An experimental evaluation of resource allocation in island plants with respect to their invertebrate herbivores

Abstract

New Zealand’s isolation and periods of marine transgression have limited its biota to an extent which can be considered depauperate, even by island standards. Endemic vertebrates are rare and prominent invertebrate families, such as the renowned forest defoliators of the Lymantriidae, are absent. The proven vulnerability of the flora to introduced vertebrates reaffirms a belief in the invasiveness of islands and fuels the contingency plans aimed at averting similar devastation from further alien invertebrate defoliators. Nothofagus is a dominant element of the climax forests of New Zealand and the larger landmasses bordering the South Pacific Ocean. Assessments of the resistance to defoliation of continental and New Zealand species of Nothofagus, and a range of other forest genera endemic to New Zealand, was undertaken using bioassays of naïve polyphagous defoliators. The bioassays were undertaken in Europe, utilising gypsy moth, Lymantria dispar (Lepidoptera: Lymantriidae) and the fall webworm, Hyphantria cunea (Lepidoptera: Arctiidae) as defoliators, fed foliage plants growing in European arboreta. In New Zealand, bioassays utilised Australian painted apple moth, Teia anartoides (Lepidoptera: Lymantriidae) and tree species from local arboreta, gardens and natural populations. Larval growth rate was the primary parameter recorded to assess plant resistance. The relevance of growth rate was investigated by comparison with other recorded parameters and resistance to a surrogate pathogen, in the form of commercially available bio-insecticide. Larval growth rate was positively correlated with survivorship, potential fecundity, mating success and resistance to disease. The growth rate of larvae fed Nothofagus was positively correlated to the species-specific leaf nitrogen content. The results of the bioassays showed that despite the accepted paradigms, New Zealand’s flora was largely resistant to exotic defoliators. As an explanation of this apparent anomaly, the Island Resource Allocation (IRA) hypothesis was developed and posits that ‘the palatability iv of a plant to invertebrate herbivores is proportional to the geographic range of the plant’. The basis for the IRA hypothesis proposes a redefinition of the fundamental ecological principle of the species: area relationship. Islands, or similarly geographically constrained ecosystems, which support lower biodiversity, have impoverished trophic levels and consequently have weaker top-down regulation of herbivores by natural enemies. The IRA hypothesis argues that island ecosystem stability is achieved through the bottom-up process of plant defence. The IRA hypothesis was tested intra-specifically using bioassays using painted apple moth in which larvae were offered foliage of specimens from naturally discontinuous populations of Nothofagus truncata. The results supported the hypothesis in that the smallest populations of N. truncata exhibited the greatest resistance to the defoliator. The IRA hypothesis and a demonstrated mechanism for a differential resistance in Nothofagus species could resolve a number of enduring debates in ecology. Habitat area appears to explain the relative strengths of top-down and bottom-up regulation of herbivores. It also predicts the strengths of reciprocal evolution within the geographic mosaic of co-evolution and highlights the influence of biodiversity in invasive ecology. It may also help to resolve the contentious and extremely relevant debate of the role of biodiversity in ecosystem function.