Inquiline predator increases nutrient-cycling efficiency of Nepenthes rafflesiana pitchers.

The modified-leaf pitchers of Nepenthes rafflesiana pitcher plants are aquatic, allochthonous ecosystems that are inhabited by specialist inquilines and sustained by the input of invertebrate prey. Detritivorous inquilines are known to increase the nutrient-cycling efficiency (NCE) of pitchers but it is unclear whether predatory inquilines that prey on these detritivores decrease the NCE of pitchers by reducing detritivore populations or increase the NCE of pitchers by processing nutrients that may otherwise be locked up in detritivore biomass. Nepenthosyrphus is a small and poorly studied genus of hoverflies and the larvae of one such species is a facultatively detritivorous predator that inhabits the pitchers of N. rafflesiana. We fitted a consumer-resource model to experimental data collected from this system. Simulations showed that systems containing the predator at equilibrium almost always had higher NCEs than those containing only prey (detritivore) species. We showed using a combination of simulated predator/prey exclusions that the processing of the resource through multiple pathways and trophic levels in this system is more efficient than that accomplished through fewer pathways and trophic levels. Our results thus support the vertical diversity hypothesis, which predicts that greater diversity across trophic levels results in greater ecosystem functioning.

Digestive mutualism in a pitcher plant supports the monotonic rather than hump-shaped stress-gradient hypothesis model.

The stress-gradient hypothesis (SGH) predicts that the strength and frequency of facilitative interactions increase monotonically with increasing environmental stress, but some empirical studies have found this decrease at extreme stress levels, suggesting a hump-shaped SGH instead. However, empirical studies of the SGH are often hindered by confounding resource and non-resource stress gradients. Nepenthes pitcher plants trap animal prey using modified-leaf pitfall traps which are also inhabited by organisms known as inquilines. Inquilines may assist pitchers in the digestion of trapped prey. This interaction is known as a digestive mutualism and is both mutualistic and facilitative by definition. Inquiline species may also facilitate each other via processing chain commensalisms. We used in vitro experiments to examine the isolated effect of resource stress on the outcomes of two facilitative interactions: (i) digestive mutualism-facilitation of pitcher nutrient sequestration by two inquiline dipteran larvae, culicids and phorids; (ii) processing chain commensalism-facilitation between these two inquiline taxa. The net nutritional benefit of phorids on N. gracilis was found to conform more to a monotonic rather than hump-shaped SGH model. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were weak. These findings provide compelling evidence that changes in facilitation along an isolated resource stress gradient conform to the predictions of the monotonic SGH model rather than that of the revised hump-shaped model, and highlight the importance of isolating stress gradients in empirical tests of the SGH.

A dipteran larva-pitcher plant digestive mutualism is dependent on prey resource digestibility.

Positive species interactions tend to be context dependent. However, it is difficult to predict how benefit in a mutualism changes in response to changing contexts. Nepenthes pitcher plants trap animal prey using leaf pitfall traps known as pitchers. Many specialized inquiline organisms inhibit these pitchers, and are known to facilitate the digestion of prey carcasses in them. Nepenthes gracilis traps diverse arthropod prey taxa, which are likely to differ greatly in the ease with which they may be digested, independently of inquilines, by plant enzymes. In this study, we used in vitro experiments to compare the nutritional benefit provided by phorid (scuttle fly) and culicid (mosquito) dipteran larvae to their host, N. gracilis, and to each other. The effects of phorids on N. gracilis nutrient sequestration were very variable, being positive for large prey which have low digestibility, but negative for small prey which are highly digestible. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were not significantly altered by prey type. These results show that a digestive mutualism is highly dependent on the digestibility of the resource-a context dependency that conforms well to the predictions of the stress-gradient hypothesis in facilitation research. Our findings have significant implications for many other digestive mutualisms, and also suggest that greater insights may be gained from the synthesis of concepts between the fields of mutualism and facilitation research.