The specialization and structure of antagonistic and mutualistic networks of beetles on rainforest canopy trees

Different kinds of species interactions can lead to different structures within ecological networks. Antagonistic interactions (such as between herbivores and host plants) often promote increasing host specificity within a compartmentalized network structure, whereas mutualistic networks (such as pollination networks) are associated with higher levels of generalization and form nested network structures. However, we recently showed that the host specificity of flower‐visiting beetles from three different feeding guilds (herbivores, fungivores, and predators) in an Australian rainforest canopy was equal to that of herbivores on leaves, suggesting that antagonistic herbivores on leaves are no more specialized than flower‐visitors. We therefore set out to test whether similarities in the host specificity of these different assemblages reflect similarities in underlying network structures. As shown before at the species level, mutualistic communities on flowers showed levels of specialization at the network scale similar to those of the antagonistic herbivore community on leaves. However, the network structure differed, with flower‐visiting assemblages displaying a significantly more nested structure than folivores, and folivores displaying a significantly more compartmentalized structure than flower‐visitors. These results, which need further testing in other forest systems, demonstrate that both antagonistic and mutualistic interactions can result in equally high levels of host specialization among beetle assemblages in tropical rainforests. If this is a widespread phenomenon, it may alter our current perceptions of food web dynamics, species diversity patterns, and co‐evolution in tropical rainforests. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 287–295.     

High densities of leaf‐tiers in open habitats are explained by host plant architecture

1. Crown architecture remains one of the least studied plant traits that influence plant–herbivore interactions. The hypotheses that dense crown architecture of mountain birches from open habitats favours leaf‐tying caterpillars through bottom‐up and/or top‐down effects associated with high leaf connectivity were tested. 2. Population densities of leaf‐tying herbivores in open (industrial barren and seashore) habitats were three times as high as in the shaded (forest) habitats. An experimental increase in leaf density by branch binding did not affect foliar consumption by free‐living herbivores but increased consumption by leaf‐tiers. 3. The specific leaf weight was lower in shaded habitats and in bound branches, but branch binding did not influence either the foliar concentrations of carbon and nitrogen or the pupal weight of the most abundant leaf‐tier, Carpatolechia proximella Hbn. (Lepidoptera: Gelechiidae). 4. Caterpillars of C. proximella build several shelters during their lifetime and spend a considerable amount of time outside the shelter, where they excrete most of their faeces. In bound branches, caterpillars built new shelters more frequently than in control branches, and consumed less foliar biomass per shelter. 5. Mortality from parasitoids in bound branches was half that in the control, presumably because the complex environment disrupted parasitoid searching behaviour and/or because of lower damage to leaves from which the shelters were built. 6. It is concluded that the crown architecture associated with high leaf connectivity decreases mortality risks from natural enemies both outside and inside the shelter. Compact and dense crowns of host plants may at least partly explain high population densities of leaf‐tiers in open habitats.     

Micro‐fungi and invertebrate herbivores on birch trees: fungal mediated plant–herbivore interactions or responses to host quality?

We studied interactions between microfungi and herbivores sharing a host tree. In a series of experiments and field observations over a 3‐year period, we compared phenotypic and genetic correlations of fungal frequencies and performance of invertebrate herbivores growing on mature half‐sib progenies of mountain birches (Betula pubescens ssp. czerepanovii) in two environments, a forested river valley and an adjacent higher‐elevation mountain birch woodland. We found little support for direct relation between fungal frequencies and performance of herbivore species. Instead, genetic correlations, particularly between autumnal moth (Epirrita autumnata) and rust fungus (Melampsoridium betulinum), suggest that herbivore performance may be caused by (1) genetic differences in plant quality for fungi and herbivores, or (2) genetic differences in responses to environmental conditions.     

Responses of insect herbivores and herbivory to habitat fragmentation: a hierarchical meta‐analysis

Loss and fragmentation of natural habitats can lead to alterations of plant–animal interactions and ecosystems functioning. Insect herbivory, an important antagonistic interaction is expected to be influenced by habitat fragmentation through direct negative effects on herbivore community richness and indirect positive effects due to losses of natural enemies. Plant community changes with habitat fragmentation added to the indirect effects but with little predictable impact. Here, we evaluated habitat fragmentation effects on both herbivory and herbivore diversity, using novel hierarchical meta‐analyses. Across 89 studies, we found a negative effect of habitat fragmentation on abundance and species richness of herbivores, but only a non‐significant trend on herbivory. Reduced area and increased isolation of remaining fragments yielded the strongest effect on abundance and species richness, while specialist herbivores were the most vulnerable to habitat fragmentation. These fragmentation effects were more pronounced in studies with large spatial extent. The strong reduction in herbivore diversity, but not herbivory, indicates how important common generalist species can be in maintaining herbivory as a major ecosystem process.     

Ecosystem engineering by leaf‐cutting ants: nests of Atta cephalotes drastically alter forest structure and microclimate

1. The role played by Atta species as ecosystem engineers remains poorly investigated despite previous evidence that their nests can impact plant assemblages. 2. In a large remnant of Atlantic forest, we compared forest structure at 36 Atta cephalotes nests to control sites and assessed shifts in microclimate along transects from nests up to 24 m into the forest (11 representative colonies). 3. Nests (average size: 55 m²) were virtually free of understorey vegetation with a high proportion of dead stems (up to 70%). 4. Canopy openness above colonies increased by roughly 40% compared with controls (5.3% at colony vs. 3.7% at control sites). 5. At nest centres, about 6% of the total radiation penetrated through the sparse canopy. Light levels declined exponentially, reaching a third (2%) in the unaffected forest understorey. 6. Likewise, maximum soil temperatures and daily amplitudes declined exponentially from 25 to 23 °C and 1.6 to 0.8 °C, respectively. Soil moisture increased significantly along transects, yet the effect was small and no differences were detected for air temperature and humidity. 7. We extrapolated that individual A. cephalotes nests modify the microclimate in an area of almost 200 m² on average. For the population, this amounts to 6% of the area along forest edges, where colonies are strongly aggregated, compared with only 0.6% in the forest interior. 8. Nests changed microclimate to an extent that has been reported to impact seed germination, plant growth, and survival of seedlings, conclusively demonstrating that leaf‐cutting ants act as ecosystem engineers.     

Stress magnitude matters: different intensities of pulsed water stress produce non‐monotonic resistance responses of host plants to insect herbivores

Abstract 1. Water stress may increase or reduce the suitability of plants for herbivores. The recently proposed ‘pulsed stress hypothesis’ suggests consideration of stress phenology (pulsed vs. continuous stress) to explain these conflicting effects of plant water stress on herbivore performance. 2. This hypothesis was tested for the effect of differing stress intensity on performance and preference of insect herbivores belonging to different feeding guilds, namely leaf‐chewing insects (Spodoptera littoralis caterpillars) and phloem‐feeding insects (Aphis pomi aphids), on apple plants (Malus domestica). The plants were non‐stressed or exposed to a low or high intensity of pulsed water stress. 3. Plant responses to the different stress levels were generally monotonic. Growth, stomatal conductance (g_s), leaf water, and old‐leaf nitrogen concentration decreased, whereas young‐leaf nitrogen concentration and leaf mass per area (LMA) increased with increasing stress intensity. The stable isotope composition of foliar carbon (δ¹³C) responded non‐monotonically to the drought treatments. The δ¹³C values were highest in low‐stress plants, intermediate in high‐stress plants, and lowest in non‐stressed plants. 4. The preference and performance responses of the caterpillars were also non‐monotonic. Non‐stressed plants were intermediately, low‐stress plants least, and high‐stress plants most attractive or suitable. Aphid population growth was highest on non‐stressed plants and lowest on low‐stress plants. 5. The results highlight the importance of water stress intensity for the outcome of interactions between herbivores and drought‐affected plants. They show that pulsed water stress may enhance or reduce insect herbivore performance and plant resistance, depending on stress intensity.     

Not all non‐natives are equally unequal: reductions in herbivore β‐diversity depend on phylogenetic similarity to native plant community

Effects of host plant α‐ and β‐diversity often confound studies of herbivore β‐diversity, hindering our ability to predict the full impact of non‐native plants on herbivores. Here, while controlling host plant diversity, we examined variation in herbivore communities between native and non‐native plants, focusing on how plant relatedness and spatial scale alter the result. We found lower absolute magnitudes of β‐diversity among tree species and among sites on non‐natives in all comparisons. However, lower relative β‐diversity only occurred for immature herbivores on phylogenetically distinct non‐natives vs. natives. Locally in that comparison, non‐native gardens had lower host specificity; while among sites, the herbivores supported were a redundant subset of species on natives. Therefore, when phylogenetically distinct non‐natives replace native plants, the community of immature herbivores is likely to be homogenised across landscapes. Differences in communities on closely related non‐natives were subtler, but displayed community shifts and increased generalisation on non‐natives within certain feeding guilds.     

From natural woodlands to cultivated land: diversity of fruit‐feeding butterflies and beetles in the mid‐Zambezi valley,northern Zimbabwe

We tested the effect of cultivation on butterfly (Nymphalidae: Charaxes) and beetle (Coleoptera: Scarabaeidae: Cetoniinae) species richness and abundance along a cultivation intensification gradient. Results showed significant differences in species richness and abundance between natural woodlands and cultivated landscapes with larger differences in areas of high cultivation intensity. The results indicate that natural woodland clearing for cultivation purposes has negative impacts on arthropod diversity, a situation more severe in highly intensified cultivated areas. Our results imply that mosaics of different land‐use units, each in a different phase of clearance‐cultivation‐abandonment‐recovery‐clearance cycle could counter the negative effects of cultivation intensity on arthropod diversity.     

Continuous exposure to the deterrents cis‐jasmone and methyl jasmonate does not alter the behavioural responses of Frankliniella occidentalis

Behavioural responses of Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), a generalist, cell sap‐feeding insect species with piercing‐sucking mouthparts, after continuous exposure to two deterrent secondary plant compounds are investigated. We compared in choice assays on bean leaf discs, the settling, feeding, and oviposition preferences of F. occidentalis females that had no experience with the two fatty acid derivatives methyl jasmonate and cis‐jasmone before testing (naïve thrips) vs. females that had been exposed to the deterrent compounds before testing (experienced thrips). The thrips were exposed to the deterrents at low or high concentrations for varied time periods and subsequently tested on bean leaf discs treated with the respective deterrent at either a low or a high concentration. Frankliniella occidentalis females avoided settling on the deterrent‐treated bean leaf discs for an observation period of 6 h, independent of their previous experience. Our results demonstrate that feeding and oviposition deterrence of the jasmonates to the thrips were not altered by continuous exposure of the thrips to the jasmonates. Habituation was not induced, neither by exposure to the low concentration of the deterrents nor by exposure to the high concentration. These results indicate that the risk of habituation to two volatile deterrent compounds after repeated exposure is not evident in F. occidentalis. This makes the two compounds potential candidates to be integrated in pest management strategies.     

Mesophyll cell‐sucking herbivores (Cicadellidae: Typhlocybinae) on rainforest trees in Papua New Guinea: local and regional diversity of a taxonomically unexplored guild

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Feeding response of Aedes aegypti and Anopheles dirus (Diptera: Culicidae) using out‐of‐date human blood in a membrane feeding apparatus

The colonization of Aedes aegypti and Anopheles dirus was performed using out‐of‐date human blood from a blood bank as a nutritional supply dispensed from a common artificial feeder. Preserved human blood was collected and used for feeding on days 5, 15, and 25 after date of expiration and dispensed from a common artificial feeder to rear the mosquitoes. Ae. aegypti had a feeding rate of 78.7, 62, and 18% at the respective intervals while An. dirus had a rate of 80, 56.8, and 7.3% on the same respective days. Direct feeding on live hamsters resulted in a rate of 96 and 90% for Ae. aegypti and An. dirus, respectively. Although egg production rates decreased from the day 5 feeding to the day 25 feeding, all of the developmental stages resulting from An. dirus fed at day 5 and 15 showed insignificant differences when compared with direct feeding on the blood of a hamster.     

Distribution and host specificity of the thistle‐feeding tortoise beetle Cassida vibex (Coleoptera: Chrysomelidae) in southwestern Hokkaido,northern Japan

The thistle‐feeding tortoise beetle Cassida vibex (Coleoptera: Chrysomelidae) is widespread in the Palearctic region. In Japan, this species has been recorded only from the largest island, Honshu. Here we report the occurrence of this species on Hokkaido, the most northern main island of Japan, along with detailed distributional records from the southwestern part of the island (southern Oshima Peninsula) and information on host plants. We also present the results of laboratory experiments on adult feeding preference and larval developmental performance to determine the specificity of C. vibex for four thistle species (Cirsium grayanum, Cir. alpicola, Cir. yezoense and Cir. aomorense; Asteraceae) common in southwestern Hokkaido. Cassida vibex was detected at only four sites among 97 sites investigated on the Oshima Peninsula, in strong contrast to the common occurrence of the congeneric thistle feeder C. rubiginosa (74 of 97 sites). Cassida vibex was found mainly on Cir. alpicola in the wild, but adult beetles showed no feeding preference for this or three other thistle species when given a choice in the laboratory. Larval performance of C. vibex was similar on Cir. alpicola, Cir. yezoense and Cir. aomorense but significantly lower on Cir. grayanum. Since Cir. alpicola often forms large clumps of individual plants, whereas Cir. yezoense and Cir. aomorense are scattered in their habitats, and C. vibex was collected only at sites where there were dense patches of Cir. alpicola, we postulate that the persistence of C. vibex is possible only where a substantial amount of suitable host plants is available.     

Comparison of early life‐stage strategies in temperate freshwater fish species: trade‐offs are directed towards first feeding of larvae in spring and early summer

Based on the analysis of 12 egg and larval variables and temperature of 65 temperate freshwater fish species, the possible relationships between oocyte diameter, larval size at hatch, time and temperature were reassessed and the main early life‐stage strategies were described and compared. Time and degree‐days required to reach hatching and mixed feeding were weakly related to oocyte diameter and strongly to temperature. These results are chiefly because oocyte diameter and yolk reserves are weakly related and temperature strongly increases tissue differentiation rate, activity of hatching glands and embryo motility. Strong positive relationships were found between larval size and oocyte diameter and degree‐days for incubation. No relationship was found between larval size and degree‐days from hatching to mixed feeding and between degree‐days for incubation and degree‐days from hatching to mixed feeding. These last two results are chiefly because the developmental stages at hatching and at the onset of exogenous feeding are not fixed in ontogeny and are not directly related to either larval size or degree‐days for incubation, but more probably are species specific. Whatever the spawning season, which can occur almost all year long, the different trade‐offs at the early life‐stages ensure that most larvae are first feeding during spring, when food size and abundance are the most appropriate.     

First molecular identification of the vertebrate hosts of Culicoides imicola in Europe and a review of its blood‐feeding patterns worldwide: implications for the transmission of bluetongue disease and African horse sickness

Culicoides (Diptera: Ceratopogonidae) are vectors of pathogens that affect wildlife, livestock and, occasionally, humans. Culicoides imicola (Kieffer, 1913) is considered to be the main vector of the pathogens that cause bluetongue disease (BT) and African horse sickness (AHS) in southern Europe. The study of blood‐feeding patterns in Culicoides is an essential step towards understanding the epidemiology of these pathogens. Molecular tools that increase the accuracy and sensitivity of traditional methods have been developed to identify the hosts of potential insect vectors. However, to the present group's knowledge, molecular studies that identify the hosts of C. imicola in Europe are lacking. The present study genetically characterizes the barcoding region of C. imicola trapped on farms in southern Spain and identifies its vertebrate hosts in the area. The report also reviews available information on the blood‐feeding patterns of C. imicola worldwide. Culicoides imicola from Spain feed on blood of six mammals that include species known to be hosts of the BT and AHS viruses. This study provides evidence of the importance of livestock as sources of bloodmeals for C. imicola and the relevance of this species in the transmission of BT and AHS viruses in Europe.     

Land‐use legacies in a central Appalachian forest: differential response of trees and herbs to historic agricultural practices

Question: Are contemporary herb and tree patterns explained by historic land use practices? If so, are observed vegetation patterns associated with life‐history characteristics, soil properties, or other environmental variables? Location: Southeastern Ohio, USA. Methods: Using archival records, currently forested sites were identified with distinct land use histories: cultivated, pasture (but not plowed), and reference sites which appear to have never been cleared. Trees were recorded by size and species on twenty 20 m × 20 m plots; percent cover was estimated for each herb species in nested 10 m × 10 m plots. Environmental characteristics were noted, and soil samples analysed for nutrient availability and organic matter. Nonmetric multidimensional scaling ordination was performed separately on both tree and herb datasets to graphically characterize community composition among plots. Life‐history traits were investigated to explain observed compositional differences. Results: Vegetation patterns were explained by current environmental gradients, especially by land‐use history. Cultivated and pasture sites had similar tree composition, distinct from reference sites. Herb composition of pasture and reference sites was similar and distinct from cultivated sites, suggesting the ‘tenacity’ of some forest herbs on formerly cleared sites. Tilling removes rhizomatous species, and disfavors species with unassisted dispersal. These life‐history traits were underrepresented on cultivated sites, although ant‐dispersed species were not. Conclusions: Historic land‐use practices accounted for as much variation in species composition as environmental gradients. Furthermore, trees and herbs responded differently to past land‐use practices. Life‐history traits of individual species interact with the nature of disturbance to influence community composition.     

Population size structure of trees in a semi‐arid African savanna: Species differ in vulnerability to a changing environment and reintroduction of elephants

The size structure of a plant population provides a snapshot of potential population trend and weak inference on past history. Size structure of 24 tree species was sampled in a savanna reserve in order to assess their potential vulnerability to local extirpation following the reintroduction of elephants. Regression of number per size class based on an expected reverse‐J structure for a healthy population was undertaken in order to classify trend of each species. On account of the weak realism of this expected structure in semi‐arid environments, log‐normal mixture models were also fitted to individual observations in order to identify states, whose distribution, size and shape can indicate potential population persistence. Population structure of 12 species approximated a reverse‐J, indicating potential future growth. Although regressions of seven species were not significant, suggesting vulnerable populations, mixture models of all except one showed multiple states, indicating variable regeneration and recruitment over time. A single state of adult plants indicates that Sclerocarya birrea was vulnerable to local extirpation if adult mortality escalates. Some apparently declining populations displayed multiple states of established individuals suggesting temporally varying regeneration and recruitment, attributed mainly to rainfall variation, but which were judged as likely to persist. Poor recent regeneration of most species was attributed to well below‐average rainfall for the preceding decade. Different savanna tree species apparently ensure persistence based on individualistic responses to population perturbations, but persistence of some following the reintroduction of elephants may be threatened if certain size classes experience escalated mortality. Subsequent monitoring has supported this concern.     

Future warmer seas: increased stress and susceptibility to grazing in seedlings of a marine habitat‐forming species

Increases in seawater temperature are expected to have negative consequences for marine organisms. Beyond individual effects, species‐specific differences in thermal tolerance are predicted to modify species interactions and increase the strength of top‐down effects, particularly in plant–herbivore interactions. Shifts in trophic interactions will be especially important when affecting habitat‐forming species such as seagrasses, as the consequences on their abundance will cascade throughout the food web. Seagrasses are a major component of coastal ecosystems offering important ecosystem services, but are threatened by multiple anthropogenic stressors, including warming. The mechanistic understanding of seagrass responses to warming at multiple scales of organization remains largely unexplored, especially in early‐life stages such as seedlings. Yet, these early‐life stages are critical for seagrass expansion processes and adaptation to climate change. In this study, we determined the effects of a 3 month experimental exposure to present and predicted mean summer SST of the Mediterranean Sea (25°C, 27°C, and 29°C) on the photophysiology, size, and ecology (i.e., plant‐herbivore interactions) of seedlings of the seagrass Posidonia oceanica. Warming resulted in increased mortality, leaf necrosis, and respiration as well as lower carbohydrate reserves in the seed, the main storage organ in seedlings. Aboveground biomass and root growth were also limited with warming, which could hamper seedling establishment success. Furthermore, warming increased the susceptibility to consumption by grazers, likely due to lower leaf fiber content and thickness. Our results indicate that warming will negatively affect seagrass seedlings through multiple direct and indirect pathways: increased stress, reduced establishment potential, lower storage of carbohydrate reserves, and increased susceptibly to consumption. This work provides a significant step forward in understanding the major mechanisms that will drive the capacity of seagrass seedlings to adapt and survive to warming, highlighting the potential additive effects that herbivory will have on ultimately determining seedling success.     

Life‐history traits related to diapause in univoltine and bivoltine populations of Ypthima multistriata (Lepidoptera: Satyridae) inhabiting similar latitudes

In most temperate insects, diapause strategies and voltinism generally exhibit latitudinal clines, supporting the concept that they represent adaptations to climate. In contrast, in the satyrine butterfly Ypthima multistriata Butler, local populations with different voltinism patterns are geographically intermingled, suggesting that life‐history traits related to diapause may differ even between geographically and phylogenetically close populations. In this study, we experimentally examined the critical photoperiod for diapause induction and the larval developmental period in two univoltine and two bivoltine populations of Y. multistriata, all of which inhabit virtually the same latitude (34.652–34.750°N). We found that the critical photoperiod for diapause induction was longer in the univoltine populations than in the bivoltine populations. Moreover, the larval period under the long day length treatment was different among populations in both sexes, although significant differences were also detected between populations with the same voltinism. These results indicate that in Y. multistriata, life‐history traits related to diapause can not be attributed merely to climatic conditions such as temperature or day length, which depend largely on latitude. Therefore, we suggest that biotic elements, such as leaf toughness, as well as abiotic elements should be taken into account in attempts to explain the enigmatic pattern of geographic variation in the diapause strategies of Y. multistriata.