Much as Rachel Carson's \"Silent Spring\" was a call to action against the pesticides that were devastating bird populations, Charles S. Elton's classic \"The Ecology of Invasions by Animals and Plants\" sounded an early warning about an environmental catastrophe that has become all too familiar today-the invasion of nonnative species. From kudzu to zebra mussels to Asian long-horned beetles, nonnative species are colonizing new habitats around the world at an alarming rate thanks to accidental and intentional human intervention. One of the leading causes of extinctions of native animals and plants, invasive species also wreak severe economic havoc, causing $79 billion worth of damage in the United States alone. Elton explains the devastating effects that invasive species can have on local ecosystems in clear, concise language and with numerous examples. The first book on invasion biology, and still the most cited, Elton's masterpiece provides an accessible, engaging introduction to one of the most important environmental crises of our time. Charles S. Elton was one of the founders of ecology, who also established and led Oxford University's Bureau of Animal Population. His work has influenced generations of ecologists and zoologists, and his publications remain central to the literature in modern biology. \"History has caught up with Charles Elton's foresight, and \"The Ecology of Invasions\" can now be seen as one of the central scientific books of our century.\"-David Quammen, from the Foreword to \"Killer Algae: The True Tale of a Biological Invasion\

The Ecology of Invasions by Animals and Plants

We DNA barcoded 2,597 parasitoid wasps belonging to 6 microgastrine braconid genera reared from parapatric tropical dry forest, cloud forest, and rain forest in Area de Conservacion Guanacaste (ACG) in northwestern Costa Rica and combined these data with records of caterpillar hosts and morphological analyses. We asked whether barcoding and morphology discover the same provisional species and whether the biological entities revealed by our analysis are congruent with wasp host specificity. Morphological analysis revealed 171 provisional species, but barcoding exposed an additional 142 provisional species; 95% of the total is likely to be undescribed. These 313 provisional species are extraordinarily host specific; more than 90% attack only 1 or 2 species of caterpillars out of more than 3,500 species sampled. The most extreme case of overlooked diversity is the morphospecies Apanteles leucostigmus. This minute black wasp with a distinctive white wing stigma was thought to parasitize 32 species of ACG hesperiid caterpillars, but barcoding revealed 36 provisional species, each attacking one or a very few closely related species of caterpillars. When host records and/or within-ACG distributions suggested that DNA barcoding had missed a species-pair, or when provisional species were separated only by slight differences in their barcodes, we examined nuclear sequences to test hypotheses of presumptive species boundaries and to further probe host specificity. Our iterative process of combining morphological analysis, ecology, and DNA barcoding and reiteratively using specimens maintained in permanent collections has resulted in a much more fine-scaled understanding of parasitoid diversity and host specificity than any one of these elements could have produced on its own.

https://www.pnas.org/content/pnas/105/34/12359.full.pdf

Extreme diversity of tropical parasitoid wasps exposed by iterative integration of natural history, DNA barcoding, morphology, and collections

Background Trees are major components of many terrestrial ecosystems and are grown in managed plantations and orchards to provide a variety of economically important products, including timber, pulp, fiber, and food. They are subject to a wide range of pests and diseases, of which the most important causative agents are viruses, bacteria, fungi, oomycetes, and insect herbivores. Research on tree pests and diseases has had a historical focus on trees of direct economic importance. However, some epidemics and infestations have damaged and killed common trees that are integral parts of natural ecosystems. These have harmed valuable landscapes and highlighted the wide-ranging consequences arising from tree pests and diseases. There is also growing concern that aspects of globalization—in particular, higher volumes and new forms of trade—may increase the risk of disease spread. A forest providing numerous ecosystem services is subject to a disease epidemic that reduces the abundance of a dominant native species, resulting in a change in forest structure. Initially, a wide range of ecosystem services (A to D) are harmed. But as trees grow to replace lost species, some (perhaps carbon storage or water purification) are regained, whereas others (perhaps the biodiversity supported by the diseased tree species) are permanently disrupted. Policy measures can both help prevent new diseases being introduced (the first stage) or improve recovery through management practices or planting resistant trees. Advances We review the challenges in maintaining tree health in natural and managed ecosystems. It is argued that it is helpful to consider explicitly the consequences of pests and diseases for the full range of ecosystem services provided by trees. In addition to forest and orchard products, tree pests and diseases can affect the ability of forests to sequester and store carbon, reduce flood risk, and purify water. They can affect the biodiversity supported by trees and the recreational and cultural values accorded to woodland by people. Many of these benefits are uncosted and enjoyed by different classes of stakeholders, which raises difficult questions about who should be responsible for measures to protect tree health. Changes in the risk of pest and disease introduction, the increasing prevalence of genetic reassortment leading to novel disease threats, and the potential role of climate change are all highlighted. Outlook Modern pest and disease management is based on an extensive science base that is rapidly developing, spurred in particular by modern molecular technologies. A research priority is to build a better understanding of why certain pathogens and insects become major pests and diseases. This will involve a better understanding of the molecular basis of pathogenicity and herbivory, as will ecological insights into why some species reach epidemic prevalence or abundance. It will also help anticipate which species may become a problem if they are transported to new geographical regions, recombine with other organisms, or experience new climatic conditions. However, identifying all species that may become pests will be impossible, and the Review stresses the importance of risk management at the “pathway of introduction” level, especially when modern trade practices provide potential new routes of entry. Last, when ecosystem services are provided by woods and forests rather than individual tree species, we need to understand better the consequences of pests and diseases that attack or feed on particular species.

https://science.sciencemag.org/content/sci/342/6160/1235773.full.pdf

The Consequence of Tree Pests and Diseases for Ecosystem Services

Community ecology is tasked with the considerable challenge of predicting the structure, and properties, of emerging ecosystems. It requires the ability to understand how and why species interact, as this will allow the development of mechanism-based predictive models, and as such to better characterize how ecological mechanisms act locally on the existence of inter-specific interactions. Here we argue that the current conceptualization of species interaction networks is ill-suited for this task. Instead, we propose that future research must start to account for the intrinsic variability of species interactions, then scale up from here onto complex networks. This can be accomplished simply by recognizing that there exists intra-specific variability, in traits or properties related to the establishment of species interactions. By shifting the scale towards population-based processes, we show that this new approach will improve our predictive ability and mechanistic understanding of how species interact over large spatial or temporal scales.


Synthesis

Although species interactions are the backbone of ecological communities, we have little insights on how (and why) they vary through space and time. In this article, we build on existing empirical literature to show that the same species may happen to interact in different ways when their local abundances vary, their trait distribution changes, or when the environment affects either of these factors. We discuss how these findings can be integrated in existing frameworks for the analysis and simulation of species interactions.

/pdf/beyond-species-why-ecological-interaction-networks-vary-3hy6s8a6bx.pdf

Beyond species: why ecological interaction networks vary through space and time

The brown planthopper (BPH), Nilaparvata lugens (Stal), which periodically erupted in tropical Asian rice before the 1960s, became a major threat after farmers adopted green revolution technologies in the 1960s. Management and policy changes in the 1980s and 1990s emphasized non-insecticidal tactics to avert BPH outbreaks. However, insecticides have resurfaced as the primary means for controlling rice insect pests and tropical Asian countries have recently experienced planthopper outbreaks in record numbers. Our review of factors that have contributed to the outbreaks points to insecticides as the most tangible outbreak factor primarily because of their harmful effects on natural enemies. BPH resistance to insecticides and especially imidacloprid has increased the probability of outbreaks as farmers have applied increasing quantities of insecticide in an attempt to combat resistant populations. Similarly, heavy use of nitrogen fertilizer, especially on hybrid rice, has increased the potential for outbreaks. Other factors triggering outbreaks are less documented, but we discuss the possibility that the high outbreak synchrony in geographically separated populations of BPH may suggest a “Moran effect” such as climate that promotes an environment favoring above-average increases in BPH populations. Also, we hypothesize that BPH functions as a metapopulation and, as such, periodic outbreaks could be a natural phenomenon requiring resupply of planthoppers into vacant areas to ensure genetic linkage among subpopulations. We conclude with a series of recommendations for research and policy changes aimed at better understanding the cause of BPH outbreaks and for developing sustainable management practices to prevent future outbreaks.

Resurrecting the ghost of green revolutions past: The brown planthopper as a recurring threat to high-yielding rice production in tropical Asia

Patterns in food-web structure have frequently been examined in static food webs, but few studies have attempted to delineate patterns that materialize in food webs under nonequilibrium conditions. Here, using one of nature's classical nonequilibrium systems as the food-web database, we test the major assumptions of recent advances in food-web theory. We show that a complex web of interactions between insect herbivores and their natural enemies displays significant architectural flexibility over a large fluctuation in the natural abundance of the major herbivore, the spruce budworm (Choristoneura fumiferana). Importantly, this flexibility operates precisely in the manner predicted by recent foraging-based food-web theories: higher-order mobile generalists respond rapidly in time and space by converging on areas of increasing prey abundance. This "birdfeeder effect" operates such that increasing budworm densities correspond to a cascade of increasing diversity and food-web complexity. Thus, by integrating foraging theory with food-web ecology and analyzing a long-term, natural data set coupled with manipulative field experiments, we are able to show that food-web structure varies in a predictable manner. Furthermore, both recent food-web theory and longstanding foraging theory suggest that this very same food-web flexibility ought to be a potent stabilizing mechanism. Interestingly, we find that this food-web flexibility tends to be greater in heterogeneous than in homogeneous forest plots. Because our results provide a plausible mechanism for boreal forest effects on populations of forest insect pests, they have implications for forest and pest management practices.

/pdf/fluctuations-in-density-of-an-outbreak-species-drive-332wqgjt9e.pdf

Fluctuations in density of an outbreak species drive diversity cascades in food webs

We conducted a 14-year intensive study of spruce budworm (Choristoneura fumiferana (Clem.)) survivorship at three study plots in largely balsam fir (Abies balsamea (L.) Mill.) stands in New Brunswick, Canada, to elucidate certain key mechanisms underlying spruce budworm outbreak cycles. The study covered a peak-to-declining phase (from 1981 and 1994) of the budworm outbreak cycle that had started in the early 1960s. Frequent sampling was carried out in each plot-year to construct a practically continuous survivorship curve, and the annual variation in population density was estimated. We found a high level of correlation between the studied phase of the outbreak cycle and annual variations in the survivorship over the postdiapause period, suggesting that postdiapause survivorship was the chief determinant of the cycle. We found the annual changes in population density in the present study to be closely similar in pattern to those from the provincial budworm surveys conducted in much larger areas. This implies that the mechanism underlying the population process found in the few study plots in largely balsam fir stands also applies to the process in much larger areas of diverse stand types.

The main source of postdiapause mortality is found to be natural enemies. The impacts of parasitoids and disease are evaluated by rearing budworm samples in the laboratory. Hymenopteran and dipteran parasitoids are by far the major sources of mortality, and microsporidians are the most prevalent pathogen. Occurrences of other entomopathogenic fungi and viruses were insignificant throughout the study. Seasonal changes in laboratory survivorship are compared with the corresponding field survivorship to estimate the effect of predation. No major mortality factor is found to singly play a predominant role in determining the outbreak cycle. Conversely, some minor factors are shown to have played significant roles. Thus, the importance of recognizing the action of natural enemies as a complex is emphasized for understanding the budworm outbreak cycle.

Finally, centered around the roles played by the chronological succession of natural enemies in the present study, the results of budworm research in New Brunswick since the mid-1940s are synthesized to outline basic mechanisms underlying the outbreak processes as a guide for further studies.

This article is protected by copyright. All rights reserved.

Mechanisms underlying spruce budworm outbreak processes as elucidated by a 14-year study in New Brunswick, Canada

An illustrated key is given to 28 genera, representing 10 families of at least 50 chalcidoid parasitoids and hyperparasitoids recorded from Choristoneura species (Tortricidae), or their primary parasitoids, in America north of Mexico. We include species reared throughout a 12-year study of the parasitoids and hyperparasitoids associated with the spruce budworm, Choristoneura fumiferana (Clemens), in New Brunswick, Canada, as well as those listed in various catalogs. Notes are provided for all the chalcidoid species, whether or not they are recorded in the literature. Nine species doubtfully or incorrectly recorded from Choristoneura species are also discussed.Primary parasitoids belonging to the Ichneumonidae, Braconidae, and Tachinidae have been recorded from 10 of the 16 Nearctic Choristoneura species but chalcidoid parasitoids are recorded, usually as hyperparasitoids, from only four of the 16 species. The species of primary parasitoids of Choristoneura checked for records of parasitism by Chalcidoidea are tabulated by host species. Also listed are species of primary parasitoids serving as hosts for each of the chalcidoid species. We report 51 new host records of primary parasitoids and hyperparasitoids from C. fumiferana and its parasitoids collected in New Brunswick.A new generic combination is given: Mesopolobus tortricis (Brues) comb.nov. from Psychophagus (proposed by Z. Boucek).

The chalcidoid parasitoids and hyperparasitoids (hymenoptera: chalcidoidea) of choristoneura species (lepidoptera: tortricidae) in america north of mexico

Nous avons examine les cocons de six especes des genres Apanteles et Dolichogenidea qui s'attaquent a la tordeuse des bourgeons de l'epinette, Choristoneura fumiferana Clemens, et aux microlepidopteres apparentes afin d'elucider les problemes taxonomiques et ecologiques, inherents a l'identification de ces especes lorsque les adultes n'arrivent pas a sortir de leur cocon. Ni la longueur du cocon, ni sa largeur, ni le rapport longueur a largeur n'ont pu etre utilises pour identifier les six especes, du fait du chevauchement considerable de ces parametres d'une espece a l'autre, et des effets exerces par la source des cocons. En employant une technique simple pour examiner les caracteristiques de tissage du cocon, nous avons constate que chaque espece presente un mode unique d'enroulement, determine par la facon dont la densite de tissage varie sur la longueur du cocon. Ce mode d'enroulement permet de caracteriser avec fiabilite chaque espece. Nous decrivons et illustrons les caracteristiques de tissage des diverses especes et proposons une cle permettant d'identifier chaque espece selon ces caracteristiques.

Identification of cocoons of Apanteles and Dolichogenidea (Hymenoptera: Braconidae) species attacking Choristoneura fumiferana (Lepidoptera: Tortricidae) and associated Microlepidoptera

Steven J. Pollock

Papers

Fluctuations in density of an outbreak species drive diversity cascades in food webs

Mechanisms underlying spruce budworm outbreak processes as elucidated by a 14-year study in New Brunswick, Canada

The chalcidoid parasitoids and hyperparasitoids (hymenoptera: chalcidoidea) of choristoneura species (lepidoptera: tortricidae) in america north of mexico

Identification of cocoons of Apanteles and Dolichogenidea (Hymenoptera: Braconidae) species attacking Choristoneura fumiferana (Lepidoptera: Tortricidae) and associated Microlepidoptera