Interspecific variation in prey capture behavior by co-occurring Nepenthes pitcher plants: evidence for resource partitioning or sampling-scheme artifacts?

Pitcher plants of the genus Nepenthes capture a wide range of arthropod prey for nutritional benefit, using complex combinations of visual and olfactory signals and gravity-driven pitfall trapping mechanisms. In many localities throughout Southeast Asia, several Nepenthes different species occur in mixed populations. Often, the species present at any given location have strongly divergent trap structures and preliminary surveys indicate that different species trap different combinations of arthropod prey, even when growing at the same locality. On this basis, it has been proposed that co-existing Nepenthes species may be engaged in niche segregation with regards to arthropod prey, avoiding direct competition with congeners by deploying traps that have modifications that enable them to target specific prey types. We examined prey capture among 3 multi-species Nepenthes populations in Borneo, finding that co-existing Nepenthes species do capture different combinations of prey, but that significant interspecific variations in arthropod prey combinations can often be detected only at sub-ordinal taxonomic ranks. In all lowland Nepenthes species examined, the dominant prey taxon is Formicidae, but montane Nepenthes trap few (or no) ants and 2 of the 3 species studied have evolved to target alternative sources of nutrition, such as tree shrew feces. Using similarity and null model analyses, we detected evidence for niche segregation with regards to formicid prey among 5 lowland, sympatric Nepenthes species in Sarawak. However, we were unable to determine whether these results provide support for the niche segregation hypothesis, or whether they simply reflect unquantified variation in heterogeneous habitats and/or ant communities in the study sites. These findings are used to propose improvements to the design of field experiments that seek to test hypotheses about targeted prey capture patterns in Nepenthes.

Capture mechanism in Palaeotropical pitcher plants (Nepenthaceae) is constrained by climate.

BACKGROUND AND AIMS: Nepenthes (Nepenthaceae, approx. 120 species) are carnivorous pitcher plants with a centre of diversity comprising the Philippines, Borneo, Sumatra and Sulawesi. Nepenthes pitchers use three main mechanisms for capturing prey: epicuticular waxes inside the pitcher; a wettable peristome (a collar-shaped structure around the opening); and viscoelastic fluid. Previous studies have provided evidence suggesting that the first mechanism may be more suited to seasonal climates, whereas the latter two might be more suited to perhumid environments. In this study, this idea was tested using climate envelope modelling. METHODS: A total of 94 species, comprising 1978 populations, were grouped by prey capture mechanism (large peristome, small peristome, waxy, waxless, viscoelastic, non-viscoelastic, 'wet' syndrome and 'dry' syndrome). Nineteen bioclimatic variables were used to model habitat suitability at approx. 1 km resolution for each group, using Maxent, a presence-only species distribution modelling program. KEY RESULTS: Prey capture groups putatively associated with perhumid conditions (large peristome, waxless, viscoelastic and 'wet' syndrome) had more restricted areas of probable habitat suitability than those associated putatively with less humid conditions (small peristome, waxy, non-viscoelastic and'dry' syndrome). Overall, the viscoelastic group showed the most restricted area of modelled suitable habitat. CONCLUSIONS: The current study is the first to demonstrate that the prey capture mechanism in a carnivorous plant is constrained by climate. Nepenthes species employing peristome-based and viscoelastic fluid-based capture are largely restricted to perhumid regions; in contrast, the wax-based mechanism allows successful capture in both perhumid and more seasonal areas. Possible reasons for the maintenance of peristome-based and viscoelastic fluid-based capture mechanisms in Nepenthes are discussed in relation to the costs and benefits associated with a given prey capture strategy.

Tuning of color contrast signals to visual sensitivity maxima of tree shrews by three Bornean highland Nepenthes species.

Three species of Nepenthes pitcher plants (Nepenthes rajah, Nepenthes lowii and Nepenthes macrophylla) specialize in harvesting nutrients from tree shrew excreta in their pitchers. In all three species, nectaries on the underside of the pitcher lid are the focus of the tree shrews' attention. Tree shrews are dichromats, with visual sensitivity in the blue and green wavebands. All three Nepenthes species were shown to produce visual signals, in which the underside of the pitcher lid (the area of highest nectar production) stood out in high contrast to the adjacent area on the pitcher (i.e., was brighter), in the blue and green wavebands visible to the tree shrews. N. rajah showed the tightest degree of "tuning," notably in the green waveband. Conversely, pitchers of Nepenthes burbidgeae, a typical insectivorous species sympatric with N. rajah, did not produce a color pattern tuned to tree shrew sensitivity maxima.

Mutualism between tree shrews and pitcher plants: perspectives and avenues for future research.

Three species of Nepenthes pitcher plants from Borneo engage in a mutualistic interaction with mountain tree shrews, the basis of which is the exchange of nutritional resources. The plants produce modified "toilet pitchers" that produce copious amounts of exudates, the latter serving as a food source for tree shrews. The exudates are only accessible to the tree shrews when they position their hindquarters over the pitcher orifice. Tree shrews mark valuable resources with faeces and regularly defecate into the pitchers when they visit them to feed. Faeces represent a valuable source of nitrogen for these Nepenthes species, but there are many facets of the mutualism that are yet to be investigated. These include, but are not limited to, seasonal variation in exudate production rates by the plants, behavioral ecology of visiting tree shrews, and the mechanism by which the plants signal to tree shrews that their pitchers represent a food source. Further research into this extraordinary animal-plant interaction is required to gain a better understanding of the benefits to the participating species.

Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size.

*Three Bornean pitcher plant species, Nepenthes lowii, N. rajah and N. macrophylla, produce modified pitchers that 'capture' tree shrew faeces for nutritional benefit. Tree shrews (Tupaia montana) feed on exudates produced by glands on the inner surfaces of the pitcher lids and defecate into the pitchers. *Here, we tested the hypothesis that pitcher geometry in these species is related to tree shrew body size by comparing the pitcher characteristics with those of five other 'typical' (arthropod-trapping) Nepenthes species. *We found that only pitchers with large orifices and lids that are concave, elongated and oriented approximately at right angles to the orifice capture faeces. The distance from the tree shrews' food source (that is, the lid nectar glands) to the front of the pitcher orifice precisely matches the head plus body length of T. montana in the faeces-trapping species, and is a function of orifice size and the angle of lid reflexion. *Substantial changes to nutrient acquisition strategies in carnivorous plants may occur through simple modifications to trap geometry. This extraordinary plant-animal interaction adds to a growing body of evidence that Nepenthes represents a candidate model for adaptive radiation with regard to nitrogen sequestration strategies.