Isolation of 55 microsatellite markers for Jatropha curcas and its closely related species

Jatropha curcas L. (physic nut) is native to Central America and now naturalized widely in many tropical and subtropical areas. Microsatellite markers were isolated and characterized. Eleven out of 55 markers showed polymorphisms, and the allelic variation was investigated using 26 accessions of J. curcas collected from several provinces in Thailand. Each marker showed 2 to 5 alleles and the average polymorphic information content (PIC) was 0.49. Thirty four markers (62 %) were also successfully amplified in J. integerrima, J. gossypifolia and J. podagrica.     

Integrated genome sequence and linkage map of physic nut (Jatropha curcas L.), a biodiesel plant

The family Euphorbiaceae includes some of the most efficient biomass accumulators. Whole genome sequencing and the development of genetic maps of these species are important components in molecular breeding and genetic improvement. Here we report the draft genome of physic nut (Jatropha curcas L.), a biodiesel plant. The assembled genome has a total length of 320.5 Mbp and contains 27 172 putative protein‐coding genes. We established a linkage map containing 1208 markers and anchored the genome assembly (81.7%) to this map to produce 11 pseudochromosomes. After gene family clustering, 15 268 families were identified, of which 13 887 existed in the castor bean genome. Analysis of the genome highlighted specific expansion and contraction of a number of gene families during the evolution of this species, including the ribosome‐inactivating proteins and oil biosynthesis pathway enzymes. The genomic sequence and linkage map provide a valuable resource not only for fundamental and applied research on physic nut but also for evolutionary and comparative genomics analysis, particularly in the Euphorbiaceae.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Interplant variation in creosotebush foliage characteristics and canopy arthropods

Summary We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.     

Evidence of biotic resistance to exotic plant invasion in degraded Bornean forests

Intact tropical forests are generally considered to be resistant to invasions by exotic species, although the shrub Clidemia hirta (Melastomataceae) is highly invasive in tropical forests outside its native range. Release from natural enemies (e.g., herbivores and pathogens) contributes to C. hirta invasion success where native melastomes are absent, and here we examine the role of enemies when C. hirta co-occurs with native Melastomataceae species and associated herbivores and pathogens. We study 21 forest sites within agricultural landscapes in Sabah, Malaysian Borneo, recording herbivory rates in C. hirta and related native Melastoma spp. plants along two 100-m transects per site that varied in canopy cover. Overall, we found evidence of enemy release; C. hirta had significantly lower herbivory (median occurrence of herbivory per plant = 79% of leaves per plant; median intensity of herbivory per leaf = 6% of leaf area) than native melastomes (93% and 20%, respectively). Herbivory on C. hirta increased when closer to native Melastoma plants with high herbivory damage, and in more shaded locations, and was associated with fewer reproductive organs on C. hirta. This suggests host-sharing by specialist Melastomataceae herbivores is occurring and may explain why invasion success of C. hirta is lower on Borneo than at locations without related native species present. Thus, natural enemy populations may provide a “biological control service” to suppress invasions of exotic species (i.e., biotic resistance). However, lower herbivory pressures in more open canopy locations may make highly degraded forests within these landscapes more susceptible to invasion.     

Analysis of the genetic diversity of physic nut, <Emphasis Type="Italic">Jatropha curcas</Emphasis> L. accessions using RAPD markers

A sum of 48 accessions of physic nut, Jatropha curcas L. were analyzed to determine the genetic diversity and association between geographical origin using RAPD-PCR markers. Eight primers generated a total of 92 fragments with an average of 11.5 amplicons per primer. Polymorphism percentages of J. curcas accessions for Selangor, Kelantan, and Terengganu states were 80.4, 50.0, and 58.7%, respectively, with an average of 63.04%. Jaccard’s genetic similarity co-efficient indicated the high level of genetic variation among the accessions which ranged between 0.06 and 0.81. According to UPGMA dendrogram, 48 J. curcas accessions were grouped into four major clusters at coefficient level 0.3 and accessions from same and near states or regions were found to be grouped together according to their geographical origin. Coefficient of genetic differentiation (G_st) value of J. curcas revealed that it is an outcrossing species.     

Biogeographical comparison of the arthropod herbivore communities associated with Lepidium draba in its native, expanded and introduced ranges

Aim To examine the composition and structure of the arthropod community on the invasive weed Lepidium draba in its native, expanded and introduced ranges, in order to elucidate the lack of a biotic constraint that may facilitate invasion. Location Europe and western North America. Methods Identical sampling protocols were used to collect data from a total of 35 populations of L. draba in its native (Eastern European), expanded (Western European) and introduced (western US) ranges. A bootstrapping analysis was used to compare herbivore richness, diversity and evenness among the regions. Core species groups (monophages, oligophages and polyphages) on the plant were defined and their abundances and host utilization patterns described. Results Species richness was greatest in the native range, while species diversity and evenness were similar in the native and expanded range, but significantly greater than in the introduced range of L. draba. Specialist herbivore abundance was greater in the native and expanded compared with the introduced range. Oligophagous Brassicaceae‐feeders were equally abundant in all three ranges, and polyphagous herbivore abundance was significantly greater in the introduced range. Overall herbivore abundance was greater in the introduced range. Host utilization was more complete in the two European ranges due to monophagous herbivores that do not exist in the introduced range. Root feeders and gall formers were completely absent from the introduced range, which was dominated by generalist sap‐sucking herbivores. However, one indigenous stem‐mining weevil, Ceutorhynchus americanus, occurred on L. draba in the introduced range. Main conclusions This is, to our knowledge, the first study documenting greater herbivore abundance on an invasive weed in its introduced, compared with its native, range. However, greater abundance does not necessarily translate to greater impact. We argue that, despite the greater total herbivore abundance in the introduced range, differences in the herbivore community structure (specialist vs. generalist herbivory) may contribute to the invasion success of L. draba in the western USA.     

Decreased resistance and increased tolerance to native herbivores of the invasive plant Sapium sebiferum

Release from natural enemies may favor invasive plants evolving traits associated with reduced herbivore‐resistance and faster‐growth in introduced ranges. Given a genetic trade‐off between resistance and tolerance, invasive plants could also become more tolerant to herbivory than conspecifics in the native range. We conducted a field common garden study in the native range of Sapium sebiferum using seeds from native Chinese populations and invasive North American populations to compare their growth and herbivory resistance. We also performed a cage‐pot experiment to compare their resistance and tolerance to Bikasha collaris beetles that are specialist feeders on S. sebiferum trees in China. Results of the common garden study showed that Sapium seedlings of invasive populations relative to native populations were more frequently attacked by native herbivores. Growth and leaf damage were significantly higher for invasive populations than for native populations. Growth of invasive populations was not significantly affected by insecticide spray, but insecticide spray benefited that of native populations. In the bioassay trial, beetles preferentially consumed leaf tissue of invasive populations compared to native populations when beetles had a choice between them. Regression of percent leaf damage on biomass showed that invasive populations tolerated herbivory more effectively than native populations. Our results suggest that S. sebiferum from the introduced range had lower resistance but higher tolerance to specialist herbivores. Both defense strategies could have evolved as a response to the escape from natural enemies in the introduced range.     

Potential impacts of climate change on patterns of insect herbivory on understorey plant species: A transplant experiment

Species‐specific responses to climate change will lead to changes in species interactions across multiple trophic levels. Interactions between plants and their insect herbivores, in particular, may become increasingly disrupted if mobile herbivores respond more rapidly to climatic change than their associated host plants. We present a multispecies transplant experiment aimed at assessing potential climatic impacts on patterns of leaf herbivory. Four shrubby understorey plant species were transplanted outside their native range into a climate 2.5°C warmer in annual mean temperature. After 12 months, we assessed the types and amount of herbivore leaf damage, compared with plants transplanted to a control site within their native range. The overall amount of foliage loss to herbivores ranged from approximately 3–10% across species and sites, a range consistent with most estimates of leaf loss in other studies. The most common types of leaf damage were sucking and chewing and this pattern was consistent for all four plant species at all sites. There were no significant differences in levels and patterns of herbivory between control and warm sites for three out of four plant species. This suggests that with moderate climate warming, most herbivory will continue to be dominated by chewers and suckers, and that the overall level of foliage loss will be similar to that experienced presently.     

Efficacy of karanjin and phorbol ester fraction against termites (Odontotermes obesus)

The non-edible oil seeds of Jatropha curcas (physic nut) and Pongamia pinnata (karanja) contain some toxic components (phorbol esters in J. curcas and karanjin in P. pinnata), which may be used as biopesticides. In this study, the active components of J. curcas and P. pinnata oil were extracted and their efficacy against the termites Odontotermes obesus (Rambur), was tested. The phorbol ester fraction of J. curcas and karanjin of P. pinnata oil were found to be effective against termites. A mortality rate of 100% was achieved in 6 h with karanjin and in 12 h with phorbol ester fraction. The LC₅₀ levels of karanjin and phorbol esters fractions were 0.038 and 0.071 g ml⁻¹, respectively, after 24 h at a 95% (0.05) confidence limit.     

Native lagomorphs suppress grass establishment in a shrub‐encroached,semiarid grassland

Shrub encroachment into arid grasslands has been associated with reduced grass abundance, increased soil erosion, and local declines in biodiversity. Livestock overgrazing and the associated reduction of fine fuels has been a primary driver of shrub encroachment in the southwestern United States, but shrublands continue to persist despite livestock removal and grassland restoration efforts. We hypothesized that an herbivory feedback from native mammals may contribute to continued suppression of grasses after the removal of livestock. Our herbivore exclusion experiment in southeastern Arizona included five treatment levels and allowed access to native mammals based on their relative body size, separating the effects of rodents, lagomorphs, and mule deer. We included two control treatments and replicated each treatment 10 times (n = 50). We introduced uniform divisions of lawn sod (Cynodon dactylon) into each exclosure for 24‐hr periods prior to (n = 2) and following (n = 2) the monsoon rains and used motion‐activated cameras to document herbivore visitations. In the pre‐monsoon trials, treatments that allowed lagomorph access had less sod biomass relative to other treatments (p < 0.001), averaging 44% (SD 36%) and 29% (SD 45%) remaining biomass after the 24‐hr trial periods. Following the onset of monsoons, differences in remaining biomass among treatments disappeared. Desert cottontails (Sylvilagus audubonii) were detected more frequently than any of the other 11 herbivore species present at the site, accounting for 83% of detections during the pre‐monsoon trials. Significantly more (p < 0.001) desert cottontails were detected during the pre‐monsoon trials (2,077) compared to the post‐monsoon trials (174), which coincided with biomass removal from lagomorph accessible treatments. We conclude that desert cottontails are significant consumers of herbaceous vegetation in shrub‐encroached arid grasslands and they, along with other native herbivores, may act as a biotic feedback contributing to the competitive advantage and persistence of shrubs.     

Test of the enemy release hypothesis: The native magpie moth prefers a native fireweed (Senecio pinnatifolius) to its introduced congener (S. madagascariensis)

Abstract The enemy release hypothesis predicts that native herbivores will either prefer or cause more damage to native than introduced plant species. We tested this using preference and performance experiments in the laboratory and surveys of leaf damage caused by the magpie moth Nyctemera amica on a co‐occuring native and introduced species of fireweed (Senecio) in eastern Australia. In the laboratory, ovipositing females and feeding larvae preferred the native S. pinnatifolius over the introduced S. madagascariensis. Larvae performed equally well on foliage of S. pinnatifolius and S. madagascariensis: pupal weights did not differ between insects reared on the two species, but growth rates were significantly faster on S. pinnatifolius. In the field, foliage damage was significantly greater on native S. pinnatifolius than introduced S. madagascariensis. These results support the enemy release hypothesis, and suggest that the failure of native consumers to switch to introduced species contributes to their invasive success. Both plant species experienced reduced, rather than increased, levels of herbivory when growing in mixed populations, as opposed to pure stands in the field; thus, there was no evidence that apparent competition occurred.     

Abundance and stratification of foliage arthropods in a lowland rain forest of Cameroon

Abstract.

• 1 Arthropod densities and apparent leaf damage were compared within the canopy ecotone and the shrub layer of a lowland rain forest in Cameroon, using a branch clipping method.
• 2 Most of the individuals collected consisted of ants (average 44%) and various herbivores (31%). Overall arthropod densities amounted to 17 individuals per sample, which, on average, consisted of 0.85 m² of foliage area. Arthropod densities were lower than on temperate foliage.
• 3 Arthropod densities were about 3 times higher in the canopy than within the shrub layer. In particular, ants and herbivores were significantly more abundant in the canopy than within the shrub layer. Usually, layer effects rather than site effects appeared to cause greater variance in arthropod abundance.
• 4 Arthropod species-richness, as estimated by the number of operational taxonomic units sorted, was higher in canopy samples than in samples obtained from the shrub layer. However, apparent leaf damage was higher within the shrub layer (10.9%) than on the canopy (5.2%).
• 5 Possible factors responsible for the high densities of ants and herbivores on the canopy and for the high leaf damage within the shrub layer are discussed.

     

Effects of bird predation on arthropod abundance and tree growth across an elevational gradient

Considerable uncertainty surrounds the conditions under which birds can cause trophic cascades. In a three‐year experiment, we studied the direct and indirect effects of insectivorous birds on arthropod abundance, herbivory, and growth of striped maple Acer pensylvanicum saplings in a northern hardwood forest of central New Hampshire, USA. We manipulated bird predation by erecting exclosures around saplings and directly manipulated herbivory by removing herbivores. We also examined how climate modifies these interactions by replicating the experiment at three locations along an elevational gradient. Effects of bird predation were variable. Overall, mean arthropod biomass was 20% greater on saplings within bird exclosures than on controls (p<0.05). The mean biomass of leaf‐chewing herbivores, primarily Lepidoptera larvae, was 25% greater within exclosures but not statistically different from controls. To a lesser degree, mean herbivore damage to foliage within exclosures exceeded that of controls but differences were not significant. We also did not detect significant treatment effects on sapling shoot growth. The high understory vegetation density relative to bird abundance, and low rate of herbivory during the study (mean 5% leaf area removed, controls), may have limited the ability of birds to affect sapling growth. Climate effects operated at multiple scales, resulting in a complex interplay of interactions within the food web. Regional synchrony of climatic conditions resulted in annual fluctuations in herbivore abundance and tree growth that were shared across elevations. At the same time, local environmental variation resulted in site differences in the plant, herbivore, and bird communities. These patterns resulted in a mosaic of top–down strengths across time and space, suggesting an overall pattern of limited effects of birds on plant growth, possibly interspersed with hotspots of trophic cascades.     

Damage levels from arthropod herbivores on <Emphasis Type="Italic">Lonicera maackii</Emphasis> suggest enemy release in its introduced range

The ‘enemy release hypothesis’ argues when a species is introduced to a novel habitat, release from regulation by herbivores results in increased vigor, abundance, and distribution. The invasive Asian shrub Lonicera maackii appears to benefit from an absence of arthropod herbivores in North America. We assessed the incidence, amount, and type of herbivory occurring on L. maackii in forest edge and interior habitats and investigated differences in timing of damage. In October 2008, leaves were sampled from shrubs in forest interior and edge habitat from 8 sites in Ohio. In 2009, sampling was repeated at 3 sites in spring, summer, and fall with a distinction made between long and short branches. Leaf area removed averaged 1.83% across the 8 populations in 2008 and 3.09% across the 3 populations in 2009, with forest edge plants receiving slightly more damage than forest interior plants in 2008. Additionally, long shoots received more damage than short shoots in 2009. Damage incidence was also higher in the edge habitat and on long shoots compared to short shoots. As measured in 2009, damage accumulated steadily throughout the season. Chewing was the most prevalent type of damage (76. 8%) and low level of pathogen infection was observed (4.81%). Results indicate that levels of herbivory experienced by L. maackii are relatively consistent across sites, vary slightly with habitat and branch identity, but are likely too low to impact fitness of shrubs. These findings indicate that low amounts of arthropod herbivory occur for L. maackii across its introduced range, which may contribute to its invasive success.     

Facilitation and herbivory during restoration of California coastal sage scrub

Using nurse plants to facilitate native plant recruitment in degraded habitats is a common restoration practice across various arid and semiarid environments. Living trees or shrubs are typically considered nurse plants, whereas dead shrubs left in the landscape from prolonged drought are understudied prospective facilitators for native plant recruitment. The interaction between nurse plants and biotic pressures, such as herbivory, on native recruitment is also not well understood in semiarid plant communities. We investigated the effects of facilitation and herbivory on native seedling germination, growth, and survival in the restoration of degraded coastal sage scrub (CSS) habitat. Native shrub seedlings (Artemisia californica and Salvia mellifera) were planted, and native annual species (Amsinckia intermedia, Deinandra fasciculata, Phacelia distans, and Pseudognaphalium californicum) were sown in three Shrub Type treatments (live shrub, dead shrub, and exposed areas), with a nested Cage treatment (no cage and cage) in each Shrub Type treatment. Annual species grew equally well in all Shrub Type treatments; shrub seedlings grew largest in exposed areas. While there was little evidence of facilitation for all species tested, there were strong positive effects of caging on growth and establishment of all species. Caging palatable native species or planting species with anti‐herbivory traits around target plants may be more strategic approaches compared to using nurse plants in restoring degraded CSS after extended drought.     

North American arthropods at risk due to widespread Fraxinus mortality caused by the Alien Emerald ash borer

Emerald ash borer (Agrilus planipennis Fairmaire) (EAB), an alien invasive wood-boring buprestid beetle, is causing large-scale decline and mortality of the most widely distributed species of ash (Fraxinus spp.) trees endemic to eastern North America. We determined which arthropod species that are associated with ash may become threatened, endangered, and co-extinct with the demise of ash as a dominant tree species. A literature survey revealed that 43 native arthropod species in six taxonomic groups (Arachnida: Acari; Hexapoda: Coleoptera, Diptera, Hemiptera, Hymenoptera, and Lepidoptera) are known to be associated only with ash trees for either feeding or breeding purposes, and thus face high risk of endangerment. Most of these species are gall-formers followed by folivores, subcortical phloem/xylem feeders, sap feeders, and seed predators. Another 30 arthropod species are associated with 1–2 host plants in addition to ash, and herbivory on these hosts may increase as these arthropods shift from declining ash trees. Extirpation of arthropods dependent upon ash may unleash multiple extinctions of affiliated species with which they may be inextricably linked. The demise of North American ash species due to EAB is expected to lead to biotic loss with cascading ecological impacts and altered processes within forested ecosystems.     

Grass-herbivore interactions altered by strains of a native endophyte

Many plants support symbiotic microbes, such as endophytic fungi, that can alter interactions with herbivores. Most endophyte research has focused on agronomically important species, with less known about the ecological roles of native endophytes in native plants. In particular, whether genetic variation among endophyte symbionts affects herbivores of plant hosts remains unresolved for most native endophytes. Here, we investigate the importance of native isolates of the endophyte Epichlo? elymi in affecting herbivory of the native grass host, Elymus hystrix. Experimental fungal isolate-plant genotype combinations and endophyte-free control plants were grown in a common garden and exposed to natural arthropod herbivory. Fungal isolates differed in their effects on two types of herbivory, chewing and scraping. Isolates exhibiting greater sexual reproduction were associated with greater herbivore damage than primarily asexual isolates. Endophyte infection also altered patterns of herbivory within plants, with stroma-bearing tillers experiencing up to 30% greater damage than nonstroma-bearing tillers. Results suggest that intraspecific genetic variation in endophytes, like plant genetic variation, can have important 'bottom-up' effects on herbivores in native systems.     

Restoration of Arthropod Assemblages in a Spartina Salt Marsh following Removal of the Invasive Plant Phragmites australis

Invasive plants are one of the most serious threats to native species assemblages and have been responsible for the degradation of natural habitats worldwide. As a result, removal of invasive species and reestablishment of natural vegetation have been attempted in order to restore biodiversity and ecosystem function. This study examined how native arthropod assemblages, an abundant and functionally important group of organisms in many ecosystems, are affected by the incursion of the invasive wetland plant Phragmites australis and if the restoration of the native vegetation in brackish Spartina alterniflora marshes results in the reestablishment of the arthropod community. The invasion of Phragmites into a coastal Spartina marsh in southern New Jersey seriously altered arthropod assemblages and trophic structure by changing the abundance of trophic groups (detritivores, herbivores, carnivores) and their taxonomic composition. Herbivore assemblages shifted from the dominance of external free‐living specialists (e.g., planthoppers) in Spartina to concealed feeders in Phragmites (stem‐feeding cecidomyiids). Moreover, free‐living arthropods in Phragmites became dominated by detritivores such as Collembola and chironomids. The dominant marsh spiders, web‐building linyphiids, were significantly reduced in Phragmites habitats, likely caused by differences in the physical environment of the invaded habitats (e.g., lower stem densities). Thus, trophic structure of arthropod assemblages in Phragmites, as seen in the large shifts in feeding guilds, was significantly different from that in Spartina. Removal of Phragmites with the herbicide glyphosate resulted in the rapid return of Spartina (≤5 yrs). Moreover, return of the dominant vegetation was accompanied by the recovery of most original habitat characteristics (e.g., live and dead plant biomass, water flow rate). The arthropod assemblage associated with Spartina also quickly returned to its preinvasion state and was not distinguishable from that in uninvaded Spartina reference sites. This study provides evidence that the reestablishment of native vegetation in areas previously altered by an invasive plant can result in the rapid recovery of the native arthropod assemblage associated with the restored habitat.