Ants in Orchards and Vineyards in the San Joaquin Valley By Devin Carroll 2014 Contents Introduction: The role of ants in SJV agriculture. - 1 List of ants found. - 4 Ant anatomy and vocabulary. - 5 SJV agricultural ant descriptions and biology, with photos. - 6 Field guide to the common species. - 21 Microscope key to the species. - 23 Bibliography. - 25 The role of ants in SJV agriculture. As any observer can attest, ants are among the most common insects in agricultural systems. They play a major and complex role in agricultural ecology, plant and soil health, and pest management. This project reflects my personal experience along with supporting information from the literature. It applies mainly to the areas, the crops, and the seasons where I work. These include grapes, tree fruit and nuts in agricultural sections of Madera County, most of Fresno County, and southern Tulare County around Delano and Earlimart. I am in the field mainly from late February to the end of September. Almond pests Ants in almonds get the most attention because two species can be very destructive of nuts on the ground between shaking and gathering16. Many ants will pick up broken pieces of almond, but only fire ants and pavements ants can chew holes in whole nuts. Scouts should take care to distinguish these two pest species from other common orchard ants such as field ants and pyramid ants. Due to the almond market, most agricultural ant baits are aimed at fire and pavement ants. They are “grease” ants, so typical baits consist of vegetable oils mixed with a carrier such as corn cob grits. These baits miss some of the other ant species. Soft tissue damage Most ants do not directly damage plants, but fire ants can be destructive of soft tissue. They can girdle young grape vines and citrus trees19. Many ants will feed on fruit that is cracked or damaged. This may actually help the farmer, because the ants may scrape out a wound before it can rot. Tenders of honeydew-producing homopterans Most ants love sugar and are attracted to honeydew-producing homopterans including aphids, soft scale, mealybugs, whiteflies, and some leafhoppers. The ants magnify infestations by chasing away or killing parasites and predators. They also move their “herd” within and between vines and trees. This makes them important indirect pests of most of our tree and vine crops. In my opinion, the most effective guards of homopterans in this area are Argentine ants, fire ants and field ants, in that order. In some cases, these ants can transform a minor pest into a major problem. Many other ant species seem to make little difference to biological control. Argentine ants are rare in SJV agriculture, so controlling fire and field ants would likely reduce our ant-induced problems to a minimum. On the good side, by cleaning up the honeydew the ants may reduce the damage from sooty mold growth. In our area, harvester ants are the only common species that do not tend homopterans. Predators Most ants will prey on almost any available arthropod that is not providing them with honeydew10. This includes some pests, some natural enemies, spiders, and other ants. The role of ants in pest management is complex as they may help control pests but may also interfere in biological control by other agents21. Most predaceous ants also consume sugar, and their numbers depend on sugar sources, particularly homopterans21. In almonds and peaches, field ants are key predators of peach twig borers, but not of oriental fruit moths, which have a spitting defense7. Their role in controlling other pests such as leafrollers, leafhoppers, katydids, and so on is not well documented but probably in some cases is substantial. Growers who successfully control their ants should be on the lookout for unexpected consequences. Soil builders Ants contribute positively to soil ecology. They are first responders to dead animals including vertebrates as well as arthropods, breaking down the bodies in a step towards converting it to soil. With their deep nests they mix upper and lower soil strata, provide aeration, and carry organic matter downwards from the surface10. Seed eating ants may drop seeds, spreading them to new places10. But they may also be so thorough as to denude patches of ground of almost all vegetation10. Their tastes might play a role in determining which weeds are more common. Ant wars Most ants fight fiercely with their neighboring colonies for territory and food10. Most often the violence is within the same species, but sometimes two species will fight6. Watching ant wars as a child in my urban yard was my first experience with entomology, and I still enjoy watching their gladiatorial combat. As a rule, advantage lies in numbers; many small ants will defeat a few large ants. The usual tactic is to grab legs or antennae and tear apart the enemy. A survivor may carry around the severed head of another ant, still clamped onto a limb. Ants around their home nests tend to be more aggressive towards other species, and intruders usually try to flee when confronted. Many ants use chemical warfare to repel other ant species10. Formicinae spray formic acid from the acidophore. Dolichoderinae use a variety of terpenoids produced by an anal gland and expelled through a slit opening at the tip of their abdomen. These chemicals produce the characteristic pungent odor of these ants when crushed. Most Myrmicinae have stingers with venom that increases their ability to injure and kill arthropods and cause pain in vertebrates. Chemicals from the venom may also be used to communicate within nests and repel foreign ants. Ant wars have interesting variants. Some species will make slaving raids to carry captives back to their own nests, or invade a nest to enslave the occupants in situ10. New colony formation Ants establish new colonies in a variety of ways depending on the species, but they may be categorized into “independent founding” in which new queens found the colonies without bringing workers, and ”swarming” or “budding” in which workers from the old colony accompany the queens10. The word “swarming” is sometimes misused to describe ant reproductives leaving the nest; the better term is “nuptial flight”. Worker rotation and sprays Ants practice an efficient rotation of work within their colonies10. The youngest workers stay deep inside, nursing juvenile stages and tending to the queens and drones. Older workers perform the more dangerous jobs outside, fighting other ants and collecting food in locations where they are exposed to predators. This insures that as older workers are lost they are replaced by others. Surface sprays that kill only the outside workers are thus only temporary6. New workers will soon appear. Colony elimination requires that the workers carry toxins back to the nest in bait or perhaps on their skin. Ants share their food with each other and with their larva (prophylaxis), so toxins spread through the colony. In many species worker ants are not capable of consuming solid food10. They feed solids to the larvae, which eat and digest it before regurgitating some back to the workers. Baits For economic reasons most baits are primarily aimed at either Argentine or fire ants. Argentines are chiefly urban pests, although they dominate in some coastal winegrowing regions in California. Their baits use sugar as attractant, usually in liquid or gel form. Typical toxins are boric acid or fipronil. The same baits are effective on many other sugar-loving household ant pests, but they are less practical in vineyards because of the difficulty of maintaining liquid bait stations. Many fire ant baits use a granular carrier soaked in vegetable oil. These baits also attract other grease lovers such as pavement ants, but they have little appeal to other species such as field ants. Sugar granules or protein baits attract both fire and field ants, but they are so far not commercially available. List of Ants North America has 10 subfamilies of ants, but all species I have found in the San Joaquin Valley belong to only three. They are differentiated at the start of the microscopic key. Those marked with asterisks are not included in this work because I have not seen them in agriculture or very rarely. Some are more common in urban settings. Some are included in Ingrid Carmean’s key to local urban ants3, and some are included in publications of the University of California by Haney et. al.9, Grape Pest Management 13, or at the UCIPM website14. Subfamily Dolichoderinae Dorymyrmex bicolor – pyramid ant Forelius pruinosus – no common name Linepithema humile – Argentine ant Tapinoma sessile - odorous House Ant * Dorymyrmex insanus – pyramid ant3 (=Conomyrma) * Liometopum occidentale – velvety tree ant9. (Oak, urban yards) Subfamily Formicinae Camponotus essigi– carpenter ant Formica aerata – California gray field ant Myrmecocystus mimicus – honeypot ant Nylanderia vividula – crazy ant Subfamily Myrmicinae Messor pergandei – Jet black harvester ant Monomorium minimum – little black ant Pogonomyrmex californicus – California harvester ant Tetramorium bicarinatum - Guinea Groove-Headed Ant (= tramp ant) Tetramorium caespitum – pavement ant Solenopsis invicta – red imported fire ant Solenopsis molesta – thief ant Solenopsis xyloni – southern fire ant * Cardiocondyla ectopia3 * Cardiocondyla mauritanica – (I found this only in my urban yard) * Monomorium pharaonis – Pharoah ant3,9 (lighter colored than M. minimum) * Pheidole californica13 – (I found 1 specimen of Pheidole on the floor of an apple orchard, another on a citrus tree.) Anatomy and Vocabulary (Right) Abdomen: Segment 1, the propodeum, is attached to the thorax. The pedicel may have either 1 or two bead-like segments, the petiole and postpetiole. The remaining abdomen is the gaster. (Tetramorium caespitum) (Below) Anal area of gaster: Dolichoderinae with slit (Dorymyrmex bicolor) Formicinae with acidophore (Formica aerata) Myrmicinae with stinger (Tetramorium caespitum) Antennae are elbowed, with a long first segment, the scape, and the remaining funiculus, which may end with a 2-3 segment expansion, the club. The clypeus is the area just above the mandibles. In addition to the compound eye, alate ants and some workers have 3 ocelli (not shown) at the top of the head. The vertical grooves are characteristic of pavement ants, Tetramorium caespitum. Ant Descriptions and Biology Pyramid ant Dorymyrmex bicolor Dolichoderinae One of our most common ants, pyramids can be confused with fire ants. They are about the same size, and have similar colors; red head and thorax with a dark abdomen. But pyramids have only a one-segmented pedicel, and the workers are all about the same size. The final test is smell; pyramid ants have a pungent odor when crushed. The typical mound is a well-formed donut-shape. When the sun is not too hot, pyramid ants run in and out of their Doughnut shaped pyramid ant mound with ant hole to add grains to the mound. Unlike “graveyard” of fire ants. fire ants, they do not rush out in mass when disturbed. Pyramids are aggressive towards any ants that pass near their nests. They compete fiercely with other species. They can block foraging of competing honeypot ants by surrounding their nest entrance and tossing in pebbles10. Many pyramid ant nests have an ant “graveyard” to the side. Usually the dead are mainly fire ants. In one case I saw mainly pavements ants, and occasionally a few harvester ants and field ants are present. Pyramids tend homopterans. I often see them on grapes with vine or grape mealybug, or European lecanium scale. They do not seem to protect mealybugs very well; I usually observe good biological control where they are present. Typically the nests are in the open outside the vineyard, with the ants following a trail in to a vine up to 7 or more yards away. These ants may also forage by scattering in all directions late in the afternoon. Pyramids are predaceous. I have seen them carrying leafhopper Worker showing the nymphs off of vines. Literature has them eating other ants and a 24 “pyramid” at the rear of variety of arthropods . the thorax. Pyramid ants will pick up bits of almond and carry them to their Photo by Alice Abela nest, but they do not chew holes in intact almonds, so they are not almond pests. Presumably their competition and aggression reduces the fire ant numbers in almond orchards. They should not be the targets of control efforts in almonds. I have seen nuptial flights from late February to mid-April. They might start earlier when I am not often in the field. Females are red and males are smaller and black. Forelius pruinosus Dolichoderinae This small ant has no common name. I have seen it in only a handful of locations. It is distinctive among our ants because the abdomen is yellowish. It has a pungent odor when crushed. The mound is doughnut shaped, similar to pyramid ant nests, but often in a cluster. There may even be ant graveyards with mainly fire ants. Forelius often forages in distinct lines like Argentine ants. The mounds are usually in the open but often near dry straw where they like to forage and pick up grass pieces. Forelius pruinosus worker. I do not have many feeding observations. They are reported to tend homopterans, but I have not seen them in grapes, and I have seen their trails bypass weeds with aphids. They also ignored fruit on the ground. They are predaceous; BugGuide.net pictures show them eating caterpillars and grasshoppers. I have seen them carry dead ants, and they are very aggressive towards other ant species. They can surround nests of competing species and keep the residents inside with repellent chemicals, and sometimes even fill the entrances with dirt10. Doughnut shaped Forelius pruinosus mound with grass seeds and ant “graveyard” of fire ants. Argentine ant Linepithema humile Dolichoderinae Argentine ants, invaders from South America, now dominate most California cities, including ours in SJV, but they are not found in most SJV vineyards and orchards. This differs from some Argentine ant worker. coastal grape districts, where Argentines also Photo by Joyce Gross dominate in vineyards13. Where present, they displace native and other introduced species12. Unlike with most ants, Argentine mated females do not fly to establish new colonies by themselves12. Also, neighboring nests do not compete and fight with each other. Instead, all the nests in a neighborhood and beyond form one inter-connecting supercolony, cooperating and sharing food, workers, immatures, and multiple queens. Multitudes of ants travel in wellestablished lines between colonies and to food sources. New nests are established by budding, with many workers digging the new holes and bringing in queens. California has only 4 known supercolonies: one covering most of the state and into Mexico, and 3 smaller supercolonies in the southern part of the state12. The lack of Argentines in most SJV farms probably relates to water. They require a steady source because they desiccate more quickly than native ants13. Because they do not fly, they cannot cross large dry expanses, or survive for long when the water is turned off. This also explains why Argentines are most often found in one corner of an orchard or vineyard, next to a building or garden or other permanent source of water. Argentines are highly effective at guarding honeydew-producing homopterans from natural enemies. For example, they can induce massive outbreaks of brown soft scale in citrus, where normally this pest is hard to find11. In grapes, good mealybug biocontrol becomes impossible with these ants5. We have two other small dark ants that resemble Argentines. Both forage solo rather than in distinct trails. The crazy ant, Nylanderia vividula, is almost the same color and size. It has dark hairs that may be visible with a hand lens. The odorous house ant, Tapinoma sessile, is black. It holds the gaster high, hiding the small petiole, which lacks the prominent dorsal scale seen on Argentine ants. Unlike many Dolichoderinae including Tapinoma, Argentines have only a mild pungent odor when crushed. Most baits for Argentines use sugar as the attractant5. Boric acid is a common toxicant because it is safe around homes and easy to register in agriculture, but control is short-lived because the supercolony repopulates quickly. Baiting in April-May is most effective because most reproductive larvae develop at that time5. Fipronil baits are now the standard in urban areas because control may last for years. Fipronil termite treatment may also eliminate Argentine ants. Fipronil is not registered in agriculture for ant baits or most other uses. It has been blamed for some cases of bee colony collapse, and a defective seed treatment caused a mass bee die-off in France24. Sugar is highly attractive, but Argentines also seek protein at times of the year when the brood is growing. For example, dog food is often infested. Oil-based baits aimed at fire ants do not attract Argentines. Odorous House Ant Tapinoma sessile Dolichoderinae This small black ant can be confused with Argentine ants. It is blacker and the petiole is small and hidden beneath the gaster, which is held high. It has a pungent odor when crushed. It forages solo. It differs from crazy ants by its more typical-sized antennae, with the scape only slightly longer than the head. Tapinoma often occurs on vines with vine or grape Odorous house ant. mealybugs; on the roots, trunk, or leaves, or in rotting bunches. Photo by Charley Eiseman The nest may be at the vine base but not obvious until disturbed. Once I found a colony holding their pupae clumped under loose vine bark. Some were carrying pupae down to the soil. An herbicide sprayer had passed not long before; perhaps the ants had moved the pupae up to escape a repellent chemical. I also found this ant on citrus. I have seen this ant west of Sanger in Fresno County, and near Earlimart in Tulare County. Odorous house ants can be a household pests14. Carpenter Ant Camponotus essigi Formicinae Carpenter ants nest in wood. Some species are all-black, but Camponotus essegi has a reddish head and thorax. I have only seen this species once, in an organic vineyard near Del Rey in Fresno County. The ants were fighting a vicious intra-species war for control of the vine. Mayhem was so widespread that I could not find any whole specimens to photograph or capture. Carpenter ants have distinct castes, with majors much larger than minor workers8. Carpenter ants forage for dead insects, extract the fluids, and usually leave the exoskeleton Carpenter ant with one antenna torn off behind24. They also tend homopterans such as 17 in battle. aphids . I found them on the same vine as vine mealybugs. California Gray Field Ant Formica aerata Formicinae The California gray field ant (AKA native gray ant) is the most widespread ant in San Joaquin Valley vineyards and orchards, present in virtually every block and dominating most13. It differs from Field ant worker. Photo by Jim Moore. our other large ants because the head and thorax are less reddish and the gaster is only a little darker, giving it a brown appearance. Camponotus essegi and Myrmecocystus mimicus both have a redder head and thorax. Formica aerata can vary in size, but it does not have distinct castes like Camponotus. Field ants love homopteran honeydew and are effective at driving away biological control13. They are a key factor in outbreaks of aphids, mealybugs, soft scale, and other pests in many tree and vine crops. They also feed at peach nectaries6 and pick up the “sugar balls” made by grape leaves. Vetch with nectaries planted between grape rows will attract F. aerata away from the vines2. Field ants can dig holes in fruit, at least where they find a crack or hole Field ant nests mounds are lopsided and not well started. Nectarines are susceptible to ant developed, often with multiple entrances. damage. In grapes, these ants might slow down rot by scrapping out the berries. I have seen serious damage to persimmons apparently caused by F. aerata. Kris Tollerup, working in Coachella Valley with Formica perpilosa, found that vine mealybug disappeared when the ants were controlled19. He found evidence that the obligate association of VMB with F. perpilosa had abiotic factors in addition to defense. Field ants carry mealybugs to new vines and move them between vine parts such as leaves, fruit, trunk, and roots. Sometimes field ants cover mealybugs on vines with piles of dirt pebbles and debris. Their colonies around vine roots are a frequent mealybug overwintering site. I have also seen these ants carrying live alate aphids on vines. Field ants seem to corral mealybugs in convenient spots such as under grape berries or on trunks. This might slow the spread of mealybugs out over the vines. Field ants prey on both pests and other predators. F. aerata is a key natural enemy of peach twig borer in almonds and peaches6. However oriental fruit moth worms have a spitting defense against the ants7. Field ants eat loopers on weeds (Pers. comm. Don Thomas) and carry leafhoppers off almond trees. They will pull apart spiders such as Pardosa and Trachelas, and wrestle down bees they can catch. Some Formica species are such effective caterpillar predators that they have been manipulated as biocontrol agents. Two species have been artificially established in European timber plantations, including F. lugubris which was also successfully transferred to eastern Canada (probably no longer allowable under stricter regulations). During outbreaks of the pine beauty moth on Europe, “green islands” surround colonies of F. polyctena. Some aphid parasites (Aphidiidae) seem to like field ants. I have seen them hopping on and off the backs of ants nowhere near aphids, as if they hope the ants will lead them to aphids. F. aerata ants often war with neighboring colonies in vineyards. Once an ant clamps its jaws Field ants often cover mealybugs with on a leg or antennae, it will not let go. Sometimes piles of pebbles and debris. Sometimes survivors will carry around severed heads still they nest in the vines. hanging onto their limbs. Sometimes they carry other adult F. aerata on their backs. Perhaps this is a sign of slaving. Field ants sometimes try to bite people, but it does not hurt much. F. aerata often nests at the base of a tree or vine. Sometimes they use pre-formed cracks in dirt. There may be multiple entrances connected by tunnels close to the surface. The mounds are not well developed. F. aerata forages solo. They have a run-stop-run behavior, often turning. They continue foraging even during hot times of the day. I have observed F. aerata nuptial flights from early May through June. The alates are brownish, similar to the workers. Fire ant baits based on grease are not attractive to Formica. Field ants are attracted to sugar and protein. Tollerup found that anchovy was the most attractive bait he studied18. Unfortunately, the most effective Formica baits are not registered in agriculture18. Fire ants compete with field ants, so controlling fire ants can allow field ants to increase6. Honeypot Ant Myrmecocystus mimicus Formicinae Honeypot ants are a little smaller than field ants and have a red head and thorax with a much darker gaster. They look similar to pyramid ants, but are larger and lack the odor when crushed. The nest is distinctive with a large entrance that appears vertical, without much mound. They typically nest in the open. The workers forage individually and have a walk-sprint-walk habit. M. mimicus tends homopterans including soft scale. Their nests are usually outside of vineyards, but their trails can extend at least 13 yards to vines inside. In my opinion they are not very effective at protecting against natural enemies. M. mimicus preys on insects including termites and bugs1. They will also steal prey from harvester ants10. I have observed M. mimicus nuptial flights from late March to early May. The all-dark males are a little larger than the workers. The females are much larger with a red head. Honeypot ant worker. Honeypot ant holes are easily recognized by their wide, vertical entrance. Crazy Ant Nylanderia vividula Formicinae Crazy ants can be confused with Argentine ants, which are about the same size and color. Nylanderia vividula forages solo without distinct trails. The dark hairs may be visible with a hand lens. The thorax is often distinctly lighter than the gaster and head. The scape is twice as long as the head. Tapinoma sessile is darker and has a pungent odor when crushed. N. vividula is native to southern United States but has spread to other continents. It is a household pest3. The name comes from their jerky movements Crazy ant worker. Note the distinct hairs. and crazy running when disturbed. . N. vividula is usually found tending homopterans, such as mealybugs on grapes and aphids on pears. They seem to be less effective than field ants at guarding. I found one in an almond on the ground. They are fairly common, but the nests are hidden . I have found this ant in Madera, Fresno, and Tulare counties. Little Black Ant Monomorium minimum Myrmicinae This extremely small ant is easily overlooked, so it is probably more common than reported. I have only seen it in one location, a raisin vineyard near Caruthers. It was tending vine mealybug. The little black ant is an invasive species that can be an urban pest17. It preys on insects in addition to tending homopterans17. Little black ant worker. Photo by Ben Coulter. California Harvester Ant Pogonomyrmex californicus Myrmicinae This ant is the largest of our common agricultural species. It is usually all red, although the gaster may be darker. It is very abundant in sandy areas but is much less common elsewhere. For example, I have not found any in the area around Earlimart. Harvester ants primarily eat seeds24. They will pick up bits of almond, but they are unable to chew holes in intact nuts, so they are not pests of California harvester ant worker. Photo by Lynn Monroe almonds. Harvesters prey on live insects and scavenge dead insects. I have seen them kill honey bees and an alate of another ant species. They eagerly scavenge on dead birds and mammals. Harvesters are among the few ants which do not tend homopterans for honeydew. I saw a mid-size wasp with yellow markings, probably Sphecidae, grab a harvester ant, land on grass for a short time, then fly off with its prey. Harvesters build a large crescentshaped mound. The entrance has a shallow slope making the ants all exit on one side. Harvesters remain active in the hot hours of the day. According to the Wheelers23, P. californicus is quick to inflict a very painful sting. The pain spreads beyond the sting site through the lymph system The nest entrance is almost horizontal. The ants exit in to points such as the groin and jaws. one direction, making a lopsided mound. The ache can last 5 hours, and swelling and itching up to a day. I have observed nuptial flights in June and early July. Jet Black Harvester Ant Messor pergandei Myrmicinae In 2005 I saw black harvester ants in an almond orchard north of Delano. I did not collect them but a good guess is that they were Messor pergandei. This species is common in the southern desserts of California and Arizona, and has been reported in the Bakersfield area. Haney lists them as Veromessor pergandei9. Red Imported Fire Ant (RIFA) Solenopsis invicta Myrmicinae This species from South America is a major pest over much of the American south20. It has been found in the San Joaquin Valley, but so far the counties have done a good job of quarantine and extermination. S. invicta looks and acts very similar to the southern fire ant, S. xyloni. Mature mounds may be larger, and the ants are even more aggressive. Under magnification S. invicta is distinguished by a longer scape which reaches to near the top corner of the face, four mandibular teeth instead of three, and a third tooth in the middle of the edge of the clypeus20. Compare also to pyramid ants, which have only one segment in the pedicel. The value of S. invicta as a beneficial predator has been demonstrated in sugar cane and cotton21. It did little harm to other predators. But it also defends homopteran pests. RIFA worker. Photo by Jon Hart RIFA worker face showing 3 teeth on the edge of the clypeus. S. xyloni lacks the middle tooth. RIFA also has longer antennae. Photo by Mario David Bazan Thief Ant Solenopsis molesta Myrmicinae Thief ants are so named because they often co-inhabit other ant nests and steal food. They will also consume host ant larvae and pupae17. Their diet is similar to fire ants and includes fats and oils, as well as high-protein foods. These tiny ants could easily be overlooked on grape vines except when they appear in mass to feed on the sap oozing from grape girdling wounds. They are fairly common but only seen by people who are searching under bark. Thief ants tend mealybugs, but it is not clear how well they defend them from predators and parasites. They are distinguished by their tiny size and reddish yellow color. Thief ant worker. Thief ants massed on recent girdling wounds. Southern Fire Ant Solenopsis xyloni – southern fire ant Myrmicinae The very common southern fire ant is our most important pest of almonds; a very effective defender of homopteran pests; capable of harming young trees and vines22; and our ant most guilty of biting and stinging humans. When fire ant nests are disturbed, the denizens rush Southern fire ant worker. out to attack. The workers come in a range of sizes; the larger ones serve especially as defensive soldiers, but all sizes fiercely sting and bite. It is a rare SJV resident who has not been victimized. Luckily, the pain is usually short-lived. Similar looking ants include pyramid ants and Guinea groove-headed ants. Fire ants can be distinguished by their aggressive behavior, their large variation in size, colony appearance, and anatomy details covered under the other species. In my experience fire ants are the most successful, after Argentine ants, at defending homopterans against predators and parasites. In vineyards where natural enemies have controlled most mealybugs, fires will often be found protecting the remaining infested vines. In grapes, where baiting is aimed at enabling biological control of mealybugs, application should be early in the season. Natural enemies need time to work. Fire ant nests usually have multiple entrances. Also, the ants like to corral the mealybugs into small areas, such as between grape bunches and wood. If ant control comes late, the mealybugs may be released to spread out in the bunch. Only fire ants and pavement ants are almond pests, and fire ants are more common in our orchards. They readily chew holes in nuts on the ground, and may even climb trees and eat the nuts there. For this reason, most agricultural ant baits are aimed at fire ants. As a “grease ant”, fires are attracted to vegetable oils, which are a typical ingredient of these baits. Except when disturbed, fire ants remain in their nests during hot hours of the day. Thus scouting for colonies is easiest in the cool of the morning. A trowel is handy for digging up suspected colonies. Scouts should take care not to count pyramid ants, which come in only one size, do not swarm out when disturbed, and have a pungent odor when crushed. Field ants are also common in almond orchards. They are larger, appear brown, vary much less in size, and are not as aggressive. Nests are most common in sunny areas. Sometimes fires nest under rocks or sticks, or around the base of weeds. Most nests have multiple entrances, and mature colonies may have multiple interconnected mounds extending over 25 feet. Partial tunnels may be built along cracks with the travelling ants intermittently visible from above. In grapes, fire ants may nest in the vine or at the base. Fires are predators and scavengers of dead arthropods, earthworms, and vertebrates. The related S. geminata reduced maize weevils in corn by 98%21. But they do not prey on twig borers6. Fires are more capable than most ants of cutting through plant surfaces. They can girdle young citrus trees and grape vines13. They sometimes feed on young leaves and twigs and can damage young citrus fruit22. On grapes, they may feed on bits of cambium exposed through holes in the bark. They also collect and store seeds14. I have seen fire ant nuptial flights in June and early July. Fire ant feeding on cambium through hole in grape bark. Fire ant mounds interconnected along soil cracks. Workers can be seen through breaks in the tunnel roof. Guinea Groove-Headed Ant (AKA Tramp Ant, Pennant Ant, Pavement Ant) Tetramorium bicarinatum Myrmicinae My single record of this ant was on a vine with vine mealybugs near Fresno on May 28, 2013. On the same vine I found pyramid ants, pavement ants, and field ants. I think that is my record for most ant species on a single vine. T. bicarinatum is in the same genus as the pavement ant, and shares the deeply grooved face. It resembles fire ants in color and size, but it is less aggressive, does not have the wide size variation, and has a 12-segmented antenna with a 3-segment club. The common name “tramp ant” refers to several unrelated species that travel around the world on ships. This invasive species can be a household pest. Diet is omnivorous, similar to pavement ants, and includes fruits, meats, and grease17. Guinea groove-headed ant worker. Photo by Graham Montgomery Pavement Ant Tetramorium caespitum Myrmicinae Pavement ants are common on farms and also in urban areas where they are pests14. Their black color distinguishes them from our similar-sized ants. Argentines, long-horned crazies, and odorous house ants are smaller. Under magnification, pavements are identified by the vertical grooves on the face. These ants move on trails in a slow walking gait and tend to ignore disturbance. They have stingers but are not very aggressive to humans. Pavement ants are the second pest of almonds, along with fire ants. Greasy fire ant baits also attract and control pavement ants. Pavements are avid tenders of homopteran pests but do not seem to be very Doughnut shaped pavement ant mound. effective at defending them. Biological control works well when they are present. They are predators of insects and also eat seeds, fruit and many other foods24. Nests in the open are donut shaped, similar to pyramid ants. Sometimes nests are at the base of vines around the roots. Pavement ants are displaced by Argentine ants, but they are among the first to return when the Argentines are controlled. They often engage in wars with neighboring colonies in the spring24. Sometimes the parasitic ant Anergates atratulus takes over T. caespitum colonies, kills the queen, and forces the workers to raise its young24. The colony lasts a couple of years until all the pavement workers die. I have not seen reproductive flights of pavement Pavement ant worker carrying a pupa of ants. the parasitic ant species Anergates atratulus. Photo by Claude Pilon. Microscopic Key Worker ants in SJV orchards and vineyards Use AntWeb California region2 for close-up photographs. 1a Pedicel with two segments (petiole and postpetiole) between thorax and gaster. A stinger may be present. Ocelli absent. Myrmicinae. - 9 1b Pedicel with one segment (petiole). No stinger. - 2 2a. End of abdomen with a circular opening (acidophore), usually surrounded by a fringe of hairs. Several of our species are large. Ocelli usually present. Formicinae. - 3 2b End of abdomen with a transverse slit opening. Our species are small to medium. No ocelli. Dolichoderinae. - 6 3a Small brown ants, about 3 mm or less. Head, thorax and antennae with long dark setae, often paired. Ocelli present. Nylanderia vividula. Crazy ant. 3b Large species, about 6 mm or longer. Thorax reddish with darker gaster. - 4 4a No fringe of hairs around acidophore. Head and thorax much redder than dark gaster. No ocelli. – Camponotus essigi. Carpenter ant. 4b Acidophore with a fringe of hairs. Ocelli present. - 5 5a Maxillary palpi about as long as head. Head and thorax much redder than dark gaster. Myrmecocystis mimicus. Honeypot ant. 5b Maxillary palpi not much larger than mandibles. Head and thorax only a little redder than gaster. Formica aerata. California gray field ant. 6a. Abdomen and thorax both dark brown or black. - 7 6b Thorax reddish, abdomen different colored. - 8 7a Petiole with a well-developed dorsal cone. First two antennal segments beyond scape about the same size. Dark brown. Linepithema humile. Argentine ant. 7b Petiole with dorsal cone barely showing, hidden under gaster. First antennal segment beyond scape longer than second. Black. Tapinoma sessile. Odorous House Ant. 8a Abdomen yellowish - Forelius pruinosus 8b Abdomen darker than thorax. Propodeum with a “pyramid”. Dorymyrmex bicolor. Pyramid ant. 9b Small or medium sized, less than 6 mm. - 10 9a Large ants, longer than 7 mm. Typically solid red or black. Long beard-like hairs under head. - 15 10a Face with many vertical grooves. Antennae with 12 segments and 3-segment club. - 11 10b Face without such grooves. - 12 11a Head and thorax black or dark brown like abdomen. Grooves cover face from edge to edge. Tetramorium caespitum. Pavement ant. 11b Head and thorax reddish, abdomen darker. Groves in middle of face, with sides reticulate. Tetramorium bicarinatum. Guinea groove-headed ant. 12a Antennae 12-segmented with 3 segment club. All black. Very tiny, less than 2 mm. Monomorium minimum. Little black ant. 12b Antennae 10-segmented with 2 segment club. Head and thorax red or reddish. Solenopsis. 13 13a Very tiny, less than 2 mm, all yellowish red. Solenopsis molesta. Thief ant. 13b Medium size, head and thorax red, gaster darker. - 14 14a Lower edge of clypeus with two teeth, one above each mandible base. Mandible with 3 teeth. Scape ends well short of upper face corner, by length of several proximal funicular segments. Mesopleuron generally smooth. Solenopsis xyloni. Southern fire ant. 14b Clypeus with a third tooth in the middle. Mandible with 4 teeth. Scape reaches near or beyond upper face corner. Mesopleuron dull with striations. Solenopsis invicta. Red imported fire ant. 15a All segments black. Messor pergandei. Jet Black Harvester Ant. 15b Head and thorax bright red. Gaster usually red, sometimes darker. Pogonomyrmex californicus. California harvester ant. Field Key Ants common in SJV orchards and vineyards. Quick checks: Pungent odor when crushed: pyramid, odorous house, Forelius. Argentine has milder odor. All segments black or dark brown: jet black harvester, pavement, odorous house, Argentine, crazy, little black. Head and thorax red, gaster much darker: California harvester (some), honeypot, carpenter, pyramid, fire. Head and thorax reddish brown, gaster a little darker: California gray field ant. All red: California harvester (most). Yellow or reddish-yellow: Forelius, thief ant. 1a 1b 1c 1d Large ants. Majority of workers 6 mm or longer. About the size of field ants or larger. - 2 Medium ants. Majority of workers 3-5 mm. About the size of typical fire ant workers. - 5 Small ants. About 2-3 mm. About the size of Argentine ants. - 8 Tiny ants. Less than 2 mm. - 11 Large Size 2a Pedicel with 2 segments. About 8 mm. Harvester ants. -3 2b Pedicel with 1 segment. About 7 mm or smaller. – 4 3a All segments black. Messor pergandei. Jet Black Harvester Ant. 3b Head and thorax bright red. Gaster may be dark, but more often bright red. Very common. Pogonomyrmex californicus. California harvester ant. 4a Head and thorax reddish brown, only a little less dark than gaster. Ants forage solo with a stop-start walking gait. Nest typically with a shallow slope at entrance, often with multiple entrances, often next to objects such as tree trunks. (Sometimes nests in vine trunks.) Some variation in size, but no distinct castes. Very common. Formica aerata. California gray field ant. 4b Head and thorax red with much darker gaster. No distinct castes. Ants near nest have a walk-sprint-walk-sprint gait. Nest with a large, circular, apparently vertical entrance, usually only one, and typically out in the open. Common. Myrmecocystis mimicus. Honeypot ant. 4c Gaster dark. Head and thorax color variable, sometimes red. Majors much larger than minor workers. Nests in wood. Rare in SJV agriculture. Camponotus essegi. Carpenter ant. Medium Size 5a Head, thorax, and gaster black or dark. - 6 5b Head and thorax reddish, gaster darker. - 7 6a Pedicel with 2 segments. No strong odor. Nest typically out in the open or next to an object. May be tending homopterans. Common. Tetramorium caespitum. Pavement ant. 6b Pedicel with 1 segment, difficult to see because the gaster is held high. Strong odor when crushed. Typically found in grape vines tending mealybugs. Less commonly seen. Tapinoma sessile. Odorous House Ant. 7a Pedicel with 1 segment. Workers all about the same size. Strong odor when smashed. Typical mound an even doughnut with a small central entrance. During active hours, workers continuously building mound. Very common. Dorymyrmex bicolor. Pyramid ant. 7b Pedicel with 2 segments. Worker castes vary greatly in size, with smaller workers and larger soldiers. Colonies excitable; workers rush out when disturbed. Nests often have multiple entrances. Odor not strong. Very common. Solenopsis xyloni. Southern fire ant. (Mature nests of Solenopsis invicta, Red imported fire ant, may be larger and higher than for S. xyloni, and the workers even more aggressive. See microscope key to distinguish these two.) Small Size (These have 1-segmented pedicel) 8a Red with yellow gaster. Nest typically in the open. Strong odor when crushed. Often foraging in grass or straw, with ants following distinct trails like Argentine ants. Forelius pruinosus. 8b Dark brown or black, or gaster darker. – 9 9a Pungent odor when crushed. Almost solid black. Petiole small and hidden beneath gaster. Tapinoma sessile. Odorous House Ant. 9b Mild odor at most. Dark brown. Petiole large and easy to see. - 10 10a Ants move between nests or forage on distinct trails which may climb vines or trees. Nests on open ground or beneath objects. Scape reaches near top of head. Very few dark hairs on body. Very common around houses, but in SJV typically found only on orchard or vineyard edges, often near buildings. Mild pungent odor. Linepithema humile. Argentine ant. 10b Ants forage solo, usually on vines tending mealybugs or aphids. Scape close to twice as long as head. Body with many long dark hairs, visible with good hand lens. Common but not often seen except under vine or tree bark, and in buildings. Nylanderia vividula. Crazy ant. Tiny Size (These have 2-segmented pedicel) 11a Reddish yellow. Common under grape bark tending mealybugs. Solenopsis molesta. Thief ant. 11b. Black. Not as common. Typically under grape bark tending mealybugs. Monomorium minimum. Little black ant. Bibliography and Resources 1. American Museum of Natural History. Ants of the Southwest. http://research.amnh.org/swrs/ants-southwest 2. AntWeb. Caltech. http://www.antweb.org/index.jsp 3. Carmean, Ingrid. 2004. Key to Most Common Ants in the Fresno Area. http://aaie.net/wp-content/uploads/2012/12/antsCarmean.pdf. 4. Clune, Maggie. 2013. Ants catch prey - and keep it. MyScience.org. http://www.myscience.org/wire/crazy_ants_catch_prey_and_keep_it-2013-sussex 5. Cooper, Monica L., Kent M. Daane, Erik H. Nelson, Lucia G. Varela, Mark Battany, Neil D. Tsutsui, & Michael K. Rust. 2008. Liquid baits control Argentine ants sustainably in coastal vineyards. Calif. Agric. 62: 177-183. 6. Daane, Kent, & Jeffery Dlott. 1998. Native gray ant has beneficial role in peach orchards. Calif. Agric. 52: 25-31. 7. Daane, Kent, Jeffery Dlott, Mathew Jones, Ingrid Peterson, & Glenn Yokata. 1995. Manipulation of the predaceous ant, Formica aerata, for control of the peach twig borer. California Tree Fruit Agreement Research Report. 8. Fisher, Brian L., & Stefan P. Cover. 2007. Ants of North America: a Guide to the Genera. Univ. Calif.. Press, Berkeley, Los Angeles, London. 9. Haney, Philip, Phil Philips, & Robert Wagner. A Key to the Most Common and/or Economically Important Ants of California, With Color Photographs. Univ. Calif. Coop. Ext. Leaflet 21433. 10. Hölldobler, Bert, & Edward O. Wilson. 1990. The Ants. Harvard Univ. Press. 11. Martin, K.W., J.A. Weeks, A.C. Hodges, and N.C. Leppla. 2012. Citrus Pests: Argentine Ant. http://idtools.org/id/citrus/pests/factsheet.php?name=Argentine+ant 12. Moffett, Mark W. 2010. Adventures among ants: a Global Safari with a Cast of Trillions. Univ. Calif. Press, Berkeley, Los Angeles, London. 13. Phillips, Philip A., Monica L. Cooper, Kent M. Daane, John H. Klotz & Mark Battany. 2013. Ants. In: Grape Pest Management Manual - Third Edition. Ed. Larry J. Bettiga; UC ANR. pp. 324-331. 14. Reynolds, C.A., M.L. Flint, M.K. Rust, P.S. Ward, R.L. Coviello, and J.H. Klotz. 2001. Key to Identifying Common Household Ants. UC Statewide IPM Program. Also available online at www.ipm.ucdavis.edu/TOOLS/ANTKEY/. 15. Rice, Eleanor. Dr. Eleanor’s Book of Common Ants. http://ants.yourwildlife.org/dr-eleanors-book-of-common-ants/ 16. Rust, M. K., & D. H. Choe. 2007. Pest Notes: Ants. UC ANR Publ. 7411 Also available online at http://ucipm.ucdavis.edu/PMG/PESTNOTES/pn7411.html. 17. Texas A&M University. Urban Pests. http://urbanentomology.tamu.edu/urban_pests.html 18. Tollerup, Kris E., Michael K. Rust, Keith W. Dorschner, Phil A. Phillips, & John H. Klotz. 2004. Low-toxicity baits control ants in citrus orchards and grape vineyards. Calif. Agric. 58: 213-217. 19. Tollerup, Kris E., Michael K. Rust, and John H. Klotz . 2007. Formica perpilosa, an Emerging Pest in Vineyards. J. Agric. & Urban Entomol. 24(3):147158. 20. Vincent, S. Bradleigh, & A. Ann Sorensen. 1986. Imported Fire Ants: Life History and Impact. Texas A&M University. 21. Way, M. J. & K. C. Khoo, 1992. Role of ants in pest management. Annu. Rev. Entomol. 37:479-503. 22. Weeks, J. A., A. C. Hodges, and N. C. Leppla. 2012. Citrus Pests: Southern Fire Ant. http://idtools.org/id/citrus/pests/factsheet.php?name=Southern+fire+ant 23. Wheeler, G.C. & J. Wheeler. 1973. Ants of Deep Canyon. Univ. Calif. Riverside. 24. Wikipedia: various articles.
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