Central Depression

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Abstract

Several populations of Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae, common name “alerce”), a threatened, long-lived conifer endemic to southern Chile and parts of Argentina, have recently been found in Chile’s Central Depression, where the species was thought to have been extirpated. The objective of this study was to determine, on eight sites in the Depression, Fitzroya’s regeneration behavior in relation to disturbance, its substrate and cover requirements for establishment, and whether regeneration is by seed or vegetative means. This objective required inspection of tree age-class distributions, analyses of microsite conditions and spatial distributions of Fitzroya individuals, and genetic analysis of Fitzroya trees using random amplified polymorphic DNA (RAPD) analysis. Age-class distributions indicated a “catastrophic” regeneration mode, in which seedling establishment follows stand-devastating disturbance, such as fire. Regeneration was both by seed and vegetative means (root sucker sprouting and layering), although overall, vegetative reproduction was more prevalent (63% of all regeneration). Regeneration from seed occurred primarily under open canopies (<40% cover) and on substrates of humus, moss beds, and coarse woody debris. Spatial analyses revealed clumping at various scales for Fitzroya seedlings, saplings, and trees. Several obvious clumps of trees were assumed to be sucker sprouts from cut stumps; however, RAPD analyses indicated they were of seed origin. Such clumping is likely due to the patchy co-occurrence of appropriate substrates and canopy covers. Despite substantial environmental differences between the Central Depression and the portions of the Coastal and Andean Cordilleras where Fitzroya is found, populations in all three regions show striking similarities in their regeneration and stand dynamics. Fitzroya’s longevity and at times abundant regeneration provide the biological basis for its persistence in Chile’s Central Depression.

Introduction

Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae, common name “alerce”) is a threatened, long-lived conifer endemic to the temperate rain forests of southern Chile and adjacent portions of Argentina. It can attain a size of up to 5 m in diameter and 50 m in height, often appearing as an emergent when mixed with other species (Lara, 1991). With a lifespan greater than 3600 years (Lara and Villalba, 1993), it is one of the longest-lived tree species worldwide. Historically, Fitzroya was found in a variety of forest types and site conditions at elevations ranging from sea level to over 1200 m. The over-exploitation of Fitzroya, owing to the beauty and decay-resistance of its wood, has left extensive areas degraded and has significantly reduced its original abundance. In addition, many former Fitzroya forests have been intentionally burned and converted to agricultural land. Fitzroya re-establishment on harvested sites, as well as on most burned sites, has been scarce or nonexistent. Because of these threats, in 1975 Fitzroya was listed on Appendix I of the Convention on International Trade in Endangered Species (CITES), which reduced harvesting pressure on the species by banning its commercial international trade. In 1976, the Chilean government declared every living Fitzroya a “national monument”, which provided in-country protection against further harvesting. More recently, the World Conservation Union, IUCN, has listed Fitzroya as endangered on their Red List of Threatened Plants (Oldfield et al., 1998).

In Chile, Fitzroya occurs in the Coastal Cordillera from ca. latitude 39°50′ to 42°35′, and in the Andean Cordillera from ca. 41° to 43°30′. It was once abundant in Chile’s Central Depression (which lies between the two Cordilleras, south and west of Llanquihue Lake; Fig. 1) before extensive logging began there in the mid-1800s. Over the last decade, several remnant Fitzroya populations have been located in Chile’s Central Depression, where the species was thought to have been extirpated (Veblen et al., 1976, Golte, 1996).

The current study was generated by field observations that Fitzroya has not re-established on much of its former habitat in the Depression, and on those sites where it currently exists, it exhibits a large amount of within- and between-site variability in seedling density. Compared to either Cordillera where Fitzroya occurs, the Depression has much lower rainfall, lower elevation, a longer growing season, and drastically different soils, which led us to speculate that its regeneration and stand dynamics may differ accordingly. Our objective was to determine the regeneration mode (Veblen (1992), the regeneration behavior in relation to disturbance), regeneration niche (Grubb (1977), the micro-environmental conditions under which a species becomes established), and regeneration mechanism (by seed or vegetative regeneration) of Fitzroya populations in the Central Depression. Given that these stands were only recently discovered, this is the first study of Fitzroya regeneration and stand dynamics in the Central Depression.

Section snippets

Study area

Chile’s Central Depression ranges between 50 and 75 km in width, and reaches nearly 200 m in elevation in its northern extreme. From Llanquihue Lake, it gradually descends southward until reaching sea level in the Gulf of Ancud. The Depression is characterized by an oceanic cool temperate climate. The nearest weather station in Puerto Montt registers a mean annual temperature of 11.5 °C and an annual rainfall of 1912 mm, with the winter months (June–August) receiving the most rain (300 mm per month

Age-class distributions

All stands except Fundo Nuñez showed single-cohort age-class distributions of Fitzroya (Fig. 2), indicating that the populations became established in one wave of regeneration that likely followed a catastrophic disturbance. Thus, Fitzroya’s regeneration mode in the Depression can be considered “catastrophic” (Veblen, 1992). The distinct cohort structure of Fitzroya results from its shade intolerance: once a stand matures and the canopy closes, recruitment of new individuals drops off

Conclusions

Earlier authors interpreted the lack of Fitzroya regeneration under its own canopy or in clear-cut areas as regeneration failure of this species (see Veblen et al. (1976) and Veblen and Ashton (1982) for discussion). More recent research has demonstrated adequate regeneration of Fitzroya in the Coastal and Andean Cordilleras given the appropriate site conditions (Veblen and Ashton, 1982, Lara, 1991, Donoso et al., 1993, Parker and Donoso, 1993, Lara et al., 1999). The results of this current

Acknowledgements

This research was funded by a UK Darwin Initiative grant administered through the Conifer Conservation Programme of the Royal Botanic Garden, Edinburgh, and the University of Edinburgh, Scotland. We thank Nelson Garcı́a, José Camilo Gómez, Alfredo Núñez, and Nelson Velázquez for granting access to their properties, Bernardo Escobar, Sebastián Dı́az, Oscar Larraı́n, Carlos Lehnebach, and Rodrigo Vergara for assistance in the field, Alexia Wolodarsky-Franke for her assistance with the processing

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    Present address: Departamento de Ecologı́a, Universidad de Salamanca, Salamanca, Spain.

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    Present address: Applied Genetics, John Innes Centre, Norwich, NR3 7UH, UK.

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    Present address: UNEP, World Conservation Monitoring Centre, 219 Huntingdon Road, Cambridge CB3 0DL, UK.

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