Fi. Pugnaire et al., FACILITATION AND SUCCESSION UNDER THE CANOPY OF A LEGUMINOUS SHRUB, RETAMA-SPHAEROCARPA, IN A SEMIARID ENVIRONMENT IN SOUTH-EAST SPAIN, Oikos, 76(3), 1996, pp. 455-464
Retama sphaerocarpa is a leafless leguminous shrub found in most regio
ns of Spain, frequently with a growth of herbs in its understorey whic
h contrasts with the surrounding areas by its higher biomass and diver
sity. We examined the relationships between the shrub and the herbs un
derneath along a gradient of shrub age. A total of fifty individuals w
ere selected to fit five age classes and shrub characteristics, soil p
roperties and flora under the canopy were examined along the estimated
chronosequence. All shrub size variables increased with time, as did
the amount of nutrients stored by the shrub, but differences were ofte
n significant only between the three oldest classes. Concentration of
nitrogen and phosphorus in photosynthetic stems remained constant. but
nutrient pools in stem biomass increased with time. The shrubs change
d the soil environment under their canopies with age by ameliorating s
oil texture, nutrient content and capacitance of water. The improvemen
t was most pronounced between the two oldest classes (IV and V), and w
as probably due to the high biomass of perennial species in the unders
torey which increased the production of litter, and the interception o
f wind-blown dust. Plant diversity in the understorey increased with s
hrub age, likely due to a greater heterogeneity under larger canopies.
Drought-resistant species, typical of the open areas between shrubs,
were displaced from the centre of the understorey by taller, more mesi
c species over the age gradient. Retama sphaerocarpa shrubs benefited
from the increase of resources in the understorey and showed a higher
reproductive output with shrub age, but decreased at the beginning of
senescence. The indirect interactions between R. sphaerocarpa and its
understorey herbs could be considered as a two-way facilitation in whi
ch both partners benefit from their association.