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The Role of Leaf Cellulose Content in Determining Host Plant Preferences of Three Defoliating Insects Present in the Andean-Patagonian Forest

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The Role of Leaf Cellulose Content in Determining Host Plant Preferences of Three Defoliating Insects Present in the Andean-Patagonian Forest Austral Ecology (2016) , – The role of leaf cellulose content in determining host plant preferences of three defoliating insects present in the Andean-Patagonian forest A. L. PIETRANTUONO, O. A. BRUZZONE AND V. FERNANDEZ-ARHEX* Estacion Experimental Agropecuaria Bariloche, CONICET – Instituto Nacional de Tecnologıa Agropecuaria, CC277, Av. Modesta Victoria 4450, 8400 San Carlos de Bariloche, Rıo Negro, Argentina (E-mail: [email protected]) Abstract Phytophagous insects choose their feeding resources according to their own requirements in addition to properties of the host plants, such as biomechanical defences. The feeding preferences of the native folivorous insects of the Andean-Patagonian forest (Argentina) have rarely been studied. These environments present a wide diversity and abundance of insects associated with trees of the Nothofagus and Lophozonia (Nothofagaceae) genera, which represent the main tree species of the forests of the southern hemisphere. In particular, Lophozonia alpina and Lophozonia obliqua are of great interest because they have a wide distribution, a high capacity for hybridization and exhibit great phenotypic plasticity. This versatility causes substantial variation in the biome- chanical properties of leaves, affecting the feeding preferences of insects. The purpose of this work was to study the food selection behaviour of three leaf-chewing insects (Polydrusus nothofagii, Polydrusus roseaus (Coleoptera: Curculionidae) and Perzelia arda (Lepidoptera: Oecophoridae)) associated with L. alpina and L. obliqua as host plants. Based on their choices, our aim was to determine a preference scale for each insect species and the vari- ables on which these preferences were based. Therefore, we selected trees of L. alpina and L. obliqua, measured several properties such as cellulose content and recorded which leaves were eaten. As a result, we determined that the three species of insects feed on both host plants but prefer the leaves of L. obliqua, with cellulose content being the main determining factor for their decisions. However, in the case of P. arda, there was a positive rela- tionship between cellulose and host plant preference, whereas there was an opposite relationship for the weevils. We conclude that during feeding selection, there are some properties of the leaves that have a more important role than others and that the same property does not exert the same behavioural response in all folivorous insects. Key words: Bayesian analysis, biomechanical properties, feeding preferences, native forest, phytophagous, Thurstone scale. INTRODUCTION preferences of an insect (Schoonhoven et al. 2005; Peeters et al. 2007). This behaviour may also vary During the process of searching for and selecting a depending on the biology, habits and developmental host plant, folivorous insects may exhibit some stage of each particular species of chewing insects in degree of selectivity among their feeding resources. the following two different ways: either by preventing This selection process involves not only choosing the the initiation of feeding by early larval instars by cre- right species of plant but also selecting an individual ating a physical obstacle and/or by causing selective plant within that species (Bernays & Chapman 1994; feeding in the smaller chewers that attempt to avoid Mayhew 1997; Price et al. 2011). Several biomechan- the toughest tissues (Ohmart et al. 1987; Choong ical characteristics of a leaf regulate insect behaviour, 1996; Wheeler et al. 1998). affecting the preferences and patterns of feeding The western area of Patagonia, a geographical (Hanson 1983; Schoonhoven et al. 2005; Hong et al. region located at the southern end of South America 2012). According to several authors, the toughness of that includes the territories of Argentina and Chile, is the leaves (determined by cell wall components: cel- characterized by the presence of the Andean forest. lulose, hemi-cellulose and lignin) is the main prop- These environments have a temperate-cold climate erty that influences the feeding behaviour and food and are dominated by trees of the Nothofagaceae (Fagales) family. Further, there is a wide diversity *Corresponding author. and abundance of folivorous insects, many of which Accepted for publication September 2016. have not been identified or studied in terms of any © 2016 Ecological Society of Australia doi:10.1111/aec.12460 2 A.L.PIETRANTUONOET AL. aspect of their behaviour (Gentili & Gentili 1988; METHODS McQuillan 1993). This family comprises approxi- mately 35 living species present in various temperate Study system forests of New Zealand, Australia, New Caledonia, New Guinea and South America (Ward 1965). In The study system is composed of three species of phy- Argentina, the following two genera are represented: tophagous insects and their host plants, which are present Nothofagus Blume and Lophozonia Heenan & Smis- in the Andean-Patagonian forest, Argentina. Polydrusus sen (2013). In particular, Lophozonia alpina ‘raulı’ nothofagii Kuschel, P. roseaus Blanchard (Coleoptera: Cur- and Lophozonia obliqua ‘roble pellın’ (previously culionidae) and Perzelia arda Clarke (Lepidoptera: named Nothofagus nervosa and Nothofagus obliqua Oecophoridae) are native insects that are commonly found respectively) are species of great importance due to feeding on the foliage of L. alpina and L. obliqua (Kuschel 1952; Giganti & Dapoto 1990; Grandon 1996; Morrone their important ecological value, excellent robustness 1999; Vergara & Jerez 2010). The Curculionidae larvae are and timber quality (Sabatier et al. 2011); it is for root-feeders, whereas the adults are skeletonizing defoliators these reasons that both species are included in that are mainly found during the months of January and domestication and conservation plans (Gallo et al. February (Gentili & Gentili 1988; Elgueta & Marvaldi 2006). Furthermore, they have the ability to grow in 2006). In the case of P. arda, the larvae cause leaf-tying a wide range of environments, including different and defoliating damage. They are commonly found from altitudes and latitudes (Ramirez 1987; Donoso et al. December to March (Cerda et al. 1982; Grandon 1996). 2004). This varied distribution reflects their high phenotypic plasticity and hybridization ability, quali- ties that often hinder their taxonomic identification Selection of host plants and experimental (Premoli 1996). These factors, in turn, are the cause insects of large variation in their biomechanical properties, including the toughness of leaves and possibly the Several individuals of L. alpina (n = 8) and L. obliqua = complex of associated insects (Donoso 1987; Donoso (n 8) were selected from a plantation in a nursery at the INTA EEA Bariloche (Instituto Nacional de Tecnologıa et al. 1990; Premoli & Mathiasen 2011; Premoli et al. ı Agropecuaria. Estacion Experimental Agropecuaria Bar- 2012; Mart nez Pastur et al. 2013). Therefore, phy- iloche, Rıo Negro, Argentina 41°7022.36″S, 71°1504.94″W) tophagous insects may show an unclear pattern of (National Institute of Agricultural Technology – Bariloche selectivity for a particular species, given that their Research Lab). The selected trees were used as a source of preferences could be more linked to the biomechani- seeds for the production of seedlings to be used for conser- cal properties of the leaves of a particular individual. vation, restoration or timber production. At the nursery, all Several studies have focused on the interactions individuals were in a healthy state, were never exposed to between Patagonian Nothofagaceae and their associ- insecticides, and experienced the same humidity, light, sub- ated native insects based on the damage caused by strate and temperature conditions. Leaves were harvested these insects using either a descriptive or quantitative from these individuals for the analysis of foliar traits and approach (Russell et al. 2000; Mazıa et al. 2004, were offered as stimuli in the experiment, ensuring that the leaves were mature, of similar size, had not been chewed 2009, 2012; Rousseaux et al. 2004; Garibaldi et al. and did not harbour fungi. At the same site, non-host 2010, 2011a,b; Paritsis & Veblen 2010; Altmann plants (Populus nigra Linneo, n = 1 and Salix nigra Mar- 2011; Chorbadjian & Francino 2013; Chaneton et al. shall, n = 1 (Malpighiales: Salicaceae)) were selected to be 2014). However, few studies have evaluated such used as controls in the experiment. interactions in terms of insect behaviour (Russell Regarding the insects, adult P. nothofagii (n = 107) and et al. 2004; Pietrantuono et al. 2014). Such studies P. roseaus (n = 51) and the last larval instar of P. arda (common in biological control trials) contribute to (n = 68) were collected from the same nursery. understanding in a variety of different ways by pro- viding knowledge regarding the ecology and natural history of native phytophagous insects and informa- Analysis of foliar traits tion that can be used for the development of conser- vation plans and even for the silvicultural For each host plant, several biomechanical properties management of different species of Nothofagaceae. related to leaf toughness and digestibility were measured (e.g. dry matter, neutral detergent fibre, neutral detergent The purpose of this work was to study the food insoluble protein, lignin, ash, total nitrogen and cellulose, selection behaviour of three species of chewing Appendix S1). To determine the amount of dry and insects associated with L. alpina and L. obliqua
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