Carnivore reintroduction is often expected to revert community and ecosystem properties to their natural states via risk effects and the direct killing of prey. Because large carnivore extirpation and reintroduction are usually believed to have symmetric and offsetting effects, fulfilling this "assumption of reciprocity" is crucial to realizing the potential of large carnivores to passively restore community structure and ecosystem function. We were unable to find any study in which the assumption of reciprocity was rigorously tested in predator-prey systems featuring large carnivores, their ungulate prey, and primary producers through a comprehensive literature search. We therefore used studies involving (1) the reintroduction of any native apex predator (including but not limited to large mammalian carnivores); and (2) the removal of any introduced apex predator (also including but not limited to large mammalian carnivores) to examine the assumption of reciprocity. Reintroduction of native apex predators did not affect any of four trophic groups (mesopredator, omnivore, herbivore, primary producer) in a positive or negative way, but removal of introduced apex predators consistently increased the abundance and biomass of mesopredators. Further, outcomes of apex predator reintroduction and removal were variable across systems, regardless of system complexity (ranging from single predator-single prey to multiple predator-multiple prey systems). We suggest that the assumption of reciprocity-in which predator extirpation and reintroduction are believed to have consistent, counterbalancing effects-is unsupported by current evidence, and perhaps unrealistic. We discuss potential directions for research that might illuminate when and why the assumption of reciprocity would be valid.
Across the globe, biological invasions have disrupted mutualisms, producing reverberating consequences for ecosystems. Although invasive species frequently trigger mutualism disruptions, few studies have quantified the demographic mechanisms by which mutualism breakdown may generate population effects. In a Kenyan savanna, the invasive big-headed ant (Pheidole megacephala) has disrupted a foundational mutualism between the monodominant whistling-thorn tree (Acacia drepanolobium) and native ants (Crematogaster spp.) that deter browsing by large mammalian herbivores. We conducted experiments to quantify the demographic consequences of this mutualism disruption in the presence and absence of large mammalian herbivores. Invasion by P. megacephala exacerbated population declines of A. drepanolobium, primarily through decreased survival and reproduction of adult trees. However, these fitness reductions were small compared to those resulting from the presence of large mammalian herbivores, which negatively impacted growth and survival. Contrary to expectation, the expulsion of metabolically costly Crematogaster mutualists by P. megacephala did not result in higher population growth rates for trees protected from large mammalian herbivores. Our results suggest that invasive P. megacephala may impose a direct metabolic cost to trees exceeding that of native mutualists while providing no protection from browsing by large mammalian herbivores. Across landscapes, we expect that invasion by P. megacephala will reduce A. drepanolobium populations, but that the magnitude and demographic pathways of this effect will hinge on the presence and abundance of browsers.
Biological invasions can lead to the reassembly of communities and understanding and predicting the impacts of exotic species on community structure and functioning are a key challenge in ecology. We investigated the impact of a predatory species of invasive ant, Pheidole megacephala, on the structure and function of a foundational mutualism between Acacia drepanolobium and its associated acacia‐ant community in an East African savanna. Invasion by P. megacephala was associated with the extirpation of three extrafloral nectar‐dependent Crematogaster acacia ant species and strong increases in the abundance of a competitively subordinate and locally rare acacia ant species, Tetraponera penzigi, which does not depend on host plant nectar. Using a combination of long‐term monitoring of invasion dynamics, observations and experiments, we demonstrate that P. megacephala directly and indirectly facilitates T. penzigi by reducing the abundance of T. penzigi’s competitors (Crematogaster spp.), imposing recruitment limitation on these competitors, and generating a landscape of low‐reward host plants that favor colonization and establishment by the strongly dispersing T. penzigi. Seasonal variation in use of host plants by P. megacephala may further increase the persistence of T. penzigi colonies in invaded habitat. The persistence of the T. penzigi–A. drepanolobium symbiosis in invaded areas afforded host plants some protection against herbivory by elephants (Loxodonta africana), a key browser that reduces tree cover. However, elephant damage on T. penzigi‐occupied trees was higher in invaded than in uninvaded areas, likely owing to reduced T. penzigi colony size in invaded habitats. Our results reveal the mechanisms underlying the disruption of this mutualism and suggest that P. megacephala invasion may drive long‐term declines in tree cover, despite the partial persistence of the ant–acacia symbiosis in invaded areas.
Quantifying tree biomass is an important research and management goal across many disciplines. For species that exhibit predictable relationships between structural metrics (e.g. diameter, height, crown breadth) and total weight, allometric calculations produce accurate estimates of above-ground biomass. However, such methods may be insufficient where inter-individual variation is large relative to individual biomass and is itself of interest (for example, variation due to herbivory). In an East African savanna bushland, we analysed photographs of small (<5 m) trees from perpendicular angles and fixed distances to estimate above-ground biomass. Pixel area of trees in photos and diameter were more strongly related to measured, above-ground biomass of destructively sampled trees than biomass estimated using a published allometric relation based on diameter alone (R2 = 0.86 versus R2 = 0.68). When tested on trees in herbivore-exclusion plots versus unfenced (open) plots, our predictive equation based on photos confirmed higher above-ground biomass in the exclusion plots than in unfenced (open) plots (P < 0.001), in contrast to no significant difference based on the allometric equation (P = 0.43). As such, our new technique based on photographs offers an accurate and cost-effective complement to existing methods for tree biomass estimation at small scales with potential application across a wide variety of settings.
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