Alternative stable states explain unpredictable biological control of Salvinia molesta in Kakadu.

Suppression of the invasive plant Salvinia molesta by the salvinia weevil is an iconic example of successful biological control. However, in the billabongs (oxbow lakes) of Kakadu National Park, Australia, control is fitful and incomplete. By fitting a process-based nonlinear model to thirteen-year data sets from four billabongs, here we show that incomplete control can be explained by alternative stable states--one state in which salvinia is suppressed and the other in which salvinia escapes weevil control. The shifts between states are associated with annual flooding events. In some years, high water flow reduces weevil populations, allowing the shift from a controlled to an uncontrolled state; in other years, benign conditions for weevils promote the return shift to the controlled state. In most described ecological examples, transitions between alternative stable states are relatively rare, facilitated by slow-moving environmental changes, such as accumulated nutrient loading or climate change. The billabongs of Kakadu give a different manifestation of alternative stable states that generate complex and seemingly unpredictable dynamics. Because shifts between alternative stable states are stochastic, they present a potential management strategy to maximize effective biological control: when the domain of attraction to the state of salvinia control is approached, augmentation of the weevil population or reduction of the salvinia biomass may allow the lower state to trap the system.

Variability and Parasitoid Foraging Efficiency: A Case Study of Pea Aphids and Aphidius ervi.

When a parasitoid is searching for hosts, not all hosts are equally likely to be attacked. This variability in attack probability may affect the parasitoid functional response. Using a collection of experiments, we quantified the functional response of Aphidius ervi (Hymenoptera: Braconidae), an insect parasitoid of the pea aphid Acyrthosiphon pisum (Homoptera: Aphididae). We measured variability in the number of hosts attacked by a foraging parasitoid both among plants and within plants. At the first scale, A. ervi, searching among plants containing different numbers of aphids, showed both aphid-density-dependent and aphid-density-independent variability in the number of aphids attacked per plant. Within plants, A. ervi selectively attacked second and third instar aphids relative to other instars. Furthermore, there was variability in the susceptibility of attack among aphids independent of instar. Variability in attack rates among aphids both among and within plants decreased parasitoid foraging efficiency, with the greatest decrease caused by among-plant variability. Furthermore, the decrease in foraging efficiency was greatest when the average number of aphids per plant was low, thereby transforming a strong Type II functional response into one approaching Type I.

Differential influence of clonal integration on morphological and growth responses to light in two invasive herbs.

BACKGROUND AND AIMS: In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions. METHODS: In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85% shade and their morphological and growth responses were assessed. KEY RESULTS: The influence of clonal integration on the light reaction norm (connection×light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection×light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mother plants invest in daughter ramets prior to their own branches. CONCLUSIONS: Clonal integration modifies light reaction norms of morphological and growth traits in a species-specific manner for A. philoxeroides and P. canescens, but it improves the establishment of juvenile ramets of both species in light-limiting environments by promoting their growth during early development. This factor may be partially responsible for their ability to successfully colonize native plant communities.