Determining the impact of scale insect honeydew, and invasive wasps and rodents, on the decomposer subsystem in a New Zealand beech forest

Relatively few studies have considered how aboveground invasive consumers influence decomposer communities. We investigated the potential effects of three types of animals on the decomposer subsystem in a floristically simple New Zealand Nothofagus forest. These animals are the native beech honeydew scale insect (Ultracoelostoma spp.) that secretes large amounts of sugar-rich honeydew that washes to the soil, invasive social wasps (Vespula spp.) that remove honeydew and prevent it from reaching the ground, and invasive rodents (the house mouse (Mus musculus) and ship rat (Rattus rattus)) that are predators of litter invertebrates. We performed a 4 years manipulative experiment involving addition of synthetic honeydew to the soil surface at amounts equal to that washed to the soil both in the absence and presence of wasps. All treatments were subjected to both exclusion and non-exclusion of rodents. Full honeydew addition influenced several components of the belowground community (both positively and negatively), and promoted fungi and fungal feeding fauna at the expense of bacteria and bacterial-feeders. The reduced addition of honeydew (representing effects of wasps) reversed some (but not all) effects of full honeydew addition. Rodents also influenced some belowground organisms, often reversing the effects of honeydew addition. The honeydew levels simulating wasp effects and the presence of rodents both greatly promoted humus carbon and nutrient storage relative to all other treatments, highlighting that invaders can alter soil carbon sequestration and nutrient capital. Our study points to invasive animals modifying the effects of a native animal on multiple components of the decomposer subsystem.     

Ecological effects of multi‐species,ant–hemipteran mutualisms in citrus

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Honeydew collection by the invasive garden ant Lasius neglectus versus the native ant L. grandis

Honeydew collection performed by the invasive ant Lasius neglectus and by the native ant L. grandis was compared. The invasive ant collected 2.09 kg of honeydew per tree while the native ant collected 0.82 kg. The aphid Lachnus roboris was visited by both ant species. In holm oaks colonized by L. neglectus, aphid abundance tended to increase and its honeydew production increased twofold. The percentage of untended aphids was lower in holm trees occupied by L. neglectus. As tending ants also prey on insects, we estimated the percentage of carried insects. The native ant workers carried more insects than the invasive ant. Both ant species preyed mainly on Psocoptera and the rarely tended aphid, Hoplocallis picta. We conclude that the higher honeydew collection achieved by L. neglectus was the consequence of (1) its greater abundance, which enabled this ant to tend more Lachnus roboris and (2) its greater level of attention towards promoting an increase of honeydew production. Handling editor: Heikki Hokkanen     

Local dynamics of worker activity of the invasive Vespula germanica and V. vulgaris (Hymenoptera: Vespidae) wasps in Argentina

1. The abundance of insects depends essentially on the reproductive success of individuals. In social insects, however, the abundance of sterile workers outside a nest depends on colony size but is also determined by ontogeny, nest demands, and local environmental factors. For invasive social wasps, the drivers of worker abundance are important because they determine the impact that these species have on the native systems, people, and their goods. 2. The aim of the present study was to understand the relative importance of endogenous and exogenous factors on the abundance of workers of populations of Vespula spp., by analysing 12 years of trap captures in NW Patagonia. This is the first attempt to model the activity levels of invasive Vespula spp. wasps over time in Argentina. 3. It was shown that between years, the worker activity of both vespids presents fluctuations, and that of V. germanica is determined by the spring mean temperatures. Within the flight season, V. germanica worker activity is affected by the relative abundance of workers in that year, whereas for V. vulgaris, activity it is affected by the relative abundance of both species that year. We found no relationship between individual weather variables and activity within a flight season for both wasps. 4. The patterns observed for Argentina are similar to those observed in all invaded temperate areas where Vespula spp. are established. This study provides useful information to understand the driving factors that affect Vespula spp. worker activity in Argentina. This could be a necessary step to develop plans to manage these invasive social insects.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Volatiles from green‐lipped mussel as a lead to vespid wasp attractants

Vespid wasps (Vespula vulgaris L. and V. germanica Fab. Hymenoptera; Vespidae) are highly abundant in 1 million ha of New Zealand's indigenous beech forests (Nothofagus spp.) and have had detrimental effects on the New Zealand native fauna. This hyperabundance is due in part to the vast supply of carbohydrate‐rich honeydew produced by scale insects Ultracoelostoma spp. native to New Zealand. Current control methods include the use of wet cat food as a protein source with insecticide as a lure‐and‐kill‐based system, but there are problems with fresh baits degrading rapidly, and a more durable formulation would enable the expansion and longevity of wasp control. Four crude protein baits were tested for vespid attraction. Green‐lipped mussels had the highest vespid catch of the crude baits tested, and aged and fresh mussels were equally attractive. From headspace analysis of the green‐lipped mussel volatiles, a series of butanoate esters, 3‐octanone and 1‐octen‐3‐ol were identified as possible attractants. These compounds were tested individually and in various blend combinations for the attraction of Vespula wasps in matagouri vegetation at the edge of beech forests. We found synergistic effects between single attractive compounds when tested in various combinations, and the multicomponent lures were more attractive to these wasps than heptyl and octyl butanoate, previously identified attractants for vespid species. The new multicomponent lures could form the basis for a new generation of attractants for social wasps that can provide sustained control methods for invasive vespid wasps.     

Invasional 'meltdown' on an oceanic island

Islands can serve as model systems for understanding how biological invasions affect community structure and ecosystem function. Here we show invasion by the alien crazy ant Anoplolepis gracilipes causes a rapid, catastrophic shift in the rain forest ecosystem of a tropical oceanic island, affecting at least three trophic levels. In invaded areas, crazy ants extirpate the red land crab, the dominant endemic consumer on the forest floor. In doing so, crazy ants indirectly release seedling recruitment, enhance species richness of seedlings, and slow litter breakdown. In the forest canopy, new associations between this invasive ant and honeydew‐secreting scale insects accelerate and diversify impacts. Sustained high densities of foraging ants on canopy trees result in high population densities of host‐generalist scale insects and growth of sooty moulds, leading to canopy dieback and even deaths of canopy trees. The indirect fallout from the displacement of a native ‘keystone’ species by an ant invader, itself abetted by introduced/cryptogenic mutualists, produces synergism in impacts to precipitate invasional ‘meltdown’ in this system.     

Foraging activity and dietary spectrum of wood ants (Formica rufa group) and their role in nutrient fluxes in boreal forests

Abstract 1. We monitored three different‐sized wood ant (Formica aquilonia Yarrow) mounds over a 3‐year period in Finnish boreal forests dominated by Norway spruce (Picea abies Karst.), to assess the seasonal temperature dependency of ant activity. Additionally, we also monitored Norway spruce trees around the mounds for descending honeydew foragers. 2. The amount of collected honeydew and prey and its composition, as well as the carbon (C), nitrogen (N), and phosphorus (P) in honeydew and invertebrate prey was also investigated. 3. The number of warm days (average temperature above 20 °C) and the amount of precipitation differed among the years. Ant activity at the mounds (but not on the trees) was highly correlated with air temperature throughout the ant‐active season (May–September), but ant activity in spring and autumn was lower than in summer at similar temperatures. During all 3 years, honeydew played a major role in wood ant nutrition (78–92% of dry mass). Invertebrate prey was mainly Diptera (on average 26.2%), Coleoptera (12.5%), Aphidina (9.3%), and Arachnoida (8.5%). 4. The total amounts of C, N, and P input brought into the ant mounds in the form of food (both honeydew and prey) on the stand level were 12.6–39.0, 1.6–4.6 and 0.1–0.4 kg ha^?1 year^?1, respectively, which is equivalent to 2–6%, 12–33% and 27–58% of the fluxes in annual needle litterfall in typical boreal Norway spruce forests. Thus, wood ants can play a significant role in short term and local N and P cycling of boreal forest ecosystems.     

Competition for honeydew between two social wasps in South Island beech forests,New Zealand

Summary Honeydew is a sugary secretion of beech scale insects (Ultracoelostoma spp.). Two introduced species of wasps forage on the sugar droplets in New Zealand beech forests. We hypothesize that competition between them may explain whyVespula germanica has become locally extinct in beech (Nothofagus) forest invaded byV. vulgaris. Changes in behaviour of the two wasp species in response to changes in the honeydew resource were monitored. Foraging and rainfall separately and together reduced the honeydew standing crop. In response to the standing crop decreasing, more wasps were found on honeydew trees, they became less active, spent more time lapping the tree surface, and ingested drops of honeydew at a slower rate.V. vulgaris was more active, and obtained drops and energy at a higher rate thanV. germanica. These behavioural differences may lead to competitive advantages affecting queen size and possibly survival.     

Attraction of the invasive social wasp,Vespula vulgaris,by volatiles from fermented brown sugar

The introduction of invasive social wasp species of the genus Vespula (Hymenoptera: Vespidae) to New Zealand has caused a major ecological problem, particularly in the beech forests (Nothofagus spp.) of the South Island, where they have destabilized the native bird and invertebrate biodiversity. New attractants are under investigation as part of a search for pest management solutions. Fermenting brown sugar has been previously reported as a social wasp attractant. This work was undertaken to identify compounds from fermented brown sugar attractive to social wasps. Raw fermented brown sugar was confirmed to be attractive in a field trial and 10 chemical compounds present in the headspace were positively identified by coupled gas chromatography‐mass spectrometry and synthetic references. During electroantennogram experiments, 3‐methyl‐1‐butanol, 3‐methylbutyl acetate, and ethyl hexanoate elicited high electrophysiological responses from Vespula vulgaris (L.) antennae. These compounds mediated attraction of V. vulgaris wasps in forest margins by trapping. A blend of these compounds could be used as a lure in a monitoring tool, or even a local suppression method if combined with a toxin.     

Comparing floral nectar and aphid honeydew diets on the longevity and nutrient levels of a parasitoid wasp

We compared the effects of floral nectar from buckwheat, Fagopyrum esculentum Moench, and honeydew produced by the soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae), on longevity, nutrient levels, and egg loads of the parasitoid Diadegma insulare Cresson (Hymenoptera: Ichneumonidae). Diadegma insulare lived for 2 days in control treatments of water or clean soybean leaves, for 6–7 days with honeydew, and in excess of 2 weeks with buckwheat nectar. Potential reasons for the superiority of buckwheat nectar over soybean aphid honeydew for extending the longevity of parasitoids include: (i) parasitoids ingest more sugars from floral sources, (ii) oligosaccharides in honeydew have a lower nutritional value than nectar sugars, and (iii) honeydew has antagonistic compounds. Overall sugar levels were lower in honeydew‐ vs. nectar‐fed female wasps, suggesting a lower feeding rate, but other explanations cannot be excluded. Diadegma insulare eclosed with high levels of lipids and glycogen, and low levels of gut and storage sugars. All carbohydrates increased over the life of both nectar‐ and honeydew‐fed wasps, but remained low or decreased in starved wasps. Lipid levels declined over the lifespan of female wasps, but females fed floral nectar showed the slowest rate of lipid decline. Diet did not affect egg load, probably because the females were not given hosts in the experiment.     

Soluble carbon production by honeydew scale insects in a New Zealand beech forest

We estimated the annual production of honeydew per unit land area of beech (Nothofagus spp.) forest by measuring the amount of honeydew produced in 24 h by scale insects (Ultracoelostoma spp.) (Hemiptera: Margarodidae) every month for 2 years. We used exclosures to prevent animals (notably Vespula wasps) removing honeydew, and we compared the standing crop of honeydew inside permanently closed exclosures with that outside exclosures. Honeydew production and the number of honeydew droplets was highly variable between individual trees, tree type, position on tree, and, exclosure type, and within and between years. The amount of honeydew available outside exclosures was significantly reduced in year 2, predominantly by Vespula wasps, even though wasp density was relatively low. Sugar composition also varied between tree type and between years. Up to 5% of the sugar was glucose, with varying proportions of fructose, sucrose and oligosaccharides. The surface area of trees infested with scale insects was estimated using allometric regression relationships between tree diameter and total surface area of tree trunk and branch material. These estimates were combined with measurements of tree diameter in 10-m radius circular plots to give a production estimate of between 3500 and 4500 kg dry weight honeydew ha^-1 year^-1. Using this data, combined with previously published estimates of carbon uptake, it was estimated that between 6 and 8% of net primary productivity was released as honeydew. Honeydew scale insects provide large amounts of biologically available carbon in the form of soluble sugar. It is a crucial resource for the above-ground system, and probably also for the below-ground system. We conclude that scale insects have the potential to function as keystone species in these forests.     

Of volcanoes and insects: the impact of the Puyehue–Cordon Caulle ash fall on populations of invasive social wasps, Vespula spp.

Volcanic eruptions have important effects on terrestrial ecosystems. The biotic effects of volcanic ash on insect populations vary from widespread and catastrophic to subtle and localized. Volcanic eruptions may increase insect mortality through the effects of the ash plume, and these vary by taxon or specific biological features. The Puyehue–Cordon Caulle Volcanic Complex is the most recent eruption in Patagonia. Here we explore and describe the effects of the eruption of this volcanic complex on invasive Vespula spp. populations. These wasps are very abundant social hymenoptera that have recently invaded Patagonia. We placed baited traps following the gradient of deposited ash, and revisited them once a week during February to April. No wasps were caught where ash deposition levels exceeded 3.0 cm. The number of wasps caught in sites with minimal ash deposits was similar to that of previous years. In locations where ash layers are intermediate, a few wasps were observed, but only during the peak of wasp abundance. We show that Vespula spp. populations were severely affected by the ash deposition and plume, caused by this eruption. These results show catastrophic effects of ash deposition over a large area, even at great distances from the eruption epicenter. We conclude that wasp mortality explained by the volcanic eruption may reach distant areas, and cause local extinction. In areas where active volcanoes are common, invasive insects may encounter an additional obstacle in their invasion success.     

Getting carbohydrates in ants: damage of the young cereal sprouts by ants of the genus Myrmica Latreille, 1804 (Hymenoptera: Formicidae), to obtain plant sap

Summary

Carbohydrate food is of high importance for survival of ant colonies. Ants are known to use sugary excretions (honeydew) of various insects, nectar of floral and extrafloral nectaries, and even sap of some trees. However, the ability of ants to use sap of herbaceous plants has not been mentioned. This is the first evidence that ants of the genus Myrmica can intentionally ‘cut off’ young cereal sprouts to obtain plant sap. The investigation was carried out in a laboratory in 2018 and 2019 and involved three ant species of the genus Myrmica [12 colonies of M. rubra (Linnaeus, 1758); eight colonies of M. ruginodis Nylander, 1846; and five colonies of M. scabrinodis Nylander, 1846]. First observations were made occasionally in 10 ant colonies during the study of ant–aphid interactions. After three days of carbohydrate starvation, ants were supplied with the plants of wheat infested with aphids of Schizaphis graminum (Rondani, 1852). Within the first day in addition to ordinary trophobiotic relations with aphids, the workers of all the studied colonies demonstrated unexpected behaviour: they ‘cut off’ some sprouts and collected sap of these plants. The experimental investigation in 15 ant colonies of various sizes (about 150, 300 and 500 workers) supplied with the plants infested or non-infested with aphids has shown that getting sap of herbs depends greatly on ant colony needs and available resources. The number of damaged plants was much higher both in the larger colonies of ants and in the absence of aphids. This way of getting carbohydrates allows ants to quickly obtain some extra food needed to maintain colony viability and seems to be one of the mechanisms promoting survival of ants in conditions of acute carbohydrate deficiency. At the same time, ants avoid using plant sap when there are more available alternative carbohydrate resources.     

Do invasive ants respond more strongly to carbohydrate availability than co‐occurring non‐invasive ants? A test along an active Anoplolepis gracilipes invasion front

Invasions by non‐native insects can have important ecological impacts, particularly on island ecosystems. However, the factors that promote the success of invaders relative to co‐occurring non‐invasive species remain unresolved. For invasive ants, access to carbohydrate resources via interactions with both extrafloral nectary‐bearing plants and honeydew‐excreting insects may accelerate the invasion process. A first step towards testing this hypothesis is to determine whether invasive ants respond to variation in the availability of carbohydrate resources, and whether this response differs from that of co‐occurring, non‐invasive ants. We investigated the effect of carbohydrate subsidies on the short‐term foraging and hemipteran‐tending behaviours of the invasive ant Anoplolepis gracilipes (Formicidae) and co‐occurring ant species on an extrafloral nectary‐bearing plant by experimentally manipulating carbohydrate levels and tracking ant recruitment. We conducted experiments in 2 years at two sites: one site was invaded by A. gracilipes prior to 2007 and the other became invaded during the course of our study, allowing pre‐ (2007) and post‐invasion (2009) comparisons. Short‐term increases in carbohydrate availability increased the density of A. gracilipes workers on plants by as much as 400% and reduced tending of honeydew‐excreting insects by this species by up to 89%, with similar responses across years. In contrast, ants at the uninvaded site in 2007 showed a weak and non‐significant forager recruitment response. Across all sites, A. gracilipes workers were the only ants that responded to carbohydrate manipulations in 2009. Furthermore, ant–carbohydrate dynamics at a site newly invaded by A. gracilipes quickly diverged from dynamics at uninvaded sites and converged on those of the site with an established invasion. These findings suggest that carbohydrate resources may be particularly important for A. gracilipes invasions, and underscore the importance of species interactions, particularly putative mutualisms, in facilitating exotic species invasions.     

Ants impact the energy reserves of natural enemies through the shared honeydew exploitation

1. Ants, as well as many species of parasitoids and predators, rely on sugar‐rich foods such as honeydew to fulfill their energetic needs. Thus, ants and natural enemies may interact through the shared honeydew exploitation. 2. Ant‐exclusion experiments were performed in a citrus orchard to test the hypothesis that ants may impact the energy reserves of predators and parasitoids through the competition for honeydew. Through the use of high‐performance liquid chromatography (HPLC) the level of ant activity with the energy reserves and feeding history of individual specimens collected in the field during representative days of spring, summer, and autumn were related. 3. Out of 145 Aphytis chrysomphali Mercet parasitoids captured in the field, 65% were classified as sugar‐fed and 24.7% as honeydew‐fed. In summer, when ant activity peaked, there was a significant negative correlation between the level of ant activity and the total sugar content and honeydew feeding incidence by A. chrysomphali. Out of 47 individuals of the predator Chrysoperla carnea sensu lato (Stephens), captured in the field, 55.3% were classified as sugar‐fed. We found a significant negative effect of the level of ant activity on the sugar‐feeding incidence by C. carnea in spring. 4. The present study provides evidence that ants can interfere with the energy reserves of natural enemies. This interaction may be widespread in various ecosystems with important consequences for the arthropod community composition and with practical implications for biological control given that absence of sugar feeding is detrimental for the fitness of many species of predators and parasitoids.     

Pollinator effectiveness and fruit set in common ivy, Hedera helix (Araliaceae)

Flowers of common ivy (Hedera helix L.) provide late season pollen and nectar for several insect groups, and its fruits are a winter and spring food source for frugivorous birds. Ivy benefits from insect pollination in order to set fruit, but it is unknown which flower-visiting insects are the most effective pollinators. Our observations suggest that Vespula wasps are potentially the most effective pollinators since they were frequent visitors, had relatively fast foraging rates, carried large numbers of pollen grains on their bodies and had the highest ‘Pollination potential PP index’ score (a measure of pollinator effectiveness) of all the insect groups examined. There was also a positive linear relationship between the proportion of ivy flowers that set fruit and wasp foraging activity in 0.5 m² quadrats. Visits by Vespula wasps may therefore be important for ensuring a supply of ivy fruits for birds.     

Aphid–ant mutualism: how honeydew sugars influence the behaviour of ant scouts

Honeydew is the keystone on which ant–aphid mutualism is built. The present study investigates how each sugar identified in Aphis fabae Scopoli honeydew acts upon the feeding and the laying of a recruitment trail by scouts of the aphid‐tending ant Lasius niger Linnaeus, and thus may enhance collective exploitation by the ant mutualists. The feeding preferences shown by L. niger for honeydew sugars are: melezitose = sucrose = raffinose > glucose = fructose > maltose = trehalose = melibiose = xylose. Although feeding is a prerequisite to the launching of trail recruitment, the reverse is not necessarily true: not all ingested sugar solutions elicit a trail‐laying behaviour among fed scouts. Trail mark laying is only triggered by raffinose, sucrose or melezitose, with the latter sugar being specific to honeydew. By comparing gustatory and recruitment responses of ant foragers to sugar food sources, the present study clarifies the role of honeydew composition both as a source of energy and as a mediator in ant–aphid interactions. Lasius niger feeding preferences can be related to the physiological suitability of each sugar (i.e. their detection by gustatory receptors as well as their ability to be digested and converted into energy). Regarding recruitment, the aphid‐synthesized oligosaccharide (melezitose) could be used by ant scouts as a cue indicative of a long‐lasting productive resource that is worthy of collective exploitation and defence against competitors or aphid predators.     

Feeding habits of Hymenoptera and Isoptera in a tropical rain forest as revealed by nitrogen and carbon isotope ratios

Despite the recognition of the functional role of Hymenoptera (ants, bees and wasps) and Isoptera (termites) in tropical ecosystems, their detailed feeding habits are not well known. To examine the feeding habits of these groups, we measured nitrogen (N) and carbon (C) stable isotope ratios (δ¹⁵N and δ¹³C) of hymenopterans (12 families, ≥16 genera and ≥32 species) and isopterans (one family and 10 species) collected in a tropical rain forest, Sarawak, Malaysia. We compared the isotopic signatures of these insects to those previously reported for other consumers collected in the same forest. The δ¹⁵N and δ¹³C values of these insects overlapped with those of the other consumers, indicating that they have access to diverse C and N sources in the forest. The δ¹⁵N values of ants and termites indicated that their feeding habits range along a continuum from herbivory (i.e. dependent on honeydew and nectar) to predation and from wood-feeders to soil-feeders, respectively. In addition, the δ¹⁵N values of wasps varied greatly from −0.1‰ (Braconidae sp.) to 8.6‰ (Bembix sp.), suggesting that their feeding habits also range from omnivory to predation. The ant species Camponotus gigas had δ¹³C values similar to those of invertebrate detritivores and omnivores rather than to those of invertebrate herbivores, although the diet of this species consists mostly of honeydew. This discrepancy suggests that the ant uses carbohydrates as an energy source, the isotopic signatures of which are not well retained in the body tissues. Values of both δ¹⁵N and δ¹³C of the predatory army ant Leptogenys diminuta and the soil-feeding termite Dicuspiditermes nemorosus did not differ significantly, indicating that both trophic level and the humification of feeding substrates can increase the isotopic signatures of terrestrial consumers.     

Spatial distribution of ants (Hymenoptera: Formicidae), vine mealybugs and mealybug parasitoids in vineyards

The mutualistic association between some ant species and honeydew‐producing Hemiptera has been shown to influence the distribution patterns and abundance of these hemipterans and their natural enemies. We studied the spatial distribution patterns of three ant species, mealybugs and mealybug parasitoids for two consecutive growing seasons on three wine grape farms in the Western Cape, South Africa. During the study period, no ant or mealybug controls were applied. Ant and mealybug monitoring was conducted on a total of 21 ha using a presence/absence sampling system, while parasitoids were collected from infested mealybug females. Spatial analysis by distance indices was used to analyse spatial distribution of insects and ArcView? was used to map the gap, patch and local association indices where significant association and disassociation occurred. Significant associations were found between some ants and parasitoids, while significant disassociations between the ants Crematogaster peringueyi and Linepithema humile; and also between Crematogaster peringueyi and Anoplolepis steingroeveri were found. Interspecific competition between ant species could play a role in the distribution of parasitoids and mealybugs. Our results stress the importance of monitoring for ants and mealybugs and further highlight the importance of restricted chemical applications against ants during the growing season.