Comparison of invertebrate herbivores on native and non‐native Senecio species: Implications for the enemy release hypothesis

The enemy release hypothesis posits that non‐native plant species may gain a competitive advantage over their native counterparts because they are liberated from co‐evolved natural enemies from their native area. The phylogenetic relationship between a non‐native plant and the native community may be important for understanding the success of some non‐native plants, because host switching by insect herbivores is more likely to occur between closely related species. We tested the enemy release hypothesis by comparing leaf damage and herbivorous insect assemblages on the invasive species Senecio madagascariensis Poir. to that on nine congeneric species, of which five are native to the study area, and four are non‐native but considered non‐invasive. Non‐native species had less leaf damage than natives overall, but we found no significant differences in the abundance, richness and Shannon diversity of herbivores between native and non‐native Senecio L. species. The herbivore assemblage and percentage abundance of herbivore guilds differed among all Senecio species, but patterns were not related to whether the species was native or not. Species‐level differences indicate that S. madagascariensis may have a greater proportion of generalist insect damage (represented by phytophagous leaf chewers) than the other Senecio species. Within a plant genus, escape from natural enemies may not be a sufficient explanation for why some non‐native species become more invasive than others.     

Effect of generalist insect herbivores on introduced Lepidium draba (Brassicaceae): implications for the enemy release hypothesis

The enemy release hypothesis (ERH) states that decreased regulation by natural enemies allows plants to increase in distribution, abundance and vigour following their introduction into an exotic range. Invasive plants rarely escape herbivory entirely, and for hoary cress [ Lepidium draba L. (Brassicaceae)] it has been demonstrated that generalist insect abundance is greater in its introduced North American range than in the native European range. We assessed the role of increased generalist herbivory on hoary cress using representatives of four important herbivore niches commonly found in the introduced range. We experimentally examined the density dependent impact of these herbivores individually and in combination on hoary cress in a series of greenhouse experiments. We found that defoliation of the oligophagous diamondback moth Plutella xylostella (L.) (Lep., Plutellidae) had the strongest and most consistent impact, while damage by the stem-mining weevil Ceutorhynchus americanus Buchanan (Col., Curculionidae) tended to have the highest per capita effect. Plant response to feeding by the oligophagous crucifer flea beetle Phyllotreta cruciferae (Goeze) (Col., Chrysomelidae) was minor despite obvious feeding damage, and the impact of the polyphagous tarnished plant bug Lygus hesperus Knight (Het., Miridae) was negligible. In multiple-species experiments, herbivore impacts were usually additive. In general, we found that hoary cress can tolerate high densities of oligophagous insect herbivory and effectively resisted attack by the polyphagous L. hesperus, but also the oligophagous C. americanus . Our results indicate that a combination of plant resistance and tolerance allows hoary cress to withstand increased generalist herbivore load in its introduced range, consistent with the predictions of the ERH.     

Larval parasitism by native insects on the introduced stem-galling moth Epiblema strenuana Walker (Lepidoptera: Tortricidae) and its implications for biological control of Parthenium hysterophorus (Asteraceae)

Abstract  The significance of natural enemies in the failure or reduced effectiveness of weed biological control agents remains largely unknown. Larval and pupal parasitism by native insects in the stem-galling moth Epiblema strenuana was studied in north Queensland, 4 (1986–87) and 16 (1998–99) years after its introduction as a biological-control agent for Parthenium hysterophorus . Specific questions answered in this study are: (i) Has parasitism increased over the years?; (ii) Are the levels of parasitism influenced by the host plant phenology?; and (iii) What are the potential implications of current parasitism levels for the effectiveness of E. strenuana ? There was no increase in the parasitoid species assemblage over the years. In 1986–87, Antrocephalus sp. and Bracon sp. were the most predominant species. The combined larval and pupal parasitism was low (2.6%) and there was no seasonal difference in the parasitism levels. In 1998–99 Bracon sp. was the most prevalent species. Parasitism by Bracon sp. in 1998–99 was several times higher (22.9%) than the combined larval and pupal parasitism in 1986–87. However, parasitism by Bracon sp. remained low (6.5–8.2%) at the beginning of the parthenium-growing season, and reached peak levels (49–53%) only at the end of the season. Parasitism by Bracon sp. was also significantly lower in rosettes (7.2%) than in flowering plants (22.8%). Galling can have a negative impact on the weed only when initiated at the rosette stage. In 1998–99 lower levels of parasitism early in the season when most plants are in the rosette stage suggest that the impact of parasitism on the effectiveness of the gall insect may not be significant.     

The snail Smaragdia bryanae (Neritopsina,Neritidae) is a specialist herbivore of the seagrass Halophila hawaiiana (Alismatidae,Hydrocharitaceae)

Abstract. The endemic Hawaiian gastropod Smaragdia bryanae is a specialized marine herbivore that uses the endemic seagrass Halophila hawaiiana as both food and habitat. These small neritids, their grazing scars, and their egg capsules are found year‐round on seagrass leaves, where they feed on protoplast contents released as the sharp outer‐lateral teeth of the snail's radula puncture leaf epidermal cells; the contents of these cells are likely swept into the mouth by the long, wispy cusps of the marginal teeth. Structural differences from the typical neritid radula include elongated outer‐lateral teeth with two sharply pointed cusps, delicate marginal teeth reduced in both size and number, and a compressed central section. Snails grazed on leaves of H. hawaiiana steadily in laboratory culture, and grew and reproduced on this diet. In laboratory choice experiments, snails did not graze the thalli of any of six macroalgal species growing near seagrass where snails were collected, and strongly preferred occupying seagrass. Seagrass samples from five field sites on Oahu and one on Maui showed from 30% to 94% of leaves damaged, with 11% of the total leaf standing area grazed. Snails are smaller (mean length 2.74±0.32 mm, mean width 2.15±0.17 mm, n=217) than the width of the leaves of H. hawaiiana (mean 3.24±1.26 mm, n=790). The snails associate constantly with their host, despite the scattered distribution, small patch size, and variability of the seagrass resource, demonstrated by a sevenfold range in the leaf area index (mean 1.11±0.61 cm² blade surface cm^?2, n=31) among samples. Damage on grazed leaves (mean 8.21±7.05 mm² per leaf, or 16.5% of leaf surface, n=511) is concentrated in the apical and central epithelia between the midrib and the marginal veins, where snails may access cells with thinner walls and few fibers. Details of the grazing interaction between these extant species in Hawai'i shed light on the ecological specialization of members of the genus Smaragdia to seagrasses over geological time.     

Complete development on Elodea canadensis (Hydrocharitaceae) eliminates Hydrellia egeriae (Diptera,Ephydridae) as a candidate biological control agent of Egeria densa (Hydrocharitaceae) in the U.S.A.

The leaf-mining fly Hydrellia egeriae Rodrigues-Junior (Diptera: Ephydridae) was evaluated under quarantine conditions as a candidate biological control agent of the invasive aquatic weed Egeria densa Planchon (Hydrocharitaceae) in California, U.S.A. The objective of this study was to test the fly’s feeding preference for E. densa as compared to the North American native Elodea canadensis Michaux (Hydrocharitaceae), an important benchmark species. No choice testing revealed that H. egeriae can complete development on E. canadensis and choice testing indicated the fly presents little oviposition discrimination between E. densa and E. canadensis. It is concluded that the host range of H. egeriae is too broad for use as a biological control agent of E. densa in the U.S.A.     

Larval survival,host plant preferences and developmental responses of the diamondback moth Plutella xylostella (Lepidoptera: Plutellidae) on wild brassicaceous species

The diamondback moth Plutella xylostella (L.) (Lepidoptera: Plutellidae) is an important pest of cultivated brassicaceous crops worldwide. The host plant preferences, developmental biology and survival and longevity of P. xylostella are relatively well understood on commercial crop species; however, its relationship with brassicaceous weeds is poorly known. Sinapis arvensis L., Erysimum cheiranthoides L. and Capsella bursa‐pastoris (L.) Medicus are among the most common brassicaceous weeds worldwide and can serve as important bridge hosts of P. xylostella. In this study, preference and performance of P. xylostella were compared on these weed species. In free‐choice situations, females deposited 5.5 and 18.8 times more eggs on S. arvensis than on E. cheiranthoides and C. bursa‐pastoris, respectively. Survival from neonate to pupa and from pupa to adult was highest on S. arvensis and E. cheiranthoides and lowest on C. bursa‐pastoris. Development was fastest, foliage consumption was greatest, pupae and silk cocoons were heaviest, adult body masses and longevities were highest and forewings were largest for both females and males when reared as larvae on S. arvensis. Realized fecundity of new generation adults was highest for individuals reared on S. arvensis compared to those reared on E. cheiranthoides or C. bursa‐pastoris. Relative growth rates of pupae and adults were highest on S. arvensis, suggesting that this plant species is a high‐quality host for P. xylostella compared with other species tested. Potential impacts of these wild brassicaceous species on P. xylostella populations are discussed.     

Infiltration of a facultative ant–plant mutualism by the introduced Argentine ant: effects on mutualist diversity and mutualism benefits

1. Ant–plant mutualisms have been the focus of considerable empirical research, but few studies have investigated how introduced ants affect these interactions. Using 2 years of survey data, this study examines how the introduced Argentine ant [Linepithema humile (Mayr)] differs from native ants with respect to its ability to protect the extrafloral nectary‐bearing coast barrel cactus (Ferocactus viridescens) in Southern California. 2. Eighteen native ant species visited cacti in uninvaded areas, but cacti in invaded areas were primarily visited by the Argentine ant. The main herbivore of the coast barrel cactus present at the study sites is a leaf‐footed bug (Narnia wilsoni). 3. Herbivore presence (the fraction of surveys in which leaf‐footed bugs were present on individual cacti) was negatively related to ant presence (the fraction of surveys in which ants were present on individual cacti). Compared with cacti in uninvaded areas, those in invaded areas were less likely to have herbivores and when they did had them less often. 4. Seed mass was negatively related to herbivore presence, and this relationship did not differ for cacti in invaded areas versus those in uninvaded areas. 5. Although the Argentine ant might provide superior protection from herbivores, invasion‐induced reductions in ant mutualist diversity could potentially compromise plant reproduction. The cumulative number of ant species on individual cacti over time was lower in invaded areas and was associated with a shortened seasonal duration of ant protection and reduced seed mass. These results support the hypothesis that multiple partners may enhance mutualism benefits.     

Larval performance in relation to oviposition site preference in olive kernel moth (Prays oleae Bern., Yponomeutidae,Praydina)

1 The tri‐voltine moth Prays oleae Bern. spends its larval stages on the native olive tree (Olea europaea L. var. sylvestris Brot. and five cultivars, Oleaceae) mining the leaves, the flowers and the fruits in each generation; it seldom switches to other native or introduced confamilial plant species. 2 In this study the pattern of oviposition of the olive moth was examined in olive fields and natural vegetation, in relation to in situ recruitment as an outcome of processes such as density dependence or risk spreading. 3 Larval body size (width of epicranial sclerites) was also examined and compared between host substrates, years and morphological, physiological, ecological and nutritional attributes of the host. 4 The factors influencing the oviposition pattern of the olive moth such as the carbon/nitrogen ratio, number of flowers, branch length and previous leaf damage were ranked differently in different cultivars. 5 ‘Hot spots’, i.e. olive trees or parts of trees receiving a high egg load, were found to be the result of in situ recruitment. 6 Physiological mortality among first instar larvae was significantly negatively correlated with the number of oviposited upon leaves; suggesting that the adult selects for oviposition the best available substrate. 7 As adult moths selected leaves with minimal probability of abscission for oviposition, leaf abscission was not a major mortality factor, although first instar larvae reduced leaf longevity. 8 Host quality did not affect all larval stages in the same way. 9 The more nutritionally poor the substrate, the longer the duration of the larval stage feeding on it. The phenological timing of the insect life stages very closely tracks the phenological phases of its host plant, primarily focusing on the most nutritious host stage in terms of larval performance.     

The population genetics of sporophytic self-incompatibility in Senecio squalidus L. (Asteraceae): the number, frequency, and dominance interactions of S alleles across its British range

Sporophytic self-incompatibility (SSI) was studied in 11 British Senecio squalidus populations to quantify mating system variation and determine how its recent colonization of the United Kingdom has influenced its mating behavior. S allele number, frequency, and dominance interactions in populations were assessed using full diallels of controlled pollinations. A mean of 5.1 S alleles per population was observed, and no population contained more than six S alleles. Numbers of S alleles within populations of S. squalidus declined with increasing distance from the center of its introduction (Oxford). Cross-classification of S alleles allowed an estimate of approximately seven and no more than 11 S alleles for the entire British S. squalidus population. The low number of S alleles observed in British S. squalidus compared to other SI species is consistent with the population bottleneck associated with S. squalidus' introduction to the Oxford Botanic Garden and subsequent colonization of Britain. Extensive S allele dominance interactions were observed to be a feature of the S. squalidus SSI system and may represent an adaptive response to improve limited mate availability imposed by the presence of so few S alleles. Multilocus allozyme genotypes were also identified for individuals in all populations and geographic patterns of S locus and allozyme loci variation investigated. Less interpopulation structure was observed for the S locus than for allozyme diversity--a finding indicative of the effects of negative frequency-dependent selection at the S locus maintaining equal S phenotypes within populations and enhancing effective migration between populations.     

High floral resource density leads to neural constraint in the generalist,floriphilic katydid,Phaneroptera brevis (Orthoptera: Phaneropterinae)

1. The neural constraint hypothesis has been investigated in the field of insect–plant interactions. It predicts that the consumer's detection ability, time spent on resources, and efficiency in resource use are affected by different aspects of resource availability. 2. The applicability of this hypothesis in florivory, however, generally lacks a mechanistic understanding of the effect of resource density on neural constraint, so using caged experiments, the (i) ability of a generalist, floriphilic katydid [(Phaneroptera brevis (Serville)] to find a floral resource (inflorescences of Bidens pilosa L.), (ii) attentiveness of the katydid on this resource after successfully detecting the floral resource, and (iii) efficiency of resource use under varying densities of the floral resource were studied. 3. The present results indicated no evidence of an obvious effect of floral resource density on the detection of resources by the florivore but an increase in floral‐resource density generally led to lower efficiency and attentiveness in the foraging katydids. 4. These findings provide the first evidence of the neural constraint hypothesis in the context of resource density and corroborate the applicability of the neural constraint hypothesis in florivory.     

The forgotten prey of an iconic predator: a review of interactions between grey wolves Canis lupus and beavers Castor spp.

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Demonstration of pollinator‐mediated competition between two native Impatiens species,Impatiens noli‐tangere and I. textori (Balsaminaceae)

Plant–plant interspecific competition via pollinators occurs when the flowering seasons of two or more plant species overlap and the pollinator fauna is shared. Negative sexual interactions between species (reproductive interference) through improper heterospecific pollen transfer have recently been reported between native and invasive species demonstrating pollination‐driven competition. We focused on two native Impatiens species (I. noli‐tangere and I. textori) found in Japan and examined whether pollinator‐mediated plant competition occurs between them. We demonstrate that I. noli‐tangere and I. textori share the same pollination niche (i.e., flowering season, pollinator fauna, and position of pollen on the pollinator's body). In addition, heterospecific pollen grains were deposited on most stigmas of both I. noli‐tangere and I. textori flowers that were situated within 2 m of flowers of the other species resulting in depressed fruit set. Further, by hand‐pollination experiments, we show that when as few as 10% of the pollen grains are heterospecific, fruit set is decreased to less than half in both species. These results show that intensive pollinator‐mediated competition occurs between I. noli‐tangere and I. textori. This study suggests that intensive pollinator‐mediated competition occurs in the wild even when interacting species are both native and not invasive.     

Eight decades of invasion by Chromolaena odorata (Asteraceae) and its biological control in West Africa: the story so far

ABSTRACT

Chromolaena odorata (L.) R.M. King and H. Robinson (Asteraceae) is a perennial weedy shrub of neotropical origin and a serious biotic threat in its invasive range. The Asian-West Africa (AWA) biotype of C. odorata present in West Africa is both morphologically and genetically different from the southern African (SA) biotype. The AWA biotype was first introduced into Nigeria in the late 1930s and rapidly spread across West Africa. Currently, 12 of the 16 countries in West Africa have been invaded, with significant negative effects on indigenous flora and fauna. However, locals in West Africa have found several uses for the weed. As chemical, physical and other conventional methods were unsustainable, costly and largely ineffective, three biological control agents, Apion brunneonigrum (Coleoptera: Brentidae), Pareuchaetes pseudoinsulata (Lepidoptera: Erebidae) and Cecidochares connexa (Diptera: Tephritidae), have been released in West Africa between the 1970s and the early 2000s. However, only C. connexa and P. pseudoinsulata established, contributing to the control of the weed, in six and four countries in West Africa respectively. Limited research funding, the absence of post-release evaluations of the established agents, and the ‘conflict of interest’ status of C. odorata (i.e. being beneficial for local use but damaging to ecosystem services and agriculture), are serious factors deterring the overall biological control effort. Here, using historical records and field surveys, we examine the invasion history, spread, impacts, and management of C. odorata in West Africa and make recommendations for the sustainable management of C. odorata in the region.     

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.