01112012STUDIES ON THE BIOSYSTEMATICS 3shodhganga.inflibnet.ac.in/bitstream/10603/24991/9/09... · 2018-07-09 · Later, in the third edition of ‘A Manual of Acarology’, Lindquist

Chapter 2

This chapter provides a review of mites associated with cereals. These are arranged as

follows:

1) General introduction – presents a concise account on the conceptualization of

Acari studies

2) Classification – enumerates major modifications in systematics

3) Acarology in India – lists major contributors to Indian acarology

4) Acari studies on cereals – entails major contributors to cereal mite studies

5) Cereal associated mites – checklisted mites have been analysed on various

parameters

6) Taxonomic review of families – cereal mites belonging to 13 families which were

collected are reviewed and details of species occurring on cereals under each family

are presented. The literature perused has been organized in terms of taxonomy,

biology and ecology.

The name and details of species enumerated in the review are presented as mentioned in

the literature perused. Also, in each sub-chapter, only the first mention of the species

name is followed by author name.

Chapter 2- Review of Literature

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2. REVIEW OF LITERATURE

2.1 Introduction

The earliest reference to the existence of mites was by Homer in 850 B.C. About

500 years later Aristotle discussed a prostigmatid mite parasite of locusts in ‘De

Animalibus Historia Libri’. A reference to ‘tick fever’ was found on an Egyptian papyrus

scroll dated 1550 B.C. Until about 1660, mites were referred to as ‘lice’, ‘beesties’ or

‘little insects’. The Latin term ‘mite’ for the Greek word ‘Akari’ was coined in 1650.

Acarology, as a discrete discipline came to existence in around 18th and 19th century.

Linnaeus (1758) used the generic name Acarus in the first edition of ‘Systema Naturae’,

and in the 10th edition recorded 29 species in this genus. Oudemans (1926, 1929a, 1937a,

1937b) recognised several mite species and in his review, ‘Kritisch Historisch Overzicht

Der Acarologie’, covered literature on mites until 1850. The importance of Acarology

was further realized in 19th and 20th century through the outstanding works of Kramer,

Megnin, Canestrine, Berlese, Vitzthum and Oudemans. The first comprehensive book in

Acarology was published by Baker and Wharton (1952). Jeppson et al. (1975) brought

out the first comprehensive book on mites injurious to economic plants. Since then, till

date many reviews, catalogues, books and monographs on mites in general have been

published by various authors.

2.2 Classification

The diversity of Acari is extraordinary and many have proposed a complex set of

taxonomic ranks to classify mites. Hence, as the phylogeny of Acari is still disputed,

several taxonomic schemes have been proposed for their classification.

Amongst the earliest literature on mites, Murray’s treatise, ‘Economic

Entomology; Aptera’ (1877), brought together specific information on mites, however,

these were compiled without adequate analysis. The foundation of a classification was

laid down by Kramer (1877) and was modified by subsequent authors. Post World War

II, Baker and Wharton, realized the need for an updated basic text book in Acarology.

The ‘Introduction to Acarology’, by Baker and Wharton (1952) gave classification till

family and a list of known genera in each family. This classification considered ‘Acarina’


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as an order under phylum Arthropoda, subphylum Chelicerata, class Arachnida. Acarina

was further divided into suborder Onychopalpida, Mesostigmata, Ixodides,

Trombidiformes and Sarcoptiformes. Baker et al. (1958) further identified supercohorts,

cohort and superfamily and provided keys and decriptions of mite groups within these

categories. Evans et al. (1961) divided the subclass Acari into seven orders, namely,

Notostigmata, Tetrastigmata, Mesostigmata, Metastigmata, Cryptostigmata, Astigmata

and Prostigmata.

Krantz (1970, 1978) in ‘A Manual of Acarology’, proposed a scheme for the

classification of the higher categories of the subclass ‘Acari’. In the classification, the

class Arachnida contained subclass ‘Acari’, within which three orders, namely,

Opilioacariformes, Parasitiformes and Acariformes were recognized. These were further

divided into suborder, supercohort, cohort and superfamily. Krantz considered

Mesostigmata as suborder in the order Parasitiformes and Prostigmata and Astigmata as

suborders under the order Acariformes. This system of classification was most accepted

and formed basis for many acarological studies.

Evans (1992) proposed another system of classification dividing subclass Acari

into three superorders, Opilioacariformes, Parasitiformes and Acariformes. Superorder

Parasitiformes contained the orders Ixodida, Mesostigmata and Holothyrida, while

superorder Acariformes was further divided into Trombidiformes containing order

Prostigmata and Sarcoptiformes containing order Oribatida and Astigmata.

Later, in the third edition of ‘A Manual of Acarology’, Lindquist et al., in Krantz

and Walter (2009) revised the classification of the higher categories of the subclass Acari

and placed four orders, Opilioacarida, Holothyrida, Ixodida and Mesostigmata under

superorders Parasitiformes and order Trombidiformes and Sarcoptiformes under

superorder Acariformes. The order Trombidiformes contained the suborders

Sphaerolichida and Prostigmata and the order Sarcoptiformes contained the suborders

Endeostigmata and Oribatida and the cohort Astigmatina.

In the present thesis the system of classification for checklist follows Krantz

(1978) and for morphological studies follows Lindquist et al., in Krantz and Walter

(2009).


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2.3 Acarology in India

Perusal of literature reveals that the first report on mites from India was in 1868

when Peal discovered red-spider mites on tea in Assam. Later, Wood Mason (1884)

identified this mite as Tetranychus bioculatus. Green (1890), Cotes (1895) and Das

(1959-1965) published their observations on several mites on tea from India. Watt (1898)

and Watt and Mann (1903) published comprehensive books on mites incorporating the

observations made by earlier workers. Hirst (1923-1926) described some spider mites

from India. Viets (1926), Walter (1928) and Lundblad (1934) enumerated water mites.

Abdussalem (1939a, 1939b, 1941) brought out publications on various

ectoparasitic mites. Rahman and Sapra (1940, 1946) described the biology of mites

belonging to family Tetranychidae. Puttarudraiah (1947a, 1947b) gave account of several

phytophagous mites. The predatory mite family Phytoseiidae was taxonomically studied

and several species were described by Narayanan et al. (1960), Narayanan and Kaur

(1960) and Ghai and Menon (1967, 1969). Several species of soil mites were described

by Haq and Clement (1995), Haq and Jaikumar (1993), Haq and Ramani (1997), Haq and

Alphonsa (2005) and Haq et al. (1983). Sanyal (1998) contributed to our knowledge of

the faunal diversity of India.

Ghai (1964) compiled and reported 65 species of mites from India.

ChannaBasavanna (1966), Chakrabarti and Mondal (1979-1982) and Mohanasundaram,

(1980-1996) further contributed to the studies on eriophyiid mites. Gupta (1985) in his

book on plant associated mites gave a comprehensive account of plant mites in India.

Herein, he included 557 species under 131 genera and 18 families. In 1986, Gupta

brought out another book exclusively dealing with mites in the family Phytoseiidae.

During the same period and thereafter, several reviews, catalogues, bibliographies and

species descriptions were made available by the concentrated efforts of Prasad (1974,

1975, 1982), ChannaBasavanna et al. (1979), ChannaBasavanna (1981) and Sadana

(1985), to name a few.


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2.4 Acari studies on cereals

Several mites have been observed, described and redescribed from cereal crops,

both in field and storage. Much information on cereal mites have been retrieved from

catalogues and family revisions. Most of the mites were either predatory, phytophagous

or storage mites mostly belonging to Mesostigmata, Prostigmata and Astigmata.

From India, the first observation of mite pests on crops was that by Hirst, who

reported Paratetranychus oryzae from paddy fields of Coimbatore. Cherian (1938) gave

an account of mite pests of crops. Some new species reported from India on cereals

include Aceria sorghi ChannaBasavanna (1966) from sorghum, Lasioseius parberlesei

Bhattacharyya (1968) from rice and Lasioseius terrestris Menon and Ghai (1968) on

wheat, Amblyseius sorghumae Gupta (1977) from maize, Oligonychus manishi Gupta

(1980), Schizotetranychus mansoni Gupta (1980), Ogmotarsonemus oryzae

Mohanasundaram, (1996) and Pronematus oryzae Menon et al., (2007a) from paddy

fields.

Prasad (1974) catalogued 769 mite species reported from India, of which he listed

the following 17 mites from cereals: Acaropsis docta (Berlese), Acarus siro (Linnaeus),

Aceria sorghi ChannaBasavanna, Amblyseius delhiensis (Narayanan and Kaur),

Amblyseius finlandicus (Oudemans), Amblyseius hibisci (Chant), Amblyseius

longispinosus (Evans), Cheyletus malaccensis Oudemans, Eutetranychus orientalis

(Klein), Glycephagus hughesi Pillai, Leidonychus krameri (Canestrini and Canestrini),

Oligonychus indicus (Hirst), Petrobia latens (Muller), Rhizoglyphus echinopus (Fumouse

and Robin), Schizotetranychus andropogoni (Hirst), Tetranychus neocaledonicus (Andre)

and Tyrophagus putrescentiae (Schrank).

Sadana (1985) compiled host and distribution records of 156 species in 29 genera

and 3 families of superfamily Tetranychoidea and listed 8 species from cereal crops,

namely, Aponychus corpuzae (Rimando), Bryobia praetiosa Koch, Oligonychus indicus,

Oligonychus oryzae (Hirst), Panonychus ulmi (Koch), Petrobia (Petrobia) latens,

Schizotetranychus andropogoni and Tetranychus neocaledonicus.


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Gupta (1985) recorded 24 species of plant mites on cereal crops, namely Aceria

sorghi, Amblyseius (Euseius) delhiensis (Narayanan and Kaur), Amblyseius (Euseius)

finlandicus (Oudemans), Amblyseius (Euseius) ovalis (Evans), Amblyseius (Neoseiulus)

fallacis (Garman), Amblyseius (Neoseiulus) indicus (Narayanan and Kaur), Amblyseius

(Neoseiulus) longispinosus (Evans), Amblyseius (Paraphytoseius) multidentatus (Swirski

and Schechter), Amblyseius (Typhlodromalus) sorghumae Gupta, Brevipalpus

californicus (Banks), Bryobia praetiosa, Cheiracus sulcatus Keifer, Chelacaropsis

moorei Baker, Eutetranychus orientalis (Klein), Lasioseius terrestris Menon and Ghai,

Oligonychus indicus, Oligonychus manishi Gupta, Oligonychus oryzae, Panonychus ulmi,

Petrobia (Petrobia) latens, Schizotetranychus andropogoni, Schizotetranychus

hindustanicus (Hirst), Schizotetranychus mansoni (Gupta) and Tetranychus

neocaledonicus Andre.

Baker and Tuttle (1994) brought out a complete guide for the identification of 216

species of tetranychid mites in 31 genera, of which they reported only 6 species from

cereals, viz., Eotetranychus yumensis (Mc Gregor), Oligonychus gramineus (Mc Gregor),

Oligonychus modestus (Banks), Oligonychus stickneyi (Mc Gregor), Schizotetranychus

elymus Mc Gregor and Tetranychus sinhai Baker.

Amrine and Stasny (1994) recorded 18 species of eriophyid mites from rice,

wheat, maize, sorghum and pearl millet, namely, Aceria bakkeri Keifer, Aceria milli Xin

and Dong, Aceria paratulipae Xin and Dong, Aceria sacchari Wang, Aceria sorghi,

Aceria tenuis (Nalepa), Aceria tosichella Keifer, Aceria zeala Keifer, Aceria zeasinis

Keifer, Abacarus hystrix (Nalepa), Abacarus oryzae Keifer, Abacarus sacchari

ChannaBasavanna, Abacarus sporoboli Keifer, Aculodes dubius (Nalepa), Aculodes

mckenziei (Keifer), Catarhinus vulgaris Kuang and Feng, Catarhinus tricholaenae Keifer

and Cheiracus sulcatus.

Rao et al. (1999) provide an excellent compilation of mites from rice-ecosystems.

This study recorded 61 species of mites from 31 genera under 14 families, known to

occur in farm-fields, storage and granaries, namely Acaridae, Tarsonemidae,

Tetranychidae, Phytoseiidae, Eriophyidae, Ascidae, Cheyletidae, Erythraeidae,


29

Pyemotidae, Laelapidae, Glyciphagidae, Uropodidae, Cunaxidae and Oribatulidae (soil

mites), the last being represented by a single genus and species.

Joshi et al. (2002) enumerated 125 species under 68 genera, 29 families and 4

suborders. In the 29 families reported, apart from the families already listed by Rao et al.,

(1999) their checklist incorporated data on 12 families of soil mites including

Oribatulidae (Carabodidae, Damaeidae, Epilohmanidae, Galumnidae, Haplozetidae,

Haplochthoniidae, Mycobatidae, Oppiidae, Oribatellidae, Oribatulidae, Scheloribatidae,

Trhypochthoniidae), encompassing 29 species in 21 genera, single family of aquatic

mites, Arrenuridae, represented by 10 species in a single genus and another 3 families,

Chortoglyphidae, Pyroglyphidae and Trombididae, each represented by a single species

and genus.

In this study, Joshi et al. (2002) listed 53 species in the order Prostigmata, 20

species in Astigmata, 29 species in the order Cryptostigmata and 13 species in

Mesostigmata. Also, of the species diversity reported from rice agroecosystems, 31

species belonging to 12 families were saprophagous, 3 species in 2 families were

parasitic, 10 species in a single family were aquatic while the rest were all generally

phytophagous.

Menon et al. (2007b) in their preliminary observations of mites associated with

rice agroecosystems reported Amblyseius alstoniae, Amblyseius delhiensis, Amblyseius

imbricatus, Amblyseius ovalis, Lasioseius terrestris, Oligonychus indicus,

Steneotarsonemus spinki and Pronematus oryzae.

Atanasov et al. (2008) studied the occurrence of mites on cereal crops in some

regions of Bulgaria and identified the following mites: Tyrophagus longior Gervais,

Penthaleus major Duges, Steneotarsonemus panshini Wainstein and Beglarov and

Steneotarsonemus phragmitidis Schlectendal. The species Steneotarsonemus panshini

and Steneotarsonemus phragmitidis were reported as new for the Bulgarian fauna.

Thus, it is evident that detailed reports of mites occurring on cereals are available

only when the individual reviews at the family level are referred and that an updated

monograph on cereal mites is still not available. Also, owing to taxonomic revisions in


30

the various orders of Acari, the systematic postioning of various families, genera and

species have changed. Thereby, a need to update the mites listed in these catalogues on

the basis of the current and most acceptable taxonomic classification is long pending.

Thus, the present study endeavours to compile relevant literature on cereal mites,

assimilate, consolidate, comprehend and provide these in a ready to use format.

2.5 Acari-crop association checklist

Perusal of literature pertaining to mites associated with cereals crops belonging, in

particular, to the suborder Mesostigmata, Prostigmata and Astigmata (sensu Krantz,

1978), revealed that most of the mites reported on cereals were either phytophagous or

predatory and were reported from both field and/or storage facilities. Thus, about 283

species of mites in 92 genera under 30 families in the suborders Mesostigmata,

Prostigmata and Astigmata of the subclass Acari have been reported from cereal

agroecosystems from across the globe (Table 1).

In Mesostigmata, 42 species of predatory mites in 17 genera and 6 families in the

superfamily Phytoseoidea and Ascoidea had been found in association with cereal crops

(Table 1). Of these, 10 species had exclusively been reported from storage facilities

infesting grains, flour and husk, 31 species had been observed from farm fields and 22

species had been observed to be predatory in field (Table 6). Of these, reports on 7

species, as potential biocontrol agents had been established. Information on applied

aspects of acarology, such as biology, ecology, was available for 15 species. Studies on

host preferences exhibited by cereal associated mesostigmatid mites revealed 23 species

associated with rice, 15 species from wheat and 13 species on maize crops (Table 5).

In Prostigmata, 203 species of mites in 56 genera, and 18 families had been found

in association with cereal crops. Of these, 31 species had been reported from storage

facilities on grains, flour and rotten straw, while others were reported from farm fields.

From the fields, 13 species were observed to be predatory, 9 species were collected from

soil, 6 species were parasitic on other crop pests, while the rest were observed as plant

infesting mites (Table 6).


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The phytophagous mites observed in fields, and in particular, all eriophyid mites,

were mostly found infesting leaves, stem, growing panicle and seeds of standing crop.

Information on applied aspects of acarology was available for 47 species. Studies on host

preferences exhibited by cereal associated prostigmatid mites revealed 90 species

associated with rice, 60 species with wheat, 54 species on maize, 30 species on sorghum

crops and only 3 species on pearl millet (Table 5).

In Astigmata, 38 species of mites in 19 genera and 6 families had been found in

association with cereal crops (Table 1). Of these, 26 species had been observed

exclusively from storage areas of grains, flour and husk (Table 6).

Information on applied aspects was available for 17 species. Studies on host

preferences exhibited by cereal mites revealed that 21 species were associated with rice,

15 species with wheat, 7 species on maize and 3 species each from sorghum and pearl

millet (Table 5).

2.6 Families

During the course of the present study, mites from cereal crops belonging to the

families Ascidae and Phytoseiidae in the order Mesostigmata, families Eriophyidae,

Cunaxidae, Cheyletidae, Neopygmephoridae, Pygmephoridae, Stigmaeidae,

Tarsonemidae, Tetranychidae, Trochometridiidae and Tydeidae in the suborder

Prostigmata and family Acaridae in the cohort Astigmatina have been collected and

studied for their species diversity and taxonomy. Hence, the review presented here

exclusively deliberates on the taxonomy, biology and ecology of the species reported

from cereals of these families only and have been arranged in the order mentioned above.

The taxonomy follows Lindquist et al. in Krantz and Walter (2009).

2.6.1 Ascidae Voigts and Oudemans, 1905: 199–252

Class: Arachnida; Subclass: Acari; Superorder: Parasitiformes; Order:

Mesostigmata; Suborder: Monogynaspida; Superfamily: Ascoidea; Family: Ascidae.

Mites belonging to the family Ascidae are distributed worldwide and are found to

inhabit soil, leaf litter, bee-hives, subcortical situations and are often associated with

Chapter 2- Review of LiteratureAscidae Voigts & Oudemans

32

other animals. Several species are obligatory associates of bees. These constitute an

important group of predators, but can also feed on fungal mycelium. They are mostly soil

inhabitants. Some are parasitic. Nematophagy is also prevalent among soil ascids

(Halliday et al., 1998).

2.6.1.1 Taxonomy

Ascidae is a family of mites in the order Mesostigmata. Lindquist and Evans (1965)

described 22 genera in this family. Several new genera have been added and revised since

then. Presently, the family comprises of more than 700 species in about 39 genera. Major

contributors include Karg (1965, 1980, 1993), Walter and Lindquist (1997), Halliday et

al. (1998) and Christian and Karg (2006).

Ascid mites belonging to the genera Blattisocius Keegan, Iphidozercon Berlese

and Lasioseius Berlese had been found in association with cereal agroecosystem. Perusal

of literature reveals that the family has undergone several generic reorganistions since its

inception. Berlese (1913), in his general classification of Acari, had originally considered

the genus Iphidozercon Berlese and some others in the family Seiidae, as proposed in

‘Acarotheca Italica.’ Later, in 1916, he transferred these genera from Seiidae to

Laelaptidae. Further he erected the genera Lasioseius for Seius Berlese.

Evans (1957) modified concepts in Mesostigmata and placed the genus

Blattisocius Keegan in the family Aceosejidae. Womersley (1956) and Lindquist and

Evans (1965) considered the families Blattisociidae Garman and Aceosejidae Baker and

Wharton synonymous with Ascidae. Lindquist and Evans (1965) placed Iphidozercon

and Lasioseius in the family Ascidae. However, Karg (1965) did not agree with Lindquist

and Evans for the reason that the classification proposed by them did not conform to the

results of phylogenetic investigations.

Karg (1993) characterized families and superfamilies of Mesostigmata based on

synapomorphies as proposed by Henning (1950, 1979) that Lasioseius belonged to the

family Podocinidae. Keys for the determination of 17 families, 13 subfamilies, 203

genera and about 1000 species were proposed (Karg, 1993; Christian and Karg, 1998)

and the Gamasina were divided into 6 superfamilies (Karg 1998a). Christian and Karg


33

(2006) in their generic revision of Lasioseius considered this genus under the family

Podocinidae, taking into account Karg’s classi

1993).

Walter and Lindquist (1997) published preliminary data on Lasioseius, wherein

they referred the genus Lasioseius under Ascidae sensu Lindquist and Evans (1965).

They opined that Karg’s family level classification was based on a restricted set of

characters, and overlooked important sources of information, especially the chaetotaxy of

legs. It was also based on a restricted range of species, which was inadequate for

formulating generic diagnoses and for distinguishing synapomorphies from

convergences.

While the position of the genus Lasioseius in the family (Ascidae or Podocinidae)

is still disputable, majority of acarologists prefer the more complex family-level

classi et al. (1998) which is based on wider set of

characters and is supported by the information on leg chaetotaxy (Evans 1963a, 1963b).

In the family Ascidae, three species in the genus Blattisocius (including one

undetermined species), one undetermined species in the genus Iphidozercon and 6 species

in the genus Lasioseius had been reported thus far from cereal crops, totaling to 10

species.

Menon and Ghai (1968) described Lasioseius terrestris as predatory mite seen in

association with Petrobia latens (Muller) from wheat fields in IARI farms, Delhi, India.

Lo and Ho (1977) reported Lasioseius sugawarai Ehara as predator on eggs and adults of

Steneotarsonemus spinki Smiley and Tarsonemus sp. from rice fields. Pai and Prabhoo

(1981) collected Lasioseius extremus (Daneshvar) from rice storage houses.

Wu and Wang (1982) described Gnorimus chaudhrii, collected on leaves and

stem of rice and observed these were predatory on Tarsonemus sp. Tseng (1984) and Lo

and Ho (1984) reported Lasioseius parberlesei Bhattacharyya and Lasioseius youcefi

Athias Henriot, from rice fields as predator of Steneotarsonemus spinki, respectively.

Mahmood (1992) reported Iphidozercon sp. and Lasioseius berlesi (Oudemans) from

storage areas of rice, wheat and barley in Iraq.


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Chant and McMurty (1994) proposed generic synonymy of Gnorimus Chaudhri

with Lasioseius, which was supported by Walter and Lindquist (1997). Walter and

Lindquist (1997) reviewed all Australian species of the porulosus group in the genus

Lasioseius Berlese, described five species, gave new collection records, diagnosis and

biological information for two previously known Australian species, presented a key to

Australian species of Lasioseius and proposed new synonyms for Lasioseius, namely

Gnorimus, Indiraseius Dansehvar, and Neolaspina Halliday.

Walter and Lindquist (1997) clarified that the genus Gnorimus, originally

ascribed to subfamily Gnoriminae under Phytoseiidae belonged to the ascid tribe

Blattisociini (sensu Lindquist and Evans, 1965), and Gnorimus was equivalent to the

phytoseioides species-group in Lasioseius and that the type species Gnorimus tabella

Chaudhri was doubtfully distinct specifically from Lasioseius phytoseioides Chant. Also,

Gnorimus was not an available name, as it was preoccupied by Gnorimus Audinet-

Serville (1825) (Coleoptera, Scarabaeidiae).

In proposing Indiraseius for the same species-group, Walter and Lindquist

believed that Dansehvar (1987) apparently was not aware of Chaudhri’s genus name and

so they synonymised the genus and provided new combinations for six species described

under the genus including Indiraseius extremus Daneshvar which was reported from rice

as Lasioseius extremus. Again, Halliday (1995) proposed a new genus of

Parholaspididae, Neolaspina; however, Walter and Lindquist (1997) concluded that the

type species Neolaspina rugosa Halliday was in fact a member of the genus Lasioseius.

Halliday et al. (1998) provided the most comprehensive account of mites in this

family known from Australia and described 14 genera and 26 species. New species were

described, new synonyms and combinations were proposed, new host and locality records

were given, lectotypes were designated for two species and key to the world genera and

Australian species was provided.

Rao et al. (1999) listed four species, namely Blattisocius keegani Fox,

Blattisocius tarsalis (Berlese), Lasioseius sugawarai and Lasioseius youcefi in their

checklist of mites associated with cereals in rice agroecosystems. Similar checklist by


35

Joshi et al. (2002) listed Blattisocius spp., Indiraseius extremus and Lasioseius berlesi

(Oudemans) mainly from storage facilities. Eliopoulos et al., (2003) reported Blattisocius

keegani and Blattisocius tarsalis from storage facilities of wheat, maize, oats and barley.

2.6.1.2 Biology

The lifecycle of Blattisocius keegani was studied by Barker (1967) on eggs of

Cryptolestes turcicus at 80°F, and 70 to 75% and 95 to 100% RH. Zhang and Lin (1991)

studied the biology of Gnorimus chaudhrii in the laboratory and concluded that the

duration from egg-adult decreased with increase in temperature.

2.6.1.3 Ecology

Barker (1967) reported that Hokbal 40% EC was the only pescticide that was least

toxic to the predatory mite Blattisocius keegani. The feeding behavior, preferred plant

part, predation and population dynamics of Lasioseius parberlesei on Steneotarsonemus

spinki Smiley in rice field were documented by Tseng (1984). This study concluded that

Lasioseius parberlesei could keep the pest mite Steneotarsonemus spinki below injurious

level in Taiwan. Lo and Ho (1984) showed that the predatory mite Lasioseius youcefi

exhibited better rate of development, food intake and reproductive output when fed on

eggs of Tarsonemus sp., than when fed on Steneotarsonemus spinki.

Zhang and Lin (1991) gave detailed account of the biology, predation potential,

description, mating behavior and population studies of Gnorimus chaudhrii. They

observed the mite to be an important natural enemy contributing to biological control of

Tarsonemus bilobatus Suski in rice fields in Fuzhou. Its population peak followed behind

that of the tarsonemid by a week or so and then there was a rapid decline in the

tarsonemid mite population. Franzolin and Baggio (2000) reported that populations of

Blattisocius tarsalis showed a greater reproductive rate during spring, summer and at the

beginning of autumn, due to high temperatures and humidity.

2.6.2 Phytoseiidae Berlese 1916a: 33

Chapter 2- Review of LiteraturePhytoseiidae Berlese

36

Class: Arachnida; Subclass: Acari; Superorder: Parasitiformes; Order:

Mesostigmata; Suborder: Monogynaspida; Superfamily: Phytoseioidea; Family:

Phytoseiidae.

Mites in the family Phytoseiidae are mainly predators of plant feeding mites and

small soft bodied insects like scales, aphids, thrips, whiteflies and their eggs (Nesbitt,

1951; Muma, 1955).

2.6.2.1 Taxonomy

Nesbitt (1951) brought out the first major taxonomic review of the family in

which they incorporated 41 species. Since then the taxonomy of phytoseiids have

advanced relatively fast with several additions to the phytoseiid fauna and various

opinions and disagreements on the system of classification. Some of the major

contributions to this family are by Beard (2001), Chant (1959, 1965), Chant and

McMurtry (2003a, 2003b, 2004, 2005a, 2005b, 2005c, 2006), Gupta (1970, 1975, 1977,

1978, 1980, 1985, 1986, 2003), Moraes and Denmark, (1999), Moraes and Flechtmann

(2008), Moraes and McMurtry (1982, 1983, 1988), Moraes and Mesa (1988), Moraes and

Oliveira (1982), Moraes et al. (1982, 1986, 1987, 1988, 1989a, 1989b, 1989c, 1990,

1991, 1994, 1997, 2000, 2001a, 2001b, 2004), Muma (1955, 1961, 1962, 1965, 1968),

Muma and Denmark (1968, 1969a) and Wu et al. (1997, 2009, 2010).

Of these, the work of Chant and McMurtry (2003-2006) is of particular

significance as these include a series of revisionary studies on the subfamily

Amblyseiinae Muma. In this study they recorded 1499 nominal species in the subfamily

Amblyseiinae, almost all of which were exclusively plant inhabiting and recorded from

every continent except Antarctica, maximum number being reported from tropical and

subtropical regions. They proposed 61 genera under 9 subtribes in 9 tribes in the

subfamily of which, Afroseiulini Chant and McMurtry, Euseiini Chant and McMurtry,

Neoseiulini Chant and McMurtry, Phytoseiulini Chant and McMurtry and

Typhlodromipsini Chant and McMurtry were newly erected. Several subgenera were

raised to generic level and subsequent new combinations were recommended for the

species under them. The paper gave clarifications for the various transformations


37

presumed to have occurred in the subfamily, provided a diagnosis and key to the

constituent tribes, and proposed a cladogram suggesting the relationships between the 9

tribes.

Wu et al. (2009) conducted studies on phytoseiid mites from China and recorded

three subfamilies, 15 genera and 307 species in total (including 36 new species) in the

Fauna Sinica volume, among which 204 species were endemic to China. Similarly, Wu

et al. (2010) reviewed the research on the systematics of the family Phytoseiidae in

China, and provided an updated checklist encompassing 304 species.

Thus, over 2500 phytoseiid mites in about 70 genera had been described

worldwide (Moraes et al., 2004) and more than 20% of these species had been found

useful in the biological control of mites and insect pests of agriculturally important crops.

So far, 24 phytoseiid species in 8 genera had been recorded from cereals.

Lo et al. (1979) observed Amblyseius taiwanicus Ehara as predatory on

tarsonemid mites on paddy crop in Taiwan. Gupta described Amblyseius

(Typhlodromalus) sorghumae from maize fields in India. Daneshvar (1987) reported

Amblyseius imbricatus Corpuz-Raros and Rimando on rice from Iran. Rao et al. (1999)

listed Amblyseius fallacis Garman, Amblyseius imbricatus, Amblyseius indicus Narayanan

and Kaur, Amblyseius longispinosus Evans, Amblyseius multidentatus Swirski and

Schechter, Amblyseius ovalis Evans and Amblyseius taiwanicus from rice

agroecosystems. Similarly, Joshi et al. (2002) listed 7 species from rice agroecosystems,

namely Amblyseius fallacis, Amblyseius longispinosus, Amblyseius ovalis, Amblyseius

paraibensis Moraes, Amblyseius multidentatus, Gnorimus chaudhrii Wu et al., and

Neoseiulus oryzacolus Daneshvar.

Gupta (1986, 2003) in his monograph on predatory mites of India listed

Amblyseius (Euseius) finlandicus (Oudemans), Amblyseius (Euseius) ovalis, Amblyseius

(Neoseiulus) fallacis, Amblyseius (Neoseiulus) paspalivorus (De Leon) and Amblyseius

(Typhlodromips) officinaria Gupta from rice; Amblyseius crotalariae Gupta, Amblyseius

(Euseius) alstoniae (Gupta), Amblyseius (Euseius) finlandicus, Amblyseius (Neoseiulus)

indicus (Narayanan and Kaur), Amblyseius (Typhlodromalus) kalimpongensis Gupta,


38

Amblyseius (Typhlodromalus) sorghumae Gupta, Amblyseius (Typhlodromips)

suknaensis Gupta, Amblyseius (Typhlodromips) syzygii Gupta from maize and

Amblyseius (Neoseiulus) indicus from wheat.

However, all these species have undergone generic revision and new

combinations and synonyms were proposed as follows: Amblyseius fallacis as Neoseiulus

fallacis by Denmark and Muma (1973); Amblyseius ovalis as Euseius ovalis by Gupta

(1978); Matthysse and Denmark 1981 synonymised Amblyseius multidentatus Swirski

and Schechter with Paraphytoseius orientalis (Narayanan, Kaur and Ghai); Chant and

McMurtry (2003a) elevated the subgenera Neoseiulus to generic status and provided the

following new combinations: Neoseiulus imbricatus, Neoseiulus indicus and Neoseiulus

taiwanicus; Moraes et al. (2004) gave the new combination Neoseiulus paraibensis

(Moraes and McMurtry) and Faraji et al. (2007b) synonymised Amblyseius oryzacolus

with Neoseiulus imbricatus.

2.6.2.2 Biology

Rock et al. (1971) reported that females of Neoseiulus fallacis (Garman)

underwent reproductive diapause. Lo et al. (1979) studied the development duration of

Amblyseius taiwanicus (Ehara) under controlled laboratory conditions and commented on

its predator potential on the rice panicle mite Steneotarsonemus spinki Smiley. Kumari

and Sadana (1991) gave an account of reproduction and development of Amblyseius

alstoniae Gupta with respect to variations in temperature and RH.

El-Laithy and Fouly (1992) studied the life history of Euseius scutalis (Athias-

Henriot) when fed on nymphs of Tetranychus urticae Koch at constant conditions and its

life tables and effeciency as biological control agents were analysed. Nomikou et al.

(2003) studied the impact of pollen and white -produced honeydew on the life history

of Euseius scutalis (Athias-Henriot) as the phytoseiid was able to suppress white

populations on single plants and was a candidate for biological control of Bemisia tabaci

(Gennadius).

Kasap and Sekeroglu (2004) determined the biology and reproductive potential

of Euseius scutalis (Athias-Henriot) at various temperatures. Momen and El-Sawi (2008)


39

reported the life history and reproductive parameters of Euseius scutalis (Athias-Henriot)

and evaluated its potential as a predator of Spodoptera littoralis, Spodoptera exigua

and Agrotis ipsilon eggs as alternative diets in the laboratory. Momen and Khalek (2009)

investigated the juvenile survival, predation and development in Euseius scutalis and

other phytoseiid mites when fed on con- and heterospecific phytoseiid immatures in the

laboratory.

Kholoud (2010) reared Euseius scutalis on immature stages (larvae and nymphs)

of the tetranychid mites, Tetranychus urticae Koch, Oligonychus afrasiaticus McGregor

and Eutetranychus orientalis Klein in the laboratory and compared the differences in

development duration and also observed its predator potential on these tetranychid mites.

2.6.2.3 Ecology

Meyerdirk and Coudriet (1986) concluded from their test studies that of the two

predaceous mite biotypes of Euseius scutalis (Athias-Henriot), collected from Lantana

sp. in Jordan and on citrus from Morocco, the Jordan variety was a better biological

control agent of Bemisia tabaci. Tanigoshi et al. (1990) observed the seasonal trend in

population of Eutetranychus orientalis and Euseius scutalis in Jordan valley on citrus

plantation and concluded that Euseius scutalis was an effective predator of Eutetranychus

orientalis.

Berry et al. (1991) concluded that humidity and temperature were critically

important in the population dynamics of Neoseiulus fallacis on maize. Noorbakhsh and

Kamali (1995) reported Neoseiulus bicaudus (Wainstein), Neoseiulus marginatus

(Wainstein), Neoseiulus zwoelferi (Dosse), Neoseiulus harrowi (Collyer) and

Proprioseiopsis messor (Wainstein) as predator and biocontrol agent of brown wheat

mite, Petrobia latens (Muller) in Iran. Likewise, Petrova et al. (2000) reported

Neoseiulus bicaudus to be potential predators of the strawberry mite Tarsonemus pallidus

(Banks) and spider mite Tetranychus urticae Koch.

Schausberger (1997, 1998, 1999) conducted a series of experiments with Euseius

finlandicus and observed its interspecific predation and cannibalism on immature forms

of Typhlodromus pyri Scheuten and Kampimodromus aberrans (Oudemans), its juvenile


40

survival, development and reproduction when fed on crawlers of the diaspidid San José

scale Quadraspidiotus perniciosus and when fed on con- and heterospecific phytoseiid

immature forms.

Kwon et al. (1998) studied the egg to adult duration, average longevity, total

fecundity, total egg consumption/day, critical temperature and effective degree-days

(DD) of Neoseiulus longispinosus and Neoseiulus fallacis under four constant

temperatures when fed on Tetranychus urticae.

Salwa et al. (2000) estimated the effect of temperature and relative humidity on

different biological aspects of the predaceous mite Euseius scutalis. Adar et al. (2012)

monitored the plant feeding behavior of Euseius scutalis and proposed that the ability to

feed on plants can be added as a cross type trait of phytoseiid life-style types.

Lawson-Balagbo et al. (2007) compared the developmental time, oviposition rate,

intrinsic rate of increase in egg deposition of Neoseiulus paspalivorus when fed with

Aceria guerreronis Keifer, Tetranychus urticae Koch, coconut pollen and

Steneotarsonemus furcatus De Leon. Kholoud (2011) tested four kinds of plant pollen as

an alternative food source for rearing Euseius scutalis and concluded that the tested

pollen food was a suitable alternative food source and met the nutritional requirements of

the phytoseiid mite.

2.6.3 Cheyletidae Leach, 1815: 399

Class: Arachnida; Subclass: Acari; Superorder: Acariformes; Order:

Trombidiformes; Suborder: Prostigmata; Superfamily: Cheyletoidea; Family:

Cheyletidae.

Cheyletidae is a family of mites in the Trombidiformes. The mites in this family

are either associated with parasitism in birds and mammals or are free-ranging predators

found in soil, forest litter, tree bark, foliage, bird-nests, animals and in house dust.

2.6.3.1 Taxonomy

The family Cheyletidae was erected by Leach (1815). Baker (1949) brought out

the first taxonomic review of the family in which they included 19 genera. Later,

Chapter 2- Review of LiteratureCheyletidae Leach

41

Lawrence (1954) reviewed the African cheyletid species. Volgin (1960, 1961, 1969)

recognized 10 tribes under 54 genera in this family. The species list was expanded to

about 186 species under 51 genera by Summers and Price (1970).

Gerson et al. (1999) further provided a key to 76 genera of Cheyletinae and listed

more than 400 species. Fain et al. (1997) studied the chaetotaxy and solenidiotaxy of

cheyletid mites.

More recently Fain and Bochkov (2001a, 2001b, 2001c) provided key to genera

and species and made important taxonomic contributions to this family. Bochkov and

Fain (2001) presented the first phylogenetic analysis of intrafamilial relationships, and

established 13 tribes which perfectly corresponded to the tribes proposed by Volgin

(1969). Presently, more than 500 species in about 80 genera of Cheyletidae are known

from the world.

Volgin (1969, English translation 1987) provided information on 172 species of

cheyletid mites listed under 54 genera. Herein, he recorded 9 cheyletids from cereals,

mainly in storage. Thereafter, several revisions on its genera and species had taken place

and many new species were added. Perusal of literature revealed that 23 mite species in

11 genera were associated with cereals- rice, wheat, maize, sorghum and pearl millet, in

particular, from field as well as storage.

Aheer et al. (1991) described two species of Cheyletidae from wheat in Pakistan,

namely, Acaropsis vitrus and Acaropsis platessa, gave key to females in the genus

Acaropsis and presented a phenogram. Aheer et al. (1994) described three species of the

genus Cheletomimus, of which Cheletomimus cambio was collected from rotten wheat

grains.

Fain and Ardeshir (2000) described Neoeucheyla iranica from floor debris in a

silo that had contained wheat grain, near Tehran, Iran. Eliopoulos and Papadoulis (2001)

gave new records for five species namely, Acaropsellina sollers (Kuzin) (collected in

floor litter in a flour mill), Cheletomorpha lepidopterorum (Shaw) (from hay, wheat,

cotton seed, trefoil, litter and maize), Cheyletus aversor Rohdendorf (from hay, cotton

seed, barley, litter, wheat, corn flour and floor litter in a flour mill), Cheyletus trouessarti


42

Oudemans (from maize, medic flour and litter in a barn) and Cheyletus trux Rohdendorf

(from cotton seed, barley, bran, litter of cattle feed, floor litter in a flour mill, maize and

floor litter in a barn) from Greece and also described Chelacheles hellenicus from floor

litter in a flour mill at Votanikos, Athens.

Fain and Bochkov (2001a) in their exemplary revision of the genus Cheyletia

synonymised 28 species, provided 4 new combinations, described a species and

considered 15 species as species inquirendae. In addition, this study clarified the

placement of Cheyletus pyriformis Banks in the genus Paracheyletia. The species

Cheyletus pyriformes was transferred to Cheyletia by Baker (1949) and later Volgin

(1969) reassigned it to Paracheyletia Volgin, designating Paracheyletia assimilis Volgin

as its type species.

Again, Fain and Bochkov (2001b) reviewed some genera of Cheyletidae,

described a new genus, 8 new species, synonymised 21 species, and considered 14

species as species inquirendae. Xia et al. (2004) reviewed Eucheyletia in China, provided

key to species and described Eucheyletia omissa from paddy.

2.6.3.2 Biology

Rizk et al. (1980) gave a detailed account on the biology of Acaropsis sollers

Kuzin attacking stored product pests. Eggs and newly hatched larvae of Tribolium

confusum Jacquelin du Val and Lasioderma serricorne (Fabricius) and adults of

Dermatophagoides farinae Hughes were provided as prey, and the development duration

of females was found to be longer than males. The sex ratio also varied according to the

age of the parent female and depended on whether the progeny were the result of first or

second mating with the male. Females lived longer than males.

The biology of Cheyletus malaccensis Oudemans was studied by Shen (1975)

using Tyrophagus putrescentiae (Schrank) as prey and later, Nangia et al. (1995) studied

its biology in laboratory using eggs of Corcyra cephalonica.

Emekci and Toros (1994) and Xia et al. (2005b) studied the biology of Cheyletus

eruditus (Schrank) in laboratory at different temperatures and humidity and observed


43

other parameters like the net reproductive rate, mean generation time, finite rates of

increase and maximum longevity.

Zhu et al. (2000) studied the development period of Cheyletus trouessarti at four

constant temperatures and deduced the developmental threshold temperature and

effective accumulative temperature.

2.6.3.3 Ecology

Barker (1991) studied the duration and mortality of immature stages of Cheyletus

eruditus at 18.5°C, 22°C, and 25°C and 76% RH. Life tables and schedules of female

eggs laid by each female per day were made from records of daily oviposition and adult

mortality. Other parameters like the net reproductive rate, mean generation time, finite

rates of increase and maximum longevity were also studied.

Mahmood (1992) studied the mite fauna of stored grain in central Iraq in a total of

108 samples of wheat, barley and rice and reported Cheyletus malaccensis on rice and

wheat. Emmanouel et al., (1994) surveyed the mites associated with wheat, maize, barley,

wheat flour, bran and residues in silos, farm stores and flour mills in Greece in 1990-91

and provided first record for Acaropsis docta (Berlese) (Acaropsellina docta). The

findings revealed that Cheyletus eruditus was the most abundant species and that farm

stores were more infested than other storage facilities.

Zhang and Lin (1996) and Zhang et al. (1997) studied predatory effect of

Cheyletus malaccensis on Lepidoglyphus destructor (Schrank) and reported each

predatory mite consuming about 10–12 prey per day and five hundred mites in its entire

life history. The study analysed the model of the relationship of systematic number

between Cheyletus malaccensis and Lepidoglyphus destructor.

Cobanoglu (1996) recorded Cheyletus trouessarti for the first time in Turkey

while studying mite pests infesting rice, wheat and sunflower seed in storage sites.

Szlendak (1998) collected samples of stored cereals including wheat, oats, maize, barley,

rye and triticale, harvested mainly in 1996, from small Polish farms located in 3 different

provinces of Central Poland. The quality of cereals was examined at the first stage of


44

grain production (sampled from farm storage) which revealed Cheyletus eruditus as the

only species of predatory mite from all the samples.

Zdarkova (1998) carried out surveys in 64 grain stores (brick stores, hangars,

silos) in the Czech Republic during spring and autumn 1996. A total of 112 samples of

wheat, 74 samples of barley and 70 samples of sweepings were examined. At an average,

moisture content was recorded between 13-13.6% in wheat and barley samples and 90%

of the samples were found infested, of which barley was more infested. A total of 32

species of mites were found, in which Cheyletus eruditus was the dominant species.

Franzolin and Baggio (2000) evaluated mite contamination rate in 23 samples of

polished rice and 53 samples of beans sold in nine municipal markets of the city of Sao

Paulo, in November 1989 - 1990. Samples were examined once a week, at intervals over

42 days at air temperature and also at 25°C and 75% RH for 28 days. No mites were

observed on the first day of analysis but were detected after incubation. Samples

incubated revealed a higher percentage of positive examinations than samples kept at air

temperature. Polished rice was more contaminated in comparison with those of beans.

There were more mites when the mean monthly temperature of the laboratory was

between 21.5°C and 22.5°C (37.8%) and RH was between 73.5% and 74.5% (31.1%).

Mite populations showed greater reproductive rate during spring, summer and at the

beginning of autumn, due to high temperatures and humidity. Cheyletus spp., amongst

others was one of the dominant species.

Ardeshir et al. (2000) conducted a survey of mites associated with stored grain in

northern Iran in December 1996. Cheyletus malaccensis Oudemans was the most

common species. The highest number of species and number of individuals were

recorded in November on rice. This study concluded that the intriguing absence of other

major species known to infest stored grain (e.g. Acarus siro Linnaeus) resulted from

predation pressure and/or chemical control during and before the period of sampling.

Putatunda (2002) in his preliminary survey on contamination of stored food

products by mites in Himachal Pradesh, observed lesser populations of predatory mite,

Cheyletus malaccensis (23%), as compared to the other acarine pests. Habibpour et al.


45

(2002) recorded Acarapsis docta (Acaropsellina docta) and Cheyletus malaccensis from

stored products in Khuzestan province, Iran, while conducting a faunistic survey of insect

and mite pests and their enemies.

Athanassiou et al. (2003) examined the seasonal abundance of insect and mite

species, in relation to their horizontal and vertical spatial distribution in the three flat

storerooms, filled with approximately 45 tonnes of wheat, stored in 1.5 m deep bulks, in

Southern Greece. Wheat samples were taken from June 2000 until March 2001, at 10-day

intervals, by using a partitioned grain tier. The surface of the grain bulk was divided into

central, edge and the corner zones. The tier samples were examined separately for the

upper, medial and lower 0.5m of the bulk. While significant differences were recorded

for moisture content among zones (P<0.0001) and among depths (P=0.0012), no

significant differences were noted for temperature among zones (P=0.2281) and depths

(P=0.3049). In general, the moisture content was higher in the upper 0.5 m of the bulk,

temperature decreased during the sampling period more vigorously from October

onwards. Eight insect species and 25 mite taxa were found during the sampling period.

The most abundant mite species was Acaropsis sollers (Acaropsellina sollers) with the

highest population being recorded during autumn. Acaropsis sollers was equally

distributed in the upper 1 m and showed an aggregated spatial pattern, as indicated by

Taylor's Power Law estimates.

Eliopoulos et al. (2003) surveyed the predatory mite fauna of stored products-

wheat, maize, oats, barley, wheat flour, maize flour, figs, raisins and sultanas in Greece

(1999-2000) stored in large concrete silos, warehouses, flat granaries, farm stores, flour

mills and household premises. The greatest presence of predatory mites was recorded in

grain, the most abundant and frequent species being Cheyletus malaccensis, followed by,

Cheyletus eruditus and Acaropsis docta (Acaropsellina docta).

Hubert et al. (2003) subjected Cheyletus malaccensis and few other mites to

mycological analysis on stored seeds (wheat, poppy, lettuce, mustard). They concluded

that the fungal transport via mites was selective and was experimentally supported by (i)

lower number of isolated fungal species from mites than from seeds; (ii) lower Shannon-

Weaver diversity index in the fungal communities isolated from mites than from seeds;


46

(iii) significant effect of mites/seeds as environmental variables on fungal presence in a

redundancy analysis (RDA) and (iv) differences in composition of isolated fungi between

mite species shown by RDA. These results supported the hypothesis that mite-fungal

interactions are dependent on mite species. Hubert et al. (2003) concluded that fungal

transport via Cheyletus malaccensis was selective.

Pascual Villalobos and Estal (2004) studied the pests and natural enemies present

in Calasparra rice stores during 2001 and 2002 by trapping and sampling methods. The

presence of Cheyletus malaccensis was associated with temperature and relative

humidity. Zd'arkova and Horak (2004) recommended the biological control of storage

mites using Cheyletus eruditus for temperatures up to 30°C. Xia et al. (2007a) studied the

predation of Cheyletus eruditus on Aleuroglyphus ovatus at six constant temperatures.

Athanassiou et al. (2005) assessed the spatio-temporal distribution of stored-

product insects and mites in a wheat granary setup, during June 2001 until March 2002

using contour analysis. Nine insect and 20 mite taxa were found; the most common

predator being Cheyletus malaccensis Oudemans. The highest population densities were

recorded during autumn and in the upper 0.5m of the bulk; Cheyletus malaccensis was

equally distributed in the upper and medial 0.5m of the bulk. The spatio-temporal

distribution during the entire experimental period notably varied according to the insect

and mite species.

Putatunda (2005) sampled food grains, food products, and animal feeds stored

under various conditions in Himachal Pradesh, for the presence of stored products mites.

Cheyletus malaccensis was recorded amongst the four predators that kept the population

of storage mites at a low level.

Hubert et al. (2006) selected 78 grain stores and sampled grain mass and residues

in order to compare concurrent mite communities in these two different habitats. This

study concluded that though the residue samples had more mites and higher species

diversity than the stored grain mass, no correlation in mite abundance and species

numbers between samples from grain residues and grain mass was found, thereby

indicating low connectivity of these two habitats. Cheyletus eruditus was seen associated


47

with grain residues and Cheyletus malaccensis was associated with both grain mass and

grain residues.

Mahgoob et al. (2006) reported Cheyletus malaccensis grain residues and mixed

flour samples collected from warehouses and mills at Great Cairo, Egypt. Palyvos and

Emmanouel (2006) evaluated population dynamics of stored-product mites and their

predators in connection with horizontal and vertical distribution in a flat store room in

central Greece, filled with approximately 60 tons of newly harvested wheat. At 15 day

interval, from July 2000 until March 2001, three replicates were taken at depths of 0-40,

40-80 and 80-120cm, with a partitioned grain tier. Twenty four mite taxa were found,

Acaropsis docta (Berlese) being one of the two dominant species. More individuals were

found in the center as compared to the periphery of the wheat bulk, and at the surface of

the bulk than at greater depths. The highest mite population density for the total mite

species was recorded during October-November and after the middle of January.

Palyvos and Emmanouel (2006) conducted population density studies on

Acaropsellina docta. Xia et al. (2007b) studied the predation of Cheletomorpha

lepidopterorum on Tyrophagus putrescentiae at six constant temperatures with maximum

predation by adult female at 28°C.

In a study of mite samples collected from stored wheat, straw and dust in silos

from flour-mills in Tehran, Karaj and Varamin, Iran, during spring 2005, Cheyletus

eruditus and four other Acarina were newly recorded from wheat stores. The most

abundant predatory mite was Acaropsellina sollers. The highest observed number of

mites was in Tehran Silo, Ard-Iran flour-mill and Jafar Joshaghani mill (Ardeshir et al.,

2008).

2.6.4 Cunaxidae Thor, 1902: 159-165


Trombidiformes; Suborder: Prostigmata; Superfamily: Bdelloidea; Family: Cunaxidae.

Family Cunaxidae was erected by Thor in 1902 to separate mites having four

segmented palp from the family Bdellidae. The family comprises of small soft-bodied

Chapter 2- Review of LiteratureCunaxidae Thor

48

mites which are very important predators of other harmful mites and small insects of

agricultural importance.

2.6.4.1 Taxonomy

Since the erection of this family by Thor in 1902, important taxonomic

contributions were made by Ewing (1917), Thor and Willmann (1941), Muma (1960) and

Smiley (1975). Baker and Hoffman (1948) and Den Heyer (1976, 1977a, 1977b, 1978,

1979a, 1979b, 1981) described and illustrated a number of cosmopolitan species. Meyer

and Ryke (1959b) discussed the cunaxid mites occurring on plants from South Africa.

Den Heyer (1978) provided keys to genera and in 1981 discussed the systematics of the

family. Tseng (1980) conducted taxonomic studies of cunaxid mites from Taiwan. Walter

and Kaplan (1991) reviewed the feeding habits of the family. Chaudhri (1977),

Inayatullah and Shahid (1993) and Muhammad and Chaudhri (1992) were amongst the

major contributors to the cunaxid fauna of Pakistan. Smiley (1992) revised the world

species with keys to subfamilies, genera and species.

Presently, the family comprises of 27 genera with more than 170 species reported

from the world (Smiley, 1992). So far, only 8 cunaxid mites in two genera, Coleoscirus

Berlese and Cunaxa Von Heyden had been reported from cereals.

Berlese (1916b) erected the genus Coleoscirus and included two species

Coleoscirus halacaroides and Coleoscirus corniculatus. Den Heyer (1979) examined

Coleoscirus corniculatus and compared them with the specimens of Scirus curtipalpis

Berlese and proved these to be conspecific. Thus, Coleoscirus corniculatus was

considered as a synonym of Scirus (= Coleoscirus) curtipalpis.

Berlese (1916) designated Coleoscirus halacaroides as type species of

Coleoscirus, despite the earlier known Coleoscirus curtipalpis Ewing. Smiley (1975)

provided the new genus name Pseudocunaxa, for species related to and including

Coleoscirus simplex (Ewing). Den Heyer (1979a) synonymised Pseudocunaxa under

Coleoscirus. In 1984, Sepasgosarian, perhaps unaware of the above change, also

mentioned that Pseudocunaxa was a synonym of Coleoscirus and included Coleoscirus

in subfamily Coleoscirinae and erected the new tribe, Coleoscirini and genus Scutascirus.


49

Three species in the genus Coleoscirus are known from cereal crops, viz.,

Coleoscirus buartsus den Heyer, new record from India, collected from soil of rice,

wheat and maize during the present study, Coleoscirus disparis Muhammad and

Chaudhri (1992) and Coleoscirus mardi Inayatulla and Shahid (1993) both reported on

paddy from Pakistan.

The genus Cunaxa is the largest genus of this family which comprises over 50

species. This genus was erected by Von Heyden in 1826, with Scirus setirostris Hermann

(1804b) as its type species. Den Heyer (1979b) created a new genus Rubroscirus for

cunaxid mites having single seta (instead of two) on coxa IV, reticulated dorsal shields

and inconspicuous macro setae on legs III-IV. Smiley (1992) considered these characters

of specific level and synonymised Rubroscirus with Cunaxa. So far, five species in this

genus had been reported from cereals.

Hermann (1804b) described Scirus setirostris from rice. This was later transferred

to Cunaxa and the new combination was proposed by Von Heyden (1826). Thor and

Willmann (1941) in their major work on the families Eupodidae, Penthalodidae,

Penthaleidae, Pachgnathidae and Cunaxidae, synonymised six species of Scirus with

Cunaxa setirostris. From India, Prasad (1974) and Gupta (1992) listed these mites in their

catalogue.

Chaudhuri (1980) described Cunaxa doxa from wheat plant. Bashir et al. (2010,

2011) described three new species from rice husk, namely, Cunaxa leuros, Cunaxa

nankanaensis and Cunaxa rafiqi from Pakistan.

2.6.4.2 Biology

Arbabi and Singh (2000) gave detailed account of the lifecycle of Cunaxa

setirostris on Tetranychus ludeni Zacher. The females of Cunaxa setirostris completed

their lifecycle with one larval stage followed by three nymphal stages while the male

attained early maturity and the lifecycle comprised of one larval stage with two nymphal

stages. The female predators were generally fast runners and were found feeding only on

active stages of prey. The maximum amount of feeding occurred during oviposition.

Webbing produced by Tetranychus ludeni on leaf lamina had no negative effect on the


50

amount of prey consumption. The total mortality during developmental period of this

predator was approximately 80% and maximum mortality (48%) was during larval

period.

2.6.5 Eriophyidae Nalepa, 1898: 5


Trombidiformes; Suborder: Prostigmata; Superfamily: Eriophyoidea; Family:

Eriophyidae.

The family Eriophyidae comprises of tiny, microscopic worm like mites which

have only two pairs of legs. Body colour may vary from being transparent, to yellow,

pink, white or purple. Natural agents like wind, rain, physical contact and insects help in

dispersal. Eriophyid mites are known to infest a wide range of plants and several are

known pests of crops causing substantial economic loss. Some species, however, are also

deployed as biological control agents.

2.6.5.1 Taxonomy

This family was erected in 1898 by Nalepa who based the etymology on the genus

Eriophyes Siebold (1850). Keifer (1938-79) described and/or redescribed majority of

economically important species in the family. Buhr (1964-65) provided key to plant gall

mites of north-western Europe. ChannaBasavanna (1966) provided a comprehensive

review of Indian eriophyid mites. This study gave details of 61 species, of which 44

species in 14 genera and 2 families were newly described.

Jeppson et al. (1975) gave an excellent introduction to morphology and

classification with key to genera. Davis et al. (1982) in their catalogue of gall-mites

recorded 1859 species assigned to 156 genera and also provided host-plant index. Keifer

et al. (1982) produced an illustrated guide to many North American gall mites along with

their life history, distribution and host data.

Manson (1984a) in his voluminous work on Eriophyoidea of New Zealand

(except Eriophyinae) enumerated 49 species under 3 families, 6 subfamilies, and 31

genera including 12 new genera. This work provided a brief historical review of the

Chapter 2- Review of LiteratureEriophyidae Nalepa

51

superfamily with notes on techniques for mounting, morphology, lifecycle, classification,

and economic importance. Further, the study described 25 new species, gave 32 new

records, enlisted 17 species from New Zealand and maintained host-plant records.

Again, Manson (1984b) published compiled information on the mites of the

subfamily Eriophyinae from New Zealand. This volume enumerated 60 species of

eriophyids, described 29 new species in 6 genera including 2 new genera, gave 30 new

records and proposed 6 new combinations alongwith host-plant records.

Boczek et al. (1989) provided key to 209 genera. Amrine and Stansy (1994)

provided an updated catalogue of Eriophyoidea, with a comprehensive species and host

index. Hong and Zhang (1996) enumerated 205 species of eriophyid mites belonging to 3

families, 9 subfamilies and 77 genera from China. Lindquist et al. (1996) provided the

most comprehensive and updated review of morphology, systematics, biology, natural

enemies and control of eriophyid mites.

Skoracka (2004) provided key to 6 genera and 29 species of eriophyids infesting

grasses in Poland. Some more major contributions on eriophyid mites pertaining to

localized regions were from Mohanasundaram, (1980a, 1980b, 1981a, 1981b, 1982a,

1982b, 1983a-c, 1984, 1996), Chakrabarti and Mondal (1980), Das and Chakrabarti

(1985), Mondal and Chakrabarti (1979, 1980, 1981, 1982a, 1982b), Mondal, Ghosh and

Chakrabarti (1982), Boczek and Chandrapatya (1989a, 1989b, 1996a, 1996b, 1998a,

1998b, 2000a-g, 2002) and Chandrapatya and Boczek (1991a-c, 1992, 1993a-b, 1996,

1997a-b, 1998, 2000a-g, 2001a-c, 2001a, 2001b).

Menon et al. (2010) compiled a checklist on the species diversity of the genus

Abacarus Keifer, discussed the salient aspects of its faunistics and reported a new species

Abacarus sorghi from sorghum crop.

Thus, about 3,600 species have been described in more than 240 genera, which is

only a tenth of the actual number of eriophyids existing in the world. So far 20 species of

eriophyid mites in 5 genera are known from cereals. However, comprehensive

information on the bionomics of these eriophyids is found lacking.


52

Nalepa first described Phytoptus tenuis in 1891 from type host Bromus

hordeaceus. Liro and Roivainen (1951) provided wheat as a new host record for Aceria

tenuis (Nalepa). Keifer in his studies of eriophyid mites described several species on

cereals, namely, Abacarus oryzae, Aceria bakkerii and Cheiracus sulcatus from Oryza

sativa in 1963a, 1969a and 1977a, respectively; Abacarus sporoboli, as probable leaf

vagrant on Sorghum halepense in 1965; Aceria tosichella from wheat, found mostly on

the upper leaf surface grooves, often causing leaf curl in 1969c; Catarhinus tricholaenae,

Aceria zeasinis and Aceria zeala from Zea mays in 1959b, 1962d and 1978, respectively.

ChannaBasavanna (1966) gave an illustrated account of all eriophyid mites

known thus far from India and described Aceria sorghi on Sorghum bicolor. Xin and

Dong (1982) described two new species, Aceria milli and Aceria paratulipae and

confirmed their status as vectors of wheat virus disease. Kuang and Feng in Kuang et al.

(1990) recorded another new species Catarhinu vulgaris on Sorghum bicolor.

Baco et al. (1991) reported sugarcane pest Aceria sacchari Wang, in Sulawesi

rice fields. Rao et al. (1999) listed Aceria sacchari Wang, Cheiracus sulcatus Keifer and

Eriophyes bakkeri Keifer from rice ecosystems. Joshi et al. (2002) in addition to the

eriophyids mentioned by Rao et al., recorded Abacarus hystrix (Nalepa) in their

checklist of eriophyid mites on rice. Skoracka (2004) reported Abacarus hystrix on

wheat, while conducting a survey of eriophyid mites on grasses in Poland. Amrine in his

unpublished catalogue on eriophyid mites reported Triticum aestivum as new host record

for Aculodes dubius (Nalepa) and Aculodes mckenziei (Keifer).

2.6.5.2 Biology

Slykhuis and Paliwal (1972) reported all stages of Abacarus hystrix as vector of

rye-grass mosaic virus. Rather (1983) observed the biology of Aceria zeasinis.

Development from egg to adult required 12-16 days in early summer and 18-25 days in

fall. Egg production ceased when temperatures went above 33°C. Three to five

generations were produced per year.

Pearis (2010) observed the developmental cycle of Aceria tosichella Keifer on

wheat under field conditions. Lifecycle comprised of eggs, immature stages and adult.


53

Eggs were placed along leaf veins. A complete generation required 8 to 10 days, on an

average. Wind helped in the natural dispersal of the mites when the hosts start drying

down.

2.6.5.3 Ecology

Gibson (1974) studied the feeding mechanism of Abacarus hystrix using scanning

electron micrographs and observed that the mites preferred the youngest leaves of

Festuca pratensis, Lolium perenne and Lolium multiflorum.

Keifer (1977a), Keifer and Hall (1977), Mohanasundaram, (1981a, 1983) and

Hong et al. (2005, 2006) reported Cheiracus sulcatus Keifer as vector of rice tungro virus

and reported its damage symptoms. Sithanantham (1979) and Barrion and Litsinger

(1991) commented on the virulent capacity and damage symptoms of Aceria sacchari

Wang, respectively. Gupta (1985) and Rao et al. (1999) conducted detailed population

density studies and observed that population peaked during pre-monsoon and dropped

during post-monsoon seasons. The mite preferred the middle of the apical 1/3rd stratum of

dorsal leaf surface and infested young greenish foliage. Lifecycle comprised of egg,

larva, protonymph and adult stages.

Frost (1997) reported wax production in Abacarus hystrix as seasonally variable

and enlarged lateral and dorsal bands of wax filaments developed in adults of summer

generations as high temperatures were found to stimulate increased rate of wax

production. The study also suggested that wax filaments enhanced buoyancy and

regulated water-loss, thereby significantly increasing the survival time of waxed mites at

low relative humidities, thereby reducing the rate of desiccation-induced mortality

Abacarus

hystrix (Nalepa) on quack grass under laboratory conditions and concluded that the mite

showed great potentiality for rapid population increase in conditions of initially low

density; life expectancy of females was more than males; reproductive output for females

was age-dependent and daily egg production reached a peak on the 5th day, and then

decreased steeply. The net reproductive rate, generation time, intrinsic rate of increase,

and the finite rate of increase were reported.

Chapter 2- Review of LiteratureNeopygmephoridae Cross

54

2.6.6 Neopygmephoridae Cross, 1965: 221


Trombidiformes; Suborder: Prostigmata; Superfamily: Pygmephoroidea; Family:

Neopygmphoridae.

The family Neopygmephoridae comprises of free-living, phoretic mites usually

ectoparasitic in nature.

2.6.6.1 Taxonomy

Cross (1965) created the tribe Neopygmephorini in the family Pyemotidae and

described four genera, namely, Acinogaster, Neopygmephorus, Pseudopygmephorus and

Parapygmephorus and five subgenera Petalomium, Sicilipes, Parapygmephorus,

Allopygmephorus and Neopygmephorus. Mahunka (1970) raised the taxonomic status of

this group to subfamily Neopygmephorinae in the family Pygmephoridae under

superfamily Pygmephoroidea. He retained in Neopygmephorinae only 4 genera.

Khaustov (2004) elevated status of subfamily Neopygmephorinae to the family

rank Neopygmephoridae Cross, based on the combination of the following

synapomorphic characters: female prodorsum with 2 pairs of setae- v2, Sc2 and a pair of

trichobothria sc1; coxisternal plates I and II with 2 pairs of setae- 1a, 1b and 2a, 2b;

hysterosoma with 2 pairs of cupules ia and ih; legs I always 4-segmented (tibia fused

with tarsus); tibiotarsus I only with 5 eupathidia (eupathidion ft” absent); femur I with 3

setae; setae dFeI hook-like. Physogastry of hysterosoma in adult females weakly

developed. Males had long aedeagus, transversely striated. He concluded that the family

Neopygmephoridae was very close to families Scutacaridae and Microodispidae and

together with Pygmephoridae (=Siteroptidae) compound the superfamily

Pygmephoroidea.

Presently this subfamily includes 16 genera: Acinogaster Cross, Aegyptophoorus

Sevastianov and Abo-Korah, Allopygmephorus Cross, Bakerdania Sasa

(=Neopygmephorus Cross), Guttacarus Mahunka, Heteropygmephorus Kurosa,

Kerdabania Khaustov, Parapygmephorus Cross, Petalomium Cross, Pseudokerdabania

Khaustov and Trach, Pseudopygmephorus Cross, Rackia Mahunka, Rhynopygmephorus

Chapter 2- Review of LiteratureNeopygmephoridae Cross

55

Kurosa, Sicilipes Cross, Singhalophorus Mahunka and. Xystrorostrum Mahunka

(=Piniphorus Sevastianov).

Khaustov (2010) listed 17 species under the genus Pseudopygmephorus including

a new species and 7 new combinations, namely Pseudopygmephorus abdominalis

(Berlese), Pseudopygmephorus agarici Zou, Gao and Ma, Pseudopygmephorus allmanni

(Krczal), Pseudopygmephorus aphodii Khaustov, Pseudopygmephorus argentiniensis

(Mahunka), Pseudopygmephorus bulbitarsus (Mahunka), Pseudopygmephorus chelatus

(Mahunka), Pseudopygmephorus chinensis, Zou, Gao and Ma, Pseudopygmephorus

madanlarae (Ramaraju and Madanlar) (from Pygmephorus), Pseudopygmephorus

magnus Mahunka, Pseudopygmephorus mileyi Hill and Deahl, Pseudopygmephorus

pappi Mahunka, Pseudopygmephorus peritrematus (Mahunka), Pseudopygmephorus

shanghaiensis Zou, Gao and Ma, Pseudopygmephorus stercoricola, (Berlese),

Pseudopygmephorus tarsalis (Hirst) and Pseudopygmephorus urlaensis (Ramaraju and

Madanlar); and transferred Pseudopygmephorus madagassicus Mahunka to the new

genus Kerdabania Khaustov.

No information on the ecology and biology of these mites are available and no

species in the family Neopygmephoridae had been reported from cereal agroecosystem.

2.6.7 Pygmephoridae Cross, 1965: 186


Trombidiformes; Suborder: Prostigmata; Superfamily: Pygmephoroidea; Family:

Pygmephoridae.

2.6.7.1 Taxonomy

The family Pygmephoridae was erected by Cross (1965) with Pygmephorus

Kramer (1877) as its type genus. More than 225 species in about 27 genera are known in

this family.

At present the taxonomy of the superfamily Pygmephoroidea sensu Lindquist

(1986) is complex and there is a general discontent with regards to the synonymy of

Siteroptidae with Pygmephoridae

Chapter 2- Review of LiteraturePygmephoridae Cross

56

The family Siteroptidae was first erected by Mahunka (1970) in the superfamily

Pygmephoroidea, and Siteroptes Amerling was recognized as a primitive taxon within

Pygmephoroidea. The subsequent discovery of intraspecific dimorphism

(phoretomorphy) among adult females in the superfamily, necessitated reevaluation of

generic and suprageneric concepts within Pygmephoroidea (Moser and Cross, 1975;

Mahunka, 1979). As a result, Mahunka (1979) synonymised Siteroptidae under

Pygmephoridae and placed Siteroptes within the subfamily Pygmephorinae, which also

was expanded to include Pediculastrinae. According to Lindquist (1986) Pygmephoroidea

comprised of the families Pygmephoridae (=Siteroptidae), Microdispidae and

Scutacaridae.

Kaliszewski et al. (1995) subdivided families previously included in the

superfamily Pygmephoroidea into two superfamilies: Pygmephoroidea with families

Pygmephoridae and Siteroptidae, and Scutacaroidea with the families Scutacaridae and

Microdispidae.

Khaustov (2004, 2009) accepted the opinion of Lindquist (1986), and

demonstrated on the basis of analysis of morphological characters that the families

Scutacaridae, Microdispidae, Pygmephoridae and Neopygmephoridae form a distinct

monophyletic group and that the family Siteroptidae was created on the basis of

plesiomorphic characters and was a synonym of the family Pygmephoridae.

Lindquist et al., in Krantz and Walter (2009) accepted the division suggested by

Kaliszewski et al. (1995), however, in the key to superfamilies provided in ‘A Manual of

Acarology’, 3rd edition, they recognized, keyed and diagonised the superfamilies

Pygmephoroidea and Scutacaroidea sensu Mahunka (1970).

Perusal of literature reveals that mites belonging to the genera Pygmephorus and

Siteroptes had been reported from cereal agroecosystems. In the genus Pygmephorus

about 52 species have been described of which Pygmephorus akermanae (Sevastianov,

Zakhida and Al Douri) and Pygmephorus nilanjana Putatunda had been reported from

wheat and rice respectively. Again, in the genus Siteroptes sensu Kaliszewski (1987), 14

species have been recognized as proposed by Khaustov and Ermilov (2011), namely,


57

Siteroptes aegyptiacus Sevastianov and Abo-Korah, Siteroptes amalae Sevastianov and

Abo-Korah, Siteroptes avenae (Muller), Siteroptes cerealium (Kirchner), Siteroptes

graminicola Mitrofanov, Shabanova and Sevastianov, Siteroptes graminisugus (Hardy),

Siteroptes huangshuiensis Su, Siteroptes kamtschatkensis Sevastianov, Siteroptes

longisetissimus Khaustov and Ermilov, Siteroptes longisomus Kaliszewski, Siteroptes

qinghaiensis Su, Siteroptes reniformis Krantz, Siteroptes triticola Su and Siteroptes

xizangensis Gao, Zou and Qin. Of these, Siteroptes avenae (Muller, 1905) had been

reported from rice and wheat and Siteroptes graminicola Mitrofanov, Shabanova and

Sevastianov (1984) had been reported from wheat.

The synonyms and nomenclature proposed for the species in the genus Siteroptes

is of much significance. Hardy (1851) described Acarus graminisugus and reported the

association of mites with silver top disease of meadow grasses in England. A decade later

Amerling (1861) introduced the generic name Siteroptes describing a similar mite from

rye in Czechoslovakia, which was named by Kirchner (1864) as Siteroptes cerealium.

Further, Reuter (1900) described Pediculoides graminum from diseased meadow grasses

in Finland, and in 1907 transferred the species to the genus Pediculopsis.

Muller (1905) and Wolcott (1908) described Pediculoides avenae from oat in

Silesia (Poland) and Pediculoides dianthophilus from diseased buds of carnation in the

state of Nebraska (U.S.A.), respectively. Reuter (1909) synonymised Pediculopsis

graminum with Pediculoides avenae Muller and Pediculoides dianthophilus Wolcott.

Krezal (1959) sustained the synonymy of Pediculopsis graminum with

Pediculoides avenae and Pediculoides dianthophilus, however, he transferred the species

into the genus Siteroptes and defined the genus in morphological terms. Siteroptes

cerealium (Kirchner) was designated as the type species of the genus.

In the revision of the family Pyemotidae, Cross (1965) considered the genus

Siteroptes to belong in the family Pyemotidae, revalued the generic criteria and divided

the genus Siteroptes into four subgenera. He synonymised Siteroptes cerealium

(Kirchner) with Siteroptes graminum (Reuter) and designated it the type-species of the

subgenus Siteroptes sensu stricto. However, Cross (1965) had not clearly expressed his


58

opinion as to the synonymy within the complex. He listed seven species in Siteroptes s.

str., including Pediculoides avenae Muller, Siteroptes cerealium (Kirchner), Pediculoides

dianthophilus Wolcott and Pediculoides graminum Reuter, while also strongly suggesting

that he considered all these specific names as being synonyms.

Suski (1973) agreed that the description and illustrations of the species presented

by Amerling (1861), which was referred to as Siteroptes cerealium by Kirchner (1864)

were similar to the description of Pediculoides graminum Reuter (1900) and that these

were synonyms as established by Cross (1965). However, the description and illustrations

of Siteroptes cerealium as presented by Krezal (1959) and the illustrations of Reuter

(1900, 1909) belonged to another species. Suski after studying the illustrations of Reuter

(1900, 1909) concluded that the males of Siteroptes avaenae (Muller) was described by

Reuter (1900) and mistakenly associated with female Siteroptes cerealium and that the

females were most probably considered as nymph of Siteroptes cerealium by Reuter

(1900, 1909) and Krezal (1959).

Also, Suski did not agree with the synonymy Pediculoides dianthophilus with

Siteroptes avenae and Pediculopsis graminum as the authors did not discriminate among

species of Siteroptes cerealium complex. Besides, Reuter had examined specimens from

the same host and locality as those used in the description of Pediculopsis dianthophilus,

but apparently from another collection date. Suski, finally concluded, based on the

morphological characters and biological studies that Siteroptes ceralium and Siteroptes

avenae were two different species. This view was also supported by Kaliszewski (1987)

on the basis of cladistic analysis and by the fact that Siteroptes cerealium lacked setae 4b

in both females and males which was present in all other members of the Siteroptes

cerealium group.

In the systematic revision of the Tarsonemina by Mahunka (1970), he erected the

family Siteroptidae and transferred the nonphoretic females of the genera Pediculaster

Vitzthum, and Pygmephorellus Cross and Moser (family Pygmephoridae Cross, 1965) to

the genus Siteroptes Amerling (family Siteroptidae Mahunka).


59

Based on rearing experiments, Moser and Cross (1975) gave evidences that some

species have two types of females - a phoretic (Pediculaster or Pygmephorellus form)

and a nonphoretic (Siteroptes form). As a consequence of this discovery, the genera

Pediculaster, Pygmephorellus and Siteroptes were synonymized under the name

Siteroptes by Smiley and Moser (1976).

Mahunka (1979) tried to reestablish the genera Pediculaster and Pygmephorellus

by distinguishing nonphoretic females and males of these genera from other species of

Siteroptes sensu Mahunka (1970). However, the combination of features which he used

to characterize nonphoretic females and males of the genera Pediculaster,

Pygmephorellus and Siteroptes seemed to be confusing.

Hence, Kaliszewski (1987) gave new diagnosis and description of the Siteroptes

species group connected with grasses, Siteroptes sensu Krezal (1959), Siteroptes

cerealium group sensu Suski (1973) and Siteroptes reniformis Krantz (1957) and called

them genus Siteroptes Amerling (1861). He discussed its systematic position and

presented genealogical relationships among Siteroptes species in the form of a cladogram

and listed the species in the genus.

Mitrofanov et al. (1984) described Siteroptes graminicola from barley in

European part of Russia. Later, Sevastianov and Abo-Korah (1984) described Siteroptes

tameri from soil of wheat in Ukraine.

Livshits et al. (1986) in their revision of the mites of the family Siteroptidae,

divided the genus Siteroptes into three subgenera: Allositeroptes Livshits, Mitrofanov and

Sharonov, with a single species, Siteroptes (Allositeroptes) primitives (Krczal), the

nominotypic subgenus Siteroptes with a single species Siteroptes cerealium (Kirchner)

and subgenus Eositeroptes Livshits, Mitrofanov and Sharonov, comprising all the rest of

the species. In addition, Siteroptes huangshuiensis Su was synonymized with Siteroptes

cerealium (Kirchner). Siteroptes tameri was transferred in the subgenus Allositeroptes.

However, Khaustov (2011) concluded that Siteroptes tameri should be excluded

from this subgenus on the basis of absence of setae 4a and presence of setae 4c (setae 4a

present and 4c absent in Allositeroptes), and the body characteristically long and thin,


60

more than two times longer than its width (not so long in Allositeroptes). He further

stated that Siteroptes tameri should be considered as a junior synonym of Siteroptes

graminicola on the basis of conspecificity.

Khaustov and Ermilov (2011) did not consider Siteroptes huangshuiensis Su a

synonym of Siteroptes cerealium as proposed by Livshits et al. (1986) and declared that

the division of the genus into subgenera based only on the number of setae on epimeres

III of females, without taking into account the leg chaetotaxy, was artificial, and that

Siteroptes (Allositeroptes) primitivus (Krezal, 1959) should be excluded from the genus

Siteroptes to form a separate genus Allositeroptes.

2.6.7.2 Biology

The biology of the species Siteroptes avenae (Muller) was studied by Suski

(1973) by rearing the mite on fungus isolated from diseased bud of carnation at room

temperature. Its lifecycle constitute of egg and adult stages only. Suski also remarked on

a characteristic morphological feature of Siteroptes avenae (Muller) referred to as

sporotheca, often filled up with granulae. There were on an average 5 granulae in each

sack, however occasionally as many as 10 granulae were counted in one sack. Similar

granulae were found scattered around sporotheca in specimens where the organ was

turned inside out due to change of internal pressure in mounting.

About 67% field collected females of Siteroptes avenae (Muller) had sporotheca

with the granular contents while 98% of laboratory collected, freshly born females

Siteroptes avenae (Muller) had sporotheca empty. It seemed very probable that

sporotheca was a specialised organ adapted for transport and dissemination of spores of

the fungus. Spores probably fell to sacks mechanically when shaken by the crawling of

mites over mycelium, and these were expelled by turning the organ inside out due to

change of internal pressure within the body.

2.6.7.3 Ecology

Siteroptes mites, partly due to their association with meadow grasses, cereal crops

and cotton plants, have been studied in many countries of the world. Members of the

Siteroptes reniformis group (Siteroptes reniformis and Siteroptes longisomus) seem to be


61

vectors of Nigrospora fungi and thus are implicated in diseases such as lint rot of cotton,

stalk rot of corn and Sorguro, and spike rot of wheat and other grains (Lindquist 1985).

Species of the Siteroptes cerealium group (Suski 1973) frequently carry spores of

Fusarium and Botrytis fungi and in this way are connected with silver top disease of

grasses.

2.6.8 Stigmaeidae Oudemans, 1931b: 251-263


Trombidiformes; Suborder: Prostigmata; Suborder: Raphignathoidea; Family:

Stigmaeidae.

Mites of the family Stigmaeidae (Acari: Prostigmata) are mainly free-living

predators (Gerson et al., 2003) but a few are parasitic on insects (Mitra and Mitra, 1953;

Swift, 1987).

2.6.8.1 Taxonomy

The family Stigmaeidae was erected by Oudemans (1931b) with Stigmaeus Koch

as its type genus. Summers (1962) revised the genus Stigmaeus. Gonzales (1965)

recognized 13 genera under this family and presented a comprehensive key. Summers

(1966) described and provided key to 14 genera. Besides these, Wood (1967, 1971,

1973), Gonzalez (1967) and Meyer (1969) made excellent contributions to advance our

knowledge in this family.

Tseng studied Stigmaeidae of Taiwan, provided key to 18 genera and enumerated

25 species. Sepasgosarian (1985, 1990) recognized more than 20 genera with 300 species

in his major work on superfamily Raphignathoidea. Faraji and Ueckermann (2005a,

2006) provided a key to Stigmaeus species known from Iran and Europe respectively.

Fan and Zhang (2005) in their exemplary revision of the superfamily

Raphignathoidea, keyed 14 genera including a new genus from the family Stigmaeidae

known from New Zealand and described 64 species including 16 new species. Fan and

Zhang (2002), Fan et al. (2003b), Fan and Walter (2005), Ehara and Ueckermann (2006),

Chapter 2- Review of LiteratureStigmaeidae Oudemans

62

Fan and Beard (2006), Faraji and Ueckermann (2005b, 2006), Faraji et al. (2007a),

Khanjani and Ueckermann (2008), Ueckermann (2008) added further to known genera

and species of Stigmaeidae.

So far more than 502 stigmaeid mites under 32 genera are known from the world.

Review of literature revealed that thus far 7 mites in 4 genera had been reported from

cereals, field and storage including the present new record of Stigmaeus unicus

Kuznetsov, from India.

Gupta (2002) reported Agistemus fleschneri Summers from rice. Later, Noei et al.

(2007) conducted a survey on the mites associated with stored rice and decayed rice bran

in northern Iran, Guilan province. Stigmaeus elongatus Berlese, Stigmaeus candidus Fan

and Li, Storchia pacifica (Summers) and Storchia robusta (Berlese), were reported new

to the fauna of Guilan province, and Storchia pacifica was new to Iran. Stigmaeus

mazandaranicus Faraji and Ueckermann was recognized as a junior synonym of

Stigmaeus candidus. Key to species of Stigmaeus found in Iran and neighboring countries

and all species of Storchia in the world was provided.

Khan et al. (2010) explored the predatory mite fauna of Punjab (Pakistan), and

described Eustigmaeus hooriaae from Ameen Pur Bangla, 20 miles away north from

district Faisalabad (Punjab) from Triticum aestivum (straw husk).

2.6.8.2 Ecology

Gupta and Gupta (1989), Chatterjee and Gupta (1996) and Gupta and Chatterjee

(1997) in their individual research observed and reported Agistemus fleschneri as an

effective predator of eriophyid and tetranychid mite.

2.6.9 Tarsonemidae Kramer, 1877: 215


Trombidiformes; Suborder: Prostigmata; Superfamily: Tarsonemoidea; Family:

Tarsonemidae.

Tarsonemidae comprise of small, rapidly moving mites with diverse feeding

habits. They exhibit a wide range of feeding behaviour. Some are predators of mite eggs,

Chapter 2- Review of LiteratureTarsonemidae Kramer

63

some are parasites of insects and other animals, while still others feed on fungi and algae.

Some of them may possess a sporotheca (a special spore-carrying structure) for carrying

a specific fungus on which they feed (Ochoa et al., 1991). However, only a limited

number of genera are found associated with plants, in particular with monocotyledonous

plants (Beer 1954; Smiley and Emmanouel 1980; Lindquist 1986; Ochoa et al. 1991;

Smiley et al. 1993); yet their infestation cause severe yield loss.

2.6.9.1 Taxonomy

The family Tarsonemidae was erected by Kramer (1877) with type genus

Tarsonemus Canestrini and Fanzago. Some of the significant contributions in this family

are by Ewing (1939), Beer (1954), Schhaarschmidt (1959), Beer and Nucifora (1965) and

Smiley (1967). Ochoa et al. (1991) gave characteristics, host symptomology and

relationships with other pathogens for 32 tarsonemid mites collected from Costa Rica,

with new descriptions of 16 species, two genera and a subgenus.

Lin and Zhang (1999) catalogued information on 93 species belonging to 15

genera in three subfamilies of the Tarsonemidae known from China. Zhang et al. (2000)

provided key to Tarsonemidae of New Zealand in which they included a total of 3

subfamilies, 14 genera and 57 species. Again, Lin and Zhang (2002) provided

geographical distribution, systematic cataloguing and an annotated bibliography along

with key to 3 subfamilies, 7 tribes and 40 genera of the world.

Till date, more than 500 species in about 45 genera are known from the world and

perusal of literature revealed that 33 tarsonemid mites in 5 genera had been recorded

from cereals. Smiley (1967) collected Steneotarsonemus spinki from rice. Wainstein and

Beglarov (1968) described wheat flower mite, Steneotarsonemus panshini.

Jeppson et al. (1975) reported Steneotarsonemus spirifex (Marchal) from cereals.

Later, Tseng and Lo (1980) reported Polyphagotarsonemus latus (Banks),

Steneotarsonemus chiaoi Tseng and Lo, Steneotarsonemus furcatus De Leon,

Steneotarsonemus mirabilis (Tseng and Lo), Steneotarsonemus spinki Smiley,

Tarsonemus brevicorpus (Tseng and Lo), Tarsonemus floricolus Canestrini and Fanzago,

and Tarsonemus smithi Ewing, mainly from rice.


64

Smiley and Emmanouel (1980) described and illustrated male, female, and larva

of Steneotarsonemus konoi along with their damage symptoms and distribution.

Emmanouel (1981) and later Emmanouel and Smiley (1985) reported Steneotarsonemus

hatzinikolsi and Steneotarsonemus hordei from wheat from Greece.

Ding and Yang (1983) recorded Steneotarsonemus phragmitidis (Schlectendal),

Tarsonemus floridanus (Attiah), Tarsonemus inornatus (Attiah) and Tarsonemus

paraunguis (Attiah) on rice from China. Ding and Yang (1984) described all stages of

Steneotarsonemus zhejiangensis on rice from Jiashan County, Zhejiang.

Lindquist (1986) in his major revisionary work on Tarsonemidae of the world,

described 31 genera including 7 new genera, 5 subgenera, 5 new species, recommended

new combinations, synonyms and provided key to genera and listed the species. In this

revisionory work, the major changes pertinent to the present dissertation include the

following: the genus Ogmotarsonemus with Ogmotarsonemus erepsis as type species was

described; Neotarsonemus Smiley was synonymised with Polyphagotarsonemus Beer

and Nucifora; Parasteneotarsonemus Beer and Nucifora and Neosteneotarsonemus Tseng

and Lo was synonymised with Steneotarsonemus Beer; Chaetotarsonemus Beer and

Nucifora, Lupotarsonemus Beer and Nucifora, Metatarsonemus Attiah,

Floridotarsonemus Attiah and Cheylotarsonemus Tseng and Lo were synonymised with

Tarsonemus Canestrini and Fanzango; the species Acarus translucens Green, Tarsonemus

phaseoli Bondar and Tarsonemus latus Banks were synonymised with

Polyphagotarsonemus latus (Banks). Tarsonemus pauperseatus Suski and Tarsonemus

setifer Ewing was synonymised with Tarsonemus waitei Banks; new combinations were

proposed for Tarsonemus wani Tseng and Lo, Neosteneotarsonemus mirabilis Tseng and

Lo, Tarsonemoides rakowiensis Kropczynska, Lupotarsonemus floridanus Attiah,

Lupotarsonemus inornatus Attiah and Lupotarsonemus paraunguis Attiah as

Xenotarsonemus wani, Steneotarsonemus mirabilis, Tarsonemus rakowiensis,

Tarsonemus floridanus, Tarsonemus inornatus and Tarsonemus paraunguis respectively

and Tarsonemus oryzae Targioni-Tozzetti and Tarsonemus phragmitidis Schlectendal

were transferred to Steneotarsonemus.


65

Mohanasundaram and Parameswaran (1988) recorded Ogmotarsonemus erepsis

Lindquist from India and gave its damage symptoms on rice. Later, Mohanasundaram,

(1996) described and illustrated Ogmotarsonemus oryzae from Tamil Nadu, India, from

rice, infesting leaf sheath and causing necrotic lesions.

Steneotarsonemus subfurcatus was described and collected from rice in Fujian,

China, in 1986 by Lin and Zhang (1990). Smiley et al. (1993) listed 14 mite species of

the genus Steneotarsonemus known to infest graminaceous plants of which

Steneotarsonemus spirifex was recorded on Zea mays. Lin and Zhang (1995) described

Steneotarsonemus guangzensis on rice from China.

Lin et al. (1995) provided a list of tarsonemid mites from Fujian province.

Similarly Lin and Zhang (1999) developed an annotated and illustrated catalogue and

bibliography of tarsonemid mites of China. This catalogue provided information on 93

species belonging to 15 genera in three subfamilies of the Tarsonemidae known from

China, of which 21 species were reported from rice. The study proposed Tarsonemus

brevicorpus Linnaeus as replacement name for Tarsonemus minutus (Tseng and Lo).

They also catalogued host details for the all the mites from China.

Rao et al. (1999) listed 19 tarsonemid species under 6 genera in their book on

mites associated with rice ecosystems. Zhang (2000) provided a user-friendly

identification key to the genera and species of the mites of Tarsonemidae in New

Zealand. A total of 3 subfamilies, 14 genera and 57 species were included in the key,

including 3 new genera. Distribution, diagnosis, list of specimens and illustrations were

provided for each species.

Joshi et al. (2002) listed 11 species of Tarsonemidae under 4 genera in their

checklist of mites in rice ecosystem. Atanasov et al. (2008) reported Steneotarsonemus

panshini and Stenoetarsonemus phragmitidis from cereal crops of Bulgaria.

2.6.9.2 Biology

Sogawa (1977) studied the biology of Steneotarsonemus spinki in field

conditions. Lifecycle studies revealed females oviposit in the tissues where they feed and

eggs hatch into an active larval stage followed by a quiescent stage. The lifecycle was


66

completed in 6 days. Chen et al. (1979, 1980), Lo and Ho (1979, 1980), Ghosh et al.

(1997), Ramos and Rodriguez (1998, 2000a, 2000b, 2001) and Lakshmi et al. (2008)

conducted individual studies on Steneotarsonemus spinki, observed its development in

controlled laboratory conditions at different temperature and recorded congruent results.

It was found in general, that as the temperature decreased, mites required more number of

days to complete its egg to adult cycle. The total number of eggs laid by a single female

varied from 59.5 (Lo and Ho, 1979) to 75 (Sogawa, 1977).

Baker (1997) and Pena and Campbell (2005) observed the lifecycle of the broad

mite, Polyphagotarsonemus latus. Adult females laid eggs and then died. The eggs hatch

into larvae that passed into a quiescent larval (nymph) stage. Quiescent female larvae

become attractive to the males which pick them up and carry them to new foliage. When

females emerge from the quiescent stage, males immediately mate with them.

2.6.9.3 Ecology

Jeppson et al. (1975) and Deheleanu (1977) discussed the mode of infestation and

the subsequent damage caused by Steneotarsonemus spirifex (Marchal) on oats and rice,

respectively. Chen et al. (1980), Fang (1980), Hsieh et al. (1980), Wei and Chow (1980),

Shikata et al. (1984), MyoungRae et al. (1999) observed that rice panicle mite,

Steneotarsonemus spinki, was the causative agent for ‘empty-head sterility’ or ‘sterile

grain syndrome’, and described its damage symptoms.

Badulin (1986) carried out investigations in USSR during 1964-1985 and reported

that Steneotarsonemus panshini was a widespread pest that caused whole or partiall

empty ears of durum wheat leading to complete crop loss. The morphology of adults and

larvae was briefly discussed and the damage characteristics were illustrated. Tetraploid

wheat and oats were infested and acted as reservoirs, the zone of constant injuriousness

coincided with that of durum wheat. Damage caused by large colonies of mites from their

entry into the flowers prior to heading until their entry into the soil before harvest was

described. The percentage of flowers damaged was also recorded and found to increase

with each subsequent year.


67

Li (1990), Zhang et al. (1995) and Ghosh et al. (1998) reported Tarsonemus

talpae Schaarschmidt as the vector of Sarcoladium oryzae. The infection from this fungii

lead to the spread of ‘rice sheath brown syndrome’. Zhang and Lin (1991) reported

Gnorimus chaudhrii Wu as dominant natural enemy of Tarsonemus bilobatus Suski and

observed its predator potentiality in the rice fields of Fujian, China.

Hallas et al. (1991) collected monthly samples of grain, straw and hay from 4

farms in Denmark and analysed these for live storage mites. This study reported the

relative abundance of species varied with the age of substrate and Tarsonemus sp. was

observed as the substrate progressed through its mid-phase.

Jiang et al. (1994) observed that in 1991-1993 damage to rice by

Steneotarsonemus spinki began during the spike-forming stage in September, in

Guangzhou, Guangdong, China. Different varieties showed different amounts of damage.

Ghosh et al. (1997) conducted field studies in India during the kharif season in

1994 and 1995, and observed Steneotarsonemus spinki infesting rice throughout the year,

the populations of Steneotarsonemus spinki reached a maximum during November and a

minimum during February. Correlation studies indicated that population increase was

favoured by low rainfall and high temperatures. MyoungRae et al. (1999) gave the first

record of Steneotarsonemus spinki from Korean Republic on rice grown in an

environment controlled greenhouse. They reported typical damage symptons for this mite

along with its description.

Jones and Brown (1983) reported that the mean number of eggs/female/day for

Polyphagotarsonemus latus (Banks) was closely related to temperature and humidity.

Navia et al. (2006) described in detail the mode of damage caused by Steneotarsonemus

furcatus De Leon and its symptoms on rice plant. Karmakar (2008) studied the damage

and loss of yield on various cultivars of rice and listed them as resistant and as

susceptible to mite attack. Among the characters studied, only chaffy grain and grain

yield had significant positive or negative correlation, respectively, with the occurrence of

the mite.

Chapter 2- Review of LiteratureTetranychidae Donnadieu

68

2.6.10 Tetranychidae Donnadieu, 1875: 9


Trombidiformes; Suborder: Prostigmata; Superfamily: Tetranychoidea; Family:

Tetranychidae.

Members of the family Tetranychidae are known to infest several hundred species

of plants. These mites spin protective silk webs mostly on the undersurface of leaves and

hence are more commonly known as ‘spider mites’. They feed by puncturing the

plant cells resulting in damage to standing crop.

2.6.10.1 Taxonomy

Donnadieu erected the family Tetranychidae in 1875. The systematics of

Tetranychidae was vague and imprecise until the work of Ewing (1913), who showed the

taxonomic value of the male genital armature, a character used by McGregor (1950) in

one of the first syntheses on this family. This study listed 102 species in 15 genera and

the number rose to 204 species in the revision by Pritchard and Baker (1955) who

proposed a system of classification based on the morphology of female empodia,

chaetotaxy of dorsum and the position of duplex setae on legs I and II and divided the

family into subfamilies, tribes and genera. Later, important taxonomic contributions were

made by Reck (1959), Baker and Pritchard (1960), Wainstein (1954, 1960a), Tuttle and

Baker (1968), Fletchmann and Baker (1970), Gutierrez (1966, 1970), Ehara (1956a,

1956b, 1956c, 1964, 1966a, 1966b, 1967, 1969, 1970, 2004), Ehara and Lee (1971),

Manson (1963, 1964, 1967a, 1967b), Meyer (1964,1974), Chaudhri et al. (1974), Tuttle

et al. (1976), Gutierrez and Schicha (1983), Tseng (1990), Toroitich et al. (2009) and

Seeman and Beard (2011). Presently the family includes about 1200 species in 71 genera

under 2 subfamilies. Perusal of literature revealed that 81 species of tetranychid mites in

12 genera had been reported on rice, wheat, maize, sorghum and pearl millet.

In a material collected from Coimbatore, Hirst (1926) isolated paddy mite,

Paratetranychus oryzae, and later named it as Oligonychus oryzae (Hirst). Kamali (1990)

conducted a faunistic survey of mites in southwestern Iran during September 1986. A

checklist of 37 genera from 21 families, with data on host plants and province was


69

prepared. Oligonychus zeae (McGregor) on sugarcane and maize and Petrobia tunisiae

Manson on wheat and barley were reported new to Iran.

Ochoa et al. (1990) described Schizotetranychus freitezi and Schizotetranychus

pseudolycurus from Costa Rica and Panama, both from rice leaves, causing yellow spots

on the surface. Flechtmann and Santana (1997) reported Catarhinus tricholaenae Keifer

in Brazil and the male was described for the first time. Oligonychus zeae was reappraised.

Catarhinus tricholaenae was found on maize and on Brachiaria plantaginea in

Minas Gerais, Oligonychus zeae was found on Brachiaria plantaginea and on Melinis

sp., a weed in maize fields in Minas Gerais. First report of paddy leaf mite, Oligonychus

oryzae, feeding on rice, was observed and reported by Singh (2001) from eastern Uttar

Pradesh, India. Leaves harbored a large population of mites and their immature stages.

Othman and Zhang (2003) described Tetranychus arifi on Zea mays from

Malaysia. Beard et al. (2003) redescribed Oligonychus oryzae and described Oligonychus

zanclopes from sugarcane and rice. Subhash Chander et al. (2003) while recording the

incidence of insect pests and diseases of rice-wheat rotation in Indo-gangetic plains of

Punjab, Haryana, Madhya Pradesh and Uttar Pradesh, during 1970-2000, reported that in

1999, Petrobia latens (Muller) was the major pest of wheat.

The host and distribution records of 68 species of Tetranychidae known from

cereals had been listed on the site http://www1.montpellier.inra.fr/CBGP/spmweb/ species.php

2.6.10.2 Biology

Cherian (1931, 1938) gave a comprehensive account of Oligonychus oryzae in

south India. The biology for Oligonychus oryzae had been extensively studied by

different authors at various time periods. Misra and Israel (1968a) conducted detailed

lifecycle studies. Parthogenetic females laid fewer eggs that developed into males.

Development involved larva, protonymph and deutonymph which emerged into greyish

white adult and each stage was interspaced by a quiescent phase. Similar observations

were made by Susan (1976) and Rao and Kulshreshtha (1985). Nayak et al. (2008) and

Radhakrishna and Ramaraju (2009) studied the biology of Oligonychus oryzae on rice


70

under laboratory conditions. The total developmental period required for the males was

found to be greater than that observed for females.

Bodenheimer (1951) and Jeppson et al. (1975) observed the development of

Eutetranychus orientalis (Klein) with respect to seasons and noted that winter eggs were

deposited on both sides of the leaves and summer eggs were deposited mainly on the

upperside of leaves. The developmental threshold, thermal constant, preoviposition

period, male and female longevity of adults in summer, spring, autumn and winter were

recorded. Gabele (1959), Jeppson et al. (1975) and Vincenzo (2010) studied the biology

of Bryobia graminum (Schrank) in Germany on apples, pears and grasses.

Boudreaux (1958) and Jeppson et al. (1975) observed that eggs of Tetranychus

gloveri Banks were always red. Mated tumid females deposited white eggs and unmated

females laid red eggs, but when adult females that develop from red eggs mated only

white eggs were deposited.

Nickel (1960) observed that Tetranychus desertorum developed from egg to adult

in about 9.5 days in California and in about a day less in Paraguay and Texas. This study

observed that females developing on heavily infested plants showed marked reductions in

life-span and fecundity, the sex ratio of eggs deposited by fertilized females changed as

the females aged, arid unfertilized females had lower fecundity and lived longer than

fertilized ones.

Later, Rivero and Vásquez (2009) also studied biological aspects and life table of

the red spider mite, Tetranychus desertorum on leaf discs of kidney bean (Phaseolus

vulgaris Linnaeus) under laboratory conditions. The results showed that developmental

stages included egg, larva, protonymph and deutonymph. Preoviposition, oviposition and

postoviposition periods, mean fecundity, mean longevity, generation time, net

reproduction rate and intrinsic natural growth were recorded.

Jeppson et al. (1975) observed the oviposition rate and average life span of

Tetranychus yusti McGregor and reported that the males required a day less than females

for completing their development. Gupta (1985) studied the biology of Oligonychus

indicus (Hirst). Rai et al. (1989) also studied the average development periods of females


71

through egg incubation, larval, protonymphal and deutonymphal stages, the pre-

oviposition period, adult female and male life span of Oligonychus indicus reared on

sorghum leaves.

Sridhar et al. (1998) conducted similar studies and also recorded the influence of

crop growth stages (flag leaf and half bloom stages) on the biology of Oligonychus

indicus in 12 grain sorghum cultivars (CSV 8R, CVS 14R, Sel 3, SPV, 913, Swati, RS

29, IS 2146, IS 2312, IS 5613, ICSV 705, Lakadi and M 35-1). This study concluded that

the effect of plant age was significant for the biology of the spider mite in all genotypes

and that lifecycle duration was longer at the flag leaf stage.

Dubitzki and Gerson (1987) studied the development of Petrobia tunisiae Manson

on various winter Gramineae, and reported that the mites produced non-diapause eggs in

the first generation and mostly diapause eggs in the second generation and a summer

diapause was also postulated for this species.

The biology of Petrobia latens was studied by Noorbakhsh and Kamali (1995),

Jat and Sharma (1999) and Nogia and Sharma (2003). On an average lifecycle completed

in 33 days at 17±1°C , 15 days at 25±1°C (Noorbakhsh and Kamali, 1995), 23.9 days at

ambient temperatures (Jat and Sharma, 1999), and 25.5 days (Nogia and Sharma, 2003)

under laboratory conditions. Under field conditions the developmental time varied

according to season as compared to that in controlled environment, the shortest duration

being 26 days (Noorbakhsh and Kamali, 1995).

Saringkaphaibul et al. (1998) studied the biology of Schizotetranychus

andropogoni (Hirst) and observed that the period of development of male from egg to the

last nymphal stage was longer than females and that male lived longer than females.

Mejia et al. (1998) observed that the development of Schizotetranychus oryzae Simons

Rossi de from egg to adult on rice required 16 days for complete development from egg

to adult. Guven and Madanlar (2000) studied biology of Tetranychus urticae in

laboratory using maize cultivars Otello and Pioneer. Silva (2002) studied the biology and

thermal requirement of Tetranychus ludeni on leaves of cotton (Gossypium hirsutum L.

r. latifolium) in Brazil.


72

Sakunwarin et al. (2003) studied life history and lifetable of Tetranychus

truncatus Ehara on mulberry leaflets under laboratory conditions. Pang-BaoPing et al.

(2004) investigated the effects of five host plants- cucumber, kidney bean, soyabean,

aubergine and maize on the development and reproduction of Tetranychus truncatus

under laboratory conditions. The duration of different developmental stages, fecundity,

egg production per day, net reproductive rate (R0), finite rate of increase ( ), intrinsic

rate of increase (rm), mean generation time (T), and number of days needed to double the

population size (DDP) significantly varied with the host plant. The time needed by

females to complete a generation on maize was 11.6 days, the oviposition time lasted

15.8 days and the survival rate on maize was 93.6. Gulati (2004) reported Tetranychus

cinnabarinus (Boisduval) completed its lifecycle in 19-29 days but at 22.8°C, lifecycle

was shortened to 10-13 days.

Noronha (2006) studied the biology of Tetranychus marianae McGregor on

Passiflora edulis f. flavicarpa under controlled environmental condition and reported that

the egg to adult time spanned approximately 11 days, with 92% survival figure. The

mean female longevity, daily mean, intrinsic rate of increase, finite rate of increase, mean

time span of one generation and net rate of reproduction (Ro) was recorded.

Kaimal and Ramani (2007) studied the post embryonic development of the

vegetable mite, Tetranychus neocaledonicus (Andre) infesting Moringa oleifera in

laboratory. Rearing of life stages of the mite was carried out following leaf flotation

technique. The lifecycle comprised both sexual and parthenogenetic generations with

slight variation in their respective durations. The duration of pre-oviposition, oviposition,

post-oviposition period, fecundity and total duration for sexual and parthenogenetic

reproduction was recorded.

Najmoon et al. (2008) studied the duration of developmental stages and fecundity

of Tetranychus urticae. The lifecycle comprised of eggs, larva, protonymph, deutonymph

and adult. Sousa et al. (2010) studied the biology of Tetranychus mexicanus (McGregor)

on soursop (Annona muricata), sweetsop (Annona squamosa) and araticum (Annona

coriaceae) in Brazil. Ehara et al. (2008) observed that the summer females of


73

Tetranychus huhhotensis Ehara et al. and Tetranychus zeae Ehara et al., were pale

greenish-yellow, while their diapausing females were pale-orange.

Chaaban (2012) observed the biology and ecology of date palm mite Oligonychus

afrasiaticus (McGregor) through regular inspection in Tunisian oases and laboratory

observations. During autumn and spring, Oligonychus afrasiaticus was found on sorghum

leaves in orchard ground cover.

A life table study in laboratory on six host plants (fruits of date palms varieties

Deglet Noor, Alig, Kentichi, Bessr, and Deglet Noor pinnae and sorghum leaves) showed

that the lifecycle of Oligonychus afrasiaticus differed among host plants with average

values being 10.9 days on sorghum leaves. Relatively, fecundity and intrinsic rate of

increase was higher while average longevity was lower on sorghum leaves.

Karami-Jamour and Shishehbor (2012) studied egg to adult development time and

found maximum population and shorter development time at higher temperatures.

2.6.10.3 Ecology

Bodenheimer (1951) and Jeppson et al. (1975) studied population dynamics of

Eutetranychus orientalis (Klein) with respect to temperature and humidity. Nickel (1960)

conducted investigations on the distribution of Tetranychus desertorum in relation to

climate. At all temperatures, high humidity resulted in longer life, lower fecundity,

immature mortality and faster development to maturity, and it was calculated that the net

reproduction rates and innate capacities for increase were generally higher at the higher

humidity.

Misra and Israel (1968a), Rao and Kulshreshtha (1985), Swamiappan (1986),

Veluswamy et al. (1987), Prakash et al. (1988) and Lakshmi et al. (2008) studied

population dynamics of Oligonychus oryzae on different rice cultivars from India. In

general, it was observed in all the studies that bright sunny weather followed by wet

seasons was most congenial for the multiplication of mites.

Misra and Israel (1968a, 1968b) developed techniques for rearing these mites on

rice leaves and reported that continuous feeding by all stages lead to yield reduction upto


74

25%. Nagarajan (1957), Gupta et al. (1972), Rai et al. (1977), Chakkaravarthy et al.

(1982), Sridharan et al. (1997), Patil et al. (1999) and Lakshmi et al. (2008) suggested

effective control measures for Oligonychus oryzae.

Pertinent information on its moulting, egg characters and preferred part of

infestation were provided by Rao and Kulshreshta (1985), Swamiappan (1986) and

Veluswamy et al. (1987), respectively. Roshan et al. (2000) reported severe damage to

rice in Jammu, India, during 1997-98 due to infestation by Oligonychus oryzae along

with other arthropods.

Davis (1969b) commented on the damage symptoms of Oligonychus araneum

Davis and Oligonychus digitatus Davis. These mites occured in very large populations,

forming conspicuous webs and yellow rings on grasses. Saba (1974) observed the

development of Tetranychus tumidus Banks at different temperature and recorded

seasonal variations in population.

Liu and Tsai (2009) observed the development, survivorship, and reproduction of

the tumid spider mite, Tetranychus tumidus on coconut palm at 6 constant temperatures

(10, 15, 20, 25, 30, and 35°C) and concluded that the development, survivorship and

reproduction of Tetranychus tumidus were significantly affected by temperatures.

Jeppson et al. (1975) studied the population variation for Eotetranychus yumensis

(McGregor) with season. Dubitzki and Gerson (1987) observed that in the fields of Israel,

Petrobia harti was most abundant during early summer, males being less than 10% of the

population and without diapause eggs.

Roy et al. (2011) studied population dynamics of Petrobia harti in relation to

seasonal variations from India. Populations were highly abundant during the summer

months, followed by rainy season and completely absent in the winter. In addition, the

fecundity rate, mortality, reproductive rate, oviposition rate, average longevity of adult

mites, total duration required to complete lifecycle was recorded.

Youngman et al. (1988) observed effect of water stress on specific life history

parameters of Tetranychus pacificus in greenhouse using potted almond trees.


75

Significantly more eggs were deposited by females on trees under variable water stress

than by females on trees under continuous water stress.

Krainacker and Carey (1990, 1991) determined the influence of week of season,

within-plant distribution and population patterns of Tetranychus urticae on field maize in

the Sacramento River Delta, California and suggested that mite sampling should focus on

the number of females on lower leaves early in the season. Wilson and Morton (1993)

studied the seasonal abundance and intra-crop distribution of Tetranychus urticae on

cotton.

Guven and Madanlar (2000) conducted population sampling from late July to

early October in 1997 to study the density of Tetranychus urticae in six maize fields in

Turkey. The highest density was determined during the first part of August and during

September.

Dhar et al. (2004) reported that temperature showed a significant positive

correlation, while relative humidity showed significant but negative correlation with the

population of mite and rainfall did not show any significant buildup of mite population.

Najmoon et al. (2008) reported that temperature played significant (P<0.001) role on the

duration of developmental stages of Tetranychus urticae. High temperature accelerated

developmental rate and reduced duration of developmental periods.

Manjunatha and Puttaswamy (1990) determined the yield losses in sorghum

caused by infestation of Oligonychus indicus in greenhouse and field experiments in

Karnataka. Kapoor et al. (1997) noted seasonal occurrence of mite pests in northern India

on maize and found them to be greatest in May and least in August-September. Predatory

mites were present throughout most of the year on both crops, with the maximum

population being recorded in August-September.

Dubey et al. (1998) used simple and multiple linear regression techniques to study

the relation between different meteorological elements (weekly data of maximum

temperature, minimum temperature, morning RH, evening RH, sunshine hours and

rainfall for 26th-52nd standard weeks) and infestation by sorghum mite pest, Oligonychus


76

indicus, and others, at Dharwar and Coimbatore. The available meterological data during

1955-80 were used for the study.

Berry et al. (1991) demonstrated the importance of humidity and temperature in

population dynamics of Oligonychus pratensis. Drought stressed maize crop resulted in

higher population. Simulation studies showed that colonization of a maize field by less

than one adult female per plant could result in mite densities sufficient to cause crop loss.

Barron and Margolies (1991) studied dispersal behaviour of Oligonychus

pratensis in within-plant distribution on corn. On an average, adult females moved up the

plant when placed on lowest leaves 1 and 2, moved down the plant when placed on

leaves 5-9 and remained on leaves 3 and 4 when placed on them. The growth stage of

maize and the initial mite density did not change the pattern of redistribution.

Li and Margolies (1991) conducted laboratory and field studies and showed that

the site on which adults of Oligonychus pratensis settled when placed on corn (maize)

leaves was significantly influenced by light, leaf surface and gravity. Stiefel et al. (1992a)

also studied the vertical distribution of Bank grass mite on 21 sorghum lines and on the

commercial variety 'Wheatland', grown in dryland and irrigated treatments in Kansas.

They also incorporated studies on leaf area covered by Oligonychus pratensis. This study

concluded vertical distribution of Banks grass mite colonies within grain sorghum was

affected by irrigation.

Another set of experiments in fields and greenhouses were conducted in

Manhattan, Kansas, to screen sorghum lines for antibiosis to Oligonychus pratensis, and

to determine if a relationship existed between different types of drought resistance and

antibiosis to Oligonychus pratensis and it was concluded that leaf temperature affected

resistance to Banks grass mite on drought-resistant grain sorghum (Stiefel et al,. 1992b).

Steifel and Margolies (1992) in an attempt to study affect of components of

colonization - dispersal cycles on offspring sex ratios of Banks grass mites, concluded

that while offspring-sex ratio did not differ when transferred from maize to sorghum or

vice versa, offspring sex-ration changed in response to deteriorating food source resulting

in more potential dispersers. Collins and Margolies (1995) reported that interspecific


77

matings between Tetranychus urticae and Oligonychus pratensis occur readily in the

laboratory but yield no female offspring.

Taha (1992) conducted field studies in Beni-Suef governorate, Egypt, to evaluate

the population dynamics of Tetranychus arabicus on different growth stages of 3 maize

varieties, and the variety Mephtah Pioneer and Giza 2 were the most tolerant. All mite

stages reached peak numbers during the flowering stage or 11 weeks from the sowing

date. Most mites were found on the lower parts of maize.

Kumar and Sharma (1993) studied population dynamics of Tetranychus ludeni

Zacher on okra and revealed significant positive correlation between the mite population

and minimum temperature, RH and rainfall. Reddy and Baskaran (2006) observed

damage potential of the spider mite Tetranychus ludeni on seedlings of four eggplant

Solanum melongena varieties.

Kaimal and Ramani (2011b) studied the feeding characteristics and damage

induced by Tetranychus ludeni on the detached leaves of the velvet bean Mucuna

deeringiana in laboratory. The results showed that Tetranychus ludeni could infest almost

all age groups of leaves though the middle aged ones showed high population densities

and reflected on the highly complex colony structure of Tetranychus ludeni through

silken webbing of the individuals.

Kaimal and Ramani (2011c) studied post embryonic development of spider mite,

Tetranychus ludeni infesting Mucuna deeringiana in the laboratory. Rearing of life stages

of the mite was carried out following leaf flotation technique. The duration of pre-

oviposition period, oviposition period, post-oviposition period, fecundity, total duration

of sexual and parthenogenetic reproduction was recorded. The results showed that

parthenogenetic development required comparatively shorter duration than sexual ones.

Wang et al. (1994) reported that population of Petrobia latens (Muller) in

Zhengzhou region of Henan, China, showed an aggregated spatial distribution as deduced

using Iwao's regression, Taylor exponent sign law and CA values. Population growth was

inhibited by low temperature.


78

Han-GuiZhong et al. (1997) reported that with an increase in population density

of Petrobia latens, hatchability of eggs decreased, propagation coefficient and longevity

of female mites declined, and mortality of larvae and migratory proportion of adult mites

increased. A density of 20-40 mites per wheat plant caused significant damage.

Bhagat (2003) observed pest incidence, infestation symptoms and mite

density/leaf of Petrobia latens on wheat foliage in irrigated and non-irrigated areas and

barley crops grown in unirrigated areas of Pura, Jammu, during February/March 1999

and 2000.

Chen et al. (1996) observed population peaks and economic damage caused by

Tetranychus truncatus Ehara on Sophora japonical in Beijing. Chen ZhiJie et al. (1999)

observed bionomics and ecological tactics of Tetranychus truncatus on maize in Shaanxi

Province, China. Fan et al. (2000a) concluded from his observations on cowpea leaves

that Tetranychus truncates required different temperatures and photoperiods at different

stages. Fan et al. (2000b) showed that temperature was much more influential than light

time on time duration of one generation of Tetranychus truncatus, and interaction

between temperature and light time on population development was not significant. Fan

et al. (2003a) concluded that the developmental threshold temperature and effective

thermal summation were different for different developmental stages of Tetranychus

truncatus.

Iraola et al. (1998) studied the population dynamics of Tetranychus turkestani

(Ugarov and Nikolskii), infesting maize in Navarra (Spain). Gotoh and Gomi (2000)

concluded that seasonal changes in plant quality was responsible for decline in

populations of Tetranychus kanzawai Kishida after every peak and it was predator

independent as observed on hydrangea (Hydrangea macrophylla) in Japan. Takafuji et al.

(2007) observed that deteriorated food conditions enhanced expression of diapause in

Teranychus kanzawai and that non-diapausing adult females could survive low

temperature conditions (3–5°C).

Zhang (2003) reported that development was faster for Tetranychus cinnabarinus

(Boisduval) at higher temperatures and showed highest oviposition rate on cucumber,

followed by pepper and tomato. This study reported diapause was lost in many


79

populations that remained on plants in winter, when induction and termination was

controlled by day length.

Gulati (2004) reported that in summer crop mites appeared in the month of April,

showed an increasing trend in May and then attained a peak in the month of June. In

Kharif crop, mite population peaked in the month of October, declining gradually.

Abiotic factors also influenced Tetranychus cinnabarinus population in different okra

varieties.

Kazak and Kibritci (2008) observed development time, reproductive rates, and

population growth parameters of Tetranychus cinnabarinus on 8 strawberry (Fragaria ×

ananassa Duchesne) varieties under laboratory conditions. Similar observations were

made on excised leaf disc of lablab bean, Dolichos lablab by Kaimal and Ramani

(2011a).

Rachna et al. (2009) while studying seasonal incidence of Tetranychus

neocaledonicus in relation to climatic factors observed that there was a significant

positive correlation with mean temperature and a significant negative correlation with

mean relative humidity.

2.6.11 Trochometridiidae Mahunka, 1970: 137–174.


Trombidiformes; Suborder: Prostigmata; Superfamily: Trochometridioidea; Family:

Trochometridiidae.

Mites of the family Trochometridiidae are associated with different kinds of

holometabolous insects, typically ground-nesting bees including families Andrenidae and

Halictidae (Lindquist 1985). They are also found on other bees, wasps, carabid and

scarabaeid beetles. The family comprises of two genera and five species only.

2.6.11.1 Taxonomy

The genus Trochometridium was described by Cross (1965) in the superfamily

Pyemotoidea, family Pyemotidae and subfamily Pyemotinae. Subsequently, Mahunka

Chapter 2- Review of LiteratureTrochometridiidae Mahunka

80

(1970) elevated the taxonomic level of this genus to subfamily, Trochometridiinae, but

placed it in the family Pygmephoridae under superfamily Pygmephoroidea.

Lindquist (1986), during reclassification of family-group taxa in Heterostigmata,

based on phylogenetic analysis, placed the genus in the newly created family

Trochometridiidae under superfamily Trochometridioidea, and recognised three species

in this genus, namely, Trochometridium tribulatum Cross from ground-nesting bees in

the United States, Trochometridium chinensis (Mahunka) from wasps in China and

Trochometridium kazachstanicum Khaustov and Eidelberg from carabid beetle in

Kazakhstan.

Hajiqanbar et al. (2009) described the genus Neotrochometridium and species

Neotrochometridium sensillum, phoretic on ventral body surfaces, grasping hairs of

abdomen and thorax of Cymbionotum semelederi and Siagona europaea Dejean and

Trochometridium iranicum, phoretic on ventral body surfaces, grasping sternite hairs of

mesosoma and metasoma of Pseudapis nilotica. Thus, no species in the family

Trochometridiidae, thus far, has been reported from cereals or cereal-associated

agroecosystems.

2.6.12 Tydeidae Kramer, 1877: 232-246


Trombidiformes; Suborder: Prostigmata; Superfamily: Tydeoidea; Family: Tydeidae.

Tydeidae is a family of acariformes mites. These are generally small mites, soft

bodied, often with striations or reticulations and eyes may or may not be present. Most of

the species in the family are considered fungivores or predators; however, a few are

known to be facultatively phytophagous.

2.6.12.1 Taxonomy

The family Tydeidae was established by Kramer (1877) on the basis of an earlier

described genus Tydeus Koch, 1835. The first monograph on Tydeoidea was published in

1933 by Thor. Subsequent revisions were given by Baker (1965, 1968a, 1968b), Andre

Chapter 2- Review of LiteratureTydeidae Kramer

81

(1979, 1980, 1981a, 1981b), Kazmierski (1990, 1996a, 1996b, 1997, 1998a, 1998b,

2000) and Andre and Fain (2000).

Zhang et al. (2001) developed a key to 12 genera, including 2 new genera and 29

species, incorporating details on distribution, diagnosis, list of specimens and

illustrations. Presently, the family consists of more than 370 species under about 60

genera. Perusal of literature revealed that these mites are under 3 genera, namely,

Metapronematus, Pronematus and Tydeus, which had been found associated with grains.

Four species had been recorded from cereals.

2.6.12.2 Ecology

Gupta and Nahar (1981) reported Pronematus fleschneri Baker as predatory mite

from paddy crop. As part of a project on the biological control of root diseases by

mycorrhizae, El-Bagoury and Momen (1988) described two new species of mites from

Egypt of which Orthotydeus longisetosus, was collected from soil under roots of wheat

crop.

However, later Khanjani and Ueckermann (2003) provided the new combination

Tydeus momeni as this was a junior homonym of Tydeus longisetosus Kuznetzov and

Zapletina. Hallas and Solberg (1989), Hallas et al. (1991) and Emmanouel et al. (1994) in

their individual survey of mites from storage reported Tydeus spp.

Zdarkova (1998) and Hubert et al. (2006) recorded Tydeus interruptus Thor from

storage in Czech Republic. Zdarkova (1998) conducted surveys in 64 grain stores (brick

stores, hangars, silos) in the Czech Republic during spring and autumn 1996. A total of

112 samples of wheat, 74 samples of barley and 70 samples of sweepings were examined.

A total of 32 species of mites were found, including Astigmata (15 species), Prostigmata

(10) and Mesostigmata (7). Barley was more infested by mites than wheat. Over 90% of

the samples of sweepings were infested. Tydeus interruptus Thor was recorded as one of

the dominant species of mite.

In a similar kind of study, Hubert et al. (2006) examined mite fauna in 78 selected

grain stores, simultaneously sampling the grain mass and residues in order to compare

concurrent mite communities in these two different habitats and reported Tydeus

Chapter 2- Review of LiteratureTydeidae Kramer

82

interruptus from residue samples. This study recorded no correlation in mite abundance

and species numbers between samples from grain residues and grain mass though residue

samples had more mites and higher species diversity than the stored grain mass, thereby

indicating low connectivity of these two habitats.

Putatunda (2002, 2005) reported Pronematus spp. from storage from India.

Menon et al. (2007a) described Pronematus oryzae from IARI paddy farm fields.

2.6.13 Acaridae Ewing and Nesbitt, 1942: 121-124


Sarcoptiformes; Suborder: Oribatida; Superfamily: Acaroidea; Family: Acaridae.

Mites belonging to the family Acaridae are economically most important owing to

their pest status. Their habitat ranges from house dust, fungal moulds to being pathogens

of human body.

2.6.13.1 Taxonomy

Major contributors to our knowledge of mites in this family are Ewing and

Nesbitt (1942), Robertson (1959), Griffiths (1964a, 1964b, 1970), Manson (1972),

Hughes (1976), Zhang and Fan (2005) and Fan and Zhang (2007). More than 400 species

are known under 115 genera, and of these 25 species in 10 genera had been reported in

association with cereal agroecosystems.

Morallo (1980) gave the new record of Caloglyphus laarmani (Samsinak) in

stored rice from south-east Asia. Baggio et al. (1987) surveyed acari in stored cereals in

Greater Sao Paulo, Brazil. Microscopic examination of 160 cereal samples subjected to

42 days of incubation at 25°C and 70% RH showed that 49% of the samples were

infested with mites, 48% were infested with Tyrophagus putrescentiae (Schrank) and

17.5% with Aleuroglyphus ovatus (Troupeau).

Haines and Lynch (1987) reported Madaglyphus javensis from rice bran residues

and broken rice in a mill in East Java, Indonesia and described female, male, tritonymph,

protonymph and larva. Rao and Prakash (1987) gave first report of Caloglyphus berlesei

Michael infesting rice seedlings and leaf sheaths and panicles of older rice plants from

Chapter 2- Review of LiteratureAcaridae Ewing and Nesbitt

83

India. It was usually found in association with another acarid, Tyrophagus palmarum

Oudemans.

The arthropod pests of stored rice were studied in Korea Republic by Kim et al.

(1988) and Acarus siro Linnaeus was reported as one of the dominant pests. Bonilla et al.

(1990) gave description of Rhizoglyphus costarricensis, a mite detected on seeds of the

rice cv- CR-1113 and CR-1821 produced in Costa Rica. Direct damage to endosperm and

embryo was observed on certified and registered seed of CR-1821, with a drastic effect

on seed germination. The fungi Aspergillus sp., Fusarium sp., and Helminthosporium sp.,

were also observed in the mite-damaged seeds.

Mohanasundaram and Parameswaran (1991) gave first record of Histiostoma

humidiatus Vitzthum on rotten sorghum crops in Tamil Nadu, India. Nakao (1991)

reported Acarus immobilis Griffiths, Rhizoglyphus robini Claparede, Tyrophagus

perniciosus Zachvatkin, Tyrophagus putrescentiae and Tyrophagus similis Volgin,

damaging rice seedling, chaff and straw from Japan.

Mahmood (1992) studied the mite fauna of stored grain in central Iraq from a total

of 108 samples of wheat, barley and rice. This study recorded the occurrence of

Tyrophagus putrescentiae on rice, barley and wheat.

Ottoboni et al. (1993) assessed the level and type of infestations by mites in 3

flour mills, a malthouse and several harbour warehouses used for temporary grain storage

at Porto Marghera, Venice, Italy. Amongst other astigmatid mites, Acarus siro was

frequently found on wheat, sorghum, barley grains and germinating barley and

Tyrophagus putrescentiae was present on maize grain and bran.

Rao et al. (1999) reported five species of acarid mites from rice ecosystem,

namely, Acarus siro, Aleuroglyphus ovatus, Caloglyphus berlesi, Tyrophagus palmarum

and Tyrophagus putrescentiae. Joshi et al. (2002) expanded this list with record of

another 10 species, namely, Acarus immobilis Griffiths, Caloglyphus laarmani

(Samsinak), Lepidoglyphus destructor (Schrank), Rhizoglyphus callae Oudemans,

Rhizoglyphus costarricensis Bonilla, Rhizoglyphus echinopus (Fumouze and Robin),

Rhizoglyphus robini, Suidasia pontifica Oudemans, Tyrophagus perniciosus Zachvatkin


84

and Tyrophagus similis. These mites were observed on rice and mostly reported as

storage mites.

Sarwar and Ashfaq (2004) studied the taxonomy of Caloglyphus clemens and

Caloglyphus cingentis from Pakistan. The distinguishing characters were compared,

similarity matrix and phenogram were constructed and a key to all known hypopods was

provided. Sousa et al. (2005) reported Caloglyphus hughesi (Samsinak) from stored

foods in supermarkets and markets of Brazil.

Ardeshir et al. (2008) collected mite samples from stored wheat, straw and dust in

silos, flour-mills and mills in Tehran, Karaj and Varamin, Iran, during spring 2005. This

study reported 25 species belonging to 11 families and 3 orders of Astigmata,

Prostigmata and Mesostigmata. Of these, Aleuroglyphus ovatus and Tyrolichus casei

Oudemans were newly recorded from wheat stores of Tehran province. Acarus siro was

one of the most abundant predator and pest mites. Atanasov et al. (2008) reported

Tyrophagus longior from cereal crops of Bulgaria. Sarwar et al. (2009) described

Caloglyphus austerus and Caloglyphus bradys from Triticum aestivum and Sorghum

vulgare, respectively.

Klimov and OConnor (2009) recently demonstrated that the taxonomic concept of

Tyrophagus putrescentiae (Schrank, 1781), involved two closely related species, and one

of them was described as Tyrophagus communis Fan and Zhang (2007). This study

observed that the prevailing usage of the name Tyrophagus putrescentiae included almost

exclusively one of these species, and this usage was not in taxonomic accord with the

neotype designated by Robertson (1959). Thus, their study proposed that this neotype

fixation for the species Tyrophagus putrescentiae be set aside under Article 75.6 (ICZN,

1999) and a neotype consistent with the prevailing usage be designated.

Further, Fan and Zhang (2007) verified the synonymy of Tyrophagus

putrescentiae (based on the new neotype) after examining the following taxa with extant

types: Tyrophagus amboinensis Oudemans, Tyrophagus americanus (Banks),

Tyrophagus australasiae (Oudemans) (tentative synonymy), Tyrophagus breviceps

(Banks), Tyrophagus castellanii (Hirst), Tyrophagus cocciphilus (Banks), Tyrophagus

communis Fan and Zhang, Tyrophagus nadinus (Lombardini) and Tyrophagus


85

neotropicus Oudemans. For the taxon Tyrophagus putrescentiae sensu Fan and Zhang

(based on the neotype designated by Robertson) a new name was proposed, Tyrophagus

fanetzhangorum. Lectotypes were designated for the following species: Tyrophagus

amboinensis, Tyrophagus americanus, Tyrophagus australasiae, Tyrophagus breviceps,

Tyrophagus castellani, Tyrophagus cocciphilus, Tyrophagus nadinus and Tyrophagus

neotropicus.

2.6.13.2 Biology

Rodriguez and Stepien (1973) and Chmielewski (2003) studied the biology and

population dynamics of Caloglyphus berlesi on artificial diet and buckwheat sprouts,

respectively. Singh and Mathur (1997) reported Suidasia nesbitti Hughes from wheat and

pearl millet. The development was studied by Shen (1988), Chmielewski (1991) and

Singh and Mathur (1997) under natural and controlled conditions. The time to complete

one lifecycle and the net mortality of different stages was higher in natural conditions

than under controlled conditions.

Kilic and Toros (1997) concluded from their experiments on Acarus siro that at

25°C, yeast was the most suitable substrate for the development of Acarus siro as

compared to maize flour, rice flour, wheat flour, dried milk, wheat starch and maize

starch. Franzolin and Baggio (2000) recorded greater reproductive rates at high

temperatures and humidity. Sanchez-Ramos et al. (2007) reported that the optimal

temperature for development of immature stages of Acarus farris was 27-28°C and

maximum mortality was at 7°C and 29.7°C.

Xia et al. (2009) and Ahmed et al. (2012) conducted studies on the total

developmental time, longevity and oviposition period of Aleuroglyphus ovatus at various

temperatures. Xia et al. (2009) reported that higher temperatures facilitated shorter

developmental cycles, while Ahmed et al. (2012) observed longer lifecycle with increase

in temperature. Similar experiments was carried out for Tyrophagus putrescentiae and a

longer duration of lifecycle and shorter adult longevity was seen associated with

increased temperature (Ahmed et al., 2012). Raut and Sarkar (1991) studied the lifecycle

of Rhizoglyphus robini under different temperatures, and conclusively proved that higher

temperature assisted in shorter developmental cycle and maximum egg-laying.


86

2.6.13.3 Ecology

Navarro et al. (1985) assayed the mortality of newly emerged adults of Acarus

siro by exposing them to different combinations of O2, N2, and CO2. The results obtained

showed that even temperatures containing 20% CO2 or 10% O2 in N2 would kill Acarus

siro efficiently at 26°C, thus proposing maintainence of modified atmospheric conditions

in storage areas for controlling mite infestation. Davis and Boczek (1988) studied thermal

acclimation of adult Acarus siro reared on wheat germ at 21±1°C and 85% RH. When

acclimated from 0°C to 33°C for 1 or 4 days, the fecundity and longevity of the mite

were affected depending on period of acclimation and the difference between the rearing

temperature of the mites and the acclimation temperature, but the sex ratio of the progeny

from the treated adults did not change.

Parkinson (1990) studied the population increase, damage to germ and endosperm

and fungal infestation by Acarus siro and Tyrophagus longior amongst other species of

mites, on wheat, at 20°C and at 90% and 75% RH, when placed in test-tubes and

examined at 20 weeks interval. Population peak for Acarus siro was more as compared to

Tyrophagus longior (14000 mites/test-tube: 2200 mites/test-tube at 90% RH; 3000

mites/test-tube: 1000mites/test-tube at 75% RH). At both humidities, visible fungus was

always less abundant on infested grain than uninfested grain. Radwan (1991) commented

on the male sperm competition during mating in Caloglyphus berlesi.

Kohli and Mathur (1994) investigated the feeding behaviour of Tyrophagus

putrescentiae on whole, broken and ground grains of wheat, pearl millet, and in

groundnut and red gram. Ground form was preferred for wheat whereas the broken grain

form was preferred for pearl millet. Radwan and Sivajothy (1996) examined and

concluded that the function of prolonged mate association in Rhizoglyphus robini was

post-copulatory mate guarding.

Skorupska and Korbas (1998) commented on fusarial infection-like symptoms by

mites on winter wheat cv. Kobra, in Poland. On prematurely ripening of plants, a partial

or total yellowing of panicle was observed, and lower parts of internodes showed brown

discoloration. A closer inspection revealed that culms and panicles were infested with

Tyrophagus longior and not infected by Fusarium. Mites contributed to decrease in grain


87

yield. Zdarkova (1998) conducted surveys in 64 grain stores in the Czech Republic

during spring and autumn 1996. A total of 112 samples of wheat, 74 samples of barley

and 70 samples of sweepings were examined. Average moisture content of wheat and

barley samples were 13-13.6%. A total of 32 species of mites were found, including

Astigmata (15 species), Prostigmata (10) and Mesostigmata (7). Barley was more

infested by mites than wheat. Over 90% of the samples of sweepings were infested.

Acarus siro was one of the dominant species.

Chaudhary and Mahla (2001) conducted survey in 39 distantly located villages

comprising different climatic zones of Haryana, India. 195 wheat samples were collected

and about 10 insect species and only one grain mite Acarus siro recorded highest damage

(7.6%) as pests of stored wheat grains in the western climatic zone. Mahla (2001) studied

the population dynamics of storage pests and their incidence in wheat grain under

different climatic zones of Haryana and recorded Acarus siro infesting stored wheat

grains in the western zone causing a damage of 7.6%.

Athanassiou et al. (2001) evaluated the abundance of insect and mite species, in a

flat storeroom in central Greece, filled with ~90 tonnes of wheat. The most abundant

acarid mite species was Acarus siro. The highest population density was recorded during

September and October. Insects and mites showed an aggregated spatial pattern, as

indicated by Iwao's Patchiness Regression. Athanassiou et al. (2003) analysed three-

dimensional distribution and sampling indices of insects and mites in horizontally-stored

wheat. Tyrophagus putrescentiae was found to be one of the most abundant species and

for majority of most abundant insect and mite species, highest population densities were

recorded during autumn, more individuals were found at the corners, central zone, and in

the upper 0.5 m of the bulk (P<0.05). All the species showed an aggregated spatial

pattern, as estimated by Taylor's Power Law. Again, Athanassiou et al. (2005), while

conducting spatiotemporal distribution of insects and mites in horizontally stored wheat

from flat storage facility in Greece, recorded Acarus siro as one of the most abundant

species.

White et al. (2003) reported that wheat cultivars, both whole and crushed, were

highly susceptible to Acarus siro and were barely susceptible to infestation by Acarus


88

farris. A second experiment of similar design was conducted to determine the infestation

rate by the two Acarus species on Columbus and Neepawa hard red spring wheat and

Tyrophagus putrescentiae along with other mites. Crushed seed of both cultivars

sustained high populations of Acarus siro and Tyrophagus putrescentiae, which increased

1059-1202x and 974-1963x, respectively. This study concluded that all stored-product

mites could survive on whole stored cereals and oilseeds to varying degrees, but Acarus

siro and Tyrophagus putrescentiae manifested most as they took advantage of damaged

cereals, where nutrients were readily available, resulting in large population increases.

Conversely, the crushing of oilseeds appeared to inhibit mite propagation, probably

because of excessive oil content (45% dry weight) within the meal.

Hubert et al. (2003) isolated fungal species from mite-infested samples of seeds

(wheat, poppy, lettuce, and mustard) and from the gut and external surface of Acarus

siro, Caloglyphus rhizoglyphoides (Zachvatkin), and Tyrophagus putrescentiae (Schrank)

amongst others. Pekar and Zdarkova (2004) developed a predator-prey model for the

prediction of the local (within-patch) population dynamics of Acarus siro and

Caloglyphus eruditus on wheat grain at 20 et al. (2004)

studied the occurrence of Tyrophagus putrescentiae and Tyrophagus similis on fresh and

rotten straws of rice in the fields in Japan and concluded that fresh straw was preferred to

rotten straw.

Abd-el-Halim et al. (2006) recorded Caloglyphus redikorzevi (Zachvatkin) as the

most dominant mite on wheat flour, macaroni, dried milk and wheat bran stored at 25°C

and 85% RH in Fayoum Governorate in Egypt, and studied its lifecycle. Palyvos and

Emmanouel (2006) studied seasonal abundance and vertical distribution of mites in flat

storage containing wheat and reported Tyrophagus putrescentiae as the dominant species,

with more individuals distributed in the center and at the surface of the wheat bulk. The

highest mite population densities for the total mite species were recorded during October-

November and after the middle of January.

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