Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Phytophagous insect assemblages and the regional species pool: patterns and asymmetries

We describe three models predicting relationships between: (a) the taxonomic composition of the regional species pool of phytophagous insects and the composition of the phytophagous insect fauna on a host taxon; and (b) the faunal composition of two host taxa. The predictions of these models were compared with empirical data representing the regional pool of phytophages in Central Europe and the faunas of two plant taxa: the cabbage plants (Brassicaceae) and the thistles (Asteraceae: Cardueae). Three important findings emerge at a general level. (1) Different taxonomic levels of insects (orders, families, genera) of the regional pool and on the investigated host taxa are well correlated in terms of species richness, but there is no consistent trend in the variance explained by this correlation across taxonomic levels. (2) The model considering evolutionary interactions and speciation processes is consistent with patterns found in the empirical data. (3) Asymmetries in sampled species numbers of insect families on both host taxa may be accounted for by reference to the biology of these insects. We conclude that the faunas of single host taxa can provide the basis for extrapolating to the regional pool, at least at high taxonomic levels.     

Plant Size as Determinant of Species Richness of Herbivores,Natural Enemies and Pollinators across 21 Brassicaceae Species

Large plants are often more conspicuous and more attractive for associated animals than small plants, e.g. due to their wider range of resources. Therefore, plant size can positively affect species richness of associated animals, as shown for single groups of herbivores, but studies usually consider intraspecific size differences of plants in unstandardised environments. As comprehensive tests of interspecific plant size differences under standardised conditions are missing so far, we investigated effects of plant size on species richness of all associated arthropods using a common garden experiment with 21 Brassicaceae species covering a broad interspecific plant size gradient from 10 to 130 cm height. We recorded plant associated ecto- and endophagous herbivores, their natural enemies and pollinators on and in each aboveground plant organ, i.e. flowers, fruits, leaves and stems. Plant size (measured as height from the ground), the number of different plant organ entities and their biomass were assessed. Increasing plant size led to increased species richness of associated herbivores, natural enemies and pollinating insects. This pattern was found for ectophagous and endophagous herbivores, their natural enemies, as well as for herbivores associated with leaves and fruits and their natural enemies, independently of the additional positive effects of resource availability (i.e. organ biomass or number of entities and, regarding natural enemies, herbivore species richness). We found a lower R² for pollinators compared to herbivores and natural enemies, probably caused by the high importance of flower characteristics for pollinator species richness besides plant size. Overall, the increase in plant height from 10 to 130 cm led to a 2.7-fold increase in predicted total arthropod species richness. In conclusion, plant size is a comprehensive driver of species richness of the plant associated arthropods, including pollinators, herbivores and their natural enemies, whether they are endophagous or ectophagous or associated with leaves or fruits.     

Do alien plants escape from natural enemies of congeneric residents? Yes but not from all

As predicted by the enemy release hypothesis, plants are supposedly less attacked by herbivores in their introduced range than in their native range. However, the nature of the natural enemies, in particular their degree of specificity may also affect the level of enemy escape. It is therefore expected that ectophagous invertebrate species, being generally considered as more generalists than endophagous species, are more prompt to colonise alien plants. In Swiss, Siberian and Russian Far East arboreta, we tested whether alien woody plants are less attacked by native herbivorous insects than native congeneric woody plant species. We also tested the hypothesis that leaf miners and gall makers show stronger preference for native woody plants than external leaf chewers. In all investigated regions, leaf miners and gall makers were more abundant and showed higher species richness on native woody plants than on congeneric alien plants. In contrast, external leaf chewers did not cause more damage to native plants than to alien plants, possibly because leaf chewers are, in general, less species specific than leaf miners and gall makers. These results, obtained over a very large number of plant-enemy systems, generally support the hypothesis that alien plants partly escape from phytophagous invertebrates but also show that different feeding guilds may react differently to the introduction of alien plants.     

Novel application of species richness estimators to predict the host range of parasites

Host range is a critical life history trait of parasites, influencing prevalence, virulence and ultimately determining their distributional extent. Current approaches to measure host range are sensitive to sampling effort, the number of known hosts increasing with more records. Here, we develop a novel application of results-based stopping rules to determine how many hosts should be sampled to yield stable estimates of the number of primary hosts within regions, then use species richness estimation to predict host ranges of parasites across their distributional ranges. We selected three mistletoe species (hemiparasitic plants in the Loranthaceae) to evaluate our approach: a strict host specialist (Amyema lucasii, dependent on a single host species), an intermediate species (Amyema quandang, dependent on hosts in one genus) and a generalist (Lysiana exocarpi, dependent on many genera across multiple families), comparing results from geographically-stratified surveys against known host lists derived from herbarium specimens. The results-based stopping rule (stop sampling bioregion once observed host richness exceeds 80% of the host richness predicted using the Abundance-based Coverage Estimator) worked well for most bioregions studied, being satisfied after three to six sampling plots (each representing 25 host trees) but was unreliable in those bioregions with high host richness or high proportions of rare hosts. Although generating stable predictions of host range with minimal variation among six estimators trialled, distribution-wide estimates fell well short of the number of hosts known from herbarium records. This mismatch, coupled with the discovery of nine previously unrecorded mistletoe-host combinations, further demonstrates the limited ecological relevance of simple host-parasite lists. By collecting estimates of host range of constrained completeness, our approach maximises sampling efficiency while generating comparable estimates of the number of primary hosts, with broad applicability to many host-parasite systems.     

The structure of the phytophagous insect fauna on the introduced weed Solidago altissima in Switzerland

Solidago altissima L. was introduced into Europe as an ornamental plant from North America more than 100 years ago and the phytophagous insect fauna of it was recently examined in Switzerland where it has become an important weed in disturbed habitats. Rhizomes and aerial parts were examined and all insects collected in summer were tested in a no-choice feeding test. 18 out of 55 phytophagous insects were found feeding on S. altissima in Switzerland and for the remaining 37 the relation with the plant was not determined. The insects that have expanded their host range to feed on S. altissima since its introduction to Switzerland are almost solely opportunistic, unspecialized ectophages not closely attuned to the growth cycle of S. altissima. Only 4% of the insects were specialists and 9% endophagous. In contrast in North America S. altissima supports 25% specialists and 17% endophages. The native Solidago virgaurea L. in Switzerland supports many more specialists (28%) and endophages (23%) than S. altissima here. Possible reasons why almost none of these have switched to S. altissima are discussed. A comparison is made between the number of insects recruited by S. altissima in Switzerland with the number recruited by the exotic plants Heracleum mantegazzianum Som. & Lev. (Apiaceae) in Switzerland and Xanthium occidentale Bertol. (Asteraceae) in Australia. Reasons for different levels of insect recruitment are discussed. The high number of phytophagous insect species found on S. altissima in North America shows that there should be no shortage of possible control agents and any insects imported into Switzerland for the biological control of S. altissima would find largely unexploited food sources awaiting them.     

Parasite richness/sampling effort/host range: the fancy three-piece jigsaw puzzle

In this article, Jean-Fran?ois Guégan and Clive Kennedy propose an alternative explanation for the confounding effects of host geographical range and sampling effort on parasite species richness using pathway analysis procedure. They suggest that much of the species richness revealed by sampling effort is also a reflection of host range. Thus, the total contribution of host range logically incorporates a contribution from sampling effort. The implications of indirect effects of host range on richness estimates have not previously been discussed, and the authors here attempt to redress the balance. The contribution of host range to richness, as derived from control of sampling effort on richness estimates, therefore, is a mathematical expression that does not take into account the cause-and-effect nature of things.     

Assemblages of endophagous insects on Asteraceae in São Paulo cerrados

A survey of the endophagous insects fauna associated to Asteraceae capitula was carried out from 2000 to 2002 in eight cerrado sensu stricto sites located in the Brazilian state of Sdo Paulo. Sixty-four endophagous species of Diptera and Lepidoptera were recorded from 49 asteracean host plants. Approximately half of the species were obtained from a single locality, with a large proportion emerging from a single sample (unicates). Thirty percent of the species were singletons (i.e. only one individual was recorded). The large proportion of rare species suggests a high species turnover among localities. Lepidopteran species were recorded on more host species than dipterans, confirming their more polyphagous food habit, also observed in other Brazilian biomes and in Europe. We conclude that the studied cerrado localities, all within S?o Paulo State, are isolated with its invertebrate fauna composed of many rare and exclusive species. We suggest that the maintenance of Asteraceae biodiversity and their endophagous insects depend on the conservation of all cerrado remnants in the state.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

Entomofauna associated to horticultural crops under organic and conventional practices in Córdoba, Argentina

Farming practices and the addition of chemical synthetic substances in conventional agroecosystems are detrimental mainly to natural enemies of phytophagous insects, diminishing the natural regulation of pest insects. On the other hand, in organic agriculture, biological processes and care of the environment are favoured, hence an increase in insect biodiversity is predicted in this type of systems. In this work, abundance, richness of insects and proportion of functional groups were compared through a single quantitative sampling of insects in horticultural crop fields, three under organic and three under conventional management practices. Insect species richness, total and for guilds (phytophagous and entomophagous insects) were significantly higher in organic orchards, and also was the abundance of entomophagous insects. Richness and abundance of all insect orders (with exception of Homoptera abundance), were higher in orchards under organic management, being significative the differences for richness of Coleoptera and richness and abundance of Hymenoptera. Similar tendencies were observed in data obtained through sweep net in weeds. These results suggest that organic practices increase the diversity of species, particularly that of natural enemies.     

Trophic associations and specialization of phytophagous beetles (Coleoptera: Chrysomeloidea,Curculionoidea) in the east of the Russian Plain

Data on the trophic associations of beetles with plants in the east of the Russian Plain are summarized and comparative analysis of host specialization of different groups of phytophagous beetles is performed. In terms of the width of the regional trophic spectrum, monophages and narrow oligophages prevail among the Curculionoidea as a whole and in the families Curculionidae and Apionidae in particular, while moderate and broad oligophages prevail in the Chrysomeloidea and in the family Chrysomelidae. Two-thirds of the regional fauna (66%) of Curculionoidea are closely associated with plants of one genus; by contrast, in Chrysomeloidea almost 40% of the species can develop on plants from different genera of one family, the fraction of the narrowly specialized forms comprising only 43%. The higher level of trophic specialization of weevils (Curculionidae, Apionidae) and seed beetles (Bruchidae), as compared to leaf beetles (Chrysomelidae), is probably due to the larval endophagy of most species of these families. Analysis of the distribution of beetles over host plants has shown that the specialized forms are associated with plants of 65 families (about 60% of the regional flora in the east of the Russian Plain). Distribution of beetles over plant families is very non-uniform. Most of the specialized forms (78%) are associated with plants of 15 families, three of which (Asteraceae, Fabaceae, and Brassicaceae) include hosts of more than onethird of the beetle species (37%). Monophages and narrow oligophages are recorded on 201 genera of plants from 59 families. Polyphagous species are recorded on plants of 58 families. The specific features of the distribution of phytophagous beetles over host plants (as compared to other insects) is a high fraction of species developing on coenophobes (particularly those of the family Brassicaceae) typical of the pioneer stages of successions with sparse herbaceous cover, and a small number of species associated with grasses and sedges. These features are most conspicuous in the fauna of Curculionidae.     

Larval habits, host-plant associations, and speciation in nematine sawflies (Hymenoptera: Tenthredinidae)

Adaptive radiations consist of two intertwined processes, diversification of species and diversification of their ecological niches, but it is unclear whether there is a causal link between the processes. In phytophagous insects, ecological diversification mainly involves shifts in host-plant associations and in larval feeding habits (internal or external) on different plant parts, and several observations indicate that speciation is facilitated by host shifts. Data on host use in individual species suggest that internal feeders are less likely to colonize new hosts than external-feeding taxa and, consequently, increases in collective host ranges and species numbers should be slowed down in endophagous lineages. We tested these related hypotheses by using phylogenetic information to reconstruct the evolutionary history of larval resource use in the sawfly subfamily Nematinae, a group of 1000 plus species with a broad range of niches: the subfamily's combined host range includes over 20 plant families, and larvae may feed externally on leaves or needles, or internally, for example, in buds, fruits, leaves, or galls. The results show that: (1) Most internally feeding groups have evolved independently from external-feeding ancestors, but several distinct internal habits have appeared convergently multiple times; (2) Shifts among host taxa are clearly more common than changes in larval habits; (3) The majority of host switches have occurred among phylogenetically close plant groups, but many shifts are manifest among distantly related, ecologically proximate hosts; (4) Although external feeding characteristic of the common ancestor of Nematinae is associated with relatively high rates of host-shifting, internal feeders are very conservative in their host use; (5) In contrast, the effect of endophagy on speciation probabilities is more variable: net speciation rates are lowered in most internal-feeding groups, but a striking exception is found in species that induce galls on Salicaceae. The loose connection between collective host ranges and species diversity provides empirical support for theoretical models suggesting that speciation rates are a function of a complex interplay between "intrinsic" niche width and resource heterogeneity.     

Geographical variation in the 'bottom-up' control of diversity: fleas and their small mammalian hosts

Aim We searched for signs of the ‘bottom‐up’ diversity effect in the association between fleas (Siphonaptera) and their small mammalian hosts (Rodentia, Insectivora and Lagomorpha). We asked (1) whether a strong dependence of flea species richness on host species richness is characteristic for both Palaeoarctic and Nearctic realms; (2) if yes, whether the ratio of host species per flea species along the host diversity gradient is similar between the Palaeoarctic and Nearctic; and (3) whether factors other than host species richness (i.e. geographical position, climate and landscape) might better explain variation in flea species richness than host species richness. Location The study used previously published data on species richness of fleas and their small mammalian hosts from 26 Palaeoarctic and 19 Nearctic regions. Methods We regressed the number of flea species on the number of small mammal species across regions, separately for Palaeoarctic and Nearctic realms, using both non‐transformed data as well as data corrected for the confounding effects of host sampling effort and sampling area. To test whether flea species richness is determined by external factors unrelated to the host, we used stepwise multiple regressions of flea species richness against host species richness and parameters describing the geographical position, climate and relief of a region. Results When non‐transformed data were analysed, flea species richness was positively correlated with host species richness in both the Palaeoarctic and Nearctic, although the slopes of the two regressions differed significantly. After removal of the confounding effects of host sampling effort and sampling area, Palaeoarctic flea species richness remained strongly positively correlated with host species richness, whereas in the Nearctic, flea species richness appeared to be completely independent of host species richness. Results of the multiple regressions using corrected data demonstrated that in the Palaeoarctic, flea species richness was correlated with both the number of host species and the mean altitude of the region, whereas in the Nearctic, flea species richness only tended to be weakly correlated with latitude (however, this correlation turned out to be non‐significant after Bonferroni correction). Main conclusions We found evidence of bottom‐up control of flea diversity in the Palaeoarctic regions only, and not in the Nearctic. We explore several potential explanations for the different patterns observed in the two biogeographical realms, including differences in (1) levels of host specialization, (2) history of host–parasite associations and (3) landscape effects on flea diversification. We conclude that these factors combine to create different macroecological patterns in different biogeographical realms, and that diversity is not governed by the same forces everywhere.     

Dynamics of host plant use and species diversity in Polygonia butterflies (Nymphalidae)

The ability of insects to utilize different host plants has been suggested to be a dynamic and transient phase. During or after this phase, species can shift to novel host plants or respecialize on ancestral ones. Expanding the range of host plants might also be a factor leading to higher levels of net speciation rates. In this paper, we have studied the possible importance of host plant range for diversification in the genus Polygonia (Nymphalidae, Nymphalini). We have compared species richness between sistergroups in order to find out if there are any differences in number of species between clades including species that utilize only the ancestral host plants ('urticalean rosids') and their sisterclades with a broader (or in some cases potentially broader) host plant repertoire. Four comparisons could be made, and although these are not all phylogenetically or statistically independent, all showed clades including butterfly species using other or additional host plants than the urticalean rosids to be more species-rich than their sisterclade restricted to the ancestral host plants. These results are consistent with the theory that expansions in host plant range are involved in the process of diversification in butterflies and other phytophagous insects, in line with the general theory that plasticity may drive speciation.     

Does host‐plant diversity explain species richness in insects? A test using Coccidae (Hemiptera)

1. The megadiverse herbivores and their host plants are a major component of biodiversity, and their interactions have been hypothesised to drive the diversification of both. 2. If plant diversity influences the diversity of insects, there is an expectation that insect species richness will be strongly correlated with host‐plant species richness. This should be observable at two levels (i) more diverse host‐plant groups should harbour more species of insects, and (ii) the species richness of a group of insects should correlate with the richness of the host groups it uses. However, such a correlation is also consistent with a hypothesis of random host use, in which insects encounter and use hosts in proportion to the diversity of host plants. Neither of these expectations has been widely tested. 3. These expectations were tested using data from a species‐rich group of insects – the Coccidae (Hemiptera). 4. Significant positive correlations were found between the species richness of coccid clades (genera) and the species richness of the host‐plant family or families upon which the clades occur. On a global scale, more closely related plant families have more similar communities of coccid genera but the correlation is weak. 5. Random host use could not be rejected for many coccids but randomisation tests and similarity of coccid communities on closely related plant families show that there is non‐random host use in some taxa. Overall, our results support the idea that plant diversity is a driver of species richness of herbivorous insects, probably via escape‐and‐radiate or oscillation‐type processes.     

Is the composition of phytophagous insects and parasitic fungi among trees predictable?

We explore the relationship between the pairwise similarity of assemblages of exploiters (phytophagous insects and parasitic fungi) and pairwise genetic distance, range overlap, niche overlap as well as habitat overlap of host trees. Presence of exploiters was extracted from published literature for 23 tree genera occurring in central Europe (6164 host records of phytophagous insects and 860 host records of parasitic fungi). Across all pairs of tree genera, we found a strong negative correlation between the pairwise similarity of assemblages and genetic distances of hosts. This close correlation is due to deep differences in the composition of assemblages on coniferous and deciduous tree genera. Range, niche and habitat overlap were always of much less importance than genetic distance to explain the variation of pairwise similarity of assemblages of exploiters, although some correlations were significant. Therefore in general host switches of exploiters between related hosts are more important that host switches between hosts co-occurring in the same habitat. We found a robust relationship of the pairwise similarity of assemblages of insects and the pairwise similarity of assemblages of fungi which points to the possibility that insects are vectors for parasitic fungi which promotes correlated switches of insects and fungi.     

Different relationships between galling and non-galling herbivore richness and plant species richness: a meta-analysis

The plant richness hypothesis (PRH) is used to explain herbivorous insect richness based on the number of plant species, predicting a positive relationship. However, the influence of plant richness on insect distribution can become stronger with greater levels of specialization of herbivores. In this meta-analysis, I tested whether there is any difference in the correlation force recorded between studies that investigated endophagous versus exophagous herbivores, and galling versus non-galling guilds, in order to determine whether more specialized groups have a stronger relationship. Furthermore, I calculated whether effect sizes were homogeneous between galling studies carried out at local and regional scales, and between tropical and temperate regions. A total of 52 correlations were analyzed between plant species richness and herbivore species richness, with 18 correlations derived from galling herbivores and 34 from non-galling herbivores. The effect sizes were significant and positive in all studies, being higher for endophages than for exophages, and for galling than for non-galling studies. These results provide evidence that groups of insects with a higher level of host specialization and specificity (e.g., endophagous and galling) exhibit a greater dependence on plant richness. There was no difference in effect sizes for galling studies between the local and regional level or between tropical and temperate groups. Despite the large variability found for galling studies, effect sizes were consistent independently of climatic region and latitudinal variation. These results suggest that the PRH for galling insects can be generalized to most ecosystem and vegetation types.     

Species richness of Macrolepidoptera on Finnish deciduous trees and shrubs

Summary Species richness of Macrolepidoptera on Finnish trees and shrubs was analysed by means of stepwise regression analysis. The explaining variables were plant frequency, geographical range, plant height, number of relatives and leaf size.Total frequency of the host plant, which correlated strongly with range, explained 57% of the observed variance of lepidopteran species richness on deciduous trees and shrubs. Height of plant and number of relatives explained significantly the residual variation and altogether these three variables explained 71% of the variance of species richness.Analyses at the plant genus level gave similar results and frequency, height and number of relatives explained 78% of the variance of species richness of Macrolepidoptera on deciduous plant genera.When conifers were included in the analysis leaf size also becomes a significant variable. Leaf size can, however, act as a dummy variable which effectively distinguishes conifers from deciduous trees.The validity of different models explaining herbivore species richness on plants is discussed. The results of this study favoured more than earlier studies the importance of relatedness of host plants as a factor which determines the species richness of herbivores.     

The genetic architecture of a niche: variation and covariation in host use traits in the Colorado potato beetle

The genetic basis of host plant use by phytophagous insects can provide insight into the evolution of ecological niches, especially phenomena such as specialization and phylogenetic conservatism. We carried out a quantitative genetic analysis of multiple host use traits, estimated on five species of host plants, in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). Mean values of all characters varied among host plants, providing evidence that adaptation to plants may require evolution of both behavioral (preference) and post-ingestive physiological (performance) characteristics. Significant additive genetic variation was detected for several characters on several hosts, but not in the capacity to use the two major hosts, a pattern that might be caused by directional selection. No negative genetic correlations across hosts were detected for any 'performance' traits, i.e. we found no evidence of trade-offs in fitness on different plants. Larval consumption was positively genetically correlated across host plants, suggesting that diet generalization might evolve as a distinct trait, rather than by independent evolution of feeding responses to each plant species, but several other traits did not show this pattern. We explored genetic correlations among traits expressed on a given plant species, in a first effort to shed light on the number of independent traits that may evolve in response to selection for host-plant utilization. Most traits were not correlated with each other, implying that adaptation to a novel potential host could be a complex, multidimensional 'character' that might constrain adaptation and contribute to the pronounced ecological specialization and the phylogenetic niche conservatism that characterize many clades of phytophagous insects.     

Helminth parasite richness among vertebrates

The numbers of intestinal helminth species (parasite richnesS) recorded from each of 488 vertebrate host species are compared using data compiled from the published literature. Associations between parasite richness, sampling effort, host size and host habitat (aquatic versus terrestrial) are assessed using a method designed to control for phylogenetic association. Parasite richness increases with the number of surveys on which each estimate of parasite richness is based (sampling effort). When the effects of sampling effort are controlled for, there remains a strong positive relationship between parasite richness and host body size. There is no tendency for aquatic hosts to harbour more parasite species than terrestrial hosts independently of differences in sampling effort and body size. The results are interpreted in the context of hosts representing habitats for parasite colonization, resource allocation between parasite species, and the age of the major mammalian radiations.