INTERNATIONAL JOURNAL OF ENVIRONMENTAL SCIENCE AND ENGINEERING (IJESE) Vol. 1: 59-72 http://www.pvamu.edu/texged Prairie View A&M University, Texas, USA

Description of life cycle stages of the broad Polyphagotarsonemus latus (Banks, 1904) (: Tarsonemidae) based on light and scanning electron microscopy

Ashraf A. Montasser1,*; Hanafy, A. R. I.2; Aleya S. Marzouk1 and Gamal M. Hassan2 1- Zoology Department, Faculty of Science, Ain Shams Univ., Egypt. 2- Plant Protection Research Institute, Agricultural Research Center, Giza, Egypt. * Corresponding author e-mail: [email protected]

ARTICLE INFO ABSTRACT Article History The broad mite, Polyphagotarsonemus latus egg, larva, nymph, Received: male and female are described using light and scanning electron Accepted: microscopy. SEM of egg revealed that its surface is sculptured by the Available online: Jan. 2011 presence of transparent sacs with irregularly shaped peripheral tubercle- ______like structures located at their bases. The structure of these sacs as seen Keywords from SEM examination suggests their role in fixation of the egg on Developmental stages plant leaves and/or respiration of the embryo. It also suggested that the Identification tubercles may act as a basal support for the sacs. SEM of larva revealed Mite the presence of whitish transverse groups of lines and hundreds of Morphology minute tubercles lying between them. The study demonstrated that the SEM nymph remains enclosed within the skin of larva until the adult is formed. Quiescent nymphs are observed to attract males. The male is observed to carry a nymph at a right angle by its genital capsule and when female emerges mating occurs. The 4th pair of legs in the male is modified into a claspers-like organ. SEM description of female P. latus showed that it has a relatively soft oval body with the gnathosoma observed broad at its base and narrow terminally like a rostrum. It also revealed downward direction of the gnathosoma and its articulation with the idiosoma. The body of the male is divided by distinct pleats between gnathosoma, prosoma, metasoma, opisthosoma and genital capsule. Detailed description and measurements of the egg as well as different body regions of larva, nymph and adults are recorded. The present study also described the fine structure of the dorsal cuticle of the larva and changes occurring during development into a quiescent nymph.

1. INTRODUCTION The broad mite, Polyphagotarsonemus latus is a minute herbivorous mite with a variety of host range and is widely distributed in tropical and subtropical areas around the world. It was first described by Banks (1904) as latus from the terminal buds of Mango in green house in Washington DC, USA (Denmark, 1980). It attacks numerous economic crops and ornamental plants belonging to more than 60 families like Solanaceae, Cucurbitaceae and Malvaceae causing severe symptoms (Zhang, 2003). ______ISSN 2156-7530 2156-7530 © 2011 TEXGED Prairie View A&M University All rights reserved. 60 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus

Primary hosts of P. latus comprise mite stages, each specimen was placed in a cotton, pepper, tomato, cucumber, citrus, drop of Hoyer's medium (50 ml distilled legumes, tea, coffee, mango, gerbera, water, 30 g gum Arabic, 200 g chloral begonia, dahlia and jasmine (Peręz-Otero et hydrate and 20 ml glycerin) and covered al., 2007). It feeds on the newest growths, with a cover slip. The slide then placed in an causing malformation of the terminal leaves. oven adjusted at 42oC for another one day to They become severely stunted, hardened and remove air bubbles near the cover edge. curl down at the edges with brownish or Slides were examined by light microscope reddish lower surfaces, depending on host and illustrated by line drawings using camera- species (Labanowski and Soika, 2006). lucida. Illustrations and measurements were Numerous studies are accessible on P. made for each stage. Body length was latus but most of them are concerned with measured from the point at which the the ecology, biology and control of the mite. gnathosoma meets the idiosoma to the A few dealt with the morphology of the life posterior edge of the body. Body width was cycle stages (Cho et al., 1993; Nucifora and measured across the widest part of the body. Vacante, 2004; Peręz-Otero et al., 2007). Changes in shape caused by preparation Zhang (2003) reported a key to family techniques, such as deformations caused by Tarsonemidae beside notes on common application of the cover slip or mounting names and distribution of its species medium, were considered to be negligible. including P. latus. However, the above 2.3. Scanning electron microscopy (SEM): studies offered incomplete data with a few Using a fine paint brush, were micrographs. Martin (1991) presented brief collected from the laboratory culture from notes on the morphology of different stages leaf pieces under stereo-binocular of P. latus using scanning electron microscope. Leaf pieces, 5×10 mm, microscopy (SEM). containing eggs or live stages of P. latus, The present work offers fine and were mounted on stubs with carbon tape, and detailed description for all life cycle stages examined under low vacuum scanning that may be useful for further taxonomical, electron microscope (JEOL/EO-JSM-5500) comparative and control studies. (Nickles et al., 2002) at Al-Azhar University, Cairo, Egypt. 2. MATERIALS AND METHODS 2.1. Collection of specimens: 3. RESULTS The broad mite P. latus was collected The morphological features of each from unsprayed sweet pepper plants grown stage (egg-adult) of the life cycle of P. latus in a plastic house in a protected cultivation are described in this study. The use of both farm belonging to the Agriculture Research light and scanning electron microscopy were Center, Giza governorate, Egypt. Mite essential in order to fully describe this pest. colonies were maintained on potted pepper 3.1. Egg stage: plants under laboratory conditions for two The egg is elongate, oval and slightly weeks before the start of experiments. Mites elevated. The egg is ~ 107 µm in length and were collected from laboratory culture using ~ 64 µm in width. The exposed surface of a fine paint brush under stereo-binocular the egg carries around 29-37 sacs more or microscope. less evenly distributed (Fig. 1a). The sacs 2.2. Light microscopy: exhibit high refractive index as they appear whitish compared to the darker background Mounted slides were made for different of the egg body. Each sac is ~10 µm long mite stages at Plant Protection Research and ~ 8.5 µm wide. SEM processing of the Institute, Dokki area, Giza governorate, egg causes the membranes of these sacs to Egypt. Specimens were cleared in lactic acid deflate and appear as a wrinkled cloth. The for one day. For permanent preparation of translucent surface of the sacs reveal Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 61 irregularly shaped tubercles arranged surround a central darker area (Fig.1b). peripherally at the bases of these sacs The latter appears finely perforated.

3.2. Larval stage: the propodosoma, two pairs on the Eggs hatch into three-legged larvae metapodosoma and four pairs on the (Fig. 2). There was no apparent sexual opisthosoma (two lateral and two median) dimorphism in the larval stage. The larva is (Fig. 2a). Propodosoma is ~ 42 µm, very small (~ 142 µm in length and ~ 72 µm metapodosoma is ~ 58 µm and opisthosoma in width). It is pear shape and more or less is 42 µm in length. The ventral side of the whitish (Fig. 2). On the dorsal surface of the larva carries seven pairs of setae; two pairs larva, there are 8 pairs of setae; two pairs on of propodosomal setae, one pair of

62 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus metapodosomal setae situated near side of like pulvilli. Legs I, II and III measured ~ 34, the coxa of leg III and four pairs of 36 and ~ 40 µm in length, respectively. opisthosomal setae (Fig. 2b) Propodosomal SEM examination revealed that the setae were longer (~ 14 µm) than cuticle of the dorsal surface is divided by metapodosomal ones (~ 10 µm). The larva three bands of marked whitish transverse possesses three pairs of legs; two pairs lines. The areas between these lines are situated more or less anterior and one sculptured by groups of minute elevations of posterior. Leg I carries a terminal claw, leg II whitish tubercle-like structures that are and III terminated with ambulatory sucker- surrounded by darker bands (Fig. 2c, d).

Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 63

3.3. Nymphal stage (quiescent): of the larva until the adult stage is formed The nymph is elliptical in shape, the (Fig. 3). The nymph is more or less sharply longest axis is ~ 120.5 µm and the widest is pointed at both ends. Two gnathosoma are ~ 75 µm. The body of the nymph is divided distinct; one more posterior for the nymph into three well-defined portions namely and another more anterior for the larva (Fig. 3). propodosoma (~ 27.3 µm long), SEM examination of the quiescent metapodosoma (~ 56.8 µm long) and nymph revealed a noted difference between opisthosoma (~36.4 µm long). The its dorsal cuticle which is in fact the cuticle gnathosoma lies between the 1st pair of legs. of the metamorphosing larva and that of the There is no apparent sexual dimorphism in preceding larval stage. Beginning of the the nymphal stage. The nymph possesses 5 molting causes the dorsal cuticle to stretch pairs of setae on the ventral surface (Fig. 3a); resulting in disappearance of the darker one pair of propodosomal setae, three pairs bands surrounding each group of tubercle- of metapodosomal setae and one pair of like structures which in turn become evenly opisthosomal setae. It carries four pairs of distributed on the body. The three bands legs that adhere tightly to the nymphal body dividing the body expand also revealing their as long as it remains enclosed within the skin true thickness and structure (Fig. 3c, d).

64 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus

3.4. Male stage: of propodosomal setae, three pairs of Sexual dimorphism is characteristically metapodosomal setae, two pairs of pronounced in adult P. latus. The male opisthosomal setae and a rounded posterior measured 138.9 ± 7.8 µm in length and 74.9 genital plate, bearing two pairs of shorter ± 1.9 µm in width (Fig. 4). It is marked by a setae. Ventrally, there are six pairs of setae; narrow posterior end and much broader in its two pair of propodosomal setae, three pairs middle region (metapodosoma) (Fig. 4). The of metapodosomal setae and one pair of male body is divided into three distinct opisthosomal setae (Fig. 4b). The portions namely propodosoma (48.2 ± 1.9 gnathosoma lying at the anterior proximity µm long), metapodosoma (64.3 ± 1.4 µm of the body measures (~ 34.2 µm in length long) and opisthosoma (26.3 ± 4.7 µm long). and ~ 31.1µm in width). The male's dorsal surface carries three pairs

The genital capsule (~ 12.5 µm in circular, located ventrally posterior to the length and ~ 15.6µm in width) is semi- opisthosoma and consists of 2-3 triangular Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 65 genital papillae (Fig. 5f). The anal plate the ventral side of the opisthosoma and surrounding the anus is large, well defined marked by three peripheral indentations and located infront of the genital capsule on (Fig. 5f).

The male has four pairs of long and postero-lateral position and the 4th pair is spindle shaped legs. The 1st and 2nd pairs of totally directed backwards (Fig. 4). Leg I, II, legs adhere closely to each other occupying III and IV measure 70 ± 2.4, 85 ± 1.4, 107.5 the anterior ⅓ of the body and are ± 4.6 and 94.6 ± 4.4 µm in length, maintained in a fronto-lateral position. The respectively. Each of first three legs consists last two pairs of legs though are similarly of five segments namely coxa, femur, genu, close to each other. The 3rd leg lies in a tibia and tarsus. Coxae III and IV of the male

66 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus are sharply and distinctively contiguous. Leg The latter in turn overlaps the genital plate. I terminates in a single claw, while legs II SEM examination of males holding the and III terminate in a pair of claws and a females in a copulation position revealed that membranous empodium. Leg IV is four- the male picks up the female nymph at a segmented namely coxa (~ 34.9 µm in right angle to the male's body (Figs. 4, 5 and length), femur (~ 17.4 µm in length), genu 7 f ). The female nymph seems to be carried (~ 24.9 µm in length) and fused tibia + tarsus on a terminal pad of the male's genital (tibio-tarsus) (~ 17.4 µm in length). The capsule (Fig. 7f). tibiotarsal segment terminates by a button 3.5. Female stage: like claw and bends into a claspers-like The female body (138.9 ± 5.5 µm long organ (Figs. 4, 5). The fourth leg possesses and 97.3 ± 5.6 µm wide) appears oval and two long setae, a dorso-lateral seta ~ 87.2 more or less elevated along its dorsal axis µm long and a ventro-lateral one ~ 57.3 µm (Fig. 6, 7). The body of female is divided long (Fig. 5). into three well-defined portions namely SEM examination of the male's dorsal propodosoma (38.9 ± 3.3 µm long), surface reveals distinct pleating of the metapodosoma (69.4 ± 1.6 µm long) and metapodosoma so as to overlap the anterior opisthosoma (33.3 ± 2.3 µm long) (Fig. 6). propodosoma and the posterior opisthosoma.

Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 67

The female has nine pairs of setae on Legs (I- IV) measured ~ 54.6, 54.3, its dorsal surface; two pairs of propodosomal 72.4 and 54.3 µm in length. The anterior two setae, three pairs of metapodosomal setae pairs of legs are widely separated from the and four pairs of opisthosomal setae (Fig. posterior ones (Fig. 7 a, d, e). The legs II, III 6a). The ventral view reveals seven pairs of and IV of female do not carry any claws setae; one pair of propodosomal setae, four (Fig. 6). pairs of metapodosomal setae and two pairs of opisthosomal setae (Fig. 6b).

Each of the first three legs consists of tibia and tarsus (Fig. 8 a-c). The leg IV has five segments namely coxa, femur, genu, three segments namely coxa (~ 11.3 µm

68 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus long), genu + femur (~ 22.6 µm long) and propodosoma (Fig. 8e). A pair of sensilla tibia + tarsus (~ 11.3 µm long). The fouth leg (trichobothria) is observed at the bases of has two long setae on the tibiotarsal segment; these organs. The female gnathosoma (~26.7 the dorso-lateral seta is ~70.2 µm long and µm long and 22.7 µm wide) is largely the ventro-lateral one is ~ 24.9 µm long. The rounded (Figs. 7, 8). The gnathosoma is posterior legs are covered with less setae observed to contain two slender stylettiform than the anterior ones (Fig. 8). A pair of chelicerae between the non segmented clavate pseudostigamtic organs is located triangular palps (Fig. 8e). Two stigmata are dorsolaterally and distally between coxae I dorso-laterally located at the gnathosomal and II near the anterior margin of the base (Fig. 8e).

SEM showed that the gnathosoma is (Fig 7 a, b). The gnathosoma appears broad held in a downward position where it is at its base and rostrum-like terminally difficult to observe from the dorsal side (Fig. 7b). The smooth dorsal surface of Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 69 female appears to be divided more or less by adhesive or non-adhesive exochorion. In P. 3 superficial lines (Fig. 7a). Ventrally a latus, the external sculpture of the egg longitudinal median groove is seen formed together with its deposition on the lower part by two lateral cuticular flaps. This groove of the leaves suggests the presence of an extends from the vicinity of the 1st two pairs adhesive layer that fixes the egg to the of legs to end between the two last pairs by a substratum similar to that reported by Alberti funnel-like structure (Fig. 7d). and Coons (1999). Other interpretations of the function of sculptured egg shells in P. 4. DISCUSSION latus as in various mites might be having a The present study extends our previous respiratory function (Callaini and Mazzini, preliminary knowledge of the morphological 1984) or serve to reduce water loss details of the egg, larva, quiescent nymph, (Witaliňski, 1993). Egg shell structure and and adults (male and female) of the mite P. function in P. latus needs further histological latus. The morphological details dealt with studies to be evaluated. are the shape, measurements, and surface Detailed description and measurements sculpturing of all different body regions in of different body regions of P. latus larva in every developmental stage using light and the present study was not documented in surface scanning electron microscopy. previous studies. The larval length in the SEM description of P. latus egg present study is ~ 142 µm versus ~ 117.5 µm showed that it is elongate, oval, slightly in description of Martin (1991). The elevated and externally sculptured. Its presence of three whitish transverse bands of exposed surface carries 29-37 knob-like sacs lines and minute tubercles scattered between arranged in evenly distributed rows. The sacs them was also observed by Martin (1991). were previously described as tubercles Using SEM at higher magnification in the (Martin, 1991; Zhang, 2003) or as tufts present study revealed triangular or (Peręz-Otero et al., 2007). The egg length polygonal arrangement of these tubercles on measures ~ 107 µm in the present study larval surface. which is more or less similar to that P. latus nymph possesses four pairs of previously mentioned by Martin (1991) who legs and remains enclosed within the skin of recorded ~ 100 µm. SEM processing in the larva until the adult is formed. The quiescent present study caused deflating of these sacs nymph is observed to attract males, carried so as to appear as a crumpled tissue. by its genital capsule to terminal new leaves Consequently, the translucent surface of and when female emerges from nymphal these sacs helped to elucidate irregularly molt, male immediately commences mating. shaped peripheral tubercles at their bases. Such contribution to mite dispersal was These tubercles may probably act as basal previously mentioned by Jeppson et al. support for the sacs holding them in an (1975), Martin (1991) and Peręz-Otero et al. upright position. SEM also revealed minute (2007). Martin (1991) suggested that the perforations in the egg surface between the male’s genital capsule might function as a tubercles of each sac. suction pad and the fourth leg of male might In P. latus, the egg shell is notably be used for initial manipulation of the sculptured by sacs or knob-like structures. female. SEM of the larval skin surrounding Witaliňski (1993) reported that egg chorion the nymph in the present study revealed in Acaridid mites of the genera Tyrophagus stretching of the integument, scattering of the (T. longior and T. perniciosus) and tubercles and widening of lines in the Aleuroglyphus is additionally coated in the transverse bands. distal portion of the oviduct by an The subdivision of P. latus body into exochorion material that forms the three portions namely propodosoma, microsculpture. He further mentioned that metapodosoma and opisthosoma mentioned eggs with microsculpture either have an

70 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus in the present study was also reported in the genital capsule as well as SEM description of general body form of family Tarsonemidae male were not presented in previous studies. (Jeppson et al., 1975). The P. latus adults are The body of female P. latus in the dimorphic where males are markedly present study appeared soft, oval and swollen different from females in their morphological with unornamented dorsal surface as features. Males are seen markedly broader at previously mentioned by Zhang (2003) and its middle region and tapers posterior similar Peręz-Otero et al. (2007). Jeppson et al. to previous findings (Zhang, 2003; (1975) reported that body of female P. latus Labanowski and Soika, 2006). The three was covered by a carapace like dorsal pairs of setae on the dorsal surface of the idiosoma and McDaniel (1979) reported that propodosoma or on the ventral surface of adults of superfamily Tarsonemoidea have metapodosoma were previously mentioned idiosoma covered by a number of by Jeppson et al. (1975), Zhang (2003) and overlapping horny plates. The current SEM Labanowski and Soika (2006). The present study revealed the presence of skin SEM description of the male showed distinct depressions on female dorsal surface which demarcation between gnathosoma, different is divided by definite suture lines. A body regions and genital plate which was not longitudinal groove appears on the female mentioned in previous light (Jeppson et al., ventral side formed by two cuticular flaps 1975; Zhang, 2003) or SEM studies (Martin, and ending in a funnel like structure. This 1991). Definite pleating was observed in the indicates that the integument of P. latus is skin of metapodosoma so as to overlap the somewhat softer than previously mentioned posterior part of the propodosoma and the in the SEM study of Martin (1991) or other anterior part of the opisthosoma. This light microscopic descriptions of the above pleating causing a kind of articulation authors. The funnel like structure observed in between body regions probably facilitates the the female may be related to the process of process of holding the female and/or mating. receiving or transporting of sperms during The male carries four pairs of long and insemination. spindle shaped legs. The 1st and 2nd pairs The anterior and posterior two pairs of extend forward, while the 3rd and 4th ones legs in female P. latus were widely separated extend more posterior (Denmark, 1980). The similar to that mentioned by Jeppson et al. fourth pair of legs in male tarsonemids was (1975). The present SEM study revealed the regarded as an accessory copulatory insertion of both the anterior and posterior appendage (Jeppson et al., 1975) since it two legs in ventrolateral grooves which were clasps the female during copulation not mentioned in previous studies. The (McDaniel, 1979). This appendage anterior two pairs of legs are more densely represents an adaptation developed for better studded with setae than the posterior ones performance of copulatory functions but similar to that reported by Jeppson et al. passively in locomotion (Jeppson et al., (1975) and Zhang (2003). The three 1975). The fourth pair of legs of the male in segments forming the fourth leg of female P. the present study was markedly enlarged, latus with the two terminal long setae on the had four segments; the last one carries two tibiotarsus, fusion between genu and femur long setae and terminates with a button like and absence of claws or empodia in that leg strong claw. Similar observations were as well as the four segments forming the previously reported by Zhang (2003). The fourth leg of the male and its termination body length of the male measured 138.9 µm with a single claw and the claspers like in the present study whereas Denmark structure were applied as a taxonomic (1980) and Peręz-Otero et al. (2007) reported features for family Tarsonemidae (Krantz, it to be 168 µm and110 µm, respectively. 1978). Female P. latus length in the present Other measurements of width, body regions, study measured 138.9 µm whereas legs, setae, leg segments, gnathosoma and previously it was reported as 160 µm by Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Pagotarsonemus latus 71

Peręz-Otero et al. (2007). The present study teratosities in plant tissues on which they added measurements of width, different body feed, presumably through injection of regions, legs, gnathosoma and genital salivary toxins. This assumption needs capsule as well as SEM description of the further histological studies to verify. female which were rarely presented in The semi-circular shape of the male previous light or SEM studies. genital capsule and its subterminal location The situation of clavate on ventral side of opisthosoma is similar to pseudostigmatic organs, stigmatal openings that described by Denmark (1980). The and trichobothrial sensilla in the anterolateral present study added that it consisted of 2-3 margin of female P. latus was previously triangular genital papillae and carried two mentioned by Zhang (2003) and represent pairs of short dorsal setae. The anal plate distinctive features for family Tarsonemidae surrounding the anus is large, well defined (Krantz, 1978). Function of pseudostigmatic and located infront of the genital capsule on organs was uncertain (Jeppson et al., 1975). the ventral side of opisthosoma and marked Jeppson and his colleagues suggested that by 3 indentations similar to Denmark (1980) these organs act as highly modified sensilla findings. or specialized sense organs since they seem The present fine morphological study to have no relationship to the tracheal provides the essential bases for identification system. Presence of stigmatal openings in of the undetectable P. latus regarded female P. latus and their absence in case of undoubtedly as the most important mite of male was also noticed by Jeppson et al. many crops in green houses. Such needed (1975) and Zhang (2003). Jeppson et al. information will help in developing an (1975) and Krantz (1978) mentioned that the integrated control system that will overcome stigmatal openings were internally connected their reproduction capacity and modes of to a tracheal system in female tarsonemid dispersal and which complies with the mites beginning from the external orifices, environmental requirements established by converge in the region of leg II and many countries. disappear inconspicuously in the opithosomal region. 5. ACKNOWLEDGMENTS Light microscope examination of male We express our deep gratitude and or female gnathosoma showed that it was appreciation to Prof. Ahmed M. Taha, Plant oval, capsulate and carried two median Protection Research Institute, Agricultural slender stylettiform chelicerae between non Research Center, Giza Governorate, Egypt, segmented triangular palps. These for his great interest, supervision and observations were previously reported by providing lab facilities throughout the study. Krantz (1978) and Zhang (2003). The present SEM study showed that the 6. REFERENCES gnathosoma was distinct from the idiosoma, Alberti, G. and Coons, L. B. (1999). Acari: completely directed downwards where it was Mites. In: Harrison, F. W. and Foelix, R. difficult to observe from the dorsal view and F. (eds). Microscopic anatomy of appeared broad at its base and narrow invertebrates, vol 8C. Chelicerate terminally like a rostrum. The latter Arthropoda. Wiley-Liss Inc, New York, observations greatly support the proper pp 515–1215. function of the chelicerae which include Banks, N. (1904). Class 111, Arachnida, piercing of plant cells and sucking the plant Order 1, Acarina, four new species of sap. Furthermore, the articulation between injurious mites. Journal of the New York gnathosoma and idiosoma helps in forward Entomological Society, 12:53-56. or downward motion of the former to Callaini, G. and Mazzini, M. (1984). Fine undergo the above functions. Krantz (1978) structure of the egg shell of Tyrophagus mentioned that P. latus may produce

72 Ashraf A. Montasser et al.: Description of life cycle stages of the broad mite Polyphagotarsonemus latus

putrescentiae (Schrank) (Acarina: Nickles, E.P.; Ghiradella, H.; Bakhru, H. and Acaridae). Acarologia, 25: 359-364. Haber, A. (2002). Egg of the Karner Blue Cho, M. R.; Chung, S. K. and Lee, W. K. Butterfly (Lycaeides Melissa samuelis): (1993). Taxonomic study on cyclamen Morphology and elemental analysis. J. mite ( pallidus) and broad Morphol., 251: 140-148. mite (Polyphagotarsonemus latus). Nucifora, A. and Vacante, V. (2004). Citrus Korean Journal of Applied Entomology, mites in Italy. VII. The family 32(4): 433-439. Tarsonemidae. Species collected and Denmark, H. A. (1980). Broad mite, notes on ecology. Acarologia, 44(1/2): Polyphagotarsonemus latus (Banks). 49-67. FDACS-DPI Bureau of Entomology Peręz-Otero, R.; Mansilla-Vazquez, J. P. and Circular No. 213. Salinero-Corral, M. C. (2007). First Jeppson, L.R.; Baker, E.W. and Keifer, H.H. report of the broad mite (1975). Mites Injurious to Economic Polyphagotarsonemus latus (Banks) on Plants. University of California Press, Camellia japonica in Spain. American Berkeley, California, 614 pp. Camellia Yearbook, pp. 52-56. Krantz, G. W. (1978). A manual of Witaliňski, W. (1993). Egg shells in mites: Acarology. 2nd ed. Oregon State Vitelline envelope and chorion in University Book Stores, Corvallis, Acaridida (Acari). Exp. Appl. Acarol., Oregon, USA, pp. 509. 17: 321-.344. Labanowski, G. and Soika, G. (2006). Zhang, Z. Q. (2003). Tarsonemid mites. Tarsonmid mites on ornamental plants in Mites of greenhouses: Identification, Poland: new data and an overview. biology and control. CABI Publishing Biological Letter, 43(2): 341-346. Co., UK, pp. 99-126. McDaniel, B. (1979). How to know the mites and . Wm. C. Brown Company Publishers. Dubuque, Iowa. Martin, N. A. (1991). Scanning electron micrographs and notes on broad mite Polyphagotarsonemus latus (Banks), (Acari: Heterostigmata: Tarsonemidae). New Zealand Journal of Zoology, 18(3): 353-356.