Family matters: effect of host plant variation in chemical and mechanical defenses on a sequestering specialist herbivore

Insect herbivores contend with various plant traits that are presumed to function as feeding deterrents. Paradoxically, some specialist insect herbivores might benefit from some of these plant traits, for example by sequestering plant chemical defenses that herbivores then use as their own defense against natural enemies. Larvae of the butterfly species Battus philenor (L.) (Papilionidae) sequester toxic alkaloids (aristolochic acids) from their Aristolochia host plants, rendering larvae and adults unpalatable to a broad range of predators. We studied the importance of two putative defensive traits in Aristolochia erecta: leaf toughness and aristolochic acid content, and we examined the effect of intra- and interplant chemical variation on the chemical phenotype of B. philenor larvae. It has been proposed that genetic variation for sequestration ability is ??invisible to natural selection?? because intra- and interindividual variation in host-plant chemistry will largely eliminate a role for herbivore genetic variation in determining an herbivore??s chemical phenotype. We found substantial intra- and interplant variation in leaf toughness and in the aristolochic acid chemistry in A. erecta. Based on field observations and laboratory experiments, we showed that first-instar larvae preferentially fed on less tough, younger leaves and avoided tougher, older leaves, and we found no evidence that aristolochic acid content influenced first-instar larval foraging. We found that the majority of variation in the amount of aristolochic acid sequestered by larvae was explained by larval family, not by host-plant aristolochic acid content. Heritable variation for sequestration is the predominant determinant of larval, and likely adult, chemical phenotype. This study shows that for these highly specialized herbivores that sequester chemical defenses, traits that offer mechanical resistance, such as leaf toughness, might be more important determinants of early-instar larval foraging behavior and development compared to plant chemical defenses.     

Ectomycorrhizal fungal communities associated to Nothofagus species in Northern Patagonia

Ectomycorrhizal fungi constitute an important component of soil biota in Nothofagus forests in Patagonia. However, ectomycorrhizal fungal community is poorly known in this region. Here, we assess biodiversity and community compositions of ectomycorrhizal fungal species associated with Nothofagus dombeyi, N. obliqua and N. alpina. We selected three monospecific Nothofagus forest sites for each species within the boundaries of the Lanin National Park in Northern Patagonia. Ectomycorrhizal fungal species were identified based on morphotyping and rDNA (ITS and 28S rDNA) sequence analysis using both universal and taxon-specific primers. Contrary to previous studies on congeneric host trees, our results showed no significant differences among Nothofagus forest types in terms of fungal biodiversity and community composition. However, altitude had a strong effect on the structure of the ectomycorrhizal fungal community associated with Nothofagus spp.     

Evidence of chloroplast capture in South American Nothofagus (subgenus Nothofagus,Nothofagaceae)

Subgenus Nothofagus, although geographically restricted at present to temperate areas of South America, has captured much attention in discussions of plant biogeography due to its widespread distribution through Gondwanan continents during the Tertiary. However, phylogenetic relationships within the subgenus Nothofagus have not yet been resolved. We examined geographic patterns of intraspecific and interspecific genetic variation to detect whether incongruences in nuclear or plastid DNA phylogenies occur, in order to better understand the evolutionary history of the subgenus Nothofagus. We conducted spatially-explicit sampling at 10 distinct locations throughout the range of austral South American forests and sampled all present Nothofagus species. We used ITS and chloroplast DNA sequences to estimate phylogenetic relationships. A phylogeny constructed from nuclear genes resolved the subgenus Nothofagus as monophyletic. We found that N. antarctica was a sister to a clade of evergreen species (N. betuloides, N. dombeyi, and N. nitida), while N. pumilio likely diverged earlier. Nine cpDNA haplotypes were distinguished in the subgenus Nothofagus which were associated to geographic locations rather than to taxonomic relationships. This species-independent cpDNA phylogeographic structures within the subgenus Nothofagus may be related to repeated chloroplast capture events over geological time in Patagonia.     

Abundance–body mass relationships among insects along a latitudinal gradient

Abstract We investigated the relationship between abundance and body size (body mass) of 162 insect herbivore species on the host plant Acacia falcata along its entire coastal latitudinal distribution (eastern Australia), spanning a gradient in mean annual temperature of 4.3°C. We extend previous research by assessing these relationships at different spatial scales (latitudes pooled, among latitudes and within latitudes) and at different taxonomic levels (insect phytophages pooled, phytophagous Coleoptera and Hemiptera, and five component suborders/superfamilies). Insect species were collected from two orders (Hemiptera and Coleoptera) and five component suborders/superfamilies. There were no consistent trends in the relationships (linear or polygonal/hump‐shaped) between abundance and body mass when latitudes were pooled, among latitudes, or when phytophagous insect species were separated into their component suborder/superfamily groups. The reason for the lack of consistent trends might be due to the insect herbivores not fully exploiting their host plant and the relative absence of competition among herbivore species for food resources. This is further assessed in relation to the lack of a consistent pattern in species richness of Coleoptera and Hemiptera herbivores from the same dataset and rates of chewing and sap‐sucking herbivory along the same latitudinal gradient. Future studies of abundance–body size relationships are discussed in relation to sampling across environmental gradients and accounting for the influence of host plant identity and insect phylogeny.     

Host-plants shape insect diversity: Phylogeny, origin, and species diversity of native Hawaiian leafhoppers (Cicadellidae: Nesophrosyne)

Herbivorous insects and the plants on which they specialize, represent the most abundant terrestrial life on earth, yet their inter-specific interactions in promoting species diversification remains unclear. This study utilizes the discreet geologic attributes of Hawai'i and one of the most diverse endemic herbivore radiations, the leafhoppers (Hemiptera: Cicadellidae: Nesophrosyne), as a model system to understand the role of host-plant use in insect diversification. A comprehensive phylogeny is reconstructed to examine the origins, species diversification, and host-plant use of the native Hawaiian leafhoppers. Results support a monophyletic Nesophrosyne, originating from the Western Pacific basin, with a sister-group relationship to the genus Orosius. Nesophrosyne is characterized by high levels of endemicity according to individual islands, volcanoes, and geologic features. Clades demonstrate extensive morphologically cryptic diversity among allopatric species, utilizing widespread host-plant lineages. Nesophrosyne species are host-plant specific, demonstrating four dominant patterns of specialization that shape species diversification: (1) diversification through host switching; (2) specialization on widespread hosts with allopatric speciation; (3) repeated, independent shifts to the same hosts; and, (4) absence or low abundance on some host. Finally, evidence suggests competing herbivore radiations limit ecological opportunity for diversifying insect herbivores. Results provide evolutionary insights into the mechanisms that drive and shape this biodiversity.     

Friend or foe? The role of leaf-inhabiting fungal pathogens and endophytes in tree-insect interactions

Trees are large organisms that structure forest ecosystems by providing an environment for an enormous diversity of animal, microbial and plant species. As these species use trees as their common hosts, many are likely to interact with each other directly or indirectly. From studies on herbaceous plant species we know that microbes can affect the interaction of plants with herbivorous insects, for example via changes in plant metabolite profiles. The consequences of fungal colonization for tree-insect interactions are, however, barely known, despite the importance of these ecological communities. In this review we explore the interaction of leaf-inhabiting pathogenic and endophytic fungi with trees and the consequences for tree-living insect herbivores. We discuss molecular, physiological, chemical, biochemical and ecological aspects of tree-fungus interactions and summarize the current knowledge on the direct and indirect effects of tree-inhabiting fungi on insect herbivores.Our mechanistic understanding of the tripartite interaction of trees with leaf-inhabiting fungi and insect herbivores is still in its infancy. We are currently facing substantial drawbacks in experimental methodology that prevent us from revealing the effect of one single fungal species on a particular insect herbivore species and vice versa. Future studies applying a versatile toolbox of modern molecular, chemical analytical and ecological techniques in combined laboratory and field experiments will unequivocally lead to a better understanding of fungus-tree-insect interactions.     

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.     

Mass loss and nutrient release from decomposing evergreen and deciduous Nothofagus litters from the Chilean Andes

Abstract Leaf litter decomposition experiments were conducted on two deciduous (Nothofagus obliqua (roble)) and Nothofagus pumilio (lenga)) and one evergreen (Nothofagus dombeyi) Nothofagus (Nothofagaceae) species from a single Chilean forest in order to understand how congeneric trees with differing leaf lifespans impact the soil in which they grow. Single‐species litter samples were decomposed in a mixed hardwood forest in Ohio and in a deciduous‐evergreen Nothofagus forest in Chile. In the Ohio forest, the two deciduous species’ litters decomposed at k ≈ 1.00 per year and the evergreen at k ≈ 0.75 per year. In Chile k ranged from k ≈ 0.06 (N. obliqua) to k ≈ 0.23 (N. pumilio) per year. In both experiments, N and P were released faster from the deciduous litters than from evergreen litter. In Ohio, evergreen litter immobilized more N and P for a longer time period than did deciduous litter. As N. dombeyi stands tend to have lower available soil N and P in this particular mixed Nothofagus forest, the increased time of N and P immobilization by N. dombeyi litter suggests a feedback role of the tree itself in perpetuating low N and P soil conditions.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

Communities of insect herbivores foraging on saplings versus mature trees of Pourouma bicolor (Cecropiaceae) in Panama

The arthropod fauna of 25 saplings and of three conspecific mature trees of Pourouma bicolor (Cecropiaceae) was surveyed for 12 months in a tropical wet forest in Panama, with particular reference to insect herbivores. A construction crane erected at the study site provided access to tree foliage in the upper canopy. A similar area of foliage (ca. 370 m²) was surveyed from both saplings and trees, but samples obtained from the latter included 3 times as much young foliage as from the former. Arthropods, including herbivores and leaf-chewing insects with a proven ability to feed on the foliage of P. bicolor were 1.6, 2.5 and 2.9 times as abundant on the foliage of trees as on that of saplings. The species richness of herbivores and proven chewers were 1.5 (n=145 species) and 3.5 (n=21) times higher on trees than on saplings, respectively. Many herbivore species preferred or were restricted to one or other of the host stages. Host stage and young foliage area in the samples explained 52% of the explained variance in the spatial distribution of herbivore species. Pseudo-replication in the two sampling universes, the saplings and trees studied, most likely decreased the magnitude of differences apparent between host stages in this forest. The higher availability of food resources, such as young foliage, in the canopy than in the understorey, perhaps combined with other factors such as resource quality and enemy-free space, may generate complex gradients of abundance and species richness of insect herbivores in wet closed tropical forests.     

Both host-plant phylogeny and chemistry have shaped the African seed-beetle radiation

For the last 40 years, many authors have attempted to characterize the main patterns of plant-insect evolutionary interactions and understand their causes. In the present work on African seed-beetles (Coleoptera: Bruchidae), we have performed a 10-year field work to sample seeds of more than 300 species of potential host-plants (from the family Fabaceae), to obtain bruchids by rearing. This seed sampling in the field was followed by the monitoring of adult emergences which gave us the opportunity to identify host-plant use accurately. Then, by using molecular phylogenetics (on a combined data set of four genes), we have investigated the relationships between host-plant preferences and insect phylogeny. Our objectives were to investigate the level of taxonomic conservatism in host-plant fidelity and host-plant chemistry. Our results indicate that phylogenetically related insects are associated with phylogenetically related host-plants but the phylogeny of the latter cannot alone explain the observed patterns. Major host shifts from Papilionoideae to Mimosoideae subfamilies have happened twice independently suggesting that feeding specialization on a given host-plant group is not always a dead end in seed-beetles. If host-plant taxonomy and chemistry in legumes generally provide consistent data, it appears that the nature of the seed secondary compounds may be the major factor driving the diversification of a large clade specializing on the subfamily Mimosoideae in which host-plant taxonomy is not consistent with chemical similarity.     

Insect herbivores associated with Baccharis dracunculifolia (Asteraceae): responses of gall-forming and free-feeding insects to latitudinal variation

The spatial heterogeneity hypothesis has been invoked to explain the increase in species diversity from the poles to the tropics: the tropics may be more diverse because they contain more habitats and micro-habitats. In this paper, the spatial heterogeneity hypothesis prediction was tested by evaluating the variation in richness of two guilds of insect herbivores (gall-formers and free-feeders) associated with Baccharis dracunculifolia (Asteraceae) along a latitudinal variation in Brazil. The seventeen populations of B. dracunculifolia selected for insect herbivores sampling were within structurally similar habitats, along the N-S distributional limit of the host plant, near the Brazilian sea coast. Thirty shrubs were surveyed in each host plant population. A total of 8 201 galls and 864 free-feeding insect herbivores belonging to 28 families and 88 species were sampled. The majority of the insects found on B. dracunculifolia were restricted to a specific site rather than having a geographic distribution mirroring that of the host plant. Species richness of free-feeding insects was not affected by latitudinal variation corroborating the spatial heterogeneity hypothesis. Species richness of gall-forming insects was positively correlated with latitude, probably because galling insect associated with Baccharris genus radiated in Southern Brazil. Other diversity indices and evenness estimated for both gall-forming and free feeding insect herbivores, did not change with latitude, suggesting a general structure for different assemblages of herbivores associated with the host plant B. dracunculifolia. Thus it is probable that, insect fauna sample in each site resulted of large scale events, as speciation, migration and coevolution, while at local level, the population of these insects is regulated by ecological forces which operate in the system.     

Colonization of a host tree by herbivorous insects under a changing climate

Climate warming has been predicted to increase the abundance of herbivorous insects. Together with concurrent poleward shifts in many insect species this may increase herbivore pressure on plants. However, the manner in which plants at higher latitudes become colonized by herbivorous insects in the future is unknown. We established a translocation experiment using 26 micropropagated silver birch Betula pendula genotypes from six populations originating from 60°N to 67°N, to study the susceptibility of the translocated birches to local herbivores. The birches were planted at three different latitudes in Finland (60°N, 62°N and 67°N). We studied the effect of source population and latitudinal translocation on herbivore density, species richness, and community composition among the genotypes growing in the same environmental conditions in two years; 2011 and 2012. The source population explained the variation in the herbivore density only in 2012, whereas latitudinal translocation did not affect herbivore density. Variation in species richness was not explained by the source population or by the latitudinal translocation. At two of the study sites, the similarity of the herbivore communities among the populations decreased with increasing latitudinal distance of the source populations, possibly because birch populations that grow geographically closer to each other are genetically more similar, and therefore support a more similar composition of the arthropod community. All birch genotypes were colonized by local herbivores, suggesting that as herbivores shift their ranges polewards, they are able to colonize novel host‐plant genotypes. This enables compositional changes in insect communities on their host plants in the future, which in turn, might affect total herbivory and eventually, plant growth.     

Arthropod assemblages in a focal tree species (Eucalyptus microcarpa) depends on the species mix in restoration plantings

The composition of arthropods in trees has long been a topic of interest, with many studies exploring ways in which arthropod communities differ among tree species or entire forests. Few studies, however, have examined arthropods from trees in restoration plantings, and little is known about how different tree plantings might lead to different biodiversity outcomes. The aim of our study was to determine if a focal tree species hosted a different arthropod fauna depending on its context. We examined arthropod assemblages from the foliage of Eucalyptus microcarpa (Grey Box) trees present in two kinds of plantings: (i) simple plantings with only Eucalyptus trees, and (ii) mixed plantings with both Eucalyptus and Acacia trees. We examined the composition of the assemblages, and looked for associations between ant and psyllid (Hemiptera) diversity across each kind of planting. We found more species of psyllids, beetles and ants in E. microcarpa trees when they were grown in simple plantings compared with the mixed species plantings. We also found that psyllid richness and abundance was positively correlated with ant richness and abundance across all plantings. Our study shows that a more diverse context (i.e. a mixed planting) is not necessarily associated with a more diverse fauna within a focal tree species, and that simple Eucalyptus-only plantings may encourage higher numbers of insect herbivores. The apparent association between psyllids and ants suggests a complex interaction that may drive patterns in tree arthropod assemblages.     

Nothofagus (Fagaceae) and its invertebrate fauna—an overview and preliminary synthesis

The invertebrates directly associated with the southern hemisphere tree genus Nothofagus are described and discussed in terms of their organization into guilds and attributes of their hostplants. Using literature records, comparisons are made of the guild composition between host species and provenances; a full invertebrate-hostplant list is appended. Sap-sucking Homoptera are particularly diverse and usually host-species specific whereas defoliating insects seem less prevalent and less specific. Few seed-eating insects are reported. Some provenances and hostplants exhibit local radiations in certain taxa, such as Eriococcidae in New Zealand, Notophorina psyllids in South America and scolytid beetles on N. dombeyi. Environmental stresses such as drought, disturbance and landslips, can induce outbreaks of certain scale insects and borers, affecting the dynamics of the forest. Generally, however, the influence of invertebrates appears benign, but occasional large-scale defoliation is observed on N. solandri var. cliffortioides, N. moorei, N. antarctica and N. pumilio. Collectively this fauna has two elements: a geographically recurrent Gondwanic element, usually monophagous, and a provenance-specific element (often polyphagous) locally recruited following vicariance of the host genus. The fauna of Nothofagus can be in stark contrast to that of adjacent forest types. In Australia, the insect profile of Nothofagus has more in common with fagaceous trees on other continents than the more recent Eucalyptus forests which surround it. There is almost no evidence of fauna transfer from Nothofagus to Eucalyptus. Relative to New Zealand, the Australian Nothofagus fauna is depauperate and may reflect truncation in Pleistocene glaciations. Typically low insect populations on these trees may partly explain the limited range of insectivorous birds and spiders observed in Australia. Much more data on the phytophagous insects of this biome is needed for comparative purposes. The fauna of the montane tropical subgenus Brassospora pollen group is almost unknown but could hold the key to the origin of fagaceous insect communities more generally. The conservation value of remaining Nothofagus forests is enhanced by recognition of their co-adapted fauna, some of which are arranged in stable functional groups likely to give valuable insight into invertebrate-hostplant interactions originating in the Cretaceous. Elucidation of its role in the processes of these unique forests remains a fertile field for endeavour in the future.     

A new subspecies and two new combinations of Nothofagus Blume (Nothofagaceae) from Chile

One new subspecies ofNothofagusBlume,Nothofagus obliquasubsp.andina, is described from Chile, and two new combinations have been madeNothofagus obliquasubsp.valdivianaandNothofagus macrocarpacomb. nov.     

Non-native plants rarely provide suitable habitat for native gall-inducing species

For insect herbivores, a critical niche requirement—possibly the critical niche requirement—is the presence of suitable host plants. Current research suggests that non-native plants are not as suitable as native plants for native herbivores, resulting in decreases in insect abundance and richness on non-native plants. Like herbivores, gall-forming insects engage in complex, species-specific interactions with host plants. Galls are plant tissue tumors (including bulbous or spindle-shaped protrusions on leaves, stems and other plant organs) that are induced by insects through physical or chemical damage (prompting plants to grow a protective tissue shell around the insect eggs and larvae). As such, we hypothesized that gall-inducing insect species richness would be higher on native than non-native plants. We also predicted higher gall-inducing insect species richness on woody than herbaceous plants. We used an extensive literature review in which we compiled gall host plant species by genus, and we assigned native or non-native (or mixed) status to each genus. We found that native plants host far more gall-inducing insect species than non-native plants; woody plants host more gall-inducing species than herbaceous plants; and native woody plants host the most gall-inducing species of all. Gall-inducing species generally are a very cryptic group, even for experts, and hence do not elicit the conservation efforts of more charismatic insects such as plant pollinators. Our results suggest that non-native plants, particularly non-native woody species, diminish suitable habitat for gall-inducing species in parallel with similar results found for other herbivores, such as Lepidopterans. Hence, the landscape-level replacement of native with non-native species, particularly woody ones, degrades taxonomically diverse gall-inducing species (and their inquilines and parasitoids), removing multiple layers of diversity from forest ecosystems.

     

The value of timber quality forests for insect conservation on Tierra del Fuego Island compared to associated non-timber quality stands

Insect community studies related to forest management focus principally on timber-quality stands, and often omit the remainder of the landscape. This study aimed mainly to compare insect communities of primary timber-quality forests (Nothofagus pumilio) with associated non-timber-quality stands (wetland, edge, riparian and N. antarctica forests), and secondarily to characterize these insect assemblages throughout the growing season and at different vertical strata to evaluate the importance of each habitat type for insect conservation. A total of 18,800 individuals belonging to 231 RTUs (recognizable taxonomic units) were identified, of which Diptera, Hymenoptera, Lepidoptera and Coleoptera were the dominant orders. Lepidoptera RTUs were mostly generalists, while the other main orders were most frequently found in timber-quality forests and included many RTUs with specific environmental requirements. Timber-quality stands had higher richness and abundance than associated non-timber-quality forests and possessed more exclusive species (18%), while 39% of RTUs were shared between all sites. The spatial heterogeneity of timber-quality stands generated different niches and favored insect diversity, which would not have been maintained by protecting non-timber-quality stands alone. Consequently, the proper management of subantarctic Nothofagus forests must include the conservation of timber-quality stands, as protection of non-timber-quality areas alone will not be sufficient for insect conservation at the landscape scale.     

A Specialist Herbivore Uses Chemical Camouflage to Overcome the Defenses of an Ant-Plant Mutualism

Many plants and ants engage in mutualisms where plants provide food and shelter to the ants in exchange for protection against herbivores and competitors. Although several species of herbivores thwart ant defenses and extract resources from the plants, the mechanisms that allow these herbivores to avoid attack are poorly understood. The specialist insect herbivore, Piezogaster reclusus (Hemiptera: Coreidae), feeds on Neotropical bull-horn acacias (Vachellia collinsii) despite the presence of Pseudomyrmex spinicola ants that nest in and aggressively defend the trees. We tested three hypotheses for how P. reclusus feeds on V. collinsii while avoiding ant attack: (1) chemical camouflage via cuticular surface compounds, (2) chemical deterrence via metathoracic defense glands, and (3) behavioral traits that reduce ant detection or attack. Our results showed that compounds from both P. reclusus cuticles and metathoracic glands reduce the number of ant attacks, but only cuticular compounds appear to be essential in allowing P. reclusus to feed on bull-horn acacia trees undisturbed. In addition, we found that ant attack rates to P. reclusus increased significantly when individuals were transferred between P. spinicola ant colonies. These results are consistent with the hypothesis that chemical mimicry of colony-specific ant or host plant odors plays a key role in allowing P. reclusus to circumvent ant defenses and gain access to important resources, including food and possibly enemy-free space. This interaction between ants, acacias, and their herbivores provides an excellent example of the ability of herbivores to adapt to ant defenses of plants and suggests that herbivores may play an important role in the evolution and maintenance of mutualisms.