Evaluation of Light Attraction for the Stored-Product Psocids, Liposcelis entomophila, Liposcelis paeta, and Liposcelis brunnea.

Psocids have become global pests of stored commodities as they can cause considerable economic losses. These insects are difficult to control because they have developed resistance to many chemical insecticides. Therefore, it is crucial to investigate alternative integrated pest management (IPM) approaches, such as the use of light attraction for monitoring and/or controlling psocids. Light attraction has been studied for Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae) but not for other psocid species. In this study, we investigated the response of adults of three psocid species (Psocoptera: Liposcelididae), Liposcelis entomophila (Enderlein), Liposcelis paeta Pearman, and Liposcelis brunnea Motschulsky, to six wavelengths of light from light-emitting diode (LED) in paired-choice pitfall tests. L. entomophila females and males were not attracted to any of the wavelengths tested. L. paeta females responded positively to two ultraviolet (UV) wavelengths (351 and 400 nm) and to green light (527 nm), while males did not respond to any light. L. brunnea females and males responded positively to all six wavelengths evaluated. Most of the LEDs that elicited positive responses to L. paeta females and L. brunnea females and males were also preferred when these lights were presented against brewer's yeast, a food attractant highly preferred by several psocid species. Females of L. paeta and L. brunnea were attracted to white light when compared with a blank, but females of L. entomophila were not attracted to white light compared to a blank.

Health Hazards Associated with Arthropod Infestation of Stored Products.

Insects and mites are common inhabitants and accidental invaders of food, including durable commodities, and their presence can have both direct and indirect effects on human health. The most common direct effect is contamination of food with arthropod fragments and related contaminants, which may be allergenic or even carcinogenic. The most important indirect effect is that their presence can change the storage microenvironment, making durable products suitable for the rapid development of fungi and other microorganisms. Some of these fungi can produce toxins (e.g., aflatoxins) that endanger human health. Insects may actively or passively contribute to the spread of microorganisms, increasing product contamination, and they may host bacteria that have developed antibiotic resistance, contributing to their spread in food. Several species also may host, attract, or transmit tapeworms, predators, or parasitoids that may affect health. This review synthesizes research on these topics and suggests directions for future research.

Sublethal exposure, insecticide resistance, and community stress.

Insecticides are an invaluable pest management tool and anthropogenic stressors of widespread environmental occurrence that are subject to biased perceptions based on the targeted application, market value of use, and regulatory requirements. As a result, short-term and simplistic efforts focusing on lethal effects toward individual species and populations prevail. Holistic and comprehensive studies exploring rather common sublethal insecticide exposures are rare, particularly considering their potential role in structuring populations and communities in diverse environmental settings and potentially interfering in a range of ecological interactions. Studies on insecticide resistance, for example, do not go beyond population-based studies, disregarding temporal and spatial effects in the associated community, and rarely considering the whole of sublethal exposure. Some of these knowledge gaps are here recognized and explored.

Evaluation of Potential Attractants for Six Species of Stored-Product Psocids (Psocoptera: Liposcelididae, Trogiidae).

Psocids have emerged as worldwide pests of stored commodities during the past two decades, and are difficult to control with conventional management tactics such as chemical insecticides. Therefore, it is necessary to investigate alternative management strategies, such as the use of attractants for monitoring and controlling psocids, which can be incorporated into integrated pest management programs for psocids. Using a two-choice pitfall test, we studied the response of adults of different ages and sexes of Liposcelis entomophila (Enderlein) (Psocoptera: Liposcelididae), Liposcelis paeta Pearman, Liposcelis decolor (Pearman), Liposcelis brunnea Motschulsky, Liposcelis corrodens (Heymons), and Lepinotus reticulatus Enderlein (Psocoptera: Trogiidae) to volatiles from different potential attractants including grains, grain-based oils, brewer's yeast, wheat germ, and commercially available kairomone lures. For all species tested, sex and age did not have a major influence on response to the different potential attractants. Brewer's yeast most consistently elicited the strongest response for psocids, but this response frequently was not different from that to wheat germ and wheat germ oil. The percentage response to brewer's yeast varied among the psocid species tested: L. decolor (73-78%), L. entomophila (62-73%), L. brunnea (64-68%), L. paeta (42-57%), Lep. reticulatus (40%), and L. corrodens (15-19%). Two psocids species (L. corrodens and Lep. reticulatus) had low responses to all the potential attractants evaluated compared with the other four species. These results show there is high potential for using these attractants in a psocid-monitoring program.

Efficacy of Methyl Bromide for Control of Different Life Stages of Stored-Product Psocids.

The psocid species Liposcelis paeta Pearman, Liposcelis entomophila (Enderlein), Liposcelis decolor (Pearman), Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae), and Lepinotus reticulatus Enderlein (Psocoptera: Trogiidae) were evaluated in laboratory bioassays to determine their susceptibility to six concentrations of methyl bromide (0.027, 0.113, 0.280, 0.393, 0.452, and 0.616 g/m3) after 48 h of exposure at 27.5°C. The life stages that were evaluated were adults (for all species), nymphs (for all species except Lep. reticulatus), and eggs (for L. entomophila, L. decolor, and L. bostrychophila). Adults and nymphs were very susceptible, and complete mortality was recorded at concentrations between 0.027 and 0.280 g/m3. In contrast, eggs were by far more tolerant than adults and nymphs for all species tested. At 0.027 g/m3, mortality did not exceed 53%, while survival was high even at 0.113 g/m3. Complete (100%) egg mortality was recorded at 0.393 g/m3 for L. decolor and at 0.452 g/m3 for L. entomophila and L. bostrychophila; concentrations estimated to give 99% mortality for eggs of these three species were 0.710, 1.044, and 0.891 g/m3, respectively. These results show that stored-product psocids are susceptible to methyl bromide, but concentrations of ≥0.452 g/m3 should be used to control all life stages.

Competition among species of stored-product psocids (Psocoptera) in stored grain.

We evaluated the competition among stored-product psocid species by conducting two series of laboratory experiments. In the first series, three species of Liposcelididae were used: Liposcelis bostrychophila, Liposcelis decolor, and Liposcelis paeta. Five adult females of these species were placed in vials containing wheat, either alone or in all possible combinations of two species. The number of adults in the vials was counted after 35, 70, 105, 140, and 175 days. These tests were performed at 25 and 30°C. At 25°C, there were no differences in numbers of L. bostrychophila when this species was reared either alone or with each of the other two species. At 30°C, L. bostrychophila was the dominant species. The presence of L. bostrychophila had a negative effect on the growth of populations of L. decolor and L. paeta. The presence of L. paeta did not affect growth of populations of L. decolor, although the presence of L. decolor occasionally reduced growth of populations of L. paeta. In the second series of tests, L. bostrychophila adult females were placed in vials of wheat either alone or with adult females of Lepinotus reticulatus, at the ratios of (L. bostrychophila: L. reticulatus) 10∶0, 9∶1, 7∶3, 5∶5, 3∶7, 1∶9, and 0∶10. These tests were carried out only at 30°C, and the observation periods were the same as for the first series of tests. Liposcelis bostrychophila was the dominant species in this case as well, regardless of the ratio of the parental females. At the end of the experimental period, L. reticulatus was present only in vials that contained this species alone. Our results showed that L. bostrychophila outcompetes the other stored-product psocid species tested.

Residual efficacy of chlorfenapyr for control of stored-product psocids (Psocoptera).

The residual effect of chlorfenapyr (Phantom) was evaluated for residual control of three stored-product psocid species: Liposcelis bostrychophila Badonnel, Liposcelis entomophila (Enderlein), and Liposcelis paeta Pearman (Psocoptera: Liposcelididae). Chlorfenapyr was applied to individual arenas with a concrete surface at rates of 0, 2.8, 13.8, 20.6, 27.5, 55, and 110 mg active ingredient (AI)/m2. Adults were exposed on the treated arenas and mortality assessed after 1, 2, and 3 d. The procedures were repeated weekly on the same treated arenas for 3 wk to assess residual efficacy. At each week, mortality of all species was low after 1 d of exposure but notably increased after 2 or 3 d. L. entomophila was the most susceptible species, with 99-100% mortality at rates of 13.8 mg/m2 or higher. Similarly, mortality of L. paeta after 3 d of exposure at the same concentration ranged from 92 to 100%. L. bostrychophila was the least susceptible species, with mortality of < 60% during the third week after application at rates < or = 27.5 mg/m2. However, even for this species, mortality after 3 wk was 90% or higher at rates > 27.5 mg/m2. Complete mortality of all species occurred after 3 d exposure at the highest rate tested of 110 mg/m2. Thus, our results show that chlorfenapyr is effective against major psocid species at the application rates evaluated in this study.

Evaluation of potential attractants for Liposcelis bostrychophila (Psocoptera: Liposcelididae).

The psocid, Liposcelis bostrychophila Badonnel (Psocoptera: Liposcelididae), can cause significant damage to stored commodities, and its pest status in the United States has been increasing over the last decade. Because L. bostrychophila is difficult to control with conventional methods, it is critical to explore alternative approaches such as the use of attractants that can be incorporated into integrated pest management programs for monitoring psocids. The orientation response of several L. bostrychophila life stages (first and second instars, third and fourth instars, 0- to 7-d-old adults, 21- to 28-d-old adults, and adults of mixed ages) to a range of potential attractants (including whole and cracked grains, grain-based oils, wheat germ, brewer's yeast, and commercially available kairomone lures) was studied using a two-choice pitfall test to identify candidates for further development as lures in traps. Among the potential attractants evaluated, the strongest response by all stages of L. bostrychophila was to brewer's yeast. Other materials for which there was consistently a strong response were psocid diet, wheat germ, and wheat germ oil. These results show the potential for developing monitoring tools for integrated pest management programs for L. bostrychophila and other psocid species.

Effect of abiotic factors on initiation of red flour beetle (Coleoptera: Tenebrionidae) flight.

Traps baited with pheromones are used to monitor the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera-Tenebrionidae), populations in flour mills to aid in making pest management decisions, but the factors that influence T. castaneum movement are not fully understood. We investigated the impact of photoperiod, light intensity, temperature, and relative humidity on flight initiation. The percentage of adults initiating flight reached a maximum at 30 -35 degrees C, and then fell to zero at 22.5 and 45 degrees C. Only 2% of beetles flew in complete darkness, and the number of beetles initiating flight increased to 41% under 18 h of light and then decreased slightly to 37% under 24 h of light. Rates of flight initiation did not vary with light intensities from 1,784 to 4,356 lux or relative humidities from 25 to 85%. Thus, temperature and photoperiod are the main abiotic factors tested that impact flight initiation in T castaneum, which have broad ranges of temperatures and photoperiods over which they can fly. The current results should be useful in helping to interpret trap catches based on abiotic conditions during the trapping period, and the results should be useful in helping to understand T. castaneum movement outside grain storages and processing facilities and their potential to infest structures.

Biology and management of psocids infesting stored products.

Previously regarded as minor nuisance pests, psocids belonging to the genus Liposcelis now pose a major problem for the effective protection of stored products worldwide. Here we examine the apparent biological and operational reasons behind this phenomenon and why conventional pest management seems to be failing. We investigate what is known about the biology, behavior, and population dynamics of major pest species to ascertain their strengths, and perhaps find weaknesses, as a basis for a rational pest management strategy. We outline the contribution of molecular techniques to clarifying species identification and understanding genetic diversity. We discuss progress in sampling and trapping and our comprehension of spatial distribution of these pests as a foundation for developing management strategies. The effectiveness of various chemical treatments and the availability and potential of nonchemical control methods are critically examined. Finally, we identify research gaps and suggest future directions for research.

Transcript analysis and expression profiling of three heat shock protein 70 genes in the ectoparasitoid Habrobracon hebetor (Hymenoptera: Braconidae).

Heat shock proteins (HSPs) are known as chaperones that help with folding of other proteins when cells are under environmental stresses. The upregulation of HSPs is essential for cold survival during insect diapause. The ectoparasitoid Habrobracon hebetor, a potential biological control agent, can enter reproductive diapause when reared at low temperature and short photoperiod. However, the expression of HSPs during diapause of H. hebetor has not been studied. In this study, we sequenced and characterized the full-length complementary DNAs of three Hsp70 genes (HhHsp70I, HhHsp70II and HhHsp70III) from H. hebetor. Their deduced amino acid sequences showed more than 80% identities to their counterparts from other insect species. However, the multiple sequence alignment among the three deduced amino acid sequences of HhHsp70s showed only 46% identities. A phylogenetic analysis of the three HhHsp70s and all other known Hsp70 sequences from Hymenoptera clustered all the Hsp70s into four groups, and the three HhHsp70s were distributed into three different groups. Real-time quantitative polymerase chain reaction analysis showed that the expression of the three HhHsp70 genes in H. hebetor reared at different conditions was quite different. HhHsp70I showed higher relative expression when H. hebetor were reared at 27.5°C than at two lower temperatures (17.5°C and 20°C) regardless of the photoperiod, whereas HhHsp70II showed higher expression when H. hebetor were reared at 20°C and 10 : 14 L : D than when reared at 17.5°C and either 16 : 8 L : D or 10 : 14 L : D. In contrast, HhHSP70III was expressed at similar levels regardless of the rearing conditions. These results may suggest functional differences among the three HhHsp70 genes in H. hebetor.

Efficacy of a combination of beta-cyfluthrin and imidacloprid and beta-cyfluthrin alone for control of stored-product insects on concrete.

The insecticidal effect of Temprid, a formulation that contains beta-cyfluthrin and imidacloprid, was tested on concrete for control of seven stored-product insect species: the rusty grain beetle, Cryptolestes ferrugineus (Stephens); the sawtoothed grain beetle, Oryzaephilus surinamensis (L.); the red flour beetle, Tribolium castaneum (Herbst); the confused flour beetle, T. confusum Jacquelin du Val; the hide beetle, Dermestes maculatus (DeGeer); and the psocids Liposcelis bostrychophila Badonnel and L. paeta Pearman. Temprid, which contains 10.5% beta-cyfluthrin and 21% imidacloprid, was tested at the rate of 17.2 ml of formulation/4.1 liters of water/100 m2. Adults were exposed for 4, 8, 14, and 24 h, and then daily for 7 d, on untreated dishes or dishes treated with Temprid. In the untreated dishes, mortality of C. ferrugineus and O. surinamensis was lower when food was present, but food did not affect mortality of the other species. Presence of food did not affect mortality of any of the species tested in the treated dishes. C. ferrugineus, O. surinamensis, and the two psocid species were very susceptible to Temprid, with mortality of 97-100% after 7 d of exposure. In contrast, D. maculatus, T. castaneum, and T. confusum were tolerant to Temprid, as mortality did not exceed 57, 25, and 17%, respectively, at the 7-d exposure. A separate series of similar bioassays with Tempo, a formulation that contains 11.8% beta-cyfluthrin alone, which was applied at the same dose rate as Temprid, was conducted using O. surinamensis and T. castaneum as the target insect species, and results showed that Tempo was at least as effective as Temprid. Our results indicate that the simultaneous use of beta-cyfluthrin with imidacloprid is not more effective on concrete than beta-cyfluthrin alone, and efficacy of both formulations varies with the target species.

Lesser grain borers, Rhyzopertha dominica, select rough rice kernels with cracked hulls for reproduction.

Tests were conducted to determine whether the lesser grain borer, Rhyzopertha dominica (F.) (Coleoptera: Bostrychidae), selects rough rice (Oryza sativa L. (Poales: Poaceae)) kernels with cracked hulls for reproduction when these kernels are mixed with intact kernels. Differing amounts of kernels with cracked hulls (0, 5, 10, and 20%) of the varieties Francis and Wells were mixed with intact kernels, and the number of adult progeny emerging from intact kernels and from kernels with cracked hulls was determined. The Wells variety had been previously classified as tolerant to R. dominica, while the Francis variety was classified as moderately susceptible. Few F 1 progeny were produced in Wells regardless of the percentage of kernels with cracked hulls, few of the kernels with cracked hulls had emergence holes, and little firass was produced from feeding damage. At 10 and 20% kernels with cracked hulls, the progeny production, number of emergence holes in kernels with cracked hulls, and the amount of firass was greater in Francis than in Wells. The proportion of progeny emerging from kernels with cracked hulls increased as the proportion of kernels with cracked hulls increased. The results indicate that R. dominica select kernels with cracked hulls for reproduction.

Induction of reproductive diapause in Habrobracon hebetor (Hymenoptera: Braconidae) when reared at different photoperiods at low temperatures.

Development of the parasitoid Habrobracon hebetor (Say) (Hymenoptera: Braconidae) at low temperatures was determined to identify rearing conditions that might result in adults that were in reproductive diapause. Diapausing adults would be expected to survive cold storage longer than nondiapausing adults for use in biological control programs. Only a few eggs were found in the ovaries when H. hebetor females were reared during the immature stages at 17.5 and 20°C with a 16-h photoperiod, and the ovaries were poorly developed and contained no eggs when females were reared with a 10-h photoperiod in these low temperatures. Rearing H. hebetor at 17.5 and 20°C did not result in diapause of immature stages, but did appear to result in possible adult reproductive diapause when the immature stages were reared with a 10-h photoperiod. Females reared during the immature stages at 17.5°C with a10-h photoperiod lived longer and took longer to lay their first eggs and to lay 50% of their eggs than those females reared at 17.5°C with a16-h photoperiod. Females reared during the immature stages at 20°C with a10-h photoperiod took longer to lay their first eggs and to lay 50% of their eggs, and they had a lower respiration rate, than those females reared at 20°C with a16-h photoperiod. Females that were reared in conditions that appeared to induce reproductive diapause resumed oviposition and their respiration rate increased soon after being transferred to a higher temperature (27.5°C). Thus, females reared at a 10-h photoperiod at 17.5 and 20°C appear to enter reproductive diapause.

Effectiveness of sulfuryl fluoride for control of different life stages of stored-product psocids (Psocoptera).

With the phase-out and impending ban of methyl bromide, sulfuryl fluoride is among the most promising alternative fumigant insecticides for control of stored-product insect pests. It has been evaluated for control of several stored-product insect pests, but there are few data available on its efficacy for control of stored-product psocids (Psocoptera). We evaluated sulfuryl fluoride for control of different life stages of the psocids Liposcelis paeta Pearman, L. entomophila (Enderlein), L. bostrychophila Badonnel, L. decolor Pearman, and Lepinotus reticulatus Enderlein (Trogiidae) in 48-hr trials at 27.5 degrees C. Adults and nymphs were susceptible to sulfuryl fluoride. Complete (100%) adult and nymphal mortality was recorded at concentrations between 4 and 8 g/m3, except for L. decolor for which all adults were only killed at 24 g/m3. Eggs were tolerant to sulfuryl fluoride. Complete egg mortality was achieved at 24 and 72 g/m3 for L. reticulatus and L. decolor, respectively. Survival of L. paeta eggs was recorded even after exposure to 96 g/m3. Given that the highest United States label concentration for sulfuryl fluoride for a 48-h exposure interval is 31.25 g/m3, our study indicates that high doses and/or longer exposures are needed for complete mortality of eggs of L. decolor and L. paeta. Moreover, the present work suggests that there is considerable variation in efficacy of sulfuryl fluoride for control of different psocid species.

Susceptibility of stored-product psocids to aerosol insecticides.

The efficacies of commercial methoprene and esfenvalerate aerosols for control of stored-product psocid pests were evaluated in simulated field studies. The efficacies of methoprene, esfenvalerate EC, the carrier Isopar-M™, and a combination of methoprene and esfenvalerate aerosols for control of Liposcelis decolor (Pearman) (Psocoptera: Liposcelididae) and Liposcelis entomophila (Enderlein) nymphs were assessed, and the effects of direct and indirect exposure of Liposcelis bostrychophila Badonnel, L. decolor, and Liposcelis paeta Pearman adults to esfenvalerate EC aerosol were evaluated. The greatest nymphal mortality attained was 76%, indicating that the four aerosols tested were ineffective against L. decolor and L. entomophila nymphs. In the direct and indirect exposure studies, the greatest adult mortalities attained for the three psocid species were 62 and 32%, respectively. Based on these data, esfenvalerate aerosol is ineffective for control of L. bostrychophila, L. decolor, L. entomophila, and L. paeta psocid species. This study shows that methoprene, esfenvalerate EC, and a combination of methoprene and esfenvalerate aerosols were ineffective against the four psocid species tested when applied at rates that are usually effective against other stored-product insect pests.

Efficacy of pyriproxyfen for control of stored-product psocids (Psocoptera) on concrete surfaces.

The insect growth regulator pyriproxyfen was evaluated as a surface treatment for control of three stored-product psocid pests Liposcelis bostrychophila Badonnel, Liposcelis decolor (Pearman), and Liposcelis paeta Pearman (Psocoptera: Liposcelididae). Nymphs were exposed for 35 d on a concrete surface treated with 2.3 mg of active ingredient/m2 pyriproxyfen. Exposure to pyriproxyfen significantly reduced the numbers of both adults and nymphs in comparison with untreated controls. In adults, the greatest reduction (> 90%) was for L. decolor and L. bostrychophila, whereas for L. paeta it was 49%. Few adults of any species were found in the pyriproxyfen treatments. The greatest numbers of nymphs were recorded for L. bostrychophila for both pyriproxyfen treatments and controls. Few adults of any species were found in the pyriproxyfen treatments. The results indicate that pyriproxyfen is effective for control of L. bostrychophila, L. decolor, and L. paeta on concrete, and although complete control was not achieved, the results warrant further long-term study to determine whether pyriproxyfen can completely eliminate psocid populations over time.

Effects of rearing density, age, sex, and food deprivation on flight initiation of the red flour beetle (Coleoptera: Tenebrionidae).

Effects of rearing density, adult density and sex ratio in the flight chamber, adult age, sex, presence or absence of food, and duration of food deprivation on rate of and time to flight initiation of the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), were studied in the laboratory. Rates of flight initiation were slightly lower at lower rearing densities, but they did not differ with age or sex of adults, presence or absence of food in the flight chamber, or duration of food deprivation. Focal adults were less likely to fly when individuals of the opposite sex were present in flight chambers. Presence of the same sex or mixed sexes and numbers of individuals in flight chambers did not affect tendency to fly of focal individuals. Mean time to flight of older beetles (7-20 d old) was less than that of younger beetles (1-4 d old). No young beetles flew during the first 24 h of flight tests. Mean time to flight did not differ with rearing density, sex, presence or absence of food, or duration of food deprivation. Our results indicate that this species is highly dispersive during the adult stage and that flight does not seem to be associated with just prereproductive or postreproductive dispersal phases, high rearing density, or short to medium periods of food deprivation. Therefore, T. castaneum level of flight activity does not seem to be associated with factors that have been shown in the literature to increase dispersal by walking for this species and to increase flight initiation in other stored-product species.

Efficacy of layer treatment with methoprene for control of Rhyzopertha dominica (Coleoptera: Bostrychidae) on wheat, rice and maize.

BACKGROUND: Insect growth regulators are promising alternatives to traditional pesticides in stored grain. The efficacy of the juvenile hormone analogue methoprene was evaluated as a layer treatment in a laboratory experiment for control of Rhyzopertha dominica (F.) (Coleoptera: Bostrychidae) in wheat, rice and maize. RESULTS: Adults of R. dominica were placed in vials containing 33, 26 and 29 g (to a depth of 6.5 cm) of wheat, rice and maize, respectively, that was entirely or partially treated with 1, 5 or 10 mg kg(-1) methoprene. In wheat and rice, the layer treatments were not as effective as the whole-grain treatment, but there was decreased progeny production as the application rate increased. However, on maize the partial treatments were as effective as the whole-grain treatment at 5 and 10 mg kg(-1) . CONCLUSIONS: The results suggest that partial layer treatments with methoprene can be used to control R. dominica on maize but may not be effective for control of this species on wheat and rice.

A computer model for simulating population development of the Indianmeal moth (Lepidoptera: Pyralidae) in stored corn.

The Indianmeal moth, Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae), is a common pest of stored corn, Zea mays L. We developed a computer model to simulate population development of the Indianmeal moth in stored corn by using previously published data describing immature developmental times and survivorship, and adult longevity and fecundity. The model accurately simulated population development of Indianmeal moths in corn stored during fall and into winter of three separate storage seasons in South Carolina. This is the period when the Indianmeal moth is a pest in stored corn in South Carolina. The model predicted that populations would increase after winter as grain temperatures rose, but observed populations in the grain bins never increased after winter. Despite this, the model should be useful from a management perspective because the corn is being sold off or used up after winter, and the observed Indianmeal moth populations never reached damaging levels after winter.