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Ant Community Dynamics in Lac Insect Agroecosystems: Conservation Benefits of a Facultative Association Between Ants and Lac Insects

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Ant Community Dynamics in Lac Insect Agroecosystems: Conservation Benefits of a Facultative Association Between Ants and Lac Insects Tropical Ecology 58(2): 283–293, 2017 ISSN 0564-3295 © International Society for Tropical Ecology www.tropecol.com Ant community dynamics in lac insect agroecosystems: conservation benefits of a facultative association between ants and lac insects 1* 1 1 2 YOUQING CHEN , SIMING WANG , ZHIXING LU & WEI ZHANG 1Research Institute of Resources Insects, Chinese Academy of Forestry (CAF), Kunming, 650224, Yunnan, China 2School of Conservation Biology, Southwest Forestry University, Kunming, 650224, Yunnan, China Abstract: Honeydew is a key resource for ants and a driver of ant community dynamics, yet we know very little about how the quantity of honeydew generates these patterns. This study investigated the effects of an ant-lac insect facultative association on ant community dynamics in a guild of ants attending Kerria yunnanensis (Hermiptera: Kerriidae) on its host plant Dalbergia obtusifolia in Yunnan Province, China. This was done by manipulating the access of ants to K. yunnanensis and the numbers of K. yunnanensis available. Regardless of the proportion of lac insect infestation (the ratio of the branch length infested by lac insects to the total branch length), there were more ant species and individuals under the ant-lac insect facultative association, and ant community composition was significantly different from that without this association. The level of branch infestation by the lac insect had a significant influence on the mean frequency of occurrence of ants; a low level of branch infestation by the lac insect led to low ant attendance. Honeydew secreted by the lac insect had a key role in structuring the ant community. There were more co-occurrences of ants under the ant-lac insect facultative association, and co-occurrence of more than three ant species only occurred on trees with lac insect infestation. The significant difference in frequency of month-by-month ant replacement was only observed in the treatment without lac insect hosting, which indicated the positive role of ant-lac insect association in stabilizing ant community. We suggest that the facultative association between ants and lac insects has conservation benefits for ant assemblages and provides a structuring mechanism for the ant community in the lac agroecosystem studied. Key words: Ant community, ant-lac insect association, co-occurrence, lac insect agroecosystem, replacement. Handling Editor: Christopher D. Beatty Introduction focus on the benefits for hemipterans, and studies investigating the advantages of the interactions to Most ant-hemipteran associations are the ants are rare (Lach et al. 2009). Experimental generally regarded as facultatively mutualistic, in studies of ant-scale interactions typically involve a that neither partner is totally dependent on the single ant species (Itioka & Inoue 1996; Rosario et other (Buckley 1987a,b; Hill & Blackmore 1980). al. 1993 other studies summarised in Gullan Studies of ant-scale insect inter-actions typically 1997). However, many hemipteran aggregations *Corresponding Author; e-mail: [email protected] 284 ANT COMMUNITY AND CO-OCCURRENCE ON LAC INSECTS have multiple ant species in attendance (i.e., action promotes ant richness and abundance, and Buckley & Gullan 1991; Chen et al. 2011; Del 2) relative low-level honeydew supply is adequate Claro & Oliveira 1999; Moya-Raygoza & Nault for maintaining this association, resulting in more 2000). ant co-occurrence and fewer ant replacements. Many ant species utilize the same food resource, and competition may occur among them Methods (Apple & Feener 2001; Oliveira et al. 1999). Competitive interactions among ants on host Study system plants may potentially shape the community This research was conducted in a lac insect structure of ant assemblages. If competition is a agroecosystem in Mojiang county, Yunnan strong determinant of the structure of ant Province, south west of China (101º43′E, 23º14′ communities, a key question becomes: what allows N，999.8–1056 m a.s.l.). The lac insect agro- multiple ant species to coexist in a given habitat ecosystem is the main form of cultivation in (Andersen 2008)? The mechanism for ant species mountainous regions between 900–1500 m a.s.l., co-existence in the ant-extrafloral nectary (EFN) with several lac insect host plant species plant system has been explored (Blüthgen et al. distributed along or among the semi-arid and 2000; Palmer et al. 2000; Young et al. 1997). It semi-humid agricultural plots. The main host tree has been found that in an Amazonian rainforest for lac insects is Dalbergia obtusifolia (Prain); canopy, competitively superior ants usually other host tree species are distributed randomly dominate the most attractive food items, while among the D. obtusifolia plantations. The lac ants with lower competitive capacity share the less insect Kerria yunnanensis is raised on trees from attractive EFNs. Ant communities that depend on the fifth year onwards, but other honeydew- honeydew and EFNs seem to be strongly shaped producing coccid insects can also be found. Many by competitive hierarchies (Blüthgen et al. 2004). ant species attend lac insects for honeydew rather Such competitive hierarchies reflect the ranking of than eating them. At the same time, these ants abundance of the particular ants in their influence the natural enemies of lac insect present community (Andersen 1995; Blüthgen et al. 2004; in the ecosystem, either directly and indirectly Blüethgen & Fiedler 2004; Davidson 1997; Leal et (Chen et al. 2013). The main crop planted by the al. 2006; but see Xu & Chen 2010). However, little farmers under the lac insect host trees or among is known of the impact on ant assemblages from the patches of trees is corn. aggregations of honeydew-producing hemipterans, especially the pattern of resource utilization in a Experimental designs resource-rich environment (i.e., one lasting at least several months). The main purpose of these designs was to Lac insects (Kerria spp., Hemiptera: Kerriidae) investigate the influence of honeydew quantity on are scale insects that occur naturally as herbivores the ant community. In each generation of K. on various host plants. They produce lac, a layer of yunnanensis, before K. yunnanensis was cultivated red resin on branches of host-trees on which they on D. obtusifolia, trees distributed within a 10 ha settle. Lac resin is natural, biodegradable and non- plot were randomly sampled. The sampling toxic, and thus is widely used in the food, textile distance between individual trees (10 m) allowed and pharmaceutical industries, in addition to its them to be considered independent samples, use as a surface coating in electrical component because most ant species forage only within a few manufacturing (Chen et al. 2010). In China, lac- metres of the nest in this study. production and farming have become integrated Kerria yunnanensis was cultivated on the into an agroforest system. The lac insect agro- sampled trees and the population of lac insect was ecosystem is a very popular agroforest planting manipulated randomly based on four categories of model due to its economical and ecological roles, lac insect load on the tree. Though we randomly and is widely distributed throughout mountainous manipulated trees into each category, the spatial areas of Southwestern China, a region noted for distribution of treatment trees was interspersed to having high conservation values (Chen et al. 2010). ensure that individual trees were true replicates. We used field manipulation of Kerria The four categories of lac insect load were 1) no yunnanensis (Ou et Hong) population levels and a K. yunnanensis on the trees, 2) small number of K. controlled ant-exclusion experiment to test the yunnanensis on the trees, 3) medium number of K. hypotheses that 1) ant-lac insect facultative inter- yunnanensis on the trees, and 4) large number of CHEN et al. 285 K. yunnanensis on the trees. The ratio of K. plantation, which is dominated by the lac plant D. yunnanensis numbers among the last three obtusifolia with a density of 1500 trees per categories was about 6:3:1. Within each category, hectare, and the total area is about 10 hectares. the number of K. yunnanensis on each tree was On the host D. obtusifolia, K. yunnanensis settles controlled by measuring the total settlement area on the main branch, the shoots and even the leaf of K. yunnanensis on all branches of a given tree stalks. In the larval stage, females and males only (same population density on the same tree species, produce a little honeydew, but after mating all so K. yunnanensis population number could be males die and only females produce honeydew, but controlled by the area of settlement). For selected the amount is huge. This honeydew can be found trees of a similar size and the same amount of K. on the body of lac insects or on the surface of yunnanensis on the trees, the proportion of leaves and branches, and even on the ground. branches with lac insects hosting D. obtusifolia Many ant species including Crematogaster was similar within each category. Thus in this macaoensis (Wheeler 1928), Dolichoderus thora- experiment, the infestation proportions were (i) cicus (Smith 1860), and Camponotus parius 0%, (ii) about 10% (ranging from 8% to 12%), (iii) (Emery 1889) visit K. yunnanensis and D. about 30% (ranging from 28% to 32%), and obtusifolia for honeydew. In this experiment, 8- (iv) about 60% (ranging from 58% to 62%). In total, year-old D. obtusifolia were selected, ranging from 120 trees were selected, with each category having 2.5 m to 2.8 m in height and from 5 cm to 7 cm in 30 trees, and four levels of lac insect infestation trunk diameter. The experiments were conducted representing four treatments levels. These from March 2009 to December 2010. treatment levels were maintained throughout the whole life cycle of one generation from May 2009 to Ant assemblage investigation October 2009 (the ratio of K. yunnanensis We randomized the order in which we observed population number among the categories was trees, and recorded the species and the individual consistent, for the lac insects live a sessile life after number of each ant species within the tree.
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