Gall‐forming insect species richness along a non‐scleromorphic vegetation rainfall gradient in South Africa: The importance of plant community composition

Abstract Currently there is no single accepted hypothesis to explain gall‐forming insect species richness at a particular locality. Hygrothermal stress, soil nutrient availability, plant species richness, plant structural complexity, plant family or genus size, and host plant geographical range size have all been implicated in the determination of gall‐forming insect species richness. Previous studies of such richness at xeric sites have included predominantly scleromorphic vegetation, usually on nutrient‐poor soils. This study is the first to investigate gall‐forming insect species richness of xeric, non‐scleromorphic vegetation. Two habitat types were sampled at each of five localities across a rainfall gradient in the savanna biome of South Africa. The habitat types differed with respect to plant species composition and topography. Gall‐forming insect species richness did not increase with increasing hygrothermal stress or decreasing soil fertility. Rather, gall‐forming insect species richness was largely dependent on the presence of Terminalia sericea as well as other members of the Combretaceae and Mimosaceae. Plots where all these taxa were present had the highest gall‐forming insect species richness, up to 15 species, whereas plots with none of these taxa had a maximum of four galling‐insect species. Despite herb, shrub and tree strata not differing in gall‐forming insect species richness, insect galls were more common on woody than non‐woody plants. Also, stem galls were more frequent than apical or leaf galls. An alternative hypothesis to explain local gall‐forming insect species richness is suggested: galling insects may preferentially select those plant species with characteristics such as chemical toxicity, mechanical strength, degree of lignification or longevity that can be manipulated to benefit the galler. Thus plant community composition should be considered when attempting to explain gall‐forming insect species richness patterns.     

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.     

Diversity of insect‐induced galls along a temperature– rainfall gradient in the tropical savannah region of the Northern Territory,Australia

Evidence regarding the effect of temperature and rainfall on gall‐inducing insects is contradictory: some studies indicate that species richness of gall‐inducing insects increases as environments become hotter and drier, while others suggest that these factors have no effect. The role of plant species richness in determining species richness of gall‐inducing insects is also controversial. These apparent inconsistencies may prove to be due to the influence of soil fertility and the uneven distribution of gall‐inducing insect species among plant taxa. The current study tested hypotheses about determinants of gall‐inducing insect species richness in a way different to previous studies. The number of gall‐inducing insect species, and the proportion of species with completely enclosed galls (more likely to give protection against heat stress and desiccation), were measured in replicate plots at five locations along a 500‐km N‐S transect in the seasonal tropics of the Northern Territory, Australia. There is a strong temperature–rainfall gradient along this transect during the wet season. Plant species lists had already been compiled for each collection plot. All plots were at low elevation in eucalypt savannah growing on infertile soils. There was no evidence to suggest that hot, dry environments in Australia have more gall‐inducing insect species than cooler, wetter environments, or that degree of enclosure of galls is related to protecting insects from heat stress and desiccation. The variable number of gall‐inducing insect species on galled plant species meant that plant species richness did not influence gall species richness. Confirmation is still required that low soil fertility does not mask temperature–rainfall effects and that galls in the study region are occupied predominantly in the wet season, when the temperature–rainfall gradient is most marked.     

Biodiversity of galling insects: historical, community and habitat effects in four neotropical savannas

Five hypotheses were tested to explain the pattern of galling insect species richness in four neotropical savanna physiognomies, 'canga ', 'campo sujo', 'cerrado' s. st., and 'cerradão', that occur in Minas Gerais, southeastern Brazil. We found 125 species of galling insects on 80 host plant species. The increase of plant species richness explained 35% of the variation in galling insect richness, corroborating the plant species richness hypothesis. Most of the galling species occurred on trees, followed by shrubs, and herbs. However, the difference in mean number of galls was only statistically significant between herbs and trees, corroborating partially the plant structural complexity hypothesis. A significant relationship was observed between galling species richness and density of herbs, and shrubs, corroborating partially the resource concentration hypothesis. Galling insect richness showed a negative correlation with magnesium, potassium, and zinc on soil, corroborating the soil fertility hypothesis. The content of magnesium, potassium, iron and CTC (T) explained 72% of the variation in galling insect richness. Plant family size positively influenced galling insect richness, corroborating the plant family size hypothesis. Overall, the results corroborate the hypothesis that predicts that habitat stress is the main factor generating the patterns of galling insect richness in Brazilian savannas.     

Spatial patterns of herbivory by gall-forming insects: a test of the soil fertility hypothesis in a Mexican tropical dry forest

It has been proposed that fertile soils reduce the incidence of gall-forming insect (GFI) species in plant communities. This is known as the soil fertility hypothesis. The main objective of this study was to analyze the spatial distribution of GFI species under different habitats in a tropical dry forest at the Chamela-Cuixmala Biosphere Reserve, Mexico. Eight habitats which differ in soil type, topography, nutrient availability and vegetation were chosen. We found that 38 GFI species specialize on their host plant species. GFI species richness was negatively correlated with phosphorous and nitrogen availability. Using phosphorous as an indicator of soil fertility, we found low frequency and density of GFI on fertile soils. Our study indicates that soil fertility is one of the factors that negatively affects the patterns of spatial distribution of species richness, incidence and abundance of GFI at the community level in two different ways: i) indirectly affecting GFI species richness in plants adapted to infertile soils and ii) directly affecting GFI responses to plant traits of hosts found in a fertility gradient.     

Effects of host plant architecture on colonization by galling insects

Abstract: To study the abundance and occurrence of herbivore insects on plants it is important to consider plant characteristics that may control the insects. In this study the following hypotheses were tested: (i) an increase of plant architecture increases species richness and abundance of gall‐inducing insects and (ii) plant architecture increases gall survival and decreases parasitism. Two hundred and forty plants of Baccharis pseudomyriocephala Teodoro (Asteraceae) were sampled, estimating the number of shoots, branches and their biomass. Species richness and abundance of galling insects were estimated per module, and mortality of the galls was assessed. Plant architecture influenced positively species richness, abundance and survival of galls. However, mortality of galling insects by parasitoids was low (13.26%) and was not affected by plant architecture, thus suggesting that other plant characteristics (a bottom‐up pressure) might influence gall‐inducing insect communities more than parasitism (a top‐down pressure). The opposite effect of herbivore insects on plant characteristics must also be considered, and such effects may only be assessed through manipulative experiments.     

Galling insect diversity patterns: the resource synchronisation hypothesis

A new hypothesis is proposed to explain the insect gall-maker species richness pattern found by Price et al. (1998; J. Biogeogr. 25: 581–591). First, it is necessary to consider an explicit model for speciation in this group of insects, and host-shifts seem to be the most likely way by which gall-makers can speciate. Since most galls are induced on young plant tissue, and mainly on leaves, the dynamics of leaf flushing in different vegetation types (how many plant species flush synchronously) can be of capital importance determining possible host-shifts for dispersing insect females. I suggest that vegetation types with synchronous leaf flushing are more likely to harbour higher insect gall-maker species richness, and this is why the pattern found by Price et al. (1998) showed low species richness of these insects for tropical rain forests and high richness for Mediterranean-like vegetation. Additionally, Mediterranean-like vegetation suffers seasonal influence of fires, and plants resprouting in response to this can provide further synchronous leaf flushes during which more host-shifts can occur. Plant diversity is also of capital importance underlying the host-shifting process in different biomes. Finally, I suggest a few possible ways of testing the hypothesis.     

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.

     

Edge effect and species–area relationships in the gall-forming insect fauna of natural forest patches in the Brazilian Pantanal

The aim of this study was to (i) measure differences in species richness between edge habitats versus interior habitats, or more precisely the edge effect, and (ii) test the species–area relationship for gall-forming insects in natural forest patches in a Brazilian floodplain (Pantanal of Mato Grosso do Sul). These patches are regionally known as capões, basically composed of woody vegetation. Twenty-seven patches were surveyed. In each patch two transects were conducted for gall sampling. One transect encircled the patches while the other was conducted in the interior of the patch, totaling 54h of sampling. Host plant and galling insect species composition differed quite characteristically between the edge and the interior of patches, but galling insect richness did not. When insect gall richness was expressed as the ratio between insect gall and host plant richness (gall per plant ratio), a weak species–area relationship was found. Our results suggest that the number of galling insects per individual plant is not affected by the size of the patch. Despite these results, the natural forest patches found in this region seem well suited for long-term studies addressing species–area relationships. With regard to herbivorous insects, these studies should be combined with research on host plant dynamics during flooding and dry seasons.     

Gall-forming and free-feeding herbivory along vertical gradients in a lowland tropical rainforest: the importance of leaf sclerophylly

In contrast to most insect guilds, gall-forming insects are thought to reach highest diversity on sclerophyllous vegetation, such as Neotropical savannas and Mediterranean vegetation types. The water and nutrient stress endured by meristems of canopy trees in tall wet tropical rainforests may cause leaf sclerophylly. Hence, the upper canopies of such ecosystems may represent a suitable habitat for gall-forming insects. At the San Lorenzo Protected Area, Panama, we estimated free-feeding herbivory and gall densities within five sites in 2003 and 2004, by surveying leaves in vertical and horizontal transects. In each sample, we recorded leaf density (mature and young foliage), free-feeding herbivore damage and number of galls, including the presence of live larvae, parasitoids or fungi. We surveyed 43 994 leaves, including 231 plants and 73 tree and liana species. We collected 5014 galls from 17 host-plant species, including 32 gall species of which 59% were restricted to the canopy (overall infestation rates: 2.4% in 2003, 5.5% in 2004). In 2003, 16% of the galls were occupied by live larvae, against 5% in 2004. About 17–20% of leaves surveyed suffered from free-feeding herbivory. Leaf sclerophylly increased significantly with sampling height, while free-feeding herbivory decreased inversely. Conversely, the number of live galls collected in the canopy was 13–16 times higher than in the understorey, a pattern consistent across sites and years. Hence, the probability of gall survivorship increased with increasing leaf sclerophylly as death by fungi, parasitoids or accidental chewing were greater in the understorey. Increasing harsh ecophysiological conditions towards the upper canopy appear favourable to galls-forming population maintenance, in support of the hypothesis of harsh environment. Hence, gall diversity and abundance in the upper canopy of tall tropical forests are perhaps among the highest in the world.     

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.     

Effects of genetic variability and habitat of Qualea parviflora (Vochysiaceae) on herbivory by free-feeding and gall-forming insects

• Background and Aims Differences in the chemical and physical traits of plants caused by both genetic and habitat characteristics may influence attack by herbivores. Leaves of Qualea parviflora (Vochysiaceae), a common tree of different habitats in the Brazilian Neotropical savannas (cerrado), are susceptible to severe attack by herbivorous free-living and gall-forming insects. Attack by free-living and gall-forming insects within and between populations of Q. parviflora were examined and it was determined to what extent genetic variability (detected by RAPD markers), phenotypic characteristics of the plants and habit traits influence the number of free-living and gall-forming insect species and individuals attacking the plants, and the intensity of attack.• Methods On four occasions in 2000, leaves were sampled from ten individual trees in each of three types of vegetation in the cerrado: campo sujo, cerrado sensu stricto and cerradão at the Ecological Station of Pirapitinga (ESP), in Três Marias, north-western Minas Gerais, Brazil. Genetic variability was detected by RAPD markers, and concentrations of nutrients, phenols and tannins, sclerophylly and pre-dawn water potential of leaves were measured. Water and nutrient contents in the soil below each tree characterized the habitat. The free-living and gall-forming herbivorous insects were determined.• Key Results Of 69 RAPD markers analysed, 41 were polymorphic and were used for analyses of genetic variability of Q. parviflora. Most of the variability occurred within habitats, accounting for 97·65 % of the genetic variability. Plants in the cerrado sensu stricto and campo sujo were the most similar. There were no significant associations between genetic similarity and the chemical and physical traits of Q. parviflora, or with habitat, nor was there significant correlation between phenotypic and habitat traits. Increasing concentrations of tannins and sulphur, C : N ratio and sclerophylly correlated with increasing percentage of leaf area damaged by herbivores. Decreased sclerophylly, concentration of tannins and C : N ratio, and increased concentration of nutrients in leaves correlated with increased severity of attack by gall-forming insects.• Conclusions Nutrient concentration in the soil had more influence, indirectly, on free-feeding insects than did composition of Q. parviflora leaves. However, gall-forming insects are affected more by leaf quality, attacking fewer sclerophyllous leaves, with larger nutrient but smaller tannin concentrations.Key words: Cerrado, genetic variability, gall attack, herbivory, insect galls, plant quality, Qualea parviflora, RAPD, Vochysiaceae     

Gall‐forming insects in a lowland tropical rainforest: low species diversity in an extremely specialised guild

1. Gall‐forming insects are a guild of endophages that exhibit a high level of fidelity to their host plants, however, their level of host specificity is seldom explicitly tested. 2. Gall‐forming insect taxa from 32 species of woody tropical plants with resolved phylogenetic relationships were collected and reared, representing 15 families from all the major clades of angiosperms, at three lowland rainforest locations in Madang, Papua New Guinea (PNG). 3. More than 8800 galled plant parts were collected from 78 gall morphospecies at an average of 2.4 per host plant. Total species richness at the sampling sites was estimated to be 83–89. All but one morphospecies were monophagous resulting in an effective specialisation of 0.98. 4. Specific leaf weight, foliar nitrogen, the presence of latex, and the successional preference of plant species all gave a phylogenetic signal, but only plant successional preference influenced the species richness of galls on analysis of phylogenetically independent contrasts. Gall species were distributed randomly among host plant species and showed no preference for any particular plant lineage. Furthermore, most gall‐forming taxa were evenly dispersed across the host plant phylogeny. 5. In the tropical rainforests of New Guinea, gall‐forming insects are ubiquitous but occur in species‐poor assemblages. Local species richness is closely tied to the diversity of angiosperms owing to very high host specificity. 6. Finally, galler species richness data from the literature across habitats and latitudes were compared and suggest that tropical rainforests may be richer in galls than previously acknowledged.     

虫瘿与其生物群落及寄主植物间的关系

虫瘿是自然界极其常见的生物现象,是植物与昆虫互作的奇特产物。本文对虫瘿生物群落多样性、虫瘿与其生物群落的关系以及虫瘿与寄主植物的关系进行概述,探讨了致瘿昆虫在虫瘿形成中的作用、植物化学对致瘿昆虫产卵交配行为的影响以及植物激素在虫瘿形成中的作用,最后对虫瘿今后的研究方向进行了讨论,为虫瘿的致瘿生物学及其瘿内生物相互关系的进一步研究奠定基础。     

Insect species richness tracking plant species richness in a diverse flora: gall-insects in the Cape Floristic Region, South Africa

The Cape Floristic Region (CFR) is one of the most plant-species-rich regions in the world. It is also a warm temperate region and hypothetically should have high gall-insect species richness, making it interesting to investigate the relationship between the insects of the region and the rich flora. The relationship between gall-insect species richness (GSR) and plant richness was investigated for the Fynbos and for representatives of vegetation of the whole CFR. Samples (of up to 600 plants per transect for Fynbos) of woody shrubs were investigated for the presence of galls. The species richness of these insects was quantified, as well as plant species richness for each transect. GSR for Fynbos was compared to global figures for GSR. Fynbos harboured significantly more gall-insect species than other CFR vegetation types. GSR was positively correlated with CFR plant richness. GSR also closely tracked plant richness in Fynbos. GSR was not significantly influenced by other variables (elevation and aspect), suggesting that plant richness per se was an important factor in generating GSR. Fynbos GSR is comparable to other sclerophyllous regions of high GSR globally, corroborating that this vegetation type is conducive to gall-insect diversification. There is likely to be a high percentage of gall-insect endemism in the Fynbos, as might be expected from the high host fidelity of this insect group. Received: 22 September 1997 / Accepted: 16 February 1998     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Plant architecture and meristem dynamics as the mechanisms determining the diversity of gall-inducing insects

Plant architecture is considered to affect herbivory intensity, but it is one of the least studied factors in plant–insect interactions, especially for gall-inducing insects. This study aimed to investigate the influence of plant architecture on the speciose fauna of gall-inducing insects associated with 17 species of Baccharis. Five architectural variables were evaluated: plant height, number of fourth-level shoots, biomass, average level and number of ramifications. The number of galling species associated with each host plant species was also determined. To test the effects of plant architecture on gall richness at the individual level, we used another data set where the number of fourth-level shoots and gall richness were determined for B. concinna, B. dracunculifolia, and B. ramosissima every 3 weeks during 1 year. The average similarity between host species based on gall fauna was low (9%), but plants with the same architectural pattern tended to support similar gall communities. The most important architectural trait influencing gall richness at the species level was the number of fourth-level shoots, which is indicative of the availability of plant meristems, a fundamental tissue for gall induction and development. This variable also showed a positive correlation with gall richness at the individual level. We propose that variations in gall richness among host species are driven by interspecific differences in plant architecture via availability of young, undifferentiated tissue, which is genetically controlled by the strength of the apical dominance. Plant architecture should have evolutionary consequences for gall communities, promoting insect radiation among architecturally similar plants through host shift and sympatric speciation. We also discuss the role of plant architecture in the global biogeography of gall-inducing insects. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of non-native plants on the native insect community of Delaware

Due to the lack of a co-evolutionary history, the novel defenses presented by introduced plants may be insurmountable to many native insects. Accordingly, non-native plants are expected to support less insect biomass than native plants. Further, native insect specialists may be more affected by introduced plants than native generalist herbivores, resulting in decreased insect diversity on non-native plants due to the loss of specialists. To test these hypotheses, we used a common garden experiment to compare native insect biomass, species richness, and the proportion of native specialist to native generalist insects supported by 45 species of woody plants. Plants were classified into three groupings, with 10 replicates of each species: 15 species native to Delaware (Natives), 15 non-native species that were congeneric with a member of the Native group (Non-native Congeners), and 15 non-native species that did not have a congener present in the United States (Aliens). Native herbivorous insects were sampled in May, June, and July of 2004 and 2005. Overall, insect biomass was greater on Natives than Non-native Congeners and Aliens, but insect biomass varied unpredictably between congeneric pair members. Counter to expectations, Aliens held more insect biomass than did Non-native Congeners. There was no difference in species richness or the number of specialist and generalist species collected among the three plant groupings in either year, although our protocol was biased against sampling specialists. If these results generalize to other studies, loss of native insect biomass due to introduced plants may negatively affect higher trophic levels of the ecosystem.     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Effect of hybridization of the Quercus crassifolia×Quercus crassipes complex on the community structure of endophagous insects

In a previous study, we showed that the geographic proximity of hybrid plants to the allopatric areas of parental species increases their morphological and genetic similarity with them. In the present work, we explored whether the endophagous fauna of hybrid plants show the same pattern. We studied the canopy species richness, diversity and composition of leaf-mining moths (Lepidoptera: Tischeridae, Citheraniidae) and gall-forming wasps (Hymenoptera: Cynipidae) associated with two species of red oaks (Quercus crassifolia and Quercus crassipes) and their interspecific hybrid (Quercus×dysophylla Benth pro sp.) in seven hybrid zones in central Mexico, during four seasons in 2 years. The study was conducted on 194 oak trees with known genetic status [identified by leaf morphology and molecular markers (random amplified polymorphic DNAs)], and the results indicate a bidirectional pattern of gene flow. Hybrid plants supported intermediate levels of infestation of gall-forming and leaf-mining insects compared to their putative parental species. The infestation level of leaf-mining insects varied significantly following the pattern: Q. crassifolia>hybrids>Q. crassipes, whereas the gall-forming insects showed an inverse pattern. A negative and significant relationship was found between these two types of insect guilds in each host taxa, when the infestation percentage was evaluated. It was found that 31.5% (n=11) of the endophagous insects were specific to Q. crassipes, 22.9% (n=8) to Q. crassifolia, and 8.6% (n=3) to hybrid individuals. The hybrid bridge hypothesis was supported in the case of 25.7% (n=9) of insects, which suggests that the presence of a hybrid intermediary plant may favor a host herbivore shift from one plant species to another. Greater genetic diversity in a hybrid zone is associated with greater diversity in the endophagous community. The geographic proximity of hybrid plants to the allopatric site of a parental species increases their similarity in terms of endophagous insects and the Eje Neovolcánico acts as a corridor favoring this pattern. Electronic Supplementary Material Supplementary material is available for this article at