Original articleFlowering, die-back and recovery of a semelparous woody bamboo in the Atlantic Forest
Highlights
► Flowering and death of bamboos may enhance tree seedlings recruitment in gaps. ► We studied the flowering pattern and seedling demography of a woody bamboo. ► The bamboo Chusquea ramosissima can re-colonize sites after die-back. ► The flowering pattern was asynchronous and occurred at different spatio-temporal scales. ► Seedling abundance of tree species was similar in flowered and non-flowered sites.
Introduction
Bamboo grasses (Bambusoideae) are important components of many forests, and influence the diversity and productivity of the ecosystems in which they are ubiquitous and abundant (Judziewicz et al., 1999). Several studies have described distinctive aspects of the role of bamboos in the structure and dynamics of different forest ecosystems (Veblen, 1982, Oliveira, 2000, Tabarelli and Mantovani, 2000, Griscom and Ashton, 2003, Griscom and Ashton, 2006, Wang et al., 2006; Campanello et al., 2007a). Not only the widespread occurrence of bamboos in woodlands and their ecological functions, but also their peculiar life cycle has drawn the attention of scientists. Most bamboo species studied so far, have long periods of vegetative growth between successive synchronized sexual reproduction events followed by the death of all plants at large spatial scale (Janzen, 1976, Makita, 1992, Makita et al., 1995, Pearson et al., 1994, Franklin, 2004). This phenomenon is known as a “bamboo flowering event”. In many cases bamboo die-back causes changes in forest dynamics and environmental conditions that can favor tree regeneration (Taylor et al., 1995, Taylor et al., 1996, Abe et al., 2001).
During bamboo flowering, large numbers of seeds are usually produced (Janzen, 1976, Gadgil and Prasad, 1984, González and Donoso, 1999). The rate of bamboo regeneration via the production of new seedlings can be spatially variable and influenced by forest canopy cover (e.g., Nakashizuka, 1988, Taylor et al., 2004). In forests where semelparous bamboos constitute an important component of the understory, flowering and die-back may have profound effects on the dynamics and structure of plant communities (Young, 1991, Taylor and Qin, 1992, González et al., 2002, Martins et al., 2004, Taylor et al., 2004, Holz and Veblen, 2006, Caccia et al., 2009), and also on the population dynamics of animal species that rely on bamboos for refuge or feeding (Janzen, 1976, Krater, 1997, Gallardo and Mercado, 1999, Kitzberger et al., 2007, Areta et al., 2009).
Despite the importance of bamboo species in tropical regions, the population dynamics of semelparous bamboos and its impacts on ecosystem structure and functioning in tropical and subtropical forests of the Neotropics are poorly known. Chusquea ramosissima Lindman is a native semelparous woody bamboo dominant in the semideciduous Atlantic Forest of northern Argentina. Its geographical distribution comprises eastern Brazil, north-eastern Argentina, eastern Paraguay, Bolivia and northern Uruguay (Judziewicz et al., 1999, Tropicos, 2011, NYBG virtual herbarium; Tropicos virtual herbarium), and is thus the most widespread species in the genus Chusquea (Judziewicz et al., 1999). The culms of this rhizomatous species can be either erect or scandent with solid internodes. It has distinctive culm leaves, and an unusual bud complement with three sizes of buds.
This bamboo species is an aggressive colonizer of disturbed sites, where it forms impenetrable thickets that inhibit gap-phase regeneration (Campanello et al., 2009, Montti et al., 2008, Montti, 2010), thus reducing tree diversity in the long term (Campanello et al., 2007a). Changes in the structure and composition may also affect other aspects of biodiversity. For example, red-brocket deer (Mazama spp.) and tapir (Tapirus terrestris) show habitat preference for forests with low densities of this bamboo (Ferrari, 2006, Gallardo et al., 2008, Di Bitetti et al., 2008).
The time necessary for bamboo populations to recover after die-back might be an important determinant of the forest dynamics where bamboo grasses are dominant in the understory (Judziewicz et al., 1999). In the year 2001 several areas with flowering culms of C. ramosissima were observed in north-eastern Argentina. The overall objective of the study was to assess the ecological significance of the flowering pattern and population recovery after die-back of the invasive C. ramosissima in the Atlantic Forest, one of the most endangered tropical ecosystems in the world (Olson and Dinerstein, 2002). The specific objectives were to (a) describe the flowering event of C. ramosissima at different spatio-temporal scales, (b) quantify bamboo seedling emergence, mortality and growth, and (c) assess the effect of flowering on the abundance of canopy tree seedlings. It was hypothesized that flowering and death of C. ramosissima opens a window of opportunity leaving space vacant for the recruitment of tree seedlings.
Section snippets
Study area
The study was conducted in the semideciduous Atlantic Forest of Misiones Province, Argentina. Mean annual precipitation in the area is about 2000 mm, evenly distributed throughout the year. Mean annual air temperature is 21 °C with monthly means of 25 °C in January and 15 °C in July, the warmest and coldest months of the year, respectively. Frost seldom occurs during winter (Gatti et al., 2008). The soils are mostly Alfisols and Ultisols derived from basaltic rocks containing high concentration
Flowering pattern
The flowering peak at La Elina occurred during 2003–2004 when more than 60% of total culms-ha−1 flowered within the 1-ha plot (Table 1). A relatively high proportion of non-flowering culms were observed every year intermingled with reproductive culms in each of the 4-m2 subplots. Flowering culms were not observed during the year 2008. The density of non-flowering culms-ha−1 for the year 2008 at La Elina was 12600, which was similar to the value for 2002 and 2003 (Table 1).
Flowering records in
Flowering at different spatio-temporal scales
The dominant bamboo C. ramosissima re-colonized areas left vacant after flowering and perpetuated in sites previously occupied by the species. At a landscape scale flowering sites in northern Misiones were interspersed with sites that did not flower from 2001 to November 2009 showing a spatially discontinuous flowering pattern. At a local scale, in any particular site (1–3 has), most of the culms flowered in a period of 3–5 years. Flowering and non-flowering culms were intermingled in small
Acknowledgments
We are grateful to F. Foletto, X. Londoño, G. Gatti and R. Ituarte for help and comments. Lynn Clark, F. Ely and Ignacio Areta made useful comments on the manuscript. We are also grateful to the NSF, USA, (grant number 02134174), Rufford Small Grant and CONICET (Argentina) for financial support and to the PNI and APSA S.A for logistic support and facilities.
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