The effects of fragmentation on foliar defensive traits and insect herbivory rates in dipterocarp seedlings

Human-modified landscapes are ubiquitous and often made up of remnant fragments of natural ecosystems nested within an agricultural or urban matrix. Understanding how species are affected by habitat degradation is a central issue in biodiversity research, yet investigations into the impacts on key ecological interactions have not kept pace. Gaining insight into the responses of ecological processes is vital in order to maximise biodiversity conservation and develop sustainable management practices in a changing world. Herbivory is a fundamental ecosystem process as it mediates the transfer of energy between primary production and higher trophic levels. In tropical rainforests herbivory is primarily carried out by insects. Here, I investigate how leaf damage changes over time by carrying out repeated measures of herbivory following fragmentation. In doing so, I build upon previous ‘snapshot’ studies which have primarily quantified leaf damage at single points in time. An experiment was established within a large-scale manipulation experiment: the Stability of Altered Forest Ecosystems (SAFE) project in the Malaysian state of Sabah, within Borneo. I used seedlings of two species of endemic Dipterocarpaceae as the study system for two main reasons. Firstly, seedlings represent the most vulnerable life stage in a tree’s life cycle, and therefore insect herbivory may be a major determinant of their growth and survival. Secondly, not only do members of the family Dipterocarpaceae dominate in lowland tropical rainforests of Southeast Asia, most are economically valued for their timber and are consequently under pressure from logging which is leading to conservation concern. Five hundred and seventy-six seedlings were planted in 12 recently isolated one hectare fragments and 12 continuous forest control sites. Eight of these control sites were located in an area of continuous forest estimated to be over one million hectares north of the experimental landscape of the SAFE project. Four further control sites were established in a virgin jungle reserve which is over 2,200 hectares and located to the south-west of the experimental area. All leaves were scanned in situ on six occasions over two field seasons (May- October) in 2015 and 2016. At the end of the experiment, traits analyses were performed to quantify three common metrics of leaf defence: total phenolic content, acid detergent fibre and leaf strength. Through this experiment I addressed the following aims; (1) to determine the initial effects of fragmentation on rates of leaf damage, (2) to investigate whether bottom-up control of herbivory was altered by fragmentation through changes in foliar defence, and (3) to assess whether responses were shared among study species or were instead individualistic. I also use data from a key published monograph to create species interaction networks between lepidopteran caterpillars and their known host plants in tropical Asia, with the aim of exploring the importance of members of the Dipterocarpaceae for insect herbivores. I found some evidence that herbivory is disrupted by fragmentation, with both species exhibiting lower levels of herbivory in fragments. Forest type was an important predictor of patterns of leaf area loss in this experiment, but overall differences between herbivory in fragments and control sites were not significant. Instead, herbivory was best explained by seedling traits, and predictors of leaf area loss varied between species indicating species-specific responses. Relaxation of phytochemical defensive traits was also detected in forest fragments, perhaps due to decreased levels of herbivory. A reduction in phytochemical defence may imply an increased vulnerability of seedlings in forest fragments which translates to differences in herbivore damage over time. I highlight the importance of repeated measures experiments when investigating a complex and dynamic ecological process such as herbivory, and propose long term monitoring to fully understand the effects of forest fragmentation. The results of this thesis contribute to understanding the effects of fragmentation on insect herbivory, which remain uncertain, and provide evidence of the extent to which this key ecosystem process is disrupted due to anthropogenic habitat modification