Hyperparasitoids Use Herbivore-Induced Plant Volatiles to Locate Their Parasitoid Host

Plants respond to herbivory with the emission of induced plant volatiles. These volatiles may attract parasitic wasps (parasitoids) that attack the herbivores. Although in this sense the emission of volatiles has been hypothesized to be beneficial to the plant, it is still debated whether this is also the case under natural conditions because other organisms such as herbivores also respond to the emitted volatiles. One important group of organisms, the enemies of parasitoids, hyperparasitoids, has not been included in this debate because little is known about their foraging behaviour. Here, we address whether hyperparasitoids use herbivore-induced plant volatiles to locate their host. We show that hyperparasitoids find their victims through herbivore-induced plant volatiles emitted in response to attack by caterpillars that in turn had been parasitized by primary parasitoids. Moreover, only one of two species of parasitoids affected herbivore-induced plant volatiles resulting in the attraction of more hyperparasitoids than volatiles from plants damaged by healthy caterpillars. This resulted in higher levels of hyperparasitism of the parasitoid that indirectly gave away its presence through its effect on plant odours induced by its caterpillar host. Here, we provide evidence for a role of compounds in the oral secretion of parasitized caterpillars that induce these changes in plant volatile emission. Our results demonstrate that the effects of herbivore-induced plant volatiles should be placed in a community-wide perspective that includes species in the fourth trophic level to improve our understanding of the ecological functions of volatile release by plants. Furthermore, these findings suggest that the impact of species in the fourth trophic level should also be considered when developing Integrated Pest Management strategies aimed at optimizing the control of insect pests using parasitoids.     

Forecasting Distributional Responses of Limber Pine to Climate Change at Management-Relevant Scales in Rocky Mountain National Park

Resource managers at parks and other protected areas are increasingly expected to factor climate change explicitly into their decision making frameworks. However, most protected areas are small relative to the geographic ranges of species being managed, so forecasts need to consider local adaptation and community dynamics that are correlated with climate and affect distributions inside protected area boundaries. Additionally, niche theory suggests that species'' physiological capacities to respond to climate change may be underestimated when forecasts fail to consider the full breadth of climates occupied by the species rangewide. Here, using correlative species distribution models that contrast estimates of climatic sensitivity inferred from the two spatial extents, we quantify the response of limber pine (Pinus flexilis) to climate change in Rocky Mountain National Park (Colorado, USA). Models are trained locally within the park where limber pine is the community dominant tree species, a distinct structural-compositional vegetation class of interest to managers, and also rangewide, as suggested by niche theory. Model forecasts through 2100 under two representative concentration pathways (RCP 4.5 and 8.5 W/m²) show that the distribution of limber pine in the park is expected to move upslope in elevation, but changes in total and core patch area remain highly uncertain. Most of this uncertainty is biological, as magnitudes of projected change are considerably more variable between the two spatial extents used in model training than they are between RCPs, and novel future climates only affect local model predictions associated with RCP 8.5 after 2091. Combined, these results illustrate the importance of accounting for unknowns in species'' climatic sensitivities when forecasting distributional scenarios that are used to inform management decisions. We discuss how our results for limber pine may be interpreted in the context of climate change vulnerability and used to help guide adaptive management.     

Mutual Use of Trail-Following Chemical Cues by a Termite Host and Its Inquiline

Termite nests are often secondarily inhabited by other termite species ( = inquilines) that cohabit with the host. To understand this association, we studied the trail-following behaviour in two Neotropical species, Constrictotermes cyphergaster (Termitidae: Nasutitermitinae) and its obligatory inquiline, Inquilinitermes microcerus (Termitidae: Termitinae). Using behavioural experiments and chemical analyses, we determined that the trail-following pheromone of C. cyphergaster is made of neocembrene and (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol. Although no specific compound was identified in I. microcerus, workers were able to follow the above compounds in behavioural bioassays. Interestingly, in choice tests, C. cyphergaster prefers conspecific over heterospecific trails while I. microcerus shows the converse behaviour. In no-choice tests with whole body extracts, C. cyphergaster showed no preference for, while I. microcerus clearly avoided heterospecific trails. This seems to agree with the hypothesis that trail-following pheromones may shape the cohabitation of C. cyphergaster and I. microcerus and reinforce the idea that their cohabitation is based on conflict-avoiding strategies.     

Peroxidase Profiling Reveals Genetic Linkage between Peroxidase Gene Clusters and Basal Host and Non-Host Resistance to Rusts and Mildew in Barley

Background

Higher plants possess a large multigene family encoding secreted class III peroxidase (Prx) proteins. Peroxidases appear to be associated with plant disease resistance based on observations of induction during disease challenge and the presence or absence of isozymes in resistant vs susceptible varieties. Despite these associations, there is no evidence that allelic variation of peroxidases directly determines levels of disease resistance.

Methodology/Principal Findings

The current study introduces a new strategy called Prx-Profiling. We showed that with this strategy a large number of peroxidase genes can be mapped on the barley genome. In order to obtain an estimate of the total number of Prx clusters we followed a re-sampling procedure, which indicated that the barley genome contains about 40 peroxidase gene clusters. We examined the association between the Prxs mapped and the QTLs for resistance of barley to homologous and heterologous rusts, and to the barley powdery mildew fungus. We report that 61% of the QTLs for partial resistance to P. hordei, 61% of the QTLs for resistance to B. graminis and 47% of the QTLs for non-host resistance to other Puccinia species co-localize with Prx based markers.

Conclusions/Significance

We conclude that Prx-Profiling was effective in finding the genetic location of Prx genes on the barley genome. The finding that QTLs for basal resistance to rusts and powdery mildew fungi tend to co-locate with Prx clusters provides a base for exploring the functional role of Prx-related genes in determining natural differences in levels of basal resistance.     

Identification of the Cues Used in the Host Finding Behavior During the Phoretic Association Ceroptera rufitarsis (Diptera: Sphaeroceridae) and Dung Beetles (Coleoptera: Scarabaeidae)

Ceroptera rufitarsis (Meigen) (Diptera: Sphaeroceridae) is a tiny fly phoretic on several roller dung beetles of the genus Scarabaeus L. (Coleoptera: Scarabaeidae). We investigated the attractiveness of Scarabaeus sacer L. and S. cicatricosus Lucas both under field and laboratory conditions. Several cues potentially responsible for the host finding behavior were tested in order to determine the basis of attraction of Ceroptera rufitarsis to Scarabaeus beetles. Visual cues, especially host movements, constituted the main factor responsible for host finding behavior and host choice while chemical cues did not play any role in host discrimination.     

The Use of Olfactory and Visual Cues in Host Choice by the Capsid Bugs Lygus rugulipennis Poppius and Liocoris tripustulatus Fabricius

Lygus rugulipennis Poppius and Liocoris tripustulatus Fabricius (Heteroptera: Miridae) are pests of glasshouse cucumber and sweet pepper crops respectively. L. rugulipennis has a wide range of foodplants, but L. tripustulatus is specialised with very few food plants. We report behavioural assessments to investigate whether either species exhibits a preference for salad over wild hosts, and whether the role of olfaction and vision in response to cues from host plants can be distinguished. Olfactory responses to leaves were tested in choice chambers. L. rugulipennis was presented nettle (wild host) and a salad leaf of cucumber or sweet pepper, where the salad leaves had higher nitrogen content. L. tripustulatus was tested with nettle and sweet pepper of two different nitrogen contents. Female L. rugulipennis spent more time on the cucumber salad host, and chose it first most often, but males showed no preference. Neither sex discriminated between sweet pepper or nettle leaves, but males made more first contacts with sweet pepper. Neither sex of L. tripustulatus discriminated between sweet pepper and nettle leaves when the sweet pepper had higher nitrogen. When the plant species contained equivalent nitrogen both sexes spent more time on nettle. There was no difference in first choice made by either sex. When visual stimuli were available, and leaves had equivalent nitrogen, L. rugulipennis showed no preference and L. tripustulatus preferred nettle leaves. We conclude that the generalist L. rugulipennis has the ability to use remote olfactory cues for host choice whereas the specialist L. tripustulatus relies mainly on contact chemosensory and gustatory cues.     

Volatile Organic Compound Analysis of Host and Non-Host Poplars for <Emphasis Type="Italic">Trypophloeus klimeschi</Emphasis> (Coleoptera: Curculionidae: Ipinae)

Trypophloeus klimeschi Eggers was first discovered in Xinjiang Province and had strong selection specificity for Populus alba var. pyramidalis Bunge. There was an outbreak of this beetle in the northwest shelter forest of China, resulting in significant economic losses and loss of ecological benefits. Based on a prior long-term field investigation, T. klimeschi had a different extent of injuries for different ages of P. alba var. pyramidalis and other Populus in the same area were not selected by T. klimeschi. To further explore the specificity volatile compounds, this study involved selecting host and non-host trees to analyse the volatile chemical profile of host and non-host poplars of T. klimeschi. The main volatile compounds of the host poplar P. alba var. pyramidalis for different physiological statuses and those of three other non-host poplars (P. alba L., P. tomentosa Carr., and P. dakuanensis Hsu) were analysed through solid-phase micro extraction (SPME) coupled with thermal desorption and gas chromatography-mass spectrometry (GC-MS). The major compound groups were aldehydes, esters, alcohols, ketones, phenols, terpenes and alkanes. Comparative analysis of the changes in the different physiological stages of P. alba var. pyramidalis and other non-host Populus volatile substances was conducted, and the results showed that 2-hydroxy-benzaldehyde, nonanal, decanal, 2-methyl-butanal, (Z)-3-hexen-1-ol benzoate, methyl benzoate, methyl salicylate, geraniol and salicyl alcohol might act as attractants for T. klimeschi, and 2-hexenal, hexanal, 2-cyclohexen-1-one, caryophyllene, eugenol, benzyl alcohol, and eucalyptol could be deterrents for T. klimeschi. These experiments may lead to the optimisation of a synthetic lure that may be used to detect and monitor T. klimeschi.     

Edaphic limitations to growth and photosynthesis in Sierran and Great Basin vegetation

Summary Soils derived from hydrothermally altered andesite support unique communities of Sierran conifers (Pinus ponderosa Laws. and P. jeffreyi Grev. and Balf.) amongst sagebrush (Artemisia tridentata Nutt.) vegetation in the western Great Basin. Plants grown in soil derived from hydrothermally altered bedrock had lower growth rates, total biomass, and net photosynthetic rates than plants grown in soil derived from unaltered andesite of the same formation. Total dry mass was 10 to 28% lower for conifers grown in altered soil whereas dry mass of Artemisia tridentata and Bromus tectorum L. was reduced by over 90%. Results from a nutrient amendment experiment indicated that low phosphorus was the dominant limitation in altered soil, and phosphorus-deficiency affected growth primarily by limiting leaf area development rather than direct inhibition of photosynthesis. The proportionately greater reduction of biomass for Artemisia and Bromus grown in altered soil supports our hypothesis that Great Basin vegetation is excluded from altered soil by intolerance to nutrient deficiency. The Sierran conifers growing on this rock type are therefore free of competition for water with Great Basin vegetation and are able to persist in an exceptionally dry climate.     

Use of Genetic and Physical Mapping to Locate the Spinal Muscular Atrophy Locus between Two New Highly Polymorphic DNA Markers

The gene for autosomal recessive forms of spinal muscular atrophy (SMA) has recently been mapped to chromosome 5ql3, within a 4-cM region between the blocks D5S465/D5S125 and MAP-1B/D5S112. We identified two new highly polymorphic microsatellite DNA markers—namely, AFM265wf5 (D5S629) and AFM281yh9 (D5S637)—which are the closest markers to the SMA locus. Multilocus analysis by the location-score method was used to establish the best estimate of the SMA gene location. Our data suggest that the most likely location for SMA is between locus D5S629 and the block D5S637/D5S351/MAP-1B/D5S112/D5S357. Genetic analysis of inbred SMA families, based on homozygosity by descent and physical mapping using mega-YACs, gave additional information for the loci order as follows: cen–D5S6–D5S125/D5S465–D5S435–D5S629–SMA–D5S637–D5S351–MAP–1B/D5S112–D5S357–D5S39–tel. These data give the direction for bidirectional walking in order to clone this interval and isolate the SMA gene.     

Challenges and limitations to native species restoration in the Great Basin,USA

The Great Basin of the western USA is an arid region characterized by high spatial and temporal variability. The region experienced high levels of ecological disturbance during the early period of Euro-American settlement, especially from about 1870–1935. The principal plant communities of the Great Basin are sagebrush steppes, dominated by various Artemisia shrubs and perennial bunchgrasses that represent the largest rangeland ecosystem in North America. In low to mid-elevation sagebrush communities, exotic annual grasses have displaced native plant species and are associated with a dramatic increase in size and frequency of wildfires. Degradation in this region is driven by processes that cause the loss of mature bunchgrasses, which, when intact, limit exotic annual grass invasion. Historically, large economic investments to restore degraded Great Basin rangeland through establishment of native bunchgrasses, principally utilizing heavily mechanized agronomic approaches, have been met with limited success. A multitude of environmental factors contribute to the lack of restoration success in this region, but seedling mortality from freezing and drought has been identified as a primary demographic limitation to successful bunchgrass establishment. Novel approaches to overcoming limitations to bunchgrass establishment will be required for restoration success. Increased national concern and a near listing of the greater sage-grouse, a steppe-obligate species, to Endangered Species status, has spurred greater regional support and collaboration across a diversity of stakeholder groups such as state and federal land and wildlife management agencies, county planners, and ranchers.     

Linking nitrogen partitioning and species abundance to invasion resistance in the Great Basin

Resource partitioning has been suggested as an important mechanism of invasion resistance. The relative importance of resource partitioning for invasion resistance, however, may depend on how species abundance is distributed in the plant community. This study had two objectives. First, we quantified the degree to which one resource, nitrogen (N), is partitioned by time, depth and chemical form among coexisting species from different functional groups by injecting ¹⁵N into soils around the study species three times during the growing season, at two soil depths and as two chemical forms. A watering treatment also was applied to evaluate the impact of soil water content on N partitioning. Second, we examined the degree to which native functional groups contributed to invasion resistance by seeding a non-native annual grass into plots where bunchgrasses, perennial forbs or annual forbs had been removed. Bunchgrasses and forbs differed in timing, depth and chemical form of N capture, and these patterns of N partitioning were not affected by soil water content. However, when we incorporated abundance (biomass) with these relative measures of N capture to determine N sequestration by the community there was no evidence suggesting that functional groups partitioned different soil N pools. Instead, dominant bunchgrasses acquired the most N from all soil N pools. Consistent with these findings we also found that bunchgrasses were the only functional group that inhibited annual grass establishment. At natural levels of species abundance, N partitioning may facilitate coexistence but may not necessarily contribute to N sequestration and invasion resistance by the plant community. This suggests that a general mechanism of invasion resistance may not be expected across systems. Instead, the key mechanism of invasion resistance within a system may depend on trait variation among coexisting species and on how species abundance is distributed in the system.     

Spatial Distribution of Mountain Pine Beetle Outbreaks in Relation to Climate and Stand Characteristics:A Dendroecological Analysis

Principal components analysis, followed by K-means cluster analysis, was used to detect variations in the timingand magnitude of Pinus contorfa Dougl. ex Loud. growth releases attributed to mountain pine beetle outbreaks in31 stands of central British Columbia. Four major growth release patterns were identified from 1970 to 2000.Variations in the timing of growth releases among clustered stands corresponded well to aerial survey dataindicating the timing of beetle outbreaks in the study area. Redundancy analysis was used to determine howvariations in the timing and magnitude of growth releases attributed to beetle outbreaks changed with variationsin climate or stand conditions over the study area. The first RDA axis, which accounted for 39% of the variations ingrowth patterns among stands, was significantly (P<0.05) correlated with gradients in the percentage of pine instands killed by mountain pine beetle, summer aridity, variation in summer precipitation, distance from initialinfestation site, average pine age, and maximum August temperatures. The second RDA axis explained 6% of thevariations and was significantly correlated with gradients in the beetle climate suitability index, extreme coldmonth temperatures, and site index. Comparisons of growth release patterns with aerial survey data and redun-dancy analyses indicated that dendrochronological techniques are useful for identifying mountain pine beetleoutbreaks in central British Columbia, particularly among stands that had a density high enough to produce agrowth release signal. Provided future studies account for interannual weather fluctuations, identification ofgrowth increases due to stand thinning caused by beetle outbreaks will be useful for reconstructing the history ofbeetle outbreaks over much longer time periods.     

Spatial Distribution of Mountain Pine Beetle Outbreaks in Relation to Climate and Stand Characteristics： A Dendroecological Analysis

Principal components analysis, followed by K-means cluster analysis, was used to detect variations in the timing and magnitude of Pinus contorta Dough ex Loud. growth releases attributed to mountain pine beetle outbreaks in 31 stands of central British Columbia. Four major growth release patterns were identified from 1970 to 2000. Variations in the timing of growth releases among clustered stands corresponded well to aerial survey data indicating the timing of beetle outbreaks in the study area. Redundancy analysis was used to determine how variations in the timing and magnitude of growth releases attributed to beetle outbreaks changed with variations in climate or stand conditions over the study area. The first RDA axis, which accounted for 39% of the variations in growth patterns among stands, was significantly （P〈0.05） correlated with gradients in the percentage of pine in stands killed by mountain pine beetle, summer aridity, variation in summer precipitation, distance from initial infestation site, average pine age, and maximum August temperatures. The second RDA axis explained 6% of the variations and was significantly correlated with gradients in the beetle climate suitability index, extreme cold month temperatures, and site index. Comparisons of growth release patterns with aerial survey data and redundancy analyses indicated that dendrochronological techniques are useful for identifying mountain pine beetle outbreaks in central British Columbia, particularly among stands that had a density high enough to produce a growth release signal. Provided future studies account for interannual weather fluctuations, identification of growth increases due to stand thinning caused by beetle outbreaks will be useful for reconstructing the history of beetle outbreaks over much longer time periods.     

Spatial Distibution of Mountain Pine Beetle Outbreaks in Relation to Climate and Stand Characteristics:A Dendroecological Analysis

Principal components analysis, followed by K-means cluster analysis, was used to detect variations in the timing and magnitude of Pinus contorta Dougl. ex Loud. growth releases attributed to mountain pine beetle outbreaks in 31 stands of central British Columbia. Four major growth release patterns were identified from 1970 to 2000.Variations in the timing of growth releases among clustered stands corresponded well to aerial survey data indicating the timing of beetle outbreaks in the study area. Redundancy analysis was used to determine how variations in the timing and magnitude of growth releases attributed to beetle outbreaks changed with variations in climate or stand conditions over the study area. The first RDA axis, which accounted for 39% of the variations in growth patterns among stands, was significantly (P＜0.05) correlated with gradients in the percentage of pine in stands killed by mountain pine beetle, summer aridity, variation in summer precipitation, distance from initial infestation site, average pine age, and maximum August temperatures. The second RDA axis explained 6% of the variations and was significantly correlated with gradients in the beetle climate suitability index, extreme cold month temperatures, and site index. Comparisons of growth release patterns with aerial survey data and redundancy analyses indicated that dendrochronological techniques are useful for identifying mountain pine beetle outbreaks in central British Columbia, particularly among stands that had a density high enough to produce a growth release signal. Provided future studies account for interannual weather fluctuations, identification of growth increases due to stand thinning caused by beetle outbreaks will be useful for reconstructing the history of beetle outbreaks over much longer time periods.     

Fine Structure of Antennal Sensilla of Paysandisia archon and Electrophysiological Responses to Volatile Compounds Associated with Host Palms

Paysandisia archon (Lepidoptera: Castniidae) is a serious pest of palm trees. A comprehensive knowledge of the insect olfactory system is essential for the development of efficient semiochemical-based control methods. The olfactory sensilla are located particularly on the antennae, and these can detect plant volatiles that provide important cues for the insects in the search for their host plants. To date, the fine structure of P. archon antennal sensilla studies and their role in host-plant perception have not been investigated in great detail. Using light microscopy and scanning and transmission electron microscopy, the antennae of both sexes of P. archon are described here in detail, according to the different types, quantities and distributions of the sensilla. Six types of sensilla were identified. The most widespread are sensilla trichoidea, sensilla basiconica and sensilla auricilica, which are associated with olfactory function. These have cuticular shafts characterised by numerous pores, and they are innervated by two or three sensory neurons. Sensilla coeloconica, sensilla chaetica and sensilla ampullacea are associated with olfactory or olfactory-thermoreception, mechano-gustatory, and thermo-hygroreception functions, respectively. Moreover, the role of P. archon antennae in locating of the host palms was evaluated using electroantennograms, to monitor responses to ester and terpene compounds previously identified as volatiles of damaged/fermenting palm tissues. P. archon showed responses to all of the synthetic chemicals tested, with greater responses in the females, providing a significant sex*dose effect. Among the compounds tested, ethyl isobutyrate elicited the strongest antenna responses. The fine structure of the cuticular and cellular components of the P. archon antenna sensory equipment is described for the first time. The results of this study form an important starting point and complement physiological and behavioural studies, to provide valuable information of practical importance for the development of efficient semiochemical-based control methods.     

Use of precise spatial data for describing spatial patterns and plant interactions in a diverse Great Basin shrub community

Community-structuring processes continue to be of great interest to plant ecologists, and plant spatial patterns have been linked to processes including disturbance, dispersal, environmental heterogeneity, and plant interactions. Under the assumption that the analysis of the spatial structure of plant communities can help to elucidate the type and importance of the predominant community-structuring processes, many studies have analyzed point pattern data on various plant species. A variety of methods have been devised to acquire point pattern data for individual plants, however, the classic tradeoff between the speed of acquisition and the precision of spatial data has meant that large and precise datasets on plant locations are difficult to obtain. The primary goal of this study was to develop a GPS-based methodology for the rapid collection of precise spatial data on plant locations in a semi-arid shrubland in the Great Basin, USA. The secondary goal was to demonstrate a potential application of this approach by using recently developed univariate and bivariate spatial statistics to test for aggregation within the shrub community, as observed in other semi-arid shrublands. We efficiently mapped 2,358 individuals of five shrub species with a spatial error of ≤0.02 m, and found strong statistical evidence of fine-scale aggregation (1) independent of species, (2) within species, and (3) between two species pairs. Our approach is useful for rapidly collecting precise point pattern data in plant communities, and has other applications related to population modeling, GIS analysis, and conservation.     

Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.