Resprouting and multi-stemming and the role of the persistence niche in the structure and dynamics of subtropical coastal dune forest in KwaZulu-Natal province, South Africa.

Abstract

Resprouting is an important means of plant regeneration especially under conditions that do not favour regeneration through seeding such as frequent disturbances, low productivity, unfavourable soil conditions, extreme cold and limited understorey light availability. Sprouts may be advantageous over seedlings because they have higher survival and growth rates than seedlings, since they use resources from parent plants unlike seedlings that have to acquire their own resources. Resprouting is well documented for ecosystems that experience severe disturbances that damage aboveground biomass. For example, resprouting is important for plant persistence against fire in fire-prone savannas and Mediterranean shrub-lands, and hurricanes and cyclones in tropical forests. In these ecosystems, resprouting often results in multi-stemming, because this dilutes the risk of damage among many stems, improving the chances of individual survival. This study was conducted at coastal dune forest at Cape Vidal in north-eastern South Africa, where there is a high incidence of multi-stemmed trees due to resprouting in response to chronic disturbances of low severity. This study examines (1) the importance of resprouting to tree survival and dynamics in an environment where disturbance severity is low but pervasive, and (2) how this resprouting strategy differs from the more familiar sprouting response to severe disturbances such as fire and hurricanes. Analysis of the relationship between multi-stemming and a number of disturbances potentially causing multi-stemming revealed that stem leaning and substrate erosion were the most important disturbances associated with multi-stemming. There were fewer multistemmed trees on dune slacks that had a stable substrate and were protected from sea winds than on dune crests and slopes that had unstable substrate and were exposed to sea winds. Trees resprouted and became multi-stemmed from an early stage to increase their chances of survival against leaning caused by strong sea winds and erosion, and occasional slumping of the unstable dune sand substrate. These low severity disturbances are persistent and are referred to as chronic disturbances in this thesis. As a result of these chronic disturbances, both single and multi-stemmed trees had short stature because taller individuals that emerged above the tree canopy would be exposed to wind damage. Under chronic disturbances plants may manifest a phylogenetically determined sprouting response. However, in this study resprouting and multi-stemming were the results of the tree-disturbance interaction and not a property of a plant or species and were not phylogenetically constrained. Because the disturbances are predominantly of low severity, leaning trees were able to regain the vertical orientation of the growing section by turning upward (a process referred to as ‘turning up’ in this study) and hence survive without resprouting. Species that were prone to turning upward had a low incidence and degree of leaning of their individuals, low frequency of loss of primary stems and high abundance of individuals. Although turning up is less costly to the individual than resprouting, it could only be used by leaning trees that had small angles of inclination and were not eroded. High intensities of the latter require that individuals resprout to survive. The form and function of resprouting varied between seedlings and juvenile and mature trees. Resprouting in seedlings resulted in a single replacement shoot, unlike sprouting in juvenile and mature trees that resulted in multi-stemmed trees. Like sprouting in juvenile and mature trees, sprouting in seedlings was not phylogenetically constrained. Resprouting in seedlings increased seedling persistence; hence species with more sprout seedlings had larger individual seedlings and seedling banks. Resprouting in seedlings increased the chances of seedling recruitment, whereas resprouting in juvenile and mature trees increased the chances of an established plant maintaining its position in the habitat. After disturbances of high severity, which destroy the photosynthesizing parts, plants resprout using carbohydrates stored below- or above ground. In this study, good resprouters stored more carbohydrates both below- and above ground than poor resprouters. The carbohydrates were mobilized for resprouting after disturbance. More carbohydrates were stored in stems than in roots because the prevailing disturbances were mostly of low severity and hence above ground resources were readily available. Similar to storage by plants in severely disturbed habitats, carbohydrates were stored by reserve formation, which competes for carbohydrates with growth and maintenance and forms permanent storage, rather than accumulation, which temporarily stores carbohydrates in excess of demands for growth and maintenance. Stored carbohydrates are not necessary for resprouting of plants after disturbances of low severity because they can resprout using resources remobilized directly from the disturbed photosynthesizing parts. However, in this study, stored carbohydrates served as a bet-hedge against occasional severe disturbances that occurred in addition to chronic disturbances. Allocation of carbohydrates to permanent storage diverts them from growth and reproduction and hence good resprouters had lower growth rates, seed output, seed size and seedling recruitment than poor resprouters. However, the costs of these traits that resulted in low recruitment from seed by good resprouters, were compensated for by high persistence of established individuals of good resprouters through recruitment of sprout stems. This study demonstrates that resprouting is not only advantageous in severely disturbed environments, but also in environments where disturbances are of low severity but nevertheless confer an advantage on individuals that persist. Thus in forest environments where aboveground biomass is seldom destroyed and individuals are relatively long-lived, resprouting can confer significant fitness and selective advantage on individuals.