Ants provide nutritional and defensive benefits to the carnivorous plant Sarracenia minor

Ants can have important, but sometimes unexpected, effects on the plants they associate with. For carnivorous plants, associating with ants may provide defensive benefits in addition to nutritional ones. We examined the effects of increased ant visitation and exclusion of insect prey from pitchers of the hooded pitcher plant Sarracenia minor, which has been hypothesized to be an ant specialist. Visitation by ants was increased by placing PVC pipes in the ground immediately adjacent to 16 of 32 pitcher plants, which created nesting/refuge sites. Insects were excluded from all pitchers of 16 of the plants by occluding the pitchers with cotton. Treatments were applied in a 2 × 2 factorial design in order to isolate the hypothesized defensive benefits from nutritional ones. We recorded visitation by ants, the mean number of ants captured, foliar nitrogen content, plant growth and size, and levels of herbivory by the pitcher plant mining moth Exyra semicrocea. Changes in ant visitation and prey capture significantly affected nitrogen content, plant height, and the number of pitchers per plant. Increased ant visitation independent of prey capture reduced herbivory and pitcher mortality, and increased the number of pitchers per plant. Results from this study show that the hooded pitcher plant derives a double benefit from attracting potential prey that are also capable of providing defense against herbivory.     

Nectar,not colour,may lure insects to their death

We experimentally demonstrate in the field that prey of the carnivorous plant Sarracenia purpurea are attracted to sugar, not to colour. Prey capture (either all taxa summed or individual common taxa considered separately) was not associated with total red area or patterning on pitchers of living pitcher plants. We separated effects of nectar availability and coloration using painted ‘pseudopitchers’, half of which were coated with sugar solution. Unsugared pseudopitchers captured virtually no prey, whereas pseudopitchers with sugar solution captured the same amount of prey as living pitchers. In contrast to a recent study that associated red coloration with prey capture but that lacked controls for nectar availability, we infer that nectar, not colour, is the primary means by which pitcher plants attract prey.     

Contribution of pitcher fragrance and fluid viscosity to high prey diversity in a Nepenthes carnivorous plant from borneo

Mechanisms that improve prey richness in carnivorous plants may involve three crucial phases of trapping:attraction, capture and retention.Nepenthes rafflesiana var. typica is an insectivorous pitcher plant that is widespread in northern Borneo.It exhibits ontogenetic pitcher dimorphism with the upper pitchers trapping more flying prey than the lower pitchers.While this difference in prey composition has been ascribed to differences in attraction,the contribution of capture and retention has been overlooked.This study focused on distinguishing between the prey trapping mechanisms, and assessing their relative contribution to prey diversity.Arthropod richness and diversity of both visitors and prey in the two types of pitchers were analysed to quantify the relative contribution of attraction to prey trapping.Rate of insect visits to the different pitcher parts and the presence or absence of a sweet fragrance was recorded to clarify the origin and mechanism of attraction.The mechanism of retention was studied by insect bioassays and measurements of fluid viscosity. Nepenthes rafflesiana was found to trap a broader prey spectrum than that previously described for any Nepenthes species,with the upper pitchers attracting and trapping a greater quantity and diversity of prey items than the lower pitchers.Capture efficiency was low compared with attraction or retention efficiency.Fragrance of the peristome,or nectar rim,accounted mainly for the observed non-specific, better prey attraction by the upper pitchers, while the retentive properties of the viscous fluid in these upper pitchers arguably explains the species richness of their flying prey.The pitchers of N. rafflesiana are therefore more than simple pitfall traps and the digestive fluid plays an important yet unsuspected role in the ecological success of the species.     

Evidence for alternative trapping strategies in two forms of the pitcher plant, Nepenthes rafflesiana

Nepenthes pitchers are specialized leaves that function as insect traps. Several pitcher components may contribute to trapping, including the pitcher fluid, slippery wax crystals and downward-pointing epidermal cells on the inner pitcher wall, and the wetness-dependent pitcher rim (peristome), but the relative importance of these traits is unclear. Mechanisms of prey capture and retention in the field were investigated by quantifying the effect of 'knock-out' manipulations of individual pitcher structures, and by testing the ability of pitcher fluids and water to retain insects. Two forms of Nepenthes rafflesiana Jack ('elongate' and 'typical') with contrasting combinations of pitcher traits were compared. Wax crystals on the inner pitcher wall were found to be the most important trapping structure in the elongate form, whereas the typical form relied primarily on the peristome. The pitcher fluids of both forms, differing markedly in the degree of viscoelasticity, retained significantly more ants than water. The present results show that pitcher plants utilize several mechanisms for prey capture and retention, varying in efficiency and relative importance between forms. It is proposed that these differences represent alternative prey capture strategies that may provide a mechanism to reduce competition and facilitate species co-existence in nutrient-limited habitats.     

Effect of pitcher age on trapping efficiency and natural prey capture in carnivorous Nepenthes rafflesiana plants

Background and Aims

Nepenthes pitchers are sophisticated traps that employ a variety of mechanisms to attract, capture and retain prey. The underlying morphological structures and physiological processes are subject to change over the lifetime of a pitcher. Here an investigation was carried out on how pitcher properties and capture efficiency change over the first 2 weeks after pitcher opening.

Methods

Prey capture, trapping efficiency, extrafloral nectar secretion, pitcher odour, as well as pH and viscoelasticity of the digestive fluid in N. rafflesiana pitchers were monitored in the natural habitat from pitcher opening up to an age of 2 weeks.

Key Results

Pitchers not only increased their attractiveness over this period by becoming more fragrant and secreting more nectar, but also gained mechanical trapping efficiency via an enhanced wettability of the upper pitcher rim (peristome). Consistently, natural prey capture was initially low and increased 3–6 d after opening. It was, however, highly variable within and among pitchers. At the same time, the pH and viscoelasticity of the digestive fluid decreased, suggesting that the latter is not essential for effective prey capture.

Conclusions

Prey capture and attraction by Nepenthes are dynamic processes strongly influenced by the changing properties of the pitcher. The results confirm insect aquaplaning on the peristome as the main capture mechanism in N. rafflesiana.Key words: Carnivorous plants, pitcher development, prey attraction, prey capture, insect aquaplaning, extrafloral nectar, Nepenthes rafflesiana     

Fenestration: a window of opportunity for carnivorous plants

A long-standing but controversial hypothesis assumes that carnivorous plants employ aggressive mimicry to increase their prey capture success. A possible mechanism is that pitcher plants use aggressive mimicry to deceive prey about the location of the pitcher''s exit. Specifically, species from unrelated families sport fenestration, i.e. transparent windows on the upper surfaces of pitchers which might function to mimic the exit of the pitcher. This hypothesis has not been evaluated against alternative hypotheses predicting that fenestration functions to attract insects from afar. By manipulating fenestration, we show that it does not increase the number of Drosophila flies or of two ant species entering pitchers in Sarracenia minor nor their retention time or a pitcher''s capture success. However, fenestration increased the number of Drosophila flies alighting on the pitcher compared with pitchers of the same plant without fenestration. We thus suggest that fenestration in S. minor is not an example of aggressive mimicry but rather functions in long-range attraction of prey. We highlight the need to evaluate aggressive mimicry relative to alternative concepts of plant–animal communication.     

With a flick of the lid: a novel trapping mechanism in Nepenthes gracilis pitcher plants

Carnivorous pitcher plants capture prey with modified leaves (pitchers), using diverse mechanisms such as 'insect aquaplaning' on the wet pitcher rim, slippery wax crystals on the inner pitcher wall, and viscoelastic retentive fluids. Here we describe a new trapping mechanism for Nepenthes gracilis which has evolved a unique, semi-slippery wax crystal surface on the underside of the pitcher lid and utilises the impact of rain drops to 'flick' insects into the trap. Depending on the experimental conditions (simulated 'rain', wet after 'rain', or dry), insects were captured mainly by the lid, the peristome, or the inner pitcher wall, respectively. The application of an anti-slip coating to the lower lid surface reduced prey capture in the field. Compared to sympatric N. rafflesiana, N. gracilis pitchers secreted more nectar under the lid and less on the peristome, thereby directing prey mainly towards the lid. The direct contribution to prey capture represents a novel function of the pitcher lid.     

Different pitcher shapes and trapping syndromes explain resource partitioning in Nepenthes species

Nepenthes pitcher plants display interspecific diversity in pitcher form and diets. This species‐rich genus might be a conspicuous candidate for an adaptive radiation. However, the pitcher traits of different species have never been quantified in a comparative study, nor have their possible adaptations to the resources they exploit been tested. In this study, we compare the pitcher features and prey composition of the seven Nepenthes taxa that grow in the heath forest of Brunei (Borneo) and investigate whether these species display different trapping syndromes that target different prey. The Nepenthes species are shown to display species‐specific combinations of pitcher shapes, volumes, rewards, attraction and capture traits, and different degrees of ontogenetic pitcher dimorphism. The prey spectra also differ among plant species and between ontogenetic morphotypes in their combinations of ants, flying insects, termites, and noninsect guilds. According to a discriminant analysis, the Nepenthes species collected at the same site differ significantly in prey abundance and composition at the level of order, showing niche segregation but with varying degrees of niche overlap according to pairwise species comparisons. Weakly carnivorous species are first characterized by an absence of attractive traits. Generalist carnivorous species have a sweet odor, a wide pitcher aperture, and an acidic pitcher fluid. Guild specializations are explained by different combinations of morpho‐functional traits. Ant captures increase with extrafloral nectar, fluid acidity, and slippery waxy walls. Termite captures increase with narrowness of pitchers, presence of a rim of edible trichomes, and symbiotic association with ants. The abundance of flying insects is primarily correlated with pitcher conicity, pitcher aperture diameter, and odor presence. Such species‐specific syndromes favoring resource partitioning may result from local character displacement by competition and/or previous adaptations to geographically distinct environments.     

Distribution of naphthoquinones, plumbagin, droserone, and 5-O-methyl droserone in chitin-induced and uninduced Nepenthes khasiana: molecular events in prey capture

Prey capture and digestion in Nepenthes spp. through their leaf-evolved biological traps involve a sequence of exciting events. Sugar-rich nectar, aroma chemicals, narcotic alkaloid secretions, slippery wax crystals, and other biochemicals take part in attracting, capturing, and digesting preys in Nepenthes pitchers. Here we report the distribution of three potent naphthoquinones in Nepenthes khasiana and their roles in prey capture. Plumbagin was first detected in N. khasiana, and its content (root: 1.33 ± 0.02%, dry wt.) was the highest found in any natural source. Chitin induction enhanced plumbagin levels in N. khasiana (root: 2.17 ± 0.02%, dry wt.). Potted N. khasiana plants with limited growth of roots and aerial parts, showed higher levels of plumbagin accumulation (root: 1.92 ± 0.02%; root, chitin induction: 3.30 ± 0.21%, dry wt.) compared with field plants. Plumbagin, a known toxin, insect ecdysis inhibitor, and antimicrobial, was also found embedded in the waxy layers at the top prey capture region of N. khasiana pitchers. Chitin induction, mimicking prey capture, produced droserone and 5-O-methyl droserone in N. khasiana pitcher fluid. Both these naphthoquinone derivatives provide antimicrobial protection to the pitcher fluid from visiting preys. A two-way barrier was found between plumbagin and its two derivatives. Plumbagin was never detected in the pitcher fluid whereas both its derivatives were only found in the pitcher fluid on chitin induction or prey capture. The three naphthoquinones, plumbagin, droserone, and 5-O-methyl droserone, act as molecular triggers in prey capture and digestion in the carnivorous plant, N. khasiana.     

Regulation of Free Calcium Concentration in the Pitchers of the Carnivorous Plant Sarracenia purpurea: A Model for Calcium in the Higher Plant Apoplast?

The pitcher of the carnivorous plant Sarracenia purpurea L.contains an entrapped body of liquid within which its prey isdigested. Free calcium in the pitcher is derived from eitherthe pitcher walls or from prey falling into the pitcher; inthe absence of exogenous (prey-derived) calcium it will dependon the active and passive calcium regulatory properties of thepitcher walls and may to some extent therefore mimic calciumin the apoplast of plant cells. Using a calcium-specific electrode,the free calcium concentration of the pitchers of Sarraceniaplants was investigated and the effect of adding a variety ofconcentrations of calcium in water determined. The mean pitcherfree calcium concentration in vivo was 2.3 x 10^–5 M±2.5x 10^–5 M; when pitchers were washed and filled with watercontaining lower calcium concentrations, the concentration inthe pitcher water rose to 1–5 x 10^–5 M. When highercalcium concentrations (up to 1 x 10^–4 M) were added,the pitcher calcium concentration declined to 1–7 x 10^–5M. Concentrations of calcium above 1 x 10^–4 M were alsoreduced, but to a lesser extent. Metabolic inhibition of activeion transport, while inhibiting pitcher acidification, did notinhibit regulation of pitcher free calcium, suggested that itoccurs as a result of calcium exchange sites in the pitcherwalls. The data are discussed in relation to the physiologyof Sarracenia pitchers and to the usefulness of the pitcheras a model for free calcium in the higher plant apoplast. Sarracenia purpurea L., carnivorous plant, pitcher, free calcium, plant apoplast     

Harmless nectar source or deadly trap: Nepenthes pitchers are activated by rain, condensation and nectar

The leaves of Nepenthes pitcher plants are specialized pitfall traps which capture and digest arthropod prey. In many species, insects become trapped by 'aquaplaning' on the wet pitcher rim (peristome). Here we investigate the ecological implications of this capture mechanism in Nepenthes rafflesiana var. typica. We combine meteorological data and continuous field measurements of peristome wetness using electrical conductance with experimental assessments of the pitchers' capture efficiency. Our results demonstrate that pitchers can be highly effective traps with capture rates as high as 80% but completely ineffective at other times. These dramatic changes are due to the wetting condition of the peristome. Variation of peristome wetness and capture efficiency was perfectly synchronous, and caused by rain, condensation and nectar secreted from peristome nectaries. The presence of nectar on the peristome increased surface wetness mainly indirectly by its hygroscopic properties. Experiments confirmed that pitchers with removed peristome nectaries remained generally drier and captured prey less efficiently than untreated controls. This role of nectar in prey capture represents a novel function of plant nectar. We propose that the intermittent and unpredictable activation of Nepenthes pitcher traps facilitates ant recruitment and constitutes a strategy to maximize prey capture.     

Plasma-membrane H+-ATPases are expressed in pitchers of the carnivorous plant Nepenthes alata Blanco

Nepenthes is a unique genus of carnivorous plants that can capture insects in trapping organs called pitchers and digest them in pitcher fluid. The pitcher fluid includes digestive enzymes and is strongly acidic. We found that the fluid pH decreased when prey accumulates in the pitcher fluid of Nepenthes alata. The pH decrease may be important for prey digestion and the absorption of prey-derived nutrients. To identify the proton pump involved in the acidification of pitcher fluid, plant proton-pump homologs were cloned and their expressions were examined. In the lower part of pitchers with natural prey, expression of one putative plasma-membrane (PM) H⁺-ATPase gene, NaPHA3, was considerably higher than that of the putative vacuolar H⁺-ATPase (subunit A) gene, NaVHA1, or the putative vacuolar H⁺-pyrophosphatase gene, NaVHP1. Expression of one PM H⁺-ATPase gene, NaPHA1, was detected in the head cells of digestive glands in the lower part of pitchers, where proton extrusion may occur. Involvement of the PM H⁺-ATPase in the acidification of pitcher fluid was also supported by experiments with proton-pump modulators; vanadate inhibited proton extrusion from the inner surface of pitchers, whereas bafilomycin A₁ did not, and fusicoccin induced proton extrusion. These results strongly suggest that the PM H⁺-ATPase is responsible for acidification of the pitcher fluid of Nepenthes. Received: 8 June 2000 / Accepted: 8 August 2000     

Fluorescent prey traps in carnivorous plants

Carnivorous plants acquire most of their nutrients by capturing ants, insects and other arthropods through their leaf‐evolved biological traps. So far, the best‐known attractants in carnivorous prey traps are nectar, colour and olfactory cues. Here, fresh prey traps of 14 Nepenthes, five Sarracenia, five Drosera, two Pinguicula species/hybrids, Dionaea muscipula and Utricularia stellaris were scanned at UV 366 nm. Fluorescence emissions of major isolates of fresh Nepenthes khasiana pitcher peristomes were recorded at an excitation wavelength of 366 nm. N. khasiana field pitcher peristomes were masked by its slippery zone extract, and prey capture rates were compared with control pitchers. We found the existence of distinct blue fluorescence emissions at the capture spots of Nepenthes, Sarracenia and Dionaea prey traps at UV 366 nm. These alluring blue emissions gradually developed with the growth of the prey traps and diminished towards their death. On excitation at 366 nm, N. khasiana peristome 3:1 CHCl₃–MeOH extract and its two major blue bands showed strong fluorescence emissions at 430–480 nm. Masking of blue emissions on peristomes drastically reduced prey capture in N. khasiana pitchers. We propose these molecular emissions as a critical factor attracting arthropods and other visitors to these carnivorous traps. Drosera, Pinguicula and Utricularia prey traps showed only red chlorophyll emissions at 366 nm.     

Adaptive significance and ontogenetic variability of the waxy zone in Nepenthes rafflesiana

Background and Aims

The slippery waxy zone in the upper part of pitchers has long been considered the key trapping structure of the Nepenthes carnivorous plants; however, the presence of wax is reported to be variable within and between species of this species-rich genus. This study raises the question of the adaptive significance of the waxy zone and investigates the basis for an ontogenetic cause of its variability and correlation with pitcher shape.

Methods

In Brunei (Borneo) the expression of the waxy zone throughout plant ontogeny was studied in two taxa of the Nepenthes rafflesiana complex, typica and elongata, which differ in pitcher shape and size. We also tested the adaptive significance of this zone by comparing the trapping efficiency and the number of prey captured of wax-bearing and wax-lacking plants.

Key Results

In elongata, the waxy zone is always well expanded and the elongated pitchers change little in form during plant development. Wax efficiently traps experimental ants but the number of captured prey in pitchers is low. In contrast, in typica, the waxy zone is reduced in successively produced pitchers until it is lost at the end of the plant''s juvenile stage. The form of pitchers thus changes continuously throughout plant ontogeny, from elongated to ovoid. In typica, the number of captured prey is greater, but the role of wax in trapping is minor compared with that of the digestive liquid, and waxy plants do not show a higher insect retention and prey abundance as compared with non-waxy plants.

Conclusions

The waxy zone is not always a key trapping structure in Nepenthes and can be lost when supplanted by more efficient features. This study points out how pitcher structure is submitted to selection, and that evolutionary changes in developmental mechanisms could play a role in the morphological diversity of Nepenthes.Key words: Carnivorous plant, developmental evolution, digestive liquid, epicuticular wax, insect trapping, heteroblasty, heterochrony, leaf form, morphological diversity, Nepenthes rafflesiana, ontogenetic change, pitcher plant     

The use of light in prey capture by the tropical pitcher plant Nepenthes aristolochioides

Nepenthes pitcher plants deploy tube-shaped pitchers to catch invertebrate prey; those of Nepenthes aristolochioides possess an unusual translucent dome. The hypothesis was tested that N. aristolochioides pitchers operate as light traps, by quantifying prey capture under three shade treatments. Flies are red-blind, with visual sensitivity maxima in the UV, blue, and green wavebands. Red celluloid filters were used to reduce the transmission of these wavebands into the interior of the pitchers. Those that were shaded at the rear showed a 3-fold reduction in Drosophila caught, relative to either unshaded control pitchers, or pitchers that were shaded at the front. Thus, light transmitted through the translucent dome is a fundamental component of N. aristolochioides' trapping mechanism.     

Role of areoles on prey abundance and diversity in the hooded pitcher plant (<Emphasis Type="Italic">Sarracenia minor</Emphasis>, <Emphasis Type="Italic">Sarraceniaceae</Emphasis>)

The hooded pitcher plant, Sarracenia minor, is a carnivorous facultative wetland species native to the southeastern USA and is listed as threatened by the state of Florida. Pitchers of S. minor possess white, semitranslucent spots (areoles), which have been hypothesized to aid in the capture of prey (= visual lures) by increasing the amount of light entering the back of the hood, which persuades insects to enter and fall into the base of the pitcher. In this study, the role of the areoles in prey capture abundance and diversity were experimentally investigated under variable lighting conditions in situ. Plants in two populations experiencing different light intensities, which varied in the amount of canopy cover and incident light reaching the plants, were experimentally manipulated by coloring varying percentages of areoles, ranging from 0 to 100 % (in increments of 25 %), with indelible ink. After 2 months, pitchers were collected and dissected to determine the number and identity of prey captured (approximately 18,000 prey were sampled). Although total prey abundance was approximately five times higher at McGirt’s Creek (sunny site) compared to UNF (shaded site), the effect of areoles on prey capture rates and biodiversity was site dependent. Reducing the number of visual lures of plants at the sunny site produced a significant decrease in the number of prey captured, but prey biodiversity (community composition) was unaffected. However, total prey capture was unaffected at the shaded site, while prey biodiversity was negatively correlated with the percent of areoles colored. Results from the current study suggest that areoles conditionally act as visual lures, but their overall importance is dependent on local environmental variables—especially canopy cover and the amount of incident sunlight reaching the plants.     

Tree shrew lavatories: a novel nitrogen sequestration strategy in a tropical pitcher plant

Nepenthes pitcher plants are typically carnivorous, producing pitchers with varying combinations of epicuticular wax crystals, viscoelastic fluids and slippery peristomes to trap arthropod prey, especially ants. However, ant densities are low in tropical montane habitats, thereby limiting the potential benefits of the carnivorous syndrome. Nepenthes lowii, a montane species from Borneo, produces two types of pitchers that differ greatly in form and function. Pitchers produced by immature plants conform to the ‘typical’ Nepenthes pattern, catching arthropod prey. However, pitchers produced by mature N. lowii plants lack the features associated with carnivory and are instead visited by tree shrews, which defaecate into them after feeding on exudates that accumulate on the pitcher lid. We tested the hypothesis that tree shrew faeces represent a significant nitrogen (N) source for N. lowii, finding that it accounts for between 57 and 100 per cent of foliar N in mature N. lowii plants. Thus, N. lowii employs a diversified N sequestration strategy, gaining access to a N source that is not available to sympatric congeners. The interaction between N. lowii and tree shrews appears to be a mutualism based on the exchange of food sources that are scarce in their montane habitat.     

A carnivorous plant fed by its ant symbiont: a unique multi-faceted nutritional mutualism

Scarcity of essential nutrients has led plants to evolve alternative nutritional strategies, such as myrmecotrophy (ant-waste-derived nutrition) and carnivory (invertebrate predation). The carnivorous plant Nepenthes bicalcarata grows in the Bornean peatswamp forests and is believed to have a mutualistic relationship with its symbiotic ant Camponotus schmitzi. However, the benefits provided by the ant have not been quantified. We tested the hypothesis of a nutritional mutualism, using foliar isotopic and reflectance analyses and by comparing fitness-related traits between ant-inhabited and uninhabited plants. Plants inhabited by C. schmitzi produced more leaves of greater area and nitrogen content than unoccupied plants. The ants were estimated to provide a 200% increase in foliar nitrogen to adult plants. Inhabited plants also produced more and larger pitchers containing higher prey biomass. C. schmitzi-occupied pitchers differed qualitatively in containing C. schmitzi wastes and captured large ants and flying insects. Pitcher abortion rates were lower in inhabited plants partly because of herbivore deterrence as herbivory-aborted buds decreased with ant occupation rate. Lower abortion was also attributed to ant nutritional service. The ants had higher δ(15)N values than any tested prey, and foliar δ(15)N increased with ant occupation rate, confirming their predatory behaviour and demonstrating their direct contribution to the plant-recycled N. We estimated that N. bicalcarata derives on average 42% of its foliar N from C. schmitzi wastes, (76% in highly-occupied plants). According to the Structure Independent Pigment Index, plants without C. schmitzi were nutrient stressed compared to both occupied plants, and pitcher-lacking plants. This attests to the physiological cost of pitcher production and poor nutrient assimilation in the absence of the symbiont. Hence C. schmitzi contributes crucially to the nutrition of N. bicalcarata, via protection of assimilatory organs, enhancement of prey capture, and myrmecotrophy. This combination of carnivory and myrmecotrophy represents an outstanding strategy of nutrient sequestration.     

Morphological correlates of necromass accumulation in the traps of an Eastern tropical pitcher plant, Nepenthes ampullaria Jack, and observations on the pitcher infauna and its reconstitution following experimental removal

I studied the trap morphology, necromass accumulation rates and pitcher infauna of an eastern tropical pitcher plant, Nepenthes ampullaria, that grew in `kerangas' heath forest in the Sungei Ingei Conservation Area, Brunei. I surveyed 164 pitchers distributed among 35 plants and extracted the necromass and larval infauna from the pitchers and then resampled the pitcher contents after 14 days. Plants varied significantly in the morphology of their pitchers, in their rate of necromass accumulation per pitcher and in the abundance and composition of the pitcher infaunas. On average, pitchers accumulated 11.5 mg dry weight over 14 days, but larger pitchers accumulated more necromass than smaller ones. Pitcher morphology explained 45% of the variation in necromass accumulation among plants. On average, pitchers initially contained 26.3 individual larval inquilines. Collectively, the larval infauna was composed of nine taxa of dipteran larvae and infrequent anuran tadpoles. These ten taxa were never found together in a single pitcher and the mean species richness per pitcher was 4.0. Of the six taxa that could be assessed, all except Toxorhynchites spp. had a contagious distribution among the pitchers. Pitcher morphology and necromass accumulation explained only 15% of the variation in inquiline abundance among plants. I found little evidence for the existence of density-dependent interactions between inquiline species: a partial correlation analysis detected only one statistically significant pairwise relationship between the abundances of inquiline taxa, which was a positive association. Fourteen days after being emptied, pitchers contained an average of 9.6 inquilines. There was no evidence that the species composition of the infauna recolonising each pitcher was related to that of its pre-removal infauna. Received: 2 June 1997 / Accepted: 9 September 1997     

Termite Prey Specialization in the Pitcher Plant Nepenthes albomarginata--Evidence from Stable Isotope Analysis

Old World pitcher plants (Nepenthes spp., Nepenthaceae) trapand digest invertebrate prey to derive nutrients, primarilynitrogen (N). In the majority of lowland Nepenthes species studiedto date, ants (Hymenoptera, Formicidae) are numerically thedominant prey taxon. Nepenthes albomarginata is unusual in showingan apparent bias towards the capture of termites (Isoptera).We tested the hypothesis that N. albomarginata derives N fromtermite capture, by comparison of foliar stable N isotope abundance(