Herbivory by an introduced Asian weevil negatively affects population growth of an invasive Brazilian shrub in Florida

The enemy release hypothesis (ERH) is often cited to explain why some plants successfully invade natural communities while others do not. This hypothesis maintains that plant populations are regulated by coevolved enemies in their native range but are relieved of this pressure where their enemies have not been co-introduced. Some studies have shown that invasive plants sustain lower levels of herbivore damage when compared to native species, but how damage affects fitness and population dynamics remains unclear. We used a system of co-occurring native and invasive Eugenia congeners in south Florida (USA) to experimentally test the ERH, addressing deficiencies in our understanding of the role of natural enemies in plant invasion at the population level. Insecticide was used to experimentally exclude insect herbivores from invasive Eugenia uniflora and its native co-occurring congeners in the field for two years. Herbivore damage, plant growth, survival, and population growth rates for the three species were then compared for control and insecticide-treated plants. Our results contradict the ERH, indicating that E. uniflora sustains more herbivore damage than its native congeners and that this damage negatively impacts stem height, survival, and population growth. In addition, most damage to E. uniflora, a native of Brazil, is carried out by Myllocerus undatus, a recently introduced weevil from Sri Lanka, and M. undatus attacks a significantly greater proportion of E. uniflora leaves than those of its native congeners. This interaction is particularly interesting because M. undatus and E. uniflora share no coevolutionary history, having arisen on two separate continents and come into contact on a third. Our study is the first to document negative population-level effects for an invasive plant as a result of the introduction of a novel herbivore. Such inhibitory interactions are likely to become more prevalent as suites of previously noninteracting species continue to accumulate and new communities assemble worldwide.     

Prey morphology constrains the feeding ecology of an aquatic generalist predator

Resource availability and accessibility are primary factors guiding the distribution and abundance of organisms. For generalists, prey availability reflects both prey abundance and differences in quality among prey taxa. Although some aspects of prey quality, such as nutritional composition, are well studied, our understanding of how prey morphology contributes to overall prey quality is limited. Because snakes cannot reduce prey size by mastication, many aspects of their feeding ecology (e.g., maximum prey size, feeding performance, and the degree of postprandial locomotor impairment) may be affected by prey shape. We conducted a uniquely comprehensive comparison of prey quality for a generalist species, the banded watersnake (Nerodia fasciata), using prey that were similar in mass and presumably similar in nutritional composition but different in shape and habitat association. Specifically, we compared nutritional composition and shape of paedomorphic salamanders (Ambystoma talpoideum) and sunfish (Lepomis MARGINATUS) and used a series of repeated-measures experiments to examine feeding performance (number of prey consumed, maximum prey size, and intra-oral transport time), digestive metabolism (specific dynamic action, SDA), and postprandial locomotor performance of snakes fed Ambystoma and Lepomis. Cost of digestion was similar between the prey types, likely reflecting their similar nutritional composition. However, snakes consumed larger Ambystoma than Lepomis and intra-oral transport time was much shorter for Ambystoma. Snakes fed Lepomis also suffered greater reduction in crawling speed than those fed Ambystoma. These differences highlight the need for behaviorally integrated approaches to understanding prey quality and support field observations of the importance of amphibian prey for juvenile watersnakes.     

Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Summary. Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450 s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores.     

Soft song in song sparrows: response of males and females to an enigmatic signal

Low-amplitude “soft song” is used by a variety of songbirds; in some species during aggressive encounters, in others during courtship, and yet others in both these contexts. In song sparrows (Melospiza melodia), soft song has thus far been observed only in aggressive encounters, where its production is a more reliable predictor of attack than any other signaling behavior. We used song playback to test the response of both male and female song sparrows to soft song. The design of the playback experiments took into account the existence of two classes of soft song: crystallized soft song, which consists of song types also found in the broadcast repertoire, and warbled soft song, which consists of less-structured song types not found in the broadcast repertoire. Female song sparrows responded with significantly less courtship display to the playback of crystallized soft song than to that of normal broadcast song, and response to warbled soft song was if anything lower than to that of crystallized soft song. Male song sparrows responded equally aggressively to normal broadcast song as to crystallized soft song, and equally aggressively to warbled soft song as to crystallized soft song. The female results support the conclusion that neither form of soft song functions in courtship. The male results suggest that the reliability of soft song as a signal of aggressive intent is not maintained by a receiver retaliation rule.     

Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

How an introduced seaweed can affect epibiota diversity in different coastal systems

Invasions by non-indigenous species have the potential to alter the biodiversity of recipient systems. The magnitude of this effect often depends on the nature of the invaded communities and the ecology of the invader. We investigated the impacts of the Japanese seaweed Sargassum muticum (Phaeophyceae, Fucales) on biodiversity in a rocky and sedimentary environment on two islands in the North Sea. In each case, we compared the epibiota of non-indigenous S. muticum with epibiota communities on taxonomically related and structurally similar native seaweed hosts. Total and average species richness on S. muticum were similar on the rocky shore (60 species and 22±6 species, respectively) and the sandy shore (64 species and 20±3 species, respectively). However, community structure and species composition differed significantly between the environments. On the rocky shore, another native fucoid seaweed, Halidrys siliquosa, supported an epibiota community very similar to that of the invader. On the sandy shore, the only other abundant native habitat-providing algal species was Fucus vesiculosus. This species supported a different and less diverse assemblage of associated taxa. We conclude that S. muticum enhances epibiota diversity in the sedimentary environment, probably by increasing the substratum availability and habitat heterogeneity. In contrast, it has negligible impacts on epibiota diversity in the rocky shore environment, where it does not represent a fundamentally new habitat component. We conclude that even within the same region, the consequences of non-indigenous species on biodiversity cannot be generalised but depend on the composition and structural complexity of the species in the recipient community. This paper is dedicated to Udo Schilling, Chief Diving Officer, Alfred Wegener Institute who trained all of the authors in the waters off Helgoland to be scientific divers. The work reported here and in previous publications has been made possible through Udo’s training and support. We remain grateful for his enthusiasm, effort and patience.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Nonredundancy in the dispersal network of a generalist tropical forest tree

Plant species with generalized dispersal mutualisms are considered to be robust to local frugivore extinctions because of redundancy between dispersal agents. However, real redundancy can only occur if frugivores have similar foraging and ranging patterns and if fruit is a limiting resource. We evaluated the quantitative and qualitative contributions of seed dispersers for an endochorus mast-fruiting species, Prunus javanica (Rosaceae) in Khao Yai National Park, Thailand, to evaluate the potential redundancy of dispersers. Data were collected from tree watches, seed/fruit traps, and seed transects under and away from fruiting trees, feeding and seed deposition by gibbons (Hylobates lar), and evaluations of seed and first-year seedling survival. We identified three clusters of dispersers within the network. Most (>80%) frugivore species observed were small birds and squirrels that were not functional dispersers, dropping most seeds under or very near the tree crown, where seedling survival was ultimately nil. Monkeys (Macaca leonina) were low-quality, short-range dispersers, but they dispersed large numbers of seeds and were responsible for 67% of surviving first-year seedlings. Gibbons and Oriental Pied Hornbills (Anthracoceros albirostris) handled few fruits, but they provided the highest quality service by carrying most seeds away from the canopy to medium and long distances, respectively. Although there was overlap in the deposition patterns of the functional dispersers, they displayed complementary, rather than redundant, roles in seed dispersal. Satiation of all functional dispersers further limited their capacity to "replace" one another. Redundancy must be evaluated at the community level because each type of disperser may shift to different species in the non-masting years of P. javanica. Our results underscore the need for research on broader spatial and temporal scales, which combines studies of dispersal and plant recruitment, to better understand mechanisms that maintain network stability.     

Effects of an invasive ant on land snails in the Ogasawara Islands

We investigated how Pheidole megacephala has affected endemic achatinellid snails because these snails are excellent indicators of the impact of ants and they have high conservation value in Ogasawara. In 2015 we surveyed the Minamizaki area of Hahajima Island of Ogasawara, designated a core zone of the World Heritage Site, for P. megacephala. In Minamizaki, we determined the distribution and density of achatinellid snails in 2015 and compared these data with their distribution and density in 2005. Land cover in the survey area was entirely forest. We also tested whether P. megacephala preyed on achatinellid snails in the laboratory. P. megacephala was present in the forested areas of Minamizaki. Achatinellid snails were absent in 19 of 39 sites where P. megacephala was present, whereas in other areas densities of the snails ranged from 2 to 228 individuals/site. In the laboratory, P. megacephala carried 6 of 7 achatinellid snails and a broken shell was found. Snail distribution and density comparisons and results of the feeding experiments suggest that the presence of P. megacephala has contributed to the decline of achatinellid snails in forests in the survey area. Yet, P. megacephala is not on the official list of invasive non‐native species. Stakeholders using the list of invasive species to develop conservation programs should recognize that invasiveness of non‐native species differs depending on the ecosystem and that official lists may not be complete.     

Predicting the economic impact of an invasive species on an ecosystem service.

Quantifying the impact of alien invasive species on ecosystem services is an essential step in developing effective practices and policy for invasive species management. Here we develop a stochastic bioeconomic model that enables the economic impact of an invasive pest to be estimated before its arrival, based on relatively poorly specified ecological and economic parameters. We developed the model by using a hypothetical invasion of the varroa bee mite (Varroa destructor) into Australia and the negative flow-on effects that it would have on pollination by reducing honey bee populations, giving rise to a loss of pollination services, reduced crop yields, and additional production costs. If the mite were to continue to be prevented from entering the country over the next 30 years, we estimate that the economic costs avoided would be U.S. $16.4-38.8 million (Aus $21.3-50.5 million) per year. We suggest that current invasion response funding arrangements in Australia, which do not acknowledge these avoided damages, require amendment.     

Social context influences cue-mediated recruitment in an invasive social wasp

Social insects often serve as model systems for communication and recruitment studies, and yet, it remains controversial whether social vespid wasps can reliably communicate resource information to nestmates. In this study, I present empirical evidence that foraging strategies depend on the initial assessment of resource size and potential competition by foraging yellowjackets. The context dependent foraging behavior of Vespula pensylvanica provides a potential explanation for the inconsistent reports of the existence of recruitment communication in vespid wasps. Furthermore, life history traits may influence yellowjacket foraging behavior; annual V. pensylvanica colonies, whose foragers routinely patrol near the nest, exhibited increased bait visitation in response to the return of successful foragers, whereas perennial colonies did not. These behavioral disparities provide insight into how foraging strategies and search patterns may shift with colony size and longevity. In experiments that investigate the effects of visual cues of conspecifics and bait dispersion, foraging decisions corresponded with expectations of yellowjackets integrating resource quantity and access into a perception of demand. When resource competition could be assessed as high, V. pensylvanica foragers quickly exploited the bait closest to their colony regardless of occupation by other wasps; however, foragers preferred visiting unoccupied baits in situations where competition could be perceived as low. Moreover, a meta-analysis revealed that context-dependent, cue-mediated recruitment was widespread in Vespidae, where such foraging behaviors changed with habitat and the potential for resource competition. Such plastic foraging strategies may contribute to the invasion success of some vespid wasps.     

Nitrogen inputs promote the spread of an invasive marsh grass.

Excess nutrient loading and large-scale invasion by nonnatives are two of the most pervasive and damaging threats to the biotic and economic integrity of our estuaries. Individually, these are potent forces, but it is important to consider their interactive impacts as well. In this study we investigated the potential limitation of a nonnative intertidal grass, Spartina alterniflora, by nitrogen (N) in estuaries of the western United States. Nitrogen fertilization experiments were conducted in three mud-flat habitats invaded by S. alterniflora in Willapa Bay, Washington, USA, that differed in sediment N. We carried out parallel experiments in San Francisco Bay, California, USA, in three habitats invaded by hybrid Spartina (S. alterniflora x S. foliosa), in previously unvegetated mud flat, and in native S. foliosa or Salicornia virginica marshes. We found similar aboveground biomass and growth rates between habitats and estuaries, but end-of-season belowground biomass was nearly five times greater in San Francisco Bay than in Willapa Bay. In Willapa Bay, aboveground biomass was significantly correlated with sediment N content. Addition of N significantly increased aboveground biomass, stem density, and the rate of spread into uninvaded habitat (as new stems per day) in virtually all habitats in both estuaries. Belowground biomass increased in Willapa Bay only, suggesting that belowground biomass is not N limited in San Francisco Bay due to species differences, N availability, or a latitudinal difference in the response of Spartina to N additions. The relative impact of added N was greater in Willapa Bay, the estuary with lower N inputs from the watershed, than in San Francisco Bay, a highly eutrophic estuary. Nitrogen fertilization also altered the competitive interaction between hybrid Spartina and Salicornia virginica in San Francisco Bay by increasing the density and biomass of the invader and decreasing the density of the native. There was no significant effect of N on the native, Spartina foliosa. Our results indicate that excess N loading to these ecosystems enhances the vulnerability of intertidal habitats to rapid invasion by nonnative Spartina sp.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Impact of contaminant exposure on resource contests in an invasive fish

There is increasing concern for the disruptive effects seen in aquatic species exposed to environmental contaminants. However, few studies have investigated the impact of such contaminants on the behavior of individuals living in exposed waters. Contaminant exposure can affect animal populations by disrupting behaviors including feeding, locomotion, and mating. In this study, we examined how living in an ecosystem polluted by combinations of polycyclic aromatic hydrocarbons, polychlorinated biphenyls, and heavy metals (arsenic, cadmium, iron, lead, zinc) impacts contest behavior in the round goby (Neogobius melanostomus). Fish collected from heavily contaminated and cleaner sites in Lake Ontario were subjected to a resource contest to determine the effect of these contaminants on aggression and the establishment of dominance hierarchies, which in turn influence access to food, shelter, and mating opportunities. Dominance establishment (a clear resource winner) was less obvious among fish from the contaminated site compared to the more stable hierarchies that formed between pairs of fish from the clean site. Pairs of fish from the contaminated site performed more assessment displays compared to fish from clean sites. These results suggest that the costs of living in an environment under exposure can shape behavioral repertoires. The altered conflict resolution strategies of contaminated fish may reflect impaired cognitive function, sensory perception, and/or higher metabolic load associated with aggression. This study provides support for the utilization of quantifiable behavioral differences as ecologically relevant measures of contaminant exposure.     

Demographic feedback between clonal growth and fragmentation in an invasive seaweed

Many abundant plants, invertebrates, and seaweed are clonal, and this allows the formation of high-density aggregations, foraging, and the placement of modules into new space, and rapid rates of expansion. For these species, population density and rates of expansion are functions of recruitment of asexual modules and post-recruitment vegetative growth and survivorship. In this study, we provide the first experimental test of the relative importance of these two processes in determining the abundance of a clonal seaweed using Caulerpa taxifolia, an invasive green alga that spreads rapidly and reaches very high abundance. We asked two main questions: What is the relative importance to abundance (biomass) of vegetative stolon growth and fragment recruitment during expansion of established patches? Does greater fragment recruitment result in greater abundance in established patches? Vegetative growth of stolons underpinned patch expansion. Plots with stolons growing into them always had a greater abundance than plots where stolons were removed, even when fragment recruitment was increased. Greater recruitment only resulted in greater abundance when stolons were absent, a situation analogous to the establishment of new populations. Although post-recruitment processes were more important in determining abundance during patch expansion, there was greater ambient fragment recruitment when stolons were present compared to when they were absent, and as the abundance of C. taxifolia increased, demonstrating an important feedback between stolon growth, abundance, and fragment recruitment. In established patches, greater fragment recruitment over six months (six levels ranging from 0 to 480 recruits x m(-2) x mo(-1)) had no effect on biomass. Our experiments demonstrate that the rapid expansion and high abundance of invasive C. taxifolia are underpinned by post-recruitment vegetative growth and, during expansion, by a feedback between vegetative growth and asexual fragmentation.     

基于食物链关系评价转Bt基因棉对寄生性天敌的非靶效应

按照“棉花 -棉大卷叶螟 (Syleptaderogata)幼虫 -螟蛉绒茧蜂 (Apantelesruficrus)”三级营养食物链关系 ,对田间转Bt基因棉苏抗 310对非靶物种螟蛉绒茧蜂种群的影响进行了调查。发现转Bt基因棉苏抗 310棉田间螟蛉绒茧蜂种群生长受到了不良影响 ,种群数量显著下降 ,棉大卷叶螟幼虫发生高峰期高龄幼虫被寄生率降低。     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

Sex allocation is an important reproductive decision for parents. However, it is often assumed that females have substantial control over sex allocation decisions, and this is particularly true in haplodiploid insects, in which females apparently determine sex by deciding whether to fertilise an egg (and produce a diploid daughter) or not (and produce a haploid son). Mechanisms by which males may influence sex allocation are not so straightforward, and their potential influence on sex ratios has been somewhat neglected. Here, we test whether males influence offspring sex ratios in the parasitoid wasp Nasonia vitripennis. We show that some of the variation in observed sex ratios can be attributed to males when comparing the affect of male strain on sex ratio. We did not find among-male variation in sex ratio with a less powerful experiment using males from only one strain or an effect of male mating environment. Our data suggest that males can influence female sex ratios and contribute to the variation around the sex ratios optimal for females. However, the influence is not large, suggesting that females have more influence on sex allocation than do males. We conclude by considering whether male influences on sex ratio represent differences in male reproductive competence or deliberate attempts by males to increase their fitness by influencing daughter production.