Size, age and composition: characteristics of plant taxa as diversity predictors of gall-midges (Diptera: Cecidomyiidae)

Many hypotheses have been proposed to explain the diversity of gall-midge insects (Diptera: Cecidomyiidae), some of them taking into account plant diversity. This study aims to test the importance of size, age and composition of host plant taxa in the diversity of Cecidomyiidae. For this we used inventories data on the diversity of galling and host plants in Brazil. We found that Asterales, Myrtales and Malpighiales, were the most important orders, with 34, 33 and 25, gall morphotypes, respectively. The most representative host families were Asteraceae (34 morphotypes), Myrtaceae (23) and Fabaceae (22). In general, the order size and the plant family were good predictors of the galling diversity, but not the taxon age. The most diverse host genera for gall-midges were Mikania, Eugenia and Styrax, with 15, 13 and nine galler species, respectively. The size of plant genera showed no significant relationship with the richness of Cecidomyiidae, contrary to the prediction of the plant taxon size hypothesis. The plant genera with the greatest diversity of galling insects are not necessarily those with the greatest number of species. These results indicate that some plant taxa have a high intrinsic richness of galling insects, suggesting that the plant species composition may be equally or more important for the diversity of gall-midges than the size or age of the host taxon.     

Diversity of galling arthropods and host plants in a subtropical forest of Porto Alegre, southern Brazil

Many hypotheses have been proposed to explain diversity patterns of galling insects. However, there are contradictory evidences on the evolutionary and ecological factors responsible for the trends. Furthermore, questions such as arthropod seasonality, sampling sufficiency and sampling team experience have been almost ignored. This study records galling arthropod diversity while paying attention to these questions. Seasonal sampling of galling arthropods and host plants were conducted in a humid subtropical forest of southern Brazil. Four transects were sampled twice per season, with two persons searching the vegetation for galls during 1h30min. After 96h.persons of sampling, 130 gall morphotypes on 84 species of host plants were recorded. An analysis of the numbers of galls and gall morphotypes found per transect along time showed that sampling team experience influences galler richness results and the interpretation of galler seasonality patterns. Different species had distinct seasonal patterns. Galling arthropod richness was bound to plant richness. Our results suggest that sampling team experience is an important factor that must be explicitly considered, as well as seasonality patterns of different galling species, at least for tropical/subtropical areas. Although sampling sufficiency was not reached, fauna heterogeneity at small spatial scales seems substantial: despite the proximity of the sampled transects (500 m), they harboured significantly specific faunas. This work adds to the literature records suggesting that both plant richness and specific composition of the vegetation have a strong influence on galler richness at least for local scales.     

Cytological and histochemical gradients on two <Emphasis Type="BoldItalic">Copaifera langsdorffii</Emphasis> Desf. (Fabaceae)—Cecidomyiidae gall systems

Previous ultrastructural and histochemical analysis proposed patterns in the accumulation of substances in galls of Diptera: Cecidomyiidae in some plant species of the temperate region. Similar analyses were done to verify the conservativeness of these patterns in the Neotropical region, where a great number of species of Cecidomyiidae is responsible for a wide diversity of morphotypes. Two gall morphotypes induced by Cecidomyiidae in a unique host plant, Copaifera langsdorffii, were studied. The gradients of carbohydrates and the activity of invertases and acid phosphatases were similar, but the cytological gradients and distribution of proteins evidenced that the sites of the induction as well as the amount of neoformed tissues may be peculiar to each gall system. The production of lipids just in the secretory cavities either in the non-galled or galled tissues indicated a potentiality of the host plant which could not be manipulated by the galling insects. Further, the absence of nucleus in the nutritive tissue, an exclusive feature of the horn-shaped galls, indicates cell death attributed to the feeding habit of the galling herbivore.     

Insect galls on Bauhinia cupulata (Fabaceae): morphotypes characterization and description of a new species of Schizomyia (Cecidomyiidae,Diptera)

Galls on Bauhinia cupulata (Fabaceae) were investigated in two physiognomies of the Cerrado, riparian and dry forest, in the municipality of Barreiras (Western Bahia, Brazil) from October 2012 to July 2014. Four insect gall morphotypes were found. They were characterized based on shape, color, indumentum, plant organ of occurrence and galling taxon. A new galling species, Schizomyia barreirensis, is described, illustrated (larva, pupa, male, female and gall) and compared to other Neotropical congeneric species.     

Evolutionary shifts between host oak sections and host-plant organs in Andricus gallwasps

Abstract.— Gall-inducing insects have especially intimate interactions with their host plants and generally show great specificity with regard to both the host-plant species and the organ (e.g. flower, leaf) galled. However, the relative roles of shifts between host species and between host-plant organs in the diversification of gall-inducers are uncertain. We employ a novel and general maximum-likelihood approach to show that shifts between host-plant organs occur at a significantly greater rate than shifts between host oak sections in European Andricus gallwasps. This suggests that speciation has more often been associated with gall location shifts than with colonization of new host-plant species, and implies that it may be easier for gall-inducers to colonize new plant organs than new plant species.
Andricus gallwasps have complex life cycles, with obligate alternation of sexual and parthenogenetic generations. Our phylogenetic analyses show that a life cycle with both generations galling white oaks (section Quercus ) is ancestral, with a single shift of the sexual generation onto black oaks (section Cerris ) to generate a clade with a novel host-alternating life cycle. This new life cycle provided the opportunity for further speciation, but may have also increased the risk of extinction of one or both generations by the demographic requirement for co-existence of both host-plant groups. In summary, it appears that Andricus gallwasp radiation may be a two-level process. Speciation events often involve shifts in gall location on the same host species. However, there are only so many ways to gall an oak, and rare shifts to new oak sections may contribute greatly to long-term diversification by opening up whole new adaptive zones.     

Morphological phylogenetics of Australian gall-inducing thrips and their allies: the evolution of host-plant affiliations, domicile use and social behaviour

A phylogeny for seventy-two species of Australian thrips in the subfamily Phlaeothripinae, based on cladistic analysis of forty morphological adult characters, is presented. We use this phylogeny to infer the evolutionary history of host-plant affiliations, gall induction and other types of domicile use, and different forms of social behaviour. Maximum parsimony analysis yielded forty-eight cladograms of length 316, and the strict consensus of these cladograms was well resolved. This phylogeny indicated that: (1) associations of thrips with their host plants tend to be evolutionarily conservative, with monophyletic groups of thrips on the host-plant genera Acacia, Casuarina and Geijera, (2) galling has evolved multiple times, on different host plants, (3) transitions in domicile use include changes between galling and living in holes or old galls, between living in glued phyllodes and living in old galls, and between leaf-feeding and galling, and (4) in three of five cases, inquiline lineages were not closely related to their host lineages and the evolution of inquilinism apparently involved a host-plant shift. However, in two cases, inquilines were very closely related to their gall-inducing hosts. Eusocial behaviour (involving soldier castes) has evolved in different lineages from those that exhibit communal behaviour (cooperation in building or defending domiciles), suggesting a lack of direct transition between the two social systems. This phylogeny serves as a framework for future molecular systematic studies, and future comparative analysis of ecology and behaviour in the Phlaeothripinae.     

Leaf gall polymorphism and molecular phylogeny of a new Bruggmanniella species (Diptera: Cecidomyiidae: Asphondyliini) associated with Litsea acuminata (Lauraceae) in Taiwan,with ecological comparisons and a species description

Two types of cecidomyiid leaf galls, cup‐shaped and umbrella‐shaped, occur on Litsea acuminata (Lauraceae) in Taiwan. Based on the concept of gall shapes as “extended phenotypes” of gall inducers, these two types could be induced by different gall midge species. However, galls with intermediate shapes between the two types were recently discovered, which implies that possible genetic exchanges occur between the gall inducers of both types. To clarify the taxonomic status of gall midges responsible for the two types of galls on L. acuminata, we undertook taxonomic, molecular phylogenetic and ecological studies. Our findings show that the two gall types are induced by the same Bruggmanniella species and the species is new to science. We describe the species forming this range of galls as Bruggmanniella litseae sp. n. , and compare their geographical distribution, galling position and morphometry. Based on our results, a possible evolutionary scenario of B. litseae sp. n. is discussed.     

EVOLUTION OF DIFFERENT GALL TYPES IN WILLOW-FEEDING SAWFLIES (HYMENOPTERA: TENTHREDINIDAE)

The sawflies that feed on the plant family Salicaceae can be divided into eight informal groups based on larval feeding habit or gall type: (1) species with free-living larvae; (2) leaf folders; (3) leaf blade gallers; (4) apical leaf gallers; (5) basal leaf gallers; (6) midrib and petiole gallers; (7) stem gallers; and (8) bud gallers. It has been proposed that the galling habit evolved from free-living larvae via leaf folders, and that the different gall types evolved gradually in the sequence mentioned above. Thus, the galling site would have “wandered” from the leaf margin toward the stem as a result of gradual changes in oviposition site preference. Allozyme data from eight informative loci were used to reconstruct the phylogeny of 18 representative sawfly species. The results suggest that indeed leaf folders seem to be a basal group; leaf blade gallers evolved independently of the other true gallers; apical and basal leaf gallers are not the ancestors of petiole and bud gallers, but they may share a common galling ancestor; bud gallers evolved from midrib/petiole gallers; and stem gallers are polyphyletic. The cause for the observed wandering of the galling site could be intraspecific competition due to a possible “nutrient shading effect” of galls situated closer to the host plant's main vascular system.     

Diversity of galling insects in Styrax pohlii (Styracaceae): edge effect and use as bioindicators

Impacts of forest fragmentation and edge effect on plant-herbivores interactions are relatively unknown, and the relationships between galling insects and their host plants are very susceptible to environmental variations. The goal of our study was to test the edge effect hypothesis for galling insects associated with Styrax pohlii (Styracaceae) host plant. Samplings were conducted at a fragment of semi-deciduous forest in Goiania, Goiás, Brazil. Thirty host plant individuals (15 at fragment edge and 15 in its interior) were sampled in July of 2007; in each plant, 10 apical branches were collected at the top, middle and bottom crown levels. Our results supported the prediction of greater richness of gall morphotypes in the edge habitat compared with remnant interior. In a similar way, gall abundance and frequency of attacked leaves were also greater in the fragment edge. These findings consequently suggest a positive response of galling insect diversity to edge effect; in the Saint-Hilaire forest, this effect probably operates through the changes in microclimatic conditions of edge habitats, which results in an increased hygrothermal stress, a determinant factor to distribution patterns of galling insects. We also concluded that these organisms could be employed as biological indicators (i) because of their host-specificity, (ii) they are sensitive to changes in plant quality, and (iii) present dissimilar and specific responses to local variation in habitat conditions.     

Direct and indirect interactions involving ants, insect herbivores, parasitoids, and the host plant Baccharis dracunculifolia (Asteraceae)

Abstract.  1. The relative importance of direct and indirect interactions in controlling organism abundance is still an unresolved question. This study investigated the role of the direct and indirect interactions involving ants, aphids, parasitoids, and the host plant Baccharis dracunculifolia (Asteraceae) on a galling herbivore Baccharopelma dracunculifoliae (Homoptera: Psyllidae).
2. The effects of these interactions on the galling herbivore's performance were evaluated by an exclusion experiment during two consecutive generations of the galling insect.
3. Ants had a direct negative effect on the performance of the galling herbivore by reducing the number of nymphs per gall. In contrast, ants had no indirect effects on gall mortality through the associated parasitoids.
4. Aphids negatively affected gall development, suggesting that galls and aphids might be partitioning photoassimilates and nutrients moving throughout host-plant tissues.
5. In addition, galls that developed during the rainy season were heavier, indicating that variation in the host plant, due to weather changes, can affect the development of B. dracunculifoliae galls. However, variation in the development of B. dracunculifoliae galls due to presence of aphids or the weather changes did not affect parasitoid attack.
6. These results suggest that direct interactions between ants and galls influenced galling insect abundance, whereas numerical indirect effects involving galling insects, ants, aphids, and host plants were less conspicuous.     

Evolution of gall morphology and host-plant relationships in willow-feeding sawflies (Hymenoptera: Tenthredinidae)

Abstract.— There are over 200 species of nematine sawflies that induce galls on willows (Salix spp.). Most of the species are monoor oligophagous, and they can be separated into seven or eight different groups based on the type of gall that they induce. We studied the evolution of different gall types and host plant associations by reconstructing the phylogeny of five outgroup and 31 ingroup species using DNA sequence data from the mitochondrial cytochrome b gene. Maximum-parsimony and maximum-likelihood analyses resulted in essentially the same phylogeny with high support for important branches. The results show that: (1) the galling species probably form a monophyletic group; (2) true closed galls evolved only once, via leaf folders; (3) with the possible exception of leaf rollers, all gall type groups are mono- or paraphyletic; (4) similar gall types are closer on the phylogeny than would be expected by a random process; (5) there is an apparent evolutionary trend in galling site from the leaf edge towards the more central parts of the host plant; and (6) many willow species have been colonized several times, which excludes the possiblity of parallel cladogenesis between willows and the gallers; however, there are signs of restrictions in the evolution of host use. Many of the patterns in the evolutionary history of nematine gallers have also been observed in earlier studies on other insect gallers, indicating convergent evolution between the independent radiations.     

Biogeographical gradients in galling species richness

Summary Five hypotheses were invoked to account for variation in galling species number per location on plants of different structural complexity, namely herbs, shrubs, and trees, both in Brazil and USA. The hypotheses were: 1) the altitudinal/latitudinal gradient hypothesis; 2) the harsh environment hypothesis; 3) the plant species richness hypothesis; 4) the host plant area hypothesis; 5) the plant structural complexity hypothesis. The altitudinal and the harsh environment hypotheses were correlated and supported with sample data in both localities, with increasing gall species number as altitude/latitude declined and as sites became hotter and drier. The two hypotheses were separated by studying riparian sites and dry hillside sites at the same elevation in Arizona. Galling species frequency was higher in dry sites than in riparian sites, supporting the harsh environment hypothesis. Of the five hypotheses tested only the harsh environment hypothesis predicted that galling insect species number should vary in response to environmental variables such as moisture and temperature. Temperate shrubs supported more galling species than did other plant types, both in dry and mesic sites. The overall difference between galling species richness for tropical and temperate latitudes was not statistically significant. Free-feeding insect herbivore species exhibited the opposite pattern of species richness to gallers, being more speciose in riparian sites. The present study corroborates the hypothesis that the gall forming habit is an adaptation to harsh or stressful environments, and we describe for the first time broad scale geographical patterns in galling insect species richness.     

Evolution of the gall wasp-host plant association

Gall wasps, or cynipids, form the second largest radiation of galling insects with more than 1300 described species. According to current views, the first cynipids were phytophagous and developed in herb stems of the Asteraceae without modifying plant growth or development. The first galls were supposedly multichambered stem swellings, and subsequent trends involved increase in gall complexity and reduction in the number of larval chambers. Gall wasps also have many of the features believed to be characteristic for phytophagous insects radiating in parallel with their host plants. We tested these hypotheses by mapping characters onto a recent estimate of higher cynipid relationships from a morphology-based analysis of exemplar taxa, controlling for phylogenetic uncertainty using bootstrapping. Characters were also mapped onto a metatree including all gall wasps, assembled from phylogenetic analyses as well as recent classifications. The results contradict many of the current hypotheses. The first cynipids with extant descendants were not Asteraceae stem feeders but induced distinct single-chambered galls in reproductive organs of herbaceous Papaveraceae, or possibly Lamiaceae. There has been a general trend toward more complex galls but the herb-stem feeders evolved from ancestors inducing distinct galls and their larval chambers are best understood as cryptic galls. Woody hosts have been colonized only three times, making the apparently irreversible transition from herbs to woody hosts one of the most conservative features of the gall wasp-host plant association. The evolution of host plant preferences is characterized by colonization of preexisting host-plant lineages rather than by parallel cladogenesis. Cynipids are mono- or oligophagous and host-plant choice is strongly phylogenetically conserved. Yet, the few major host shifts have involved remarkably distantly related plants. Many shifts have been onto plant species already exploited by other gall wasps, suggesting that interspecific parasitism among cynipids facilitates colonization of novel host plants.     

Bottom‐up manipulations alter the community structures of galling insects and gall morphs on Neoboutonia macrocalyx trees in a moist tropical rainforest

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Global patterns in local number of insect galling species

Abstract. We evaluate a three-part hypothesis explaining why gall-inducing insect species richness is so high in scleromorphic vegetation: (1) persistence of low nutrient status scleromorphic leaves facilitates the galling habit in warm temperate latitudes; (2) favourable colonization sites for gallers result from reduced hygrothermal stress, high phenolics in the outer cortex of the gall, and reduced carnivore and fungal attack in the gall; and (3) in more mesic sites, mortality is high due to carnivore attack and invasion of galls by fungi. Over 280 samples of local species of galling herbivorous insects from fourteen countries on all continents except Antarctica revealed a strong pattern of highest richness in warm temperate latitudes, or their altitudinal equivalents. The peak of galling species richness on the latitudinal gradient from the equator into the Arctic was between 25 to 38° N or S. Galling species were particularly diverse in sclerophyllous vegetation, which commonly had greater than twelve species per local sample. In mesic, non-sclerophyllous vegetation types the number of galling species was lower with twelve or fewer species present. Many sites in sclerophyllous vegetation supported between thirteen and forty-six galling species locally, including campina islands in Amazonia, cerrado savanna in central Brazil, the Sonoran Desert in Arizona and Mexico, shrubland in Israel, fynbos in South Africa and coastal scleromorphic vegetation in Australia. At the same latitude, or its elevational equivalent, galling species richness was significantly higher in relatively xeric sites when compared to riparian or otherwise mesic habitats, even when scleromorphic vegetation dominated the mesic sites. The results were consistent with the hypothesis and extend to a more general level the patterns and predictions on the biogeography of gall-inducing insects.     

Determinants of assemblage size for the parasitoids of Cecidomyiidae (Diptera)

Summary The parasitoids known to attack 191 phytophagous species of gall midges (Cecidomyiidae) were used to examine factors influencing parasitoid assemblage size. The number of parasitoid species a midge species supports was tested against nine variables describing geographical, biological and ecological attributes of hosts. The apparency of midge larvae was found to have the greatest influence on parasitoid assemblage size; highly visible species support more parasitoids than less visible ones. Pupation site and midge voltinism also significantly affect associated parasitoids, at least for highly apparent hosts. Biogeographic region, host-plant architecture and the plant parts infested were found to be of secondary importance. The surface texture of infested plant parts, the number of midge larvae occupying galls and the diversity of plant tissues infested have minimal apparent effects on parasitoid richness. Parasitoid assemblage size and total parasitism rates were also found to be positively correlated for 73 galling and nongalling midge species, and gallers typically suffer higher levels of parasitism than non-gallers. Using these data to test the enemy hypothesis, which proposes that the galling habit has evolved to escape attack from parasitoids, we conclude that parasitoid pressure cannot account for the presence of galls in the Cecidomyiidae.     

Interactions among three trophic levels and diversity of parasitoids: a case of top-down processes in Mexican tropical dry forest

The objective of this study was to analyze the relationship between plant hosts, galling insects, and their parasitoids in a tropical dry forest at Chamela-Cuixmala Biosphere Reserve in western Mexico. In 120 transects of 30 by 5 m (60 in deciduous forest and 60 in riparian habitats), 29 galling insects species were found and represented in the following order: Diptera (Cecidomyiidae, which induced the greatest abundance of galls with 22 species; 76%), Homoptera (Psylloidea, 6.9%; Psyllidae, 6.9%; Triozidae, 3.4%), Hymenoptera (Tanaostigmatidae, 3.4%; which were rare), and one unidentified morphospecies (3.4%). In all cases, there was a great specificity between galling insect species and their host plant species; one galling insect species was associated with one specific plant species. In contrast, there was no specificity between parasitoid species and their host galling insect species. Only 11 species of parasitoids were associated with 29 galling insect species represented in the following families: Torymidae (18.2%), Eurytomidae (18.2%), Eulophidae (18.2%), Eupelmidae (9.1%), Pteromalidae (9.1%), family Braconidae (9.1%), Platygastridae (9.1%), and one unidentified (9.1%). Most parasitoid species parasitized several gall species (Torymus sp.: 51.1%, Eurytoma sp.: 49.7%, Torymoides sp.: 46.9%). Therefore, the effects of variation in plant defenses do not extend to the third trophic level, because a few species of parasitoids can determine the community structure and composition of galling insect species in tropical plants, and instead, top-down processes seem to be regulating trophic interactions of galling insect species in tropical gall communities.     

The gall-inducing habit has evolved multiple times among the eriococcid scale insects (Sternorrhyncha: Coccoidea: Eriococcidae)

The habit of inducing plant galls has evolved multiple times among insects but most species diversity occurs in only a few groups, such as gall midges and gall wasps. This phylogenetic clustering may reflect adaptive radiations in insect groups in which the trait has evolved. Alternatively, multiple independent origins of galling may suggest a selective advantage to the habit. We use DNA sequence data to examine the origins of galling among the most speciose group of gall-inducing scale insects, the eriococcids. We determine that the galling habit has evolved multiple times, including four times in Australian taxa, suggesting that there has been a selective advantage to galling in Australia. Additionally, although most gall-inducing eriococcid species occur on Myrtaceae, we found that lineages feeding on Myrtaceae are no more likely to have evolved the galling habit than those feeding on other plant groups. However, most gall-inducing species-richness is clustered in only two clades ( Apiomorpha and Lachnodius  +  Opisthoscelis ), all of which occur exclusively on Eucalyptus s.s . The Eriococcidae and the large genus Eriococcus were determined to be non-monophyletic and each will require revision.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 441–452.     

Phenotypic plasticity and similarity among gall morphotypes on a superhost,Baccharis reticularia (Asteraceae)

Understanding factors that modulate plant development is still a challenging task in plant biology. Although research has highlighted the role of abiotic and biotic factors in determining final plant structure, we know little of how these factors combine to produce specific developmental patterns. Here, we studied patterns of cell and tissue organisation in galled and non‐galled organs of Baccharis reticularia, a Neotropical shrub that hosts over ten species of galling insects. We employed qualitative and quantitative approaches to understand patterns of growth and differentiation in its four most abundant gall morphotypes. We compared two leaf galls induced by sap‐sucking Hemiptera and stem galls induced by a Lepidopteran and a Dipteran, Cecidomyiidae. The hypotheses tested were: (i) the more complex the galls, the more distinct they are from their non‐galled host; (ii) galls induced on less plastic host organs, e.g. stems, develop under more morphogenetic constraints and, therefore, should be more similar among themselves than galls induced on more plastic organs. We also evaluated the plant sex preference of gall‐inducing insects for oviposition. Simple galls were qualitative and quantitatively more similar to non‐galled organs than complex galls, thereby supporting the first hypothesis. Unexpectedly, stem galls had more similarities between them than to their host organ, hence only partially supporting the second hypothesis. Similarity among stem galls may be caused by the restrictive pattern of host stems. The opposite trend was observed for host leaves, which generate either similar or distinct gall morphotypes due to their higher phenotypic plasticity. The Relative Distance of Plasticity Index for non‐galled stems and stem galls ranged from 0.02 to 0.42. Our results strongly suggest that both tissue plasticity and gall inducer identity interact to determine plant developmental patterns, and therefore, final gall structure.     

Comparing galling insect richness among Neotropical savannas: effects of plant richness, vegetation structure and super-host presence

Plant species diversity maintains the stability of ecosystems and the diversity of consumer species such as insect herbivores. Considering that gall-inducing insects are highly specialized on their host plants and dependent on the occurrence, abundance and distribution of plants, we evaluated the diversity patterns of gall-inducing insect along Brazilian Neotropical savannas and the potential role of plant species richness, vegetation structure and super-host presence on determining these patterns. We found 1,882 individual plants that belonged to 64 different host plant species grouped in 31 families, associated to 112 galling insect species. The galling richness was positively influenced by plant species richness and the presence of the super-host genus Qualea (Vochysiaceae). Plant species richness explained 48 % of the galling richness and areas with presence of super-hosts had more than twice of galling species than areas where they were absent. On the other hand, we found no evidence that larger plants hosted more species of galling insects. We observed that for the diversity of galling insects in the Brazilian Cerrado, vegetation structure explained almost the same portion as plant richness, because structural variables did not have an effect on residuals of galling richness and plant richness regression. Our findings suggests that plant richness has a more important role on the mitigation of natural enemies and adaptive radiation of galling species, while structural aspects of the vegetation does not seem to have that effect. Furthermore, we show that the super-host taxa provide an increment in local galling richness because they present a great diversity of local number of gall morphospecies (i.e. alpha diversity) and the high turnover of morphospecies among different localities (i.e. beta diversity). Therefore we argue that the quality of resources (richness and super host presence) appears to be a most important factor for the diversity of galling insects in Neotropical systems, than the amount of resources.