Local and Regional-Scale Food Web Structure in Nepenthes alata Pitchers1

Tropical Nepenthes pitcher plants provide small, isolated aquatic habitats. We examined inter-pitcher variation in the community structure of the inhabitants of Nepenthes alata Blanco in West Sumatra, focusing on the conditions of the pitchers, bacterial density in the pitcher fluid, density and biomass of metazoan inhabitants, and the frequencies of interspecific encounters. Older pitchers contained more insect carcasses. The bacterial density increased with the age of the pitchers, but decreased in withered pitchers that contained finely decomposed detritus. In live pitchers, the bacterial density, the density, mass and species richness of metazoa, and the number of trophic levels per pitcher were positively correlated with detrital mass, which was correlated with volume of pitcher fluid. The metazoan fauna from N. alata consisted of 4 predators and 12 saprophages, among the richest known for Nepenthes species. However, each individual pitcher harbored a limited numbers of species, owing to (1) the low incidence of many species, and (2) the aggregated distribution and different temporal colonization pattern of major species. Six dipteran taxa (one predator and five saprophages) accounted for the bulk of metazoan inhabitant biomass. Of 48 combinations of predator-prey encountered, only four occurred frequently (in > 30% of pitchers), which included two predators and three saprophages. Thus, individual pitchers harbored relatively simple communities despite the regional species richness, and only limited kinds of predator-prey encounters seemed to occur frequently in the regional food web. The local-scale properties of the subdivided communities presented here provide the basic information for understanding the maintenance of regional species richness and food web complexity.     

Why are Aedes mosquitoes rare colonisers of Nepenthes pitcher plants?

1. Nepenthes pitcher plants produce fluid‐containing animal traps that are colonised by a variety of specialised arthropods, especially dipterans. However, container‐breeding vector mosquitoes, such as Aedes albopictus Skuse have rarely been recorded from pitchers. Increasing overlap in the geographical ranges of Nepenthes and Ae. albopictus in urban parts of Southeast Asia owing to urbanisation highlights a growing need to investigate the potential role of pitchers as larval habitats for vector mosquitoes. 2. The ability of Ae. albopictus larvae to survive in three common lowland Nepenthes in Peninsular Malaysia that are most likely to co‐occur with Ae. albopictus [viz., Nepenthes ampullaria Jack, Nepenthes gracilis Korth., and Nepenthes mirabilis (Lour.) Druce] was investigated. 3. The larval survival rates of Ae. albopictus in pitcher fluids of the three Nepenthes species were determined, then the effects of low pH, larvicidal agents (such as microbes, predators, and chemical compounds) through manipulative experiments were investigated. 4. It was found that pitchers represent a hostile environment to Ae. albopictus, but that the principal cause of larval mortality varies among Nepenthes species (i.e. low fluid pH in N. gracilis, predation by Toxorhynchites acaudatus Leicester larvae in N. ampullaria, and microbial activity in N. mirabilis). It was concluded that Nepenthes pitchers are generally not suitable larval habitats for Ae. albopictus. However, the pitcher environment of N. ampullaria is worthy of further study, as pitchers that lack predators are nevertheless rarely colonised by Ae. albopictus, indicating that other aspects of the host pitcher environment inhibit oviposition or larval survivorship.     

Nepenthesin Protease Activity Indicates Digestive Fluid Dynamics in Carnivorous Nepenthes Plants

Carnivorous plants use different morphological features to attract, trap and digest prey, mainly insects. Plants from the genus Nepenthes possess specialized leaves called pitchers that function as pitfall-traps. These pitchers are filled with a digestive fluid that is generated by the plants themselves. In order to digest caught prey in their pitchers, Nepenthes plants produce various hydrolytic enzymes including aspartic proteases, nepenthesins (Nep). Knowledge about the generation and induction of these proteases is limited. Here, by employing a FRET (fluorescent resonance energy transfer)-based technique that uses a synthetic fluorescent substrate an easy and rapid detection of protease activities in the digestive fluids of various Nepenthes species was feasible. Biochemical studies and the heterologously expressed Nep II from Nepenthes mirabilis proved that the proteolytic activity relied on aspartic proteases, however an acid-mediated auto-activation mechanism was necessary. Employing the FRET-based approach, the induction and dynamics of nepenthesin in the digestive pitcher fluid of various Nepenthes plants could be studied directly with insect (Drosophila melanogaster) prey or plant material. Moreover, we observed that proteolytic activity was induced by the phytohormone jasmonic acid but not by salicylic acid suggesting that jasmonate-dependent signaling pathways are involved in plant carnivory.     

Reinigung der Proteinase aus Nepenthes-Kannensaft

Zusammenfassung Der sterile Kannensaft von drei Nepenthes-Arten wurde chromatographisch und elektrophoretisch untersucht. Bei Auftrennungen durch Adsorptionschromatographie an Ecteola-Cellulose und durch Elektrophorese wurde jeweils nur eine proteolytische Enzymfraktion erhalten. In gereinigten, kohlenhydratfreien Enzymfraktionen wurde ein pH-Optimum bei 2,2 und ein Temperaturoptimum bei 50°C gefunden.

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Purification of the proteinase from Nepenthes pitcher secretion

Summary The proteolytic enzymes from the sterile secretion of the pitchers of 3 Nepenthes species were purified to electrophoretic homogenity by chromatography on Ecteola cellulose. The properties of the purified enzymes were investigated.

     

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.     

The nitrogen supply from soils and insects during growth of the pitcher plants Nepenthes mirabilis, Cephalotus follicularis and Darlingtonia californica

This study investigated the nitrogen (N) acquisition from soil and insect capture during the growth of three species of pitcher plants, Nepenthes mirabilis, Cephalotus follicularis and Darlingtonia californica. ¹⁵N/¹⁴N natural abundance ratios (δ¹⁵N) of plants and pitchers of different age, non-carnivorous reference plants, and insect prey were used to estimate proportional contributions of insects to the N content of leaves and whole plants. Young Nepenthes leaves (phyllodes) carrying closed pitchers comprised major sinks for N and developed mainly from insect N captured elsewhere on the plant. Their δ¹⁵N values of up to 7.2‰ were higher than the average δ¹⁵N value of captured insects (mean δ¹⁵N value = 5.3‰). In leaves carrying old pitchers that are acting as a N source, the δ¹⁵N decreased to 3.0‰ indicating either an increasing contribution of soil N to those plant parts which in fact captured the insects or N gain from N₂ fixation by microorganisms which may exist in old pitchers. The δ¹⁵N value of N in water collected from old pitchers was 1.2‰ and contained free amino acids. The fraction of insect N in young and old pitchers and their associated leaves decreased from 1.0 to 0.3 mg g⁻¹. This fraction decreased further with the size of the investigated tiller. Nepenthes contained on average 61.5 ± 7.6% (mean ± SD, range 50–71%) insect N based on the N content of a whole tiller. In the absence of suitable non-carnivorous reference plants for Cephalotus, δ¹⁵N values were assessed across a developmental sequence from young plants lacking pitchers to large adults with up to 38 pitchers. The data indicated dependence on soil N until 4 pitchers had opened. Beyond that stage, plant size increased with the number of catching pitchers but the fraction of soil N remained high. Large Cephalotus plants were estimated to derive 26 ± 5.9% (mean ± SD of the three largest plants; range: 19–30%) of the N from insects. In Cephalotus we observed an increased δ¹⁵N value in sink versus source pitchers of about 1.2‰ on average. Source and sink pitchers of Darlingtonia had a similar δ¹⁵N value, but plant N in this species showed δ¹⁵N signals closer to that of insect N than in either Cephalotus or Nepenthes. Insect N contributed 76.4 ± 8.4% (range 57–90%) to total pitcher N content. The data suggest complex patterns of partitioning of insect and soil-derived N between source and sink regions in pitcher plants and possibly higher dependence on insect N than recorded elsewhere for Drosera species. Received: 14 April 1997 / Accepted: 18 August 1997     

Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five <Emphasis Type="Italic">Nepenthes</Emphasis> species and hybrids

Plants of the carnivorous genus Nepenthes efficiently trap insects in leaf pitchers, mostly employing epicuticular wax crystals on the pitcher walls to make them slippery for the prey. In the present study, the compositions and micromorphologies of the wax crystals of five Nepenthes species and hybrids were analysed in order to test whether the chemical principles underlying this ecological function are widespread within the genus. Three wax layers could be distinguished within the Nepenthes pitcher cuticles: (1) the outermost part of the crystals forming the platelets visible in standard scanning electron microscopy, (2) the bottom portion of the epicuticular wax crystals, and (3) an intracuticular wax layer. The composition of the intracuticular wax differed significantly from that of the neighbouring epicuticular layer. The compositions of corresponding wax mixtures from all five Nepenthes species and hybrids were very similar, with almost equal amounts of very long chain aldehydes and primary alcohols. While triacontanal (C₃₀ aldehyde) was prevailing in the epicuticular crystals of Nepenthes albomarginata and Nepenthes x intermedia, Nepenthes x superba and Nepenthes x henriana were found to have especially high percentages of dotriacontanal (C₃₂ aldehyde). Nepenthes “khasiana” had an intermediate aldehyde composition with almost equal amounts of both chain lengths.     

Local‐ to continental‐scale variation in the richness and composition of an aquatic food web

Aim We investigated patterns of species richness and composition of the aquatic food web found in the liquid‐filled leaves of the North American purple pitcher plant, Sarracenia purpurea (Sarraceniaceae), from local to continental scales. Location We sampled 20 pitcher‐plant communities at each of 39 sites spanning the geographic range of S. purpurea– from northern Florida to Newfoundland and westward to eastern British Columbia. Methods Environmental predictors of variation in species composition and species richness were measured at two different spatial scales: among pitchers within sites and among sites. Hierarchical Bayesian models were used to examine correlates and similarities of species richness and abundance within and among sites. Results Ninety‐two taxa of arthropods, protozoa and bacteria were identified in the 780 pitcher samples. The variation in the species composition of this multi‐trophic level community across the broad geographic range of the host plant was lower than the variation among pitchers within host‐plant populations. Variation among food webs in richness and composition was related to climate, pore‐water chemistry, pitcher‐plant morphology and leaf age. Variation in the abundance of the five most common invertebrates was also strongly related to pitcher morphology and site‐specific climatic and other environmental variables. Main conclusions The surprising result that these communities are more variable within their host‐plant populations than across North America suggests that the food web in S. purpurea leaves consists of two groups of species: (1) a core group of mostly obligate pitcher‐plant residents that have evolved strong requirements for the host plant and that co‐occur consistently across North America, and (2) a larger set of relatively uncommon, generalist taxa that co‐occur patchily.     

Organic matter sources and size structuring in stream invertebrate food webs across a tropical to temperate gradient

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Consequences of consumer origin and omnivory on stability in experimental food web modules

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Aspects of Pitcher Morphology and Spectral Characteristics of Six BorneanNepenthesPitcher Plant Species: Implications for Prey Capture

Pitcher plants of the genusNepenthesattract and trap invertebrateprey using nectar-secreting pitchers. Pitcher morphology andspectral reflectance characteristics were investigated for sixNepenthesspeciesfrom northwest Borneo (N. albomarginata, N. ampullaria, N. bicalcarata,N. gracilis, N. mirabilisvar.echinostomaandN. rafflesiana).Morphological measurements focused on the size of the pitcherrim (or peristome, the site of the major nectaries) in relationto pitcher length. The results show considerable interspecificvariation in morphology. Spectral reflectance measurements quantifiedthe degree of colour contrast between the peristome and pitcherbody, from ultraviolet (UV) to red wavelengths. The contrastmaxima for each species were compared with insect visual sensitivitymaxima. The six species showed a wide range of reflectance patterns,with pitchers ofN. rafflesianapossessing the greatest degreeof ‘fit’ between contrast maxima and insect sensitivitymaxima, in the UV, blue and green regions of the spectrum. Basedon the morphological and reflectance analyses, we hypothesizedthat pitchers ofN. rafflesianawould be more attractive to anthophilous(flower-visiting) invertebrates than the sympatricN. gracilis.Analysis of prey contents generally supported the hypothesis,suggesting possible interspecific resource partitioning. Morphologicaland spectral characteristics of the other species are discussedin relation to published studies on prey capture by those species.Copyright1999 Annals of Botany Company Borneo, carnivory, colour,Nepenthesspp., pitcher plants, prey attraction.     

Photosynthetic characterization of Australian pitcher plant <Emphasis Type="Italic">Cephalotus follicularis</Emphasis>

Australian carnivorous pitcher plant Cephalotus follicularis Labill. produces two types of leaves. During the spring time, the plant produces a foliage type of noncarnivorous leaf called lamina. Later, the second type of leaf is produced — carnivorous pitcher. Using simultaneous measurements of gas exchange and chlorophyll (Chl) fluorescence photosynthetic efficiency of these two distinct forms of leaves were compared. In addition stomatal density, an important component of gas exchange, and Chl concentration were also determined. Pitcher trap had lower net photosynthetic rate (P _N) in comparison to noncarnivorous lamina, whereas the rate of respiration (R _D) was not significantly different. This was in accordance with lower stomatal density and Chl concentration in the pitcher trap. On the other hand maximum quantum yield of PSII (F_v/F_m) and effective quantum yield of photochemical energy conversion in PSII (Φ_PSII) was not significantly different. Nonphotochemical quenching (NPQ) was significantly higher in the lamina at higher irradiance. These data are in accordance with hypothesis that changing the leaf shape in carnivorous plants to make it a better trap generally makes it less efficient at photosynthesis. However, the pitcher of Cephalotus had much higher P _N than it was expected from the data set of the genus Nepenthes. Because it is not possible to optimize for contrasting function such as photosynthesis and carnivory, it is hypothesized that Cephalotus pitchers are less elaborated for carnivorous function than the pitchers of Nepenthes.     

Wetland isolation facilitates larval mosquito density through the reduction of predators

• 1 Wetlands harbour high biodiversity and offer important ecosystem services, but they are also a habitat for mosquito larvae (Diptera: Culicidae), which are important disease vectors.
• 2 Isolation among remnant, or newly created wetlands and ponds, and their consequent density in the landscape, is a key factor that can influence a variety of food web processes, including effects on mosquitoes which are important prey to many predators.
• 3 We assess the impact of habitat isolation on the density of pond‐breeding mosquitoes (several Anopheles and Culex species) both directly and indirectly through the food web.
• 4 Results from structural equation modelling of survey data shows that larval mosquitoes are denser in ponds that are more isolated from one another, and that this result was primarily driven indirectly by a reduction of larval mosquito predators (e.g. predaceous insects and amphibians). Furthermore, results from a long‐term mesocosm experiment factorially manipulating isolation and predator reduction show that the effect of isolation on mosquito density was eliminated when predators were experimentally reduced.
• 5 It is concluded that metacommunity processes, both directly and indirectly mediated through predators, can play an important role in the local abundance of wetland breeding mosquitoes and possibly the diseases they spread.

     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Trophic opportunism of central Amazon floodplain fish

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Apparent competition leaves no detectable imprint on patterns of community composition: observations from a natural experiment

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Food‐web structure and mercury dynamics in a large subarctic lake following multiple species introductions

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Bacterial communities associated with the pitcher fluids of three Nepenthes (Nepenthaceae) pitcher plant species growing in the wild

Nepenthes pitcher plants produce modified jug-shaped leaves to attract, trap and digest insect prey. We used 16S rDNA cloning and sequencing to compare bacterial communities in pitcher fluids of each of three species, namely Nepenthes ampullaria, Nepenthes gracilis and Nepenthes mirabilis, growing in the wild. In contrast to previous greenhouse-based studies, we found that both opened and unopened pitchers harbored bacterial DNA. Pitchers of N. mirabilis had higher bacterial diversity as compared to other Nepenthes species. The composition of the bacterial communities could be different between pitcher types for N. mirabilis (ANOSIM: R = 0.340, p < 0.05). Other Nepenthes species had similar bacterial composition between pitcher types. SIMPER showed that more than 50 % of the bacterial taxa identified from the open pitchers of N. mirabilis were not found in other groups. Our study suggests that bacteria in N. mirabilis are divided into native and nonnative groups.     

The facilitation of the native bluegill sunfish by the invasive bighead carp

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