Rodrigo J. Mercader

Dispersal of the emerald ash borer, Agrilus planipennis, in newly-colonized sites.

1 Emerald ash borer Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an invasive forest insect pest threatening more than 8 billion ash (Fraxinus spp.) trees in North America. Development of effective survey methods and strategies to slow the spread of A. planipennis requires an understanding of dispersal, particularly in recently established satellite populations. 2 We assessed the dispersal of A. planipennis beetles over a single generation at two sites by intensively sampling ash trees at known distances from infested ash logs, the point source of the infestations. Larval density was recorded from more than 100 trees at each site. 3 Density of A. planipennis larvae by distance for one site was fit to the Ricker function, inverse power function, and the negative exponential function using a maximum likelihood approach. The prediction of the best model, a negative exponential function, was compared with the results from both sites. 4 The present study demonstrates that larval densities rapidly declined with distance, and that most larvae (88.9 and 90.3%) were on trees within 100 m of the emergence point of the adults at each site. The larval distribution pattern observed at both sites was adequately described by the negative exponential function.

Evaluation of potential strategies to SLow Ash Mortality (SLAM) caused by emerald ash borer (Agrilus planipennis): SLAM in an urban forest.

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive pest native to Asia, has killed millions of ash (Fraxinus spp.) trees in North America since it was first discovered there in 2002. As of autumn 2011, A. planipennis has been detected in 15 US states and two Canadian provinces. A pilot project to slow the onset and progression of ash mortality, termed SLAM (SLow Ash Mortality), has been implemented in localized A. planipennis populations. Here we use spatially explicit simulations to evaluate the potential of a recently developed systemic insecticide to protect the ash resource in urban forests as a component of the SLAM approach. Over a 10-year horizon, simulations showed ash survival varied depending on: (i) how soon insecticide treatment began after the A. planipennis introduction; (ii) the proportion of trees treated; and (iii) the distribution of treated trees relative to the A. planipennis introduction point. Annual treatment of 20% of ash trees annually protected 99% of trees after 10 years, and the cumulative costs of treatment were substantially lower than costs of removing dead or severely declining ash trees.

The influence of satellite populations of emerald ash borer on projected economic costs in U.S. communities, 2010-2020.

The invasion spread of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) is characterized by the formation of satellite populations that expand and coalesce with the continuously invading population front. As of January 2010, satellite infestations have been detected in 13 states and two Canadian provinces. Understanding how newly established satellite populations may affect economic costs can help program managers to justify and design prevention and control strategies. We estimate the economic costs caused by EAB for the 10-yr period from 2010 to 2020 for scenarios of fewer EAB satellite populations than those found from 2005 to 2010 and slower expansion of satellite populations found in 2009. We measure the projected discounted cost of treatment, removal, and replacement of ash trees (Fraxinus spp.) growing in managed landscapes in U.S. communities. Estimated costs for the base scenario with the full complement of satellites in 2005-2010 and no program to mitigate spread is

Influence of foraging behavior and host spatial distribution on the localized spread of the emerald ash borer, Agrilus planipennis

12.5 billion. Fewer EAB satellites from 2005 to 2010 delay economic costs of

Allochronic isolation and incipient hybrid speciation in tiger swallowtail butterflies

1.0 to 7.4 billion. Slower expansion of 2009 satellite populations delays economic costs of

Hybridization leads to host-use divergence in a polyphagous butterfly sibling species pair

0.1 to 0.7 billion. Satellite populations that are both distant from the core EAB infestation and close to large urban areas caused more economic costs in our simulations than did other satellites. Our estimates of delayed economic costs suggest that spending on activities that prevent establishment of new satellite EAB populations or slow expansion of existing populations can be cost-effective and that continued research on the cost and effectiveness of prevention and control activities is warranted.

Diversification of host use in two polyphagous butterflies: differences in oviposition specificity or host rank hierarchy?

Management programs for invasive species are often developed at a regional or national level, but physical intervention generally takes place over relatively small areas occupied by newly founded, isolated populations. The ability to predict how local habitat variation affects the expansion of such newly founded populations is essential for efficiently targeting resources to slow the spread of an invasive species. We assembled a coupled map lattice model that simulates the local spread of newly founded colonies of the emerald ash borer (Agrilus planipennis Fairmaire), a devastating forest insect pest of ash (Fraxinus spp.) trees. Using this model, we investigated the spread of A. planipennis in environments with different Fraxinus spp. distributions, and explored the consequences of ovipositional foraging behavior on the local spread of A. planipennis. Simulations indicate that increased larval density, resulting from lower host tree density or higher initial population sizes, can increase the spread rate during the first few years after colonization by increasing a density-dependent developmental rate and via host resource depletion. Both the radial spread rate and population size were greatly influenced by ovipositional foraging behavior. Two known behaviors of ovipositing A. planipennis females, attraction towards areas with high ash tree density and attraction to stressed trees, had opposing effects on spread. Results from this model illustrate the significant influence of resource distribution and foraging behavior on localized spread, and the importance of these factors when formulating strategies to monitor and manage invasive pests.

Divergence in the ovipositional behavior of the Papilio glaucus group

Hybridization leading to reproductively isolated, novel genotypes is poorly understood as a means of speciation and few empirical examples have been studied. In 1999, a previously non-existent delayed flight of what appeared to be the Canadian tiger swallowtail butterfly, Papilio canadensis, was observed in the Battenkill River Valley, USA. Allozyme frequencies and morphology suggest that this delayed flight was the product of hybridization between Papilio canadensis and its sibling species Papilio glaucus. The mitochondrial DNA (mtDNA) restriction fragment length polymorphisms presented here indicate that only P. canadensis-like mtDNA occurs in this population, suggesting that introgression likely occurred from hybrid males mating with P. canadensis females. Preliminary studies of this population indicated that delayed post-diapause pupal emergence in this hybrid genotype was the root cause behind the observed delayed flight, which suggests a potential empirical example of a mechanism leading to reproductive isolation. Here we provide further evidence of the role of adult pupal emergence as a reproductive barrier likely leading to reproductive isolation. In particular, we present results from pupal emergence studies using four different spring and two different winter temperature treatments. The results indicate a clear separation of adult emergences between the hybrid population and both parental species. However, our results indicate that exceptionally hot springs are likely to lead to greater potential for overlap between the local parental species, P. canadensis, and this delayed population with hybrid origins. Conversely, our results also show that warmer winters are likely to increase the temporal separation of the hybrid population and the parental species. Finally, we report recently collected evidence that this hybrid population remains morphologically distinct.

Cost of potential emerald ash borer damage in U.S communities, 2009-2019

Climate warming has lead to increased genetic introgression across a narrow hybrid zone separating the eastern and Canadian tiger swallowtails (Papilio glaucus and Papilio canadensis). This situation has led to the formation of an allochronically separated hybrid population with a delayed emerging phenotype or “late flight”. Here, we assess how the recombination of the parental genomes that lead to this phenotype may have facilitated another major ecological shift, host-use divergence. We first contrast the ovipositional profiles of the late flight population to that of the parental species P. glaucus and P. canadensis. Subsequently we contrast the larval survival and growth of the late flight, a P. canadensis and a P. glaucus population, and a population from the northern edge of the hybrid zone on five hosts. Our results indicate that the ovipositional preference of this hybrid swarm is identical to that of the introgressing parental species, P. glaucus. Due to the absence of the preferred hosts of P. glaucus (Liriodendron tulipifera L. and Ptelea trifoliata L.) where the late flight occurs, this ovipositional pattern implies a functional specialization onto a secondary host of both parental species, Fraxinus americana L. In contrast, the larval host-use abilities represent a mixture of P. glaucus and P. canadensis, indicating divergence in larval host-use abilities has not taken place. However, high genetic variability (genetic coefficient of variation) is present for growth on F. americana in the late flight hybrid swarm and tradeoffs for larval performance on the preferred hosts of the parental species are evident; indicating a strong potential for future specialization in larval host-use abilities. This current scenario represents an instance where a shift in a major ecological trait, host use, is likely occurring as a byproduct of a shift in an unrelated trait (delayed emergence) leading to partial reproductive isolation.

Cost of potential emerald ash borer damage in U.S

Novel host usage may represent an initial step towards diversification or radiation onto novel hosts within an evolutionary lineage, particularly if a shift in host plant preference ranking takes place. Polyphagous stages of evolutionary lineages may represent transitional states in which novel host associations are more likely to develop, but may be more difficult to detect experimentally. The polyphagous sister species Papilio glaucus L. and Papilio canadensis (Lepidoptera: Papilionidae; these Papilio = Pterourus) are known to exhibit differences in host‐plant use, despite significant overlap in host‐use abilities, providing an opportunity to examine how host shifts in polyphagous species may occur and what the implications for future divergence may be. In particular, we were interested in (i) determining whether differences in oviposition behavior of these species were due to changes in specificity or shifts in host‐plant hierarchy, (ii) whether the varying preference for primary hosts also affected the preference for secondary hosts, and (iii) what the oviposition preferences of a new hybrid swarm population are. We examined more than 40 000 oviposition bouts from more than 400 P. glaucus, P. canadensis, and hybrid females placed in seven‐, three‐, or two‐choice assays. In each of the choice assays, leaves from plants in different plant families of varying suitability for P. glaucus and P. canadensis larvae were used. We found the primary difference between P. glaucus and P. canadensis to be limited to a Z‐linked shift in host rank hierarchy due to an acceptance of Populus tremuloides Michx. (Salicaceae) and reduced specificity for Liriodendron tulipifera L. (Magnoliaceae) in P. canadensis. In addition, we found the absence of the Z‐linked oviposition acceptance of P. tremuloides in a recently formed allochronically separated hybrid swarm population found in P. canadensis territory at the northern border of the P. glaucus and P. canadensis hybrid zone.