Morphology, Classification and Control of Mites

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1. Morphology, Classification and Control of Mites Muhammad Zeeshan Nazar 1 Glenn B. Stracher 2 Zahid Mahmood Sarwar 3 1 Faculty of Agriculture Science and Technology Department of Agriculture Entomology Bahauddin Zakariya University, Multan, Pakistan 2 School of Science and Mathematics East Georgia State College, University System of Georgia Swainsboro, Georgia 30401 USA 3 Faculty of Agriculture Science and Technology Department of Agriculture Entomology Bahauddin Zakariya University, Multan, Pakistan 1

2. 2 Taxonomy Kingdom: Animalia Phylum: Arthropoda Class: Arachnida Order: Acari

3. Mites and ticks collectively form the most diverse group of class Arachnida Mites are the 2 nd most diverse group of living organisms after insects Until 1999, about 50,000 species of mites have been identified all over the world (Walter and Proctor, 1999). Up to 2011, a total of 54,617 mites and ticks species has been reported (Zhang, 2011). There is an estimate that half a million species are still to be identified. Distribution

4. Mites are minute to small range from 300 to 500 µm in body length. The largest Acari (Red velvet mites: Trombidiidea ) may reach lengths 10 – 20 mm i.e., 0.4 – 0.8 inches ( Weygoldt , 1998) . The small in size and strongest in evolutionary flexibility of the Acari have permitted mites to colonize most of the aquatic and terrestrial habitats. As human being, the Acari form a major component of the fauna of the cultivated crops and forests either as pest or as biological control agent (Kiefer et al , 1982 and Gerson et al , 2003).

5. Moreover, this little animal play an important role in litter, grasslands and agricultural soils, where they recycle minerals ( Balogh , 1972). Walter and Proctor (1999) summarized very effectively the ecological importance and the economic role of mites as “ The average mite is minuscule, barley perceptible to even the sharpest eyes. When enough are present, mites can exert efforts disproportionate to their size.

6. Mites are microscopic in nature and world wide in distribution and successfully colonized the terrestrial and aquatic habitats. They are present in all types of abiotic and biotic habitats like planes, mountains, desert, fresh water, salt water and springs, oceans, organic matter and lotter. Mites are present in large number in soils where they constitute up to 7% of total weight of invertebrate fauna.

7. Morphology of Mites

8. The mite are too small in size to be studied like insects. Therefore, they have been ignored by zoologies and Entomologists. At the time of Linnaeus, only 30 species had been known. Mites can be found as ecto -parasites of other invertebrate and vertebrate animals. Morphology

9. In the mites, the developing integument initially appears as undifferentiated tissues which is covered by thin layer of cuticulin and separated from the epidermis layer by ‘Schmidt layer’. During developmental process, base of the integument undergoes more conversion and shows from the outside inwards. An epiculicular layer an underlying Schmidt layer and finally, a basal lamina below the epidermis ( Alberti et al , 1981 and Norton et al, 1997) . Integument

10. In addition to micropores , in integument, the body surface of mites also have variety of macropores that play a functional role in secretory and sensory process ( Henriot , 1969). Cement layer Basal lamina Epidermis Nucleus Schimdt layer Endoculticle Evolving pore canal Exocuticle Pore canal Inner epicuticle Outer epicuticle Wax layer (According to Norton et al , 1997)

11. Mostly mites have oval-shaped bodies with two body regions that may appear fused together. Mostly mites have piercing sucking mouthparts such as Phytophagous and Predatory mites. Some mites have chewing mouthparts as Stored grain mites. Body Division

12. Mites can easily be distinguished from their sister class Insecta by the following characters S. No. Features Insects Mites 1. Body Division Head, thorax and abdomen Gnathosoma and Idiosoma 2. Antenna Present Absent 3. Wings Present Absent 4. Legs 3 pairs 4 pairs

13. The mites lack the true head and conspicuous body segmentation. The body of mites divided into Gnathosoma Idiosoma The anterior part of the mite body is called as ‘ gnathosoma ’ that is moveably connected to the idiosoma . The idiosoma is divided into the anterior podosoma and the posterior opisthosoma .

14. Podosoma Prosoma (Gnathosoma + Podosoma) Sejugal furrow disjugal furrow Propodosoma Idiosoma Hysterosoma Gnathosoma Opisthosoma

15. Gnathosoma

16. Gnathosoma considered the mouth parts which are mainly concerned with feeding and for sensatory purpose. It is generally located anterior to the body but in some cases, it may be hidden under the propodosoma . The gnathosoma having the mouthparts differ from a true head in a sense that is lacks the eyes, antennae and brain. If eyes present, they are located in idiosoma . It generally consists of chelicerae and pedipals ( Krantz , 2009). Gnathosoma

17. Chelicerae Chelicerae are the main food getting organs . They are placed dorsally in relation to the opening mouth and commonly consist of three segments known as cheliceral base, digitus fixus and digitus mobilis . First segment is basal and bears the digitus fixus which is articulated with the distal digitus mobilis dorsally ( Grandjean , 1947). Both digits are provided with teeth on the opposite side of each other .

18. The chelicerae are used for the cutting and piercing of food. They vary from three segmented and pincer like appendages of mesostigmata to slender prostigmata . In phytophagous and parasitic mites, there are modification in chelicerae that result in styliform , hood-like or finely toothed like adapted to pierce plant or animal tissues or attacked on bacterial film ( Krantz , 2009).

19. Bases of the chelicerae may be fused partially or completely to each other to form stylophore as in Tetranychoidea . The digitus mobilis is modified into a stylet that can pierce the pant cell wall. Extrusion and retraction movements depends on contraction of dorsoventral idiosomatic muscles and action of retractor muscles respectively. In some groups, the chelicerae are involved in sperm transfer ( Krantz , 2009).

20. Pedipalp The pedipals are usually called simply “pulp” have five segments beyond coax, usually resembling legs but shorter and primarily sensory in nature rather than locomotors . The coax portions border the cheliceral bases and form the side walls of the gnathosoma . Their distal segments bear many setae and sometimes claws. 21

21. Their function is related to searching for and handling food, with the distal segments usually leading to chemosensory and thigmotactic sensory receptors which act as support for feeding activity. The number of papli vary from species to species as one or two in Astigmata and Prostigmata , three or four in Ixodida , and five in Mesostigmata and Oribatida ( Krantz , 2009). In Tetranychidae , the palpi contain part of silk gland ( Alberti and Crooker, 1985) .

22. Idiosoma

23. Idiosoma The second division of the acarine body is the idiosoma . Often it is ovoid or sac like but occasionally worm like. A simple pattern divides the idiosoma into anterior propodosoma and the posterior hysterosoma , distinct or not by sejugal furrow. The region bearing the legs that are derives from embryonal somites known as podosoma ( Coineau , 1974 and Krantz , 2009). 25

24. The first two somites includes the first two pair of legs known as propodosoma , while last part bearing two other pairs of legs known as metapodosoma ( Coineau , 1974 and Krantz , 2009). O rgans for digestion, excretion and reproduction are present in idiosoma. Ocili if present they are also present in idiosoma. Sclerotized shields or plates that are heavily tanned are present on the cuticle of the idiosoma , normally protect the upper surface of the idiosoma . 26

25. Setae are also present on the idiosoma . The number, type, shape, distance and pattern of distribution of these setae on idiosoma is important are used to classify many groups. 27 Setae

26. Legs Adult and nymphs of mites, except in some Prostigmata and Astigmata , have four pairs of jointed legs and the larval instar has three pairs of legs. Typically the legs consists of seven segments. Their names are coxa, trochanter, femur, genu, tibia, tarsus and apotele . According to systematic group, the coxae may be free or fused with the ventral podosoma and femur may be divided into basifemur and telofemur . 28

27. One or, more segments of leg 2 and sometimes leg 5 are spurred in males of the Mesostigmata are used to grasp the female during mating. Different segments of legs bear a number of setae which are arranged in whorls i.e., in rings around the circumference of legs (Evans, 1963). 29

28. The idiosoma has different types of sensory receptors known as chemoreceptors, mechanoreceptors and photoreceptors with setal structure (Evans, 1992 and Coons, 1999). The cuticular surface has different pore-like openings, which have a sensory function ( Krantz , 2009). These pores have various shaped but mostly small membrane covered clefts Sensory Receptors

29. Secretory Organs Pore-like openings located in the cuticle of mites and connected b a ducts to suncuticular gland cells ( Alberti and Seeman , 2005). The nature of the secretory products is heterogenous and varies from cement like to waxy (Evans, 1992 and Coons, 1999). The secretory products contain vary chemical composition, having monoterpenes, hydrocarbons, esters, aromatics etc and may have various functions that act as pheromones ( Mizoguchi et al , 2003).

30. Life History of Mites

31. The life cycle of Acari develops through the eggs and six biological instars: Eggs Prelarva Larva Protonymph Deutonymph Tritonymph Adult The two resting stages occur between larva and nymph and between nymph and adult known as nymphochrysalis and imagochrysalis respectively (Manson and Oldfiels , 1996). Life cycle

32. Life cycle of mites depending on the temperature and availability of food. This cycle usually takes 17 days at 20 o C Female deposit 5 to 6 eggs per day, with total of 60 to 100 eggs underside of the leaves. Mostly eggs are oval in shape and reddish, orange or whitish in colour. Eggs hatch in 3 to 6 days depending on environmental condition. Eggs Spider mite eggs by Scot Nelson

33. The prelarva also known as prolarva or deutovum and represents a non-feeding stage and lack legs, known as calyptostasis . It develops inside the egg chorion and consume yolk. In some groups the prelarva has three pairs of legs, mouthparts and setae, known as elattostasis ( Coineau , 1974). A calyptostasis is reported for Tetranychus urticae and elattostasis for Panthaleidae (Andre and Van Impe, 2012). Prelarva

34. This is active, feeding instar, a weak, miniature stage. After eclosion , larvae shed both skin and chorion. Larva is typical six-legged instar and lack external genitalia. In some groups, larva directly develops into nymph, known as nymphochrysalis while in other case, larva first changes into protonymph b efore nymph, known as protochrysalis (Oku et al, 2003). Larva

35. They represent the first eight-legged nymphal instar that is free living and active. The protonymph of the Tetranychus kanzawai develops into the deutonymph through a resting stage known as deutochrysalis (Oku et al, 2003). This second nymphal instar in eight legged and resembles t the adult but lack external sexual organ . Protonymph Deutonymph

36. When tritonymph (third instar) is absent, the deutonymph develops into the adult through resting stage known as teliochrysalis (Oku et al, 2003 ). Tritonymph This third nymphal instar is eight legged and uncommon. When present, it is active and free living. It is absent in Mesostigmata and in many Prostigmata . Larva and nymphs complete development in 4 to 9 days depending on temperature .

37. Generally, the adult instar concludes the biological life cycle. It is eight legged and sexually functional and live about 30 days. In T. urticae , the adult instar is absent and is replaced by tritonymph instar, which is able to reproduce by pedogensis (Andre and Van Impe, 2012). The male and female sex ration vary from species to species as in eriophyoid mites, the percentage of female ranges from 51 to 95 % ( Sabelis and Bruin, 1996). Adult

38. Generally , mites tend to increase in population during the summer as the prefer hot day conditions . Reproduction in mites is characterized by a haploid-diploid genetic system. The main genetic systems are diplodiploidy , haplodiploidy and thelytoky . All these systems are sexual. Mites have very small chromosomes, less than 0.03 mm long. They have 2 to 12 numbers of chromosomes varies from group to group ( Wrensch et al, 1994)

39. Life stages of  Chaetodactylus krombeini  ( Astigmata ) by Ron Ochoa and Gary Bauchan , USDA-ARS

40. Categories of Mites

41. The mites can be divided into four main groups according to their economic importance i.e., Phytophagous mites ( plant feeding mites) Predatory mites. Stored grain and stored product mites. Parasites mites (mites of medical and veterinary importance) Types

42. Phytophagous Mites

43. Phytophagous mites infest and damage cultivated crops, vegetables, orchards, ornamental plants, forest trees and also wild vegetation. Worldwide, around 7500 species of phytophagous mites are known which damage plants. They are usually opaque white, translucent, slow moving and short legged. They infest the leaves, inflorescence and developing tissue by sucking the sap with their stylet like chelicerae. Phytophagous Mites

44. These mites cause both qualitative and quantitative losses. In various crops like rice, sugarcane, brinjal , okra and chillies , 10-30% losses are reported due to spider mites. The damage is more severe in case of mangoes where it may reach up to 50-80 % ( Chhillar et al., 2007). Tetranychus urticae can damage 18-22 cells in a minute ( Liesering 1960) . Their feeding results in leaf stippling, blotching, curling and twisting .

45. They also modify the developing tissue by forming galls and injecting toxins ( Jeppson , Baker and Keifer , 1975 ) . The symptoms may be irregular deformities of growth pattern, rosette type growth, irregular leaf or fruit growth, total destruction of growing tips etc. All stages of mites except eggs can transmit virus and cause diseases. Different viral diseases caused are wheat streak mosaic, fig mosaic, potato virus, tobacco ring spot and tobacco mosaic virus etc .

46. Certain soil inhabiting mites carry fungal spores of various root crop diseases like fungal rot of garlic and onion (Evans, 1992). The damage pattern of mites belonging to these families is as follows

47. 1. Tetranychidae Both nymphs and adults feed on the leaf surface. White spots are formed on the leaves in later stages of infestation and general chlorosis occurs in patches. Tetranychid mites secrete certain substances into plant cells. Puncturing of new cells proceeds from one spot to another in the form of a circle, which results in the formation of small rounded chlorotic spots.

48. . It is estimated that roughly 50 per cent of the mass of an adult female spider mite is eaten per mite per hour. The numbers of photosynthetically active leaf cells that are punctured and emptied per mite, are 100 cells per minute. At the macroscopic level, damage from mite feeding can cause leaf bronzing, stippling or scorching as well as extensive webbing on leaf surface and black fecal dots are seen on the leaf surface.

49. Severe spider mite infestation cause major reductions in plant growth rates, flower formation and yield. Penetration of cells by mite stylets and injection of saliva cause both mechanical damage and changes in cell cytology, physiological and biochemical processes of non-punctured adjacent cells. In case of severe infestation, plants show yellowing and general drying of leaves, which drop prematurely.

50. 2. Eriophyidae : The mites occur on all parts of a plant and may or may not exhibit the symptoms of damage. Based on type of injury, they have been classified as under: Gall Formers Due to feeding on various plant parts hypertrophy of cells occur. It results into formation of galls on leaves, flower buds and stem. Different types of galls like pouch galls ( Pongamia sp .), bead galls ( Ficus sp .), finger galls ( Pongamia s p.)

51. Leaf Rollers These mites roll the whole leaves or only edges of leaves and feed within the rolls. Erineum Formers Due to feeding by mites, epidermal layer of cells produce hair like out growths, which produce the erineum . Blister Mites Some species cause formation of blisters on the leaf sheath and feed within.

52. Leaf Rollers Erineum Formers Blister Mites

53.   3. Tenuipalpidae : These mites generally feed on the ventral surface of leaves near the midrib or veins. There is bronzing and rusting symptoms on the lower surface of leaves due to feeding of nymphs and adults. Some species form galls on the leaves and stems of plants.

54. 4. Tarsonemidae : They usually infest the tender portion of plants and suck the sap from buds, leaves, shoots, flowers and stem sheath. They cause curling, crinkling and brittleness of foliage but shows little leaf symptoms. The injury caused by this group is often mistaken as a disease symptoms caused by pathogenic microorganisms.

55. 5. Tuckrellidae : This is the smallest phytophagous family, which includes four species. These are brightly colored having fan like dorsal body setae and long whip like caudal setae. The mites do not have much importance, as they do not cause any economic damage ( Chhillar et al., 2007).

56. Predatory Mites

57. Predatory Mites There is a large group of predatory mites which feed on other harmful mites, small soft bodied insects, their eggs and occasionally on nematodes (Krantz, 1978). These mites are usually red, yellow and green, long legged and fast moving.

58. Phytoseiidae This group has received maximum attention globally as they are reported as predators of phytophagous mites and small insects. These mites are whitish/ creamish / reddish or light brownish in color, fast moving and abundant in nature. They have very long legs, which help them to run very fast. They have wide range of food habits from carnivores to non-animal food (pollen, honey, nectar, plant sap) eaters.

59. These mites have several advantages over other predatory mites because of High fecundity Abundant availability Good searching ability Dispersal rate Adaptability to different ecological niches High degree of prey specificity

60. Stigmaeidae These are probably next to phytoseiidae as far as predatory efficiency is concerned. These are yellowish/reddish/light brownish in color Ovoid or elongated in shape which occupy various habitats. These mites cannot run very fast hence these are used for controlling slow moving mites and for destruction of mite eggs. Agistemus spp . are found to be of high promise as biocontrol agents.

61. Ventral view Dorsal view

62. Anystidae These are round, long legged, reddish and soft bodied mites. They are very fast moving and start making whirling movement as soon as touched.

63. Bdellidae These are fast moving, medium sized and reddish in colour . They prey upon phytophagous mites and small insects like collembola . So far, no species of high predatory potentiality has been noticed in field.

64. Cheyletidae These are free living predators of phytophagous, stored and house dust mites. Comb like or sickle like setae are present on pedipalp tarsus. Cheletogenes , Cheyletus , Hemicheyletia , Paracheyletia are important genera whose interaction with phytophagous mites are well documented ( Chhillar et al., 2007).

65. Cunaxidae These are reddish/yellowish/ brownish mites, fairly fast moving and are known to be active predators of phytophagous mites. They have very strong and thorny mouthparts but their number is limited in nature. Some of the mites are known to fasten their preys with silken threads secreted by their mouthparts.

66. Stored Grain and Stored Product Mites

67. Stored Grain and Stored Product Mites Mites of stored grain and stored product are of great economic importance. These mites infest and cause damage to goods in following ways: Stored grain cereals. Seeds of all kind Bulbs, tubers and decaying material Fresh, cultivated and stored mushrooms Dried fruits of all kind 71

68. Direct Damages Some free living mites live in the stored grains and their products where they multiply rapidly and attain the status of pests. Food products, especially the cereals are liable to be infested directly. The mites penetrate the seeds through cracks, tear the outer covering of embryo and eat away cavities where they develop and multiply. 72

69. These mites feed on the grain itself or fungi growing on it. These mites have blunt chelicerae for scraping and gouging the food. Majority of these mites feed on the embryo while some can feed on cotyledons as well. Due to the attack of these mites quality of the stored grains is affected more badly as compared to the quantity. 73

70. Indirect Damages These mites cause damage to stored grains products by raising the moisture content and generating sufficient heat which favors growth and infections of pathogens. They contaminate the space between the grains with their dead bodies, cast skins and excrement hereby hindering the circulation of the air in the stock. They also act a vectors of fungal and bacterial diseases and spread those through out the whole mass. 74

71. The flour which is prepared from infested grains is more acidic in nature, bitter taste, stagnant smell, more hygroscopic and has a tendency to stick togather. They are clumsy, slow in movement and almost incapable of covering large distance by themselves. They can undergo a transitory quiescent stage which is very difficult to control. The mites of stored of stored grain include following important families 75

72. Acaridae Inhalation or contact on the skin or mucus membranes of the eyes can induce allergic reactions. These mites also occur in bread, pancakes, cakes, pizza, pasta, and bread made from ingredients contaminated with mites. Humans have had anaphylactic reactions after eating these mite contaminated foods.

73. This mite also known as flour mite, which is pale greyish white in color with pink legs. The males are from 0.33–0.43 mm in length and the female is from 0.36–0.66 mm in length . Flour mites contaminate grain and flour by allergens and they transfer pathogenic microorganisms. Foodstuffs acquire a sickly sweet smell and an unpalatable taste. Tyroglyphidae

74. When infested feeds fed by animals , they show reduced feed intake, diarrhea, inflammation of the small intestine, and impaired growth. If a person is bitten from a flour mite they might suffer a reaction called Baker's itch.

75. Carpoglyphus lactis is a stored product mite infesting saccharide rich stored commodities including dried fruits, wine, beer, milk products, jams and honey. The association with micro-organisms can improve the survival of mites on dried fruits. Carpoglyphidae

76. This mite also known as Furniture mite, is found in foods and grains in warehouses. In homes, it flourishes in infested foodstuffs and in damp areas. It has a soft, cream-white body. For both sexes, the body bristles are very long and feathery, 0.3 – 0.7 mm in length. Its main food sources are flour, cereals, other cereal products and fungi. Glycyphagidae

77. Pyroglyphidae These mites feed on stored products such as grain, cereals, nuts, dried fruit, cheeses and pet foods, but only in conditions of high relative humidity. Each species is the source of multiple potent allergens that sensitize and trigger allergic reactions cause perennial rhinitis, asthma and atopic dermatitis.

78. Parasitic Mites

79. Parasitic Mites Some mites are parasites of man, animal, other arthropods ,poultry birds and cause many disease in them. These are either ecto-parasites and endo-parasites. They have tearing and piercing type of that cause scaling and crushing around the legs of poultry birds. Northern fowl mite and red chicken mite are very important parasites of poultry birds all over the world. 83

80. Psoroptes species (non burrowing mites) cause sheep scab. They damage wool and sometimes the loss of animal may also occur. Scrub typhus is caused by chigger mites Even human beings are not free from their effect Sarcoptes scabies causes itching in human. Trombicula akamushi and dust mite cause lung disease like asthma. 84 Scrub typhus

81. Trombicula akamushi and dust mite cause lung disease like asthma. These mites are also vectors of internal parasites like tape worm and filarial worms. e.g , Family Oribatidae , Acrididae and Pyemotidae etc. Gamasid mites are considered as vectors of epidemic hemorrhagic fever virus. Leptotombidiumni akamushi and L. deliense are transmitters of scrub typhus in man. 85

82. Biology Mites associated with mammals and birds are usually translucent or whitish in color so they are difficult to detect. After feed on host’s blood their color changes to reddish brown. Life cycle include egg, larva , two nymphal stages and adult. 86

83. Nature of Damage These mites act as causative agents for anaemia . Pneumonia crusting of skin, hair loss, decreased production, death of host occurs in severe case of inflammation. Inflammation, scratching and irrition lead to secondary infection. Sarcoptes scabei in case of heavy infestation may cause death of the animal. The life cycle of these mites generally require 2 to 17 days. 87

84. Mites associated with honeybees There are certain mites which are associated with honey bees. These mites maybe external parasites like varroa destructor , varroa jacobsoni and internal parasites like Acarapis woodi . Some mites which cause the bees as carries to move from one place to another are also present around the bee hives. 88

85. Families of Parasitic mites Family sarcoptidae mites are tiny arachinids that are parasites of mammals and humans These cause infection and the mites spend their life in the epidermis of the skin of their host causing various skin disorders. 89

86. 1. Psoroptidae Well-known sheep mange mites causing serious damage to fleece and can even cause deaths. 90

87. 2. Knemidocoptidae Species of this family burrow in the non-feathered areas around the break, eyes, vent and legs of birds causing tiny non-itching wart-like lesions. 91

88. 3. Pyroglyphidae Members of the family are the well-known house dust mites causing asthma, rhinitis and allergies in human due to an antigen they produce. Where the humidity is very high. 92

89. 4. Demodicidae Members of this family cause symptoms in mammals characterized by itching, inflammation and other skin disorders. 93

90. Management o f Mites Monitoring Mites are small and difficult to see with the naked eye. Using a 10x hand lens will enhance your ability to see mites and their eggs. Spider mites can be detected by infestations such as cast skins and webbing. The mites, eggs and cast skins can seen under surfaces of the leaves. Mites can also be sampled using the " beat method " whereby plant parts are beaten onto an white paper. This method works particularly well for evergreens and small-leaved plants.

91. Cultural Control Use of clean, pest free plants and cuttings is essential. Knowledge of mite prone species/ varieties can enable the grower to avoid these plants or to monitor these most closely as "indicator" plants. Watering practices affect spider mite populations. Drought-stressed plants are most susceptible to mite outbreaks Overhead sprinkler systems are less favorable for mite outbreaks.

92. Biological Control A number of predatory mite species are available for mites control. Phytoseiulus persimilis , Mesoseiulus longipes , Metaseiulus occidentalis and Neoseilus californicus have been marketed for released into greenhouses.  Occasionally they are applied as Biotic Insecticides using an Inundative Release to try to bring down an existing  population.

93. Physical Control High-volume, high pressure water sprays through some application devices such as the Water Wand and Jet All-Water Wand can displace many mites from foliage and reduced mites population for short period of time. Chemical Control Different miticides have different performance characteristics. For example: Avid penetrates into treated foliar plant cells. Pentac is relatively slow acting and has ovicidal activity Sulfur, registered as a Miticide on some vegetable crops but highly phytotoxic.

94. Resistance Management: Use miticides only when mites or plant injury they cause is first detected. Use the lowest effective miticide rates initially. Use long rotation of miticides with different modes of activity. Use of tank mixtures containing two or more products with different modes of action on the mite's nervous system.

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