title: The nitrogen supply from soils and insects during growth of the pitcher plants Nepenthes mirabilis, Cephalotus follicularis and Darlingtonia californica.
abstract: 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 N2fixation 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.
authors.affiliation: Plant Ecology, University Bayreuth, D-95440 Bayreuth, Germany, , , , , , DE.
authors.forename: W
authors.initials: W
authors.lastname: Schulze
authors.affiliation: Plant Ecology, University Bayreuth, D-95440 Bayreuth, Germany, , , , , , DE.
authors.forename: E D
authors.initials: ED
authors.lastname: Schulze
authors.affiliation: Dept. of Botany, University of Western Australia, Nedlands WA 6009 Australia, , , , , , AU.
authors.forename: J S
authors.initials: JS
authors.lastname: Pate
authors.affiliation: Center for International Forestry Research, P.O. Box 6596 JKPWB, Jakarta, Indonesia, , , , , , ID.
authors.forename: A N
authors.initials: AN
authors.lastname: Gillison
keywords: Insect nitrogen; Key words Carnivorous plants; Nitrogen partitioning; Pitcher plants