Invasive Plant Species and Biomass Production in Savannas
Abstract
Savannas are the second largest biome accounting for c. 30% of terrestrial production. Tropical savannas are distributed largely in Africa, Australia and South America occurring between tropical forests and deserts. It is the coexistence of trees and grasses that make savannas unique. The structure of savannas or the ratio of trees to grasses which has
important implications on ecosystem productivity is determined by resource availability (rainfall and soil nutrients) and disturbances (fire and herbivory) also referred to as ‘drivers’. Resources influence the distribution and productivity of savanna vegetation while fire can alter vegetation structure via effects on the woody layer. Herbivory influences savannas structure and composition through its effects on nutrient cycling, seed dispersal and physical defoliation effects and may lead to expansion of the shrub layer. While ecologists agree the four drivers determine tree-grass balance the exact mechanisms are still debated with one school of thought emphasizing the importance of resources as ‘primary determinants’ in what are referred to as ‘competition models’ which basically invoke the classic niche separation mechanisms in resource acquisition. The other school of thought referred to as ‘demographic bottleneck models’ emphasizes the role of disturbances as the primary determinants through their effects on life history stages of trees. It’s been shown
however that at low levels of mean annual rainfall, precipitation governs the cover of trees and above a critical value disturbances prevent trees from forming a closed canopy.
Invasive species are considered to be non-native species that have been introduced outside
their normal range and are expanding in range causing ecological and economic harm and
can drastically alter the structure and composition of savannas. Most non-native species
introduced in savannas were for well intended commercial and ecological purposes such as
pasture and fodder improvement or rehabilitation of degraded areas. Even though patterns
of invasion can not be easily generalized, a trend is that African C4 grasses such as Melinis
minutiflora and Andropogon gayanus make up the most obnoxious invaders in the South
American and Australian savannas while in contrast neotropical trees and shrubs are among
the most successful invaders of African and Australian savannas such as Prosopis spp and
Lantana camara. Ecologists have persistently attempted to answer the question ‘what makes a
community susceptible to invasion’? Plant characteristics of the invader is an important
factor, plants introduced in savannas for improvement of pasture/fodder are generally
selected for aggressiveness/competitiveness compared to native species. Selected shrubs for
example tend to have fast growth, easy to propagate and often N fixers while grasses display aspects of higher resource use efficiency and greater tolerance to grazing. Ecological
disturbances such as heavy grazing can destroy native vegetation and favor unpalatable
invaders through effects on resource availability. Among other factors thought to enhance
invasibility is climate change and its synergistic interactions with elevated CO2 since most
invasive species have traits that allow them to respond strongly to elevated CO2.
Productivity levels of savannas are on a broad scale related to the relative proportion of
trees to grasses while precipitation is the most important factor with an almost linear
relationship to biomass production. Gaps and inconsistencies in savanna Net Primary
Productivity data collected over the years make spatial and temporal comparison difficult.
This paucity arises from the ‘evolution’ of methodologies in Net Primary Productivity (NPP)
determination from the earlier commonly used ‘peak biomass’ methods that grossly
underestimated NPP, through improvements incorporated in International Biological
Programme (IBP) studies in the 1970’s to further refinements in the United Nations
Environmental Programme (UNEP) grassland studies that made corrections for a wide
range of losses during the growth phase previously unaccounted for. Further gaps in data
are because most savanna productivity studies have focused on single species within the
community of study or lumped several species and rarely included both tree and grass
components. Comparison of non-native and native species prior to introduction was often
made through screening trials where the fodder trees were largely evaluated for
productivity, digestibility, nutritional value and soil amelioration among others. Selected
non-native woody species invariably had superior performance in growth parameters e.g
Prosopis juliflora produced up to 188% more in aboveground biomass than the valuable
indigenous Acacia tortilis in Senegal. Many screening trials also showed that despite slow
growth native tree species in most trials had other positive attributes and not all were
outperformed by non-natives and moreover only a small proportion of selected non-natives
became invasive. African C4 grasses introduced in the neotropics and Australia on account
of higher productivity have also altered fire regimes, hydrology and nutrient cycling for
example Andropogon gaynus invasion in Australia which can lead to a biomass load of over
300% compared to native species but has resulted in fires eight times more intense on
average. Invasive herbs just like grasses and trees can have negative impacts such as the bi-
annual unpalatable Ipomoea hildebrandtii which depresses native grass biomass production in
addition to changes in site hydrologic and nutrient dynamics patterns.
Can invasive species in savannas increase carbon sequestration? Given the rapid increase in
coverage of invasive species e.g Prosopis juliflora is already estimated to cover 500,000 and
700,000ha in Kenya and Ethiopia while vast areas in Columbia, Venezuela, Brazil and
Australia are dominated by higher yielding African C4 invasive grasses. An assessment of
several studies in forests, grasslands and wetlands showed that ecosystem productivity was
higher in invaded ecosystems. In savannas above ground carbon (C) stocks increases as the
proportion of trees increases relative to grasses. Soil carbon constitutes over two-thirds of
the global carbon found in terrestrial ecosystems. Net soil carbon stock in savannas is
regulated by inputs from primary productivity and heavy losses due to herbivory and fire.
It follows alteration of the C and N cycles by invasive species can vary carbon sequestration.
Alteration of the C cycle components in savannas is attributed to differences in
ecophysiological traits between the invasive and indigenous species. Some invasive species
traits that lead to increased sequestration include faster relative growth, deep rooting,
herbivore defense traits, faster litter decomposition and N fixation. However not all invasive
species have these traits some decrease sequestration by depressing N mineralization and having lower litter decomposition, more studies to enable the quantification of this process
in savannas are required.