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Kenneth F. Raffa

Publications and source records attributed to Kenneth F. Raffa.

7 recordsLinked to original sources

What is a specialist? Quantifying host breadth enables impact prediction for invasive herbivores

Herbivores are commonly classified as host specialists or generalists for various purposes, yet the definitions of these terms, and their intermediates, are often imprecise and ambiguous. We quantified host breadth for 240 non-native, tree-feeding insects in North America using phylogenetic diversity. We demonstrated that a partitioning of host breadth: (1) causes 67% of non-native insects to shift from a generalist to specialist category, (2) displays a reduction in host breadth from the native to introduced range, (3) identifies an inflection point in a model predicting the likelihood of non-native insect ecological impact, with a corresponding change in behaviour associated with specialists versus generalists, and (4) enables three models for strong prediction of whether a non-native forest insect will cause high impacts. Together, these results highlight the primacy of how herbivore host recognition and plant defences mediate whether novel host interactions will result in high impact after invasion.

Ecology Letters

Phylogenetic risk assessment is robust for forecasting the impact of European insects on North American conifers

Some introduced species cause severe damage, although the majority have little impact. Robust predictions of which species are most likely to cause substantial impacts could focus efforts to mitigate those impacts or prevent certain invasions entirely. Introduced herbivorous insects can reduce crop yield, fundamentally alter natural and managed forest ecosystems, and are unique among invasive species in that they require certain host plants to succeed. Recent studies have demonstrated that understanding the evolutionary history of introduced herbivores and their host plants can provide robust predictions of impact. Specifically, divergence times between hosts in the native and introduced ranges of a nonnative insect can be used to predict the potential impact of the insect should it establish in a novel ecosystem. However, divergence time estimates vary among published phylogenetic datasets, making it crucial to understand if and how the choice of phylogeny affects prediction of impact. Here, we tested the robustness of impact prediction to variation in host phylogeny by using insects that feed on conifers and predicting the likelihood of high impact using four different published phylogenies. Our analyses ranked 62 insects that are not established in North America and 47 North American conifer species according to overall risk and vulnerability, respectively. We found that results were robust to the choice of phylogeny. Although published vascular plant phylogenies continue to be refined, our analysis indicates that those differences are not substantial enough to alter the predictions of invader impact. Our results can assist in focusing biosecurity programs for conifer pests and can be more generally applied to nonnative insects and their potential hosts by prioritizing surveillance for those insects most likely to be damaging invaders.

Ecological Applications

Predicting non-native insect impact: Focusing on the trees to see the forest

Non-native organisms have invaded novel ecosystems for centuries, yet we have only a limited understanding of why their impacts vary widely from minor to severe. Predicting the impact of non-established or newly detected species could help focus biosecurity measures on species with the highest potential to cause widespread damage. However, predictive models require an understanding of potential drivers of impact and the appropriate level at which these drivers should be evaluated. Here, we used non-native, specialist herbivorous insects of forest ecosystems to test which factors drive impact and if there were differences based on whether they used woody angiosperms or conifers as hosts. We identified convergent and divergent patterns between the two host types indicating fundamental similarities and differences in their interactions with non-native insects. Evolutionary divergence time between native and novel hosts was a significant driver of insect impact for both host types but was modulated by different factors in the two systems. Beetles in the subfamily Scolytinae posed the highest risk to woody angiosperms, and different host traits influenced impact of specialists on conifers and woody angiosperms. Tree wood density was a significant predictor of host impact for woody angiosperms with intermediate densities (0.5–0.6 mg/mm 3 ) associated with highest risk, whereas risk of impact was highest for conifers that coupled shade tolerance with drought intolerance. These results underscore the importance of identifying the relevant levels of biological organization and ecological interactions needed to develop accurate risk models for species that may arrive in novel ecosystems.

Biological Invasions

Growth and defense characteristics of whitebark pine (Pinus albicaulis) and lodgepole pine (Pinus contorta var latifolia) in a high-elevation, disturbance-prone mixed-conifer forest in northwestern Montana, USA

Recent, widespread tree mortality in the western U.S. resulting from changes in climate, pathogens, insect activity, and forest management practices has led to concerns for many ecologically and culturally important species. Within conifers, resin-based defenses have long been recognized as a primary defense mechanism against a variety of insects and pathogens. Oleoresin produced by trees contain complex mixtures of terpenoids that have numerous insecticidal and fungicidal properties. Research has also identified links between resin duct characteristics and increased probability of survival during bark beetle outbreaks. Whitebark pine ( Pinus albicaulis ) is a culturally significant high elevation species that provides numerous ecological services within subalpine and alpine ecosystems. Whitebark pine has co-evolved with a suite of biotic and abiotic disturbances. Individual trees allocate resources towards growth and resin-based defenses, making it a good candidate species to evaluate growth and defense relationships and tradeoffs. In this study we compared constitutive resin chemistry, tree growth and resin duct anatomy between similarly aged whitebark and lodgepole pine ( P. contorta var latifolia ) growing in proximity within a disturbance-prone, mixed-conifer forest in northwestern Montana. These two host species have varying degrees of historical exposure to mountain pine beetle. Our research yields four important findings. First, we did not find evidence of a tradeoff between tree growth and tree defenses (resin duct morphology and resin chemistry). This suggests that trees growing under favorable field conditions can experience high growth rates and still allocate ample resources towards defense. Second, we found that resin ducts and constitutive mono- and sesqui- terpenes were not correlated in lodgepole pine while duct production and area were positively related to constitutive monoterpenes, and duct size and area were positively related to constitutive sesquiterpenes, in whitebark pine. The lack of distinct, consistent relationships between these defensive features suggests that both whitebark and lodgepole pine trees present beetles with numerous, complex combinations of resin-based defenses. Third, based on constitutive terpene profiles, bark beetles are more likely to enter lodgepole pine but more likely to successfully elicit mass attacks in whitebark pine, which agrees with beetle attack and success patterns observed in the field. Fourth, overstory competition, particularly by Engelmann spruce ( Picea engelmannii ), can influence tree defenses, specifically by reducing constitutive terpene concentrations in lodgepole and whitebark pine. Competitive tree interactions could lead to altered bark beetle-conifer interactions as host and nonhost species migrate in response to changing climate. Our results suggest that strategies designed to support whitebark pine populations can benefit from better understanding interactions among growth, competition and physical and chemical defenses in response to multiple disturbance.

Montana

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

Scientific Data

The impact is in the details: Evaluating a standardized protocol and scale for determining non-native insect impact

Assessing the ecological and economic impacts of non-native species is crucial to providing managers and policymakers with the information necessary to respond effectively. Most non-native species have minimal impacts on the environment in which they are introduced, but a small fraction are highly deleterious. The definition of ‘damaging’ or ‘high-impact’ varies based on the factors determined to be valuable by an individual or group, but interpretations of whether non-native species meet particular definitions can be influenced by the interpreter’s bias or level of expertise, or lack of group consensus. Uncertainty or disagreement about an impact classification may delay or otherwise adversely affect policymaking on management strategies. One way to prevent these issues would be to have a detailed, nine-point impact scale that would leave little room for interpretation and then divide the scale into agreed upon categories, such as low, medium, and high impact. Following a previously conducted, exhaustive search regarding non-native, conifer-specialist insects, the authors independently read the same sources and scored the impact of 41 conifer-specialist insects to determine if any variation among assessors existed when using a detailed impact scale. Each of the authors, who were selected to participate in the working group associated with this study because of their diverse backgrounds, also provided their level of expertise and uncertainty for each insect evaluated. We observed 85% congruence in impact rating among assessors, with 27% of the insects having perfect inter-rater agreement. Variance in assessment peaked in insects with a moderate impact level, perhaps due to ambiguous information or prior assessor perceptions of these specific insect species. The authors also participated in a joint fact-finding discussion of two insects with the most divergent impact scores to isolate potential sources of variation in assessor impact scores. We identified four themes that could be experienced by impact assessors: ambiguous information, discounted details, observed versus potential impact, and prior knowledge. To improve consistency in impact decision-making, we encourage groups to establish a detailed scale that would allow all observed and published impacts to fall under a particular score, provide clear, reproducible guidelines and training, and use consensus-building techniques when necessary.

NeoBiota

Evolutionary history predicts high-impact invasions by herbivorous insects

A long‐standing goal of invasion biology is to identify factors driving highly variable impacts of non‐native species. Although hypotheses exist that emphasize the role of evolutionary history (e.g., enemy release hypothesis & defense‐free space hypothesis), predicting the impact of non‐native herbivorous insects has eluded scientists for over a century. Using a census of all 58 non‐native conifer‐specialist insects in North America, we quantified the contribution of over 25 factors that could affect the impact they have on their novel hosts, including insect traits (fecundity, voltinism, native range, etc.), host traits (shade tolerance, growth rate, wood density, etc.), and evolutionary relationships (between native and novel hosts and insects). We discovered that divergence times between native and novel hosts, the shade and drought tolerance of the novel host, and the presence of a coevolved congener on a shared host, were more predictive of impact than the traits of the invading insect. These factors built upon each other to strengthen our ability to predict the risk of a non‐native insect becoming invasive. This research is the first to empirically support historically assumed hypotheses about the importance of evolutionary history as a major driver of impact of non‐native herbivorous insects. Our novel, integrated model predicts whether a non‐native insect not yet present in North America will have a one in 6.5 to a one in 2,858 chance of causing widespread mortality of a conifer species if established (R2 = 0.91) Synthesis and applications. With this advancement, the risk to other conifer host species and regions can be assessed, and regulatory and pest management efforts can be more efficiently prioritized.

Ecology and Evolution