Evaluation of Biorational Insecticides for Management of Euproctis chrysorrhoea (Lepidoptera: Erebidae).

The browntail moth (Euproctis chrysorrhoea L.) is an invasive species which over the past five years, has been undergoing outbreaks on a scale not seen in the northeastern U.S. in over 100 years. Browntail moth larvae feed on and defoliate a number of deciduous tree species, but the health issues caused by contact with the toxic urticating hairs of the overwintered larvae have resulted in very low tolerance for this pest amongst homeowners and land managers. Few recent studies have been conducted to assess management options for browntail moth, which is abundant in ecologically sensitive areas along coastal waters, and around people's homes. We investigated the potential to manage overwintered larvae with currently available biorational insecticides. Laboratory bioassays revealed susceptibility to Beauveria bassiana (Balsamo) Vuillemin (Hypocreales: Cordycipitaceae), Bacillus thuringiesis kurstaki (Berliner), azadirachtin, and spinosad products. A field trial was conducted to assess efficacy of B. bassiana, Btk, and azadirachtin. All treatments reduced the abundance of larvae compared with the control, but only two applications of Btk and single application of a tank mix of B. bassiana and Btk reduced pupal nest abundance. A laboratory experiment revealed that temperature did not affect the feeding and survival of larvae exposed to the field trial foliage from the Btk and the Btk/B. bassiana tank mix treatments, whereas slower feeding rates and increased time to death were observed with the control and B. bassiana alone treatment.

Bacterial transfer from Pristionchus entomophagus nematodes to the invasive ant Myrmica rubra and the potential for colony mortality in coastal Maine.

The necromenic nematode Pristionchus entomophagus has been frequently found in nests of the invasive European ant Myrmica rubra in coastal Maine, United States, and may contribute to ant mortality and collapse of colonies by transferring environmental bacteria. Paenibacillus and several other bacterial species were found in the digestive tracts of nematodes harvested from collapsed ant colonies. Serratia marcescens, Serratia nematodiphila, and Pseudomonas fluorescens were collected from the hemolymph of nematode-infected wax moth (Galleria mellonella) larvae. Virulence against waxworms varied by the site of origin of the nematodes. In adult nematodes, bacteria were highly concentrated in the digestive tract with none observed on the cuticle. In contrast, juveniles had more on the cuticle than in the digestive tract. Host species was the primary factor affecting bacterial community profiles, but Spiroplasma sp. and Serratia marcescens sequences were shared across ants, nematodes, and nematode-exposed G. mellonella larvae.

Factors Influencing the Population Fluctuations of Euproctis chrysorrhoea (Lepidoptera: Erebidae) in Maine.

The browntail moth (Euproctis chrysorrhoea (L.)) is a forest pest that was accidentally introduced in the late 1800's and spread throughout New England in the early part of the 20th Century. At its peak range expansion in 1915 it encompassed an area of 150,000 km2 after which populations declined. By the 1960s, its distribution had receded to relic populations on outer Cape Cod, MA, and islands in Casco Bay, ME. In 1989 browntail moth resurged in Maine, with periodic, moderate outbreaks before a dramatic increase of the population occurred in 2016. We examined the pattern of annual defoliation by browntail moth since its resurgence in the 1990s as well as variation in populations throughout infested areas in Maine during three years of the recent outbreak, 2016-2018, relative to differences in weather, parasitism and habitat characteristics. Levels of defoliation over 24 yr were predicted by the preceding spring precipitation (-, negative effect) and the year's previous late summer and early fall temperatures (+, positive effect) when first to third instar larvae feed and then construct winter hibernacula. Late summer temperatures predicted the abundance of hibernacula across outbreak areas (+). Early spring temperatures (+) and early and late spring precipitation (-) predicted early summer larval and pupal nest abundance. Warmer fall temperatures result in more mature populations coming out of winter hibernacula in the spring, whereas spring precipitation drives epizootic outbreaks of Entomophaga aulicae (Reichardt in Bail) Humber (Entomophthorales: Entomophthoraceae). with parasitoids playing a lesser role. Climate trends indicate continued increases in fall temperatures since browntail moth resurgence.

Interactions between sympatric invasive European fire ants (Myrmica rubra) and blacklegged ticks (Ixodes scapularis).

The blacklegged tick (Ixodes scapularis) and the invasive European fire ant (Myrmica rubra) are both expanding throughout their sympatric range in coastal New England. Ixodes scapularis is the primary vector of the bacterium Borrelia burgdorferi, which is the causative agent of Lyme disease, and Mount Desert Island, Maine, home to Acadia National Park, currently is affected by a high Lyme disease burden. Ticks have many natural predators, including ants, although no previous studies have investigated interactions between these two species. To test the hypothesis that the presence of M. rubra alters I. scapularis abundance, we collected ticks by drag-sampling at eight ant-infested sites and eight uninfested control sites in Acadia National Park. We found that nymph density was significantly higher at ant-infested sites, while larval density was significantly higher at control sites. In addition, we conducted a laboratory bioassay to measure M. rubra aggression against I. scapularis larvae, nymphs, and adults and Dermacentor variabilis adults, and found that ant aggression was significantly higher against D. variabilis adults than I. scapularis adults. Our findings support the hypothesis that M. rubra has divergent effects across I. scapularis life stages, and we discuss possible ecological mechanisms, including optimal microclimate and predation, that could promote density of nymphs while inhibiting density of larvae.

Evaluation of Four Plant Extract Repellents for Management of the European Red Ant Myrmica rubra (Hymenoptera: Formicidae).

Spearmint oil, peppermint oil, neem oil, and d-limonene were tested as nest site repellents against the colonization of the invasive European red ant, Myrmica rubra (L.) in both laboratory and field trials. In a laboratory assay, a 10% (v/v) solution of each extract repelled M. rubra colonies from nesting in plant pots filled with moist soil compared to water-treated controls, when applied as a dip to pots. Extracts also repelled colonies compared to a water control in a second laboratory experiment, where pots were dipped 15 d prior to the start of the experiment. In a 2008 field comparison, 20 plant pots, filled with moist potting soil, were dipped in either 10% (v/v) spearmint oil or peppermint oil, 70% (v/v) neem oil, or a water control and left in infested sites. Seven control pots were colonized over 3 mo, while none of the extract-treated pots were colonized. This field trial was repeated for a 15-wk duration in 2009 with the addition of a 10% (v/v) d-limonene solution as a treatment, and a change in neem oil concentration to 10% (v/v). Spearmint and peppermint oils repelled M. rubra colonies for the duration of the experiment. Neem oil and d-limonene repelled colonies for 3 and 4 wk, respectively. These extracts, especially mint oils, show potential as low-hazard repellents against M. rubra in greenhouse and nursery settings, and could reduce the number of new infestations incurred by the transport of plant stock.

Host Plant Effects on Winter Moth (Lepidoptera: Geometridae) Larval Development and Survival.

The winter moth, Operophtera brumata (L.) is an invasive forest and agricultural pest in North America that causes severe defoliation to a wide range of host species. This study examines the differential larval densities, development, and survival on seven host species in midcoast Maine: red oak (Quercus rubra L., Fagales: Fagaceae), apple (Malus domestica L., Rosales: Rosaceae) and crab apple (Malus sp. L., Rosales: Rosaceae), red maple (Acer rubrum L., Sapindales: Sapindaceae), pin cherry (Prunus pensylvanica L., Rosales: Rosaceae), white birch (Betula papyrifera L., Fagales: Betulaceae), wild lowbush blueberry (Vaccinium angustiflolium L., Ericales: Ericaceae), and highbush blueberry (Vaccinium corymbosum L., Ericales: Ericaceae). We also explore the degree of synchrony between selected host plants and larval hatch and its effect on survival. We found that densities, development, and survival were significantly greater on red oak (Quercus rubra) and apple (Malus sp.) than on all other target species and were lowest on pin cherry (Prunus pennsylvanica). We found low larval densities in open, wild lowbush blueberry fields; however, larvae successfully fed and developed on wild lowbush blueberry in a laboratory setting. This suggests that winter moth is a potential pest to wild lowbush blueberry in Maine if the outbreak expands to include areas with wild lowbush blueberry production.

Impacts of the Invasive European Red Ant (Myrmica rubra (L.): Hymenoptera; Formicidae) on a Myrmecochorous System in the Northeastern United States.

We investigated the impact of an invasive ant species from Europe, Myrmica rubra (L.), on a myrmecochorous system (seeds dispersed by ants) in its invaded range in North America. We assessed: 1) how M. rubra process the myrmecochorous diapsores (seeds and elaiosome as a single dispersal unit transported by ants) in comparison with native ants; 2) its preference for common native and invasive diaspore species relative to native ants; 3) how far they disperse diaspores in the field; and 4) the diaspore removal rate by invertebrates and vertebrates in infested areas compared to noninvaded sites. Field experiments demonstrated higher diaspore removal rates over a 10-min and 24-h period by M. rubra compared to native ants. M. rubra's diaspore dispersal distance was 40% greater compared to native ants. In two of three laboratory studies and one field study, there was no significant difference between the seed species which M. rubra and native ants selected. Our data suggest no long-term deleterious effects of M. rubra's invasion on diaspore dispersal in the Maine plant community that is comprised of both native and invasive species. This implies that M. rubra benefits from the myrmechorous plant species' diaspores by increasing their dispersal range away from the parent plant and potentially reducing seed predation. However, it is not known whether the fact that the native ant fauna and M. rubra are attracted to the same plant species' diaspores creates a high level of competition between the ants with deleterious effects on the native ant community.

Ophiocordyceps myrmicarum, a new species infecting invasive Myrmica rubra in Maine.

Myrmica rubra L. (Formicidae: Hymenoptera), an invasive ant species in Maine, has been problematic in coastal communities over the past 15years and the subject of studies on its ecology, effects, and management. In recent years we observed a fungus growing from moribund M. rubra (in vitro) and have isolated multiple cultures of this previously unreported fungus over 2 y. We analyzed morphology and DNA molecular sequences of this fungus and identified it as a member of the genus Hirsutella Pat. (Ophiocordycipitaceae; Hypocreales), the use of which is suppressed in favor of a monophyletic Ophiocordyceps clade. Molecular and morphological characters support the description of this fungus as a new species, O. myrmicarum, which is the first species in this genus to be isolated from North American M. rubra.

Phylogeny of Hirsutella species (Ophiocordycipitaceae) from the USA: remedying the paucity of Hirsutella sequence data.

Hirsutella (Ophiocordycipitaceae: Hypocreales) is a genus of insect, mite, and nematode pathogens with an asexual morph, which generally produce a mucilaginous cluster of one or several conidia on phialides that are basally subulate and taper to a fine neck. The generic name Hirsutella has been proposed for suppression in favour of Ophiocordyceps as a consequence of the ending of dual nomenclature for different morphs of pleomorphic fungi in 2011. Though the generic name is well established, geographically dispersed, and speciose, exceptionally few sequences are available in online databases. We examined 46 isolates of 23 Hirsutella species from the USA, curated by the USDA-ARS Collection of Entomopathogenic Fungal Cultures (ARSEF Culture Collection), that previously had not been molecularly characterized and produced a phylogeny of these organisms; we included previously published Hirsutella and Ophiocordyceps taxa. In producing the largest phylogeny of Hirsutella isolates so far, we provide: (1) context for discussing previously-hypothesized relationships; (2) evidence for revisions as taxonomic transitions move forward; and (3) available molecular data to be incorporated into further evolutionary studies of Ophiocordycipitaceae.

Mechanisms of competitive displacement of native ant fauna by invading Myrmica rubra (Hymenoptera: Formicidae) populations.

Exotic ants have become invasive in many regions around the world, with variable ecological impacts. Postinvasion, native ant communities are often found to be depauperate, though the causes of this apparent lack of coexistence are rarely well known. Myrmica rubra (L.), a Palearctic Myrmecine ant, is currently expanding its range as an invasive in North America. This aggressive ant forms dense, patchy local infestations and appears to aggressively displace native ant fauna. We measured behavioral interactions and rates of recruitment in experimental field assays pitting native foragers against captive colonies of M. rubra at tuna-jelly or aphid baits in uninfested areas of Mt. Desert Island, ME. Behavioral interactions were idiosyncratic with respect to the native opponent, but M. rubra generally showed significantly higher levels of recruitment, aggression, and displacement of native foragers. As with other invasive ant species shown to have broken the "dominance-discovery trade-off," M. rubra was consistently faster to discover baits and disproportionately displaced native foragers, providing a plausible proximate mechanism for native ant exclusion. Finally, we surveyed ant recruitment at baits for 24 h in August 2004 at four sites with varying M. rubra abundance but found little evidence of temporal niche partitioning. Taken together, these results indicate competitive superiority by M. rubra with respect to native ant communities of the northeastern North America and suggest direct aggression and competitive exclusion at food resources can lead to local native displacement.

Homopterans and an invasive red ant, Myrmica rubra (L.), in Maine.

Myrmica rubra (L.), is an invasive ant that is spreading across eastern North America. It is presently found in over 40 communities in Maine and areas in Vermont, New Hampshire, Massachusetts, New York, and several provinces in the Canadian Maritimes and Ontario. In addition to disrupting native ant faunas, invasive ants also have been shown to influence homopteran abundance and species composition. We conducted surveys of Homoptera in infested and noninfested sites and conducted manipulative experiments to quantify the effects of M. rubra on homopteran abundance and composition in the summers of 2003, 2006, and 2007 on Mount Desert Island, ME. In 2003, Homoptera family-level richness was higher in infested sites compared with noninfested sites with two out of three sampling methods. Homopteran abundance in infested compared with noninfested sites depended upon the site. The sites with the highest population of M. rubra were associated with significant differences in Homoptera population abundance. In 2006 and 2007, two out of three host plants sampled had significantly higher abundances of the aphids, Aphis spiraephila Patch and Prociphilus tessellatus Fitch. An ant exclusion field experiment on the native plant, meadowsweet (Spiraea alba Du Roi), resulted in higher abundances of A. spiraephila with M. rubra tending compared with native ant tending. A predator exclusion field experiment was conducted on meadowsweet using adult ladybeetles, Hippodamia convergens Guérin-Méneville, larval green lacewings, Chyrsoperla carnea Stephens, and no predators. Predator impacts on aphid populations were reduced in the presence of M. rubra with C. carnea and moderately reduced with H. convergens.

New fungal pathogens of the red ant, Myrmica rubra, from the UK and implications for ant invasions in the USA.

The red ant, Myrmica rubra, is an increasingly invasive pest species in north-eastern USA, where it is known as the European fire ant. During surveys for natural enemies in part of its native range in the UK, three previously unreported fungal pathogens developed on ants when incubated in the laboratory. These are described and illustrated: Paraisaria myrmicarum sp. nov., Hirsutella stilbelliformis var. myrmicarum var. nov., and Hirsutella subramanianii var. myrmicarum var. nov. Based on analyses of the protein coding region EF-1α and LSU rDNA, all three described taxa are shown to be affiliated with the hypocrealean family Ophiocordycipitaceae. The implications for the management of M. rubra in its exotic North American range using classical biological control are discussed.

Ant diversity and distribution in Acadia National Park, Maine.

Exotic ant species are a primary threat to ant biological diversity, posing a negative impact to native ant communities. In this study, we examine species richness of ants (family Formicidae) in Acadia National Park, ME, as a fundamental step toward understanding the present impact of the exotic species Myrmica rubra on native ant species. Twelve habitat types were sampled, along six transects, with pitfall traps, visual searching, bait traps, and leaf litter extraction, and the aid of 34 volunteers. We report 42 species of ants in Acadia National Park, comprising five subfamilies (Amblyoponinae, Dolichoderinae, Formicinae, Myrmicinae, and Ponerinae) and 15 genera; the cataloged species represents 75% of the species originally recorded in the area by Procter (1946). Our findings suggest M. rubra is currently not a dominant species throughout the entire island. However, where this species has invaded locally, few competing native species coexist. The species Lasius alienus, Formica subsericea, Myrmica detritinodis, Camponotus herculeanus, Formica argentea, Formica aserva, and Tapinoma sessile occurred most often in our survey. We report the ant species Amblyopone pallipes and Dolichoderus mariae as two new records for the state of Maine.

Intercolony aggression within and among local populations of the invasive ant, Myrmica rubra (Hymenoptera: Formicidae), in coastal Maine.

Myrmica rubra L. was introduced into New England in the early 20th century, and at present, has a patchy distribution in parts of northeastern North America, including records from 31 communities in Maine. M. rubra is highly polygynous, and colonies reproduce vegetatively, forming dense local populations where conditions are favorable. Using mobile nests and baited arenas in a series of field aggression bioassays, we tested patterns of internest tolerance within and among local populations on Mt. Desert Island, ME. We found that foragers originating from fragments of the same colony or from neighboring nests retained a high level of intraspecific tolerance over several months, whereas significant intercolony aggression among workers was present between colonies within the same local patch separated by approximately 10 m. Within populations, aggression score values were found to increase linearly with internest distance within a site. Aggression was highest between colonies from spatially different populations on the island and was higher still when nests were assayed against colonies at an off-island site 70 km away in Castine, ME. These data strongly suggest a multicolonial organization within and among local populations of M. rubra in parts of its introduced range. These findings contradict the loss of intraspecific aggression and unicolonial social structure over large geographic areas that have previously been observed in other invasive ant species, particularly Linepithema humile Mayr.

Distribution of an invasive ant, Myrmica rubra (Hymenoptera: Formicidae), in Maine.

Introduced populations of the north temperate ant species, Myrmica rubra (L.), have become pestiferous in various locations in the northeastern United States, particularly in coastal communities in Maine. Native populations of this ant are widely distributed throughout northern Europe and western Asia; however, nest densities in its native range do not usually reach the high levels observed for many introduced populations. This aggressive ant readily stings, and because of its high densities, homeowners continually encounter nests at a frequency that interferes with use of their properties. Surveys were conducted in Maine from 2001 through 2004 to determine the statewide extent of current infestations. Nests in established populations in coastal Maine were sampled from April through September to assess seasonal changes in the density and composition of colonies. Similarities and differences between introduced populations of M. rubra in Maine and published reports of this species in Europe are discussed. Museum records of this species in North America were also researched and are presented.

Potential of Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) for biological control of European corn borer (Lepidoptera: Crambidae) in solanaceous crops.

We assessed the ability of Trichogramma ostriniae (Peng & Chen) to locate and parasitize Ostrinia nubilalis (Hübner) eggs in crops other than corn, and we evaluated the efficacy of inundative releases of the parasitoid in two solanaceous crops, pepper and potato. Despite a greater plant surface area to search, parasitism of O. nubilalis eggs was consistently higher in sweet corn than dicotyledonous crops such as pepper, snap bean, broccoli, potato, and melon, in choice and no-choice experiments. Nonetheless, in 2002 and 2003, we made four to five separate inundative releases of approximately 30,000-50,000 T. ostriniae per 0.02 ha in nine pepper fields in Virginia, Pennsylvania, and Massachusetts and compared O. nubilalis egg parasitization and fruit damage in those plots with spatially isolated nonrelease plots. Egg parasitization averaged 48.7% in T. ostriniae release plots, which was significantly higher than in nonrelease plots (1.9%). Also, cumulative pepper fruit damage averaged 8.7% in release plots, which was significantly less than in nonrelease plots (27.3%). In potatoes in 2002 and 2003, we made two releases of approximately 75,000 T. ostriniae per 0.2 ha in nine fields in Maine and Virginia and compared O. nubilalis damage in those plots with that in nonrelease plots. T. ostriniae releases significantly reduced the number of tunnel holes and number of O. nubilalis larvae in potato stems. We conclude that this parasitoid has great potential as a biocontrol agent for O. nubilalis in solanaceous crops.

Starvation induced stress and the susceptibility of the Colorado potato beetle, Leptinotarsa decemlineata, to infection by Beauveria bassiana.

Starvation of second instar Colorado potato beetle larvae for 24h immediately after treatment with Beauveria bassiana conidia increased susceptibility to the pathogen and subsequent sporulation of cadavers but decreased time to larval death. In feeding studies, B. bassiana-treatment had no effect on subsequent larval development, and mortality occurred 5-6 days after treatment. Twenty-four hours of starvation alone retarded subsequent larval development but did not affect mortality. Mortality of B. bassiana-treated starvation stressed larvae occurred 4-5 days after treatment. Both B. bassiana treatment and 24h starvation significantly reduced total foliage consumption and daily weight gains. On the day of treatment, B. bassiana had no effect on the efficiency with which food was converted to biomass (ECI). ECI was not affected by B. bassiana or starvation alone on the day following treatment but was significantly affected by a combination of both. When larvae were exposed to a range of limited food quantities, ECI decreased with decreasing food availability but only extreme stress (starvation for 24h) increased susceptibility to B. bassiana. Topical application of Dacryodes excelsa resin (an antifeedant) to potato leaves caused a concentration dependent reduction in foliage consumption and weight gain by second instar larvae but did not affect larval mortality. When larvae were exposed to a fixed concentration of B. bassiana and a range of antifeedant concentrations there were significant linear relationships between 24h larval weight gain and mortality and 24h larval weight gain and sporulation. The interaction between starvation stress and the susceptibility to B. bassiana infection is discussed and its possible implications in pest management considered.