Presumed virus particles mostly occur in parenchyma cells of the lesion in affected orange leaves, fruits or stems. Particles are short, bacilliform, 120-130 nm long (occasionally up to 300 nm) and 50-55 nm wide. They occur within the lumen of the endoplasmic reticulum (Kitajima et al., 1974, Colariccio et al., 1995). There is a report of similar but unenveloped particles in the nucleoplasm (Kitajima et al., 1972).;In addition to the presence of the rhabdovirus-like particles within the endoplasmic reticulum of tissues from the lesion, dense viroplasm-like material is commonly found in the cytoplasm, near the particles. Small vesicle-containing fibrillar materials are frequently present in the vacuole, associated with the tonoplast, next to the dense material (Kitajima et al., 1972, Colariccio et al., 1995).;Chloroplasts are usually affected with a disorganized hypertrophied lamella system (Kitajima et al., 1972, Rodrigues, 1995). There is a report in which rod-like particles, considered to be naked rhabdovirus particles accumulate in the nucleoplasm associated with the nuclear envelope (Kitajima et al., 1972).
Round to elliptical local lesions are seen on fruits, leaves and twigs. The severity of the lesions varies with the type of citrus and the region of origin. Leaf symptoms are usually round with a dark-brown central spot about 2-3 mm diameter, surrounded by a chlorotic halo, in which 1-3 brownish rings frequently appear surrounding the central spot, the overall lesion size varies from 10 to 30 mm, though larger lesions may form by the fusion of 2 or more adjacent lesions.;On fruits, lesions are necrotic spots 10-20 mm in diameter, with a necrotic centre. Gum exudation is occasionally observed on the lesion. On green fruits, the lesions are initially yellowish, becoming more brown or black, sometimes depressed, and reducing the market value of the fruits.;On stems, lesions may be protuberant, cortical, grey or brown. Lesions may coalesce when present in large numbers, leading to the death of the twig. In extreme cases observed in different places (JCV Rodrigues, personal communication), as described initially in 'lepra explosiva' in Argentina, severe defoliation and fruit fall may occur (Frezzi, 1940, Bitancourt, 1955, Rossetti et al., 1969).;Citrus leprosis lesions are usually very characteristic, but may sometimes be mistaken for lesions of citrus canker caused by the bacterium Xanthomonas axonopodis pv. axonopodis, or zonate chlorosis (Rossetti, 1980). Zonate chlorosis, which is associated with infestation by the same mites, does not become necrotic. Symptoms are essentially concentric green and chlorotic rings (Bitancourt, 1934).;Other viral diseases are vectored by Brevipalpus phoenicis in Brazil. Coffee ringspot virus in Coffea arabica (Chagas, 1978), Ligustrum ringspot virus in Ligustrum lucidum (Rodrigues et al., 1995), and green spot of passion fruit in Passiflora edulis (Kitajima et al., 1997). In addition, Brevipalpus californicus is a vector of Orchid fleck virus in orchids (Maeda et al., 1998). However, cross-transmission was not described among these viruses and leprosis.
P. glandulosa exhibits high levels of variability in morphological characters. Variations are observed principally in native populations. In invading populations, clinal variations are obscured because of the rapid and widespread dispersal of diverse genetic material by humans and animals over a range of site and climatic conditions. The following description is adapted from Burkart (1976).
P. glandulosa is a tree up to 9 m tall, with a trunk diameter up to 1 m, though larger specimens are recorded. Spines are axillary, uninodal, 1-4.5 cm long, mostly solitary, sometimes very few, solitary or paired, sometimes with solitary and paired thorns on different nodes of the same twig. Leaves bipinnate, glabrous, 1-2 pairs of pinnae per leaf, up to 15 cm long. Pinnae 6-17 cm long, each with 6-17 pairs of leaflets. Leaflets distant on the rachis, 2-6.3 cm long by 1.5-4.5 mm broad, linear or oblong, 5-15 times as long as broad, obtuse, glabrous, subcoriaceous, prominently veined below. Flowers yellow, racemes 5-14 cm long, multiflorous, petals 2.5-3.5 mm long, ovary stalked, villous. Legume straight, 8-20 cm long by 0.7-1.3 cm broad, rarely subfalcate, compressed to subterete, submoniliform, glabrous, straw coloured or tinged with violet, short-stalked, with strong, varyingly acuminate. There are 5-18 seeds per pod, seeds 6-7 mm long, oblique to longitudinal. P. glandulosa var. torreyana has a similar habit to P. glandulosa var. glandulosa but with generally shorter pinnae and shorter leaflets that are less distant on the rachis. P. glandulosa var. prostrata is similar in foliar and floral morphology to var. glandulosa but differs in its habit, being generally a low-growing shrub.
P. glandulosa has been widely introduced and planted as a fuel and fodder tree. Seed are spread widely by grazing animals from established plantations or single trees around houses or water-holes, and will persist for long periods in the seed bank. It has shown itself to be a very aggressive invader, especially in sub-tropical arid and semi-arid natural grasslands, both in its native range and where introduced. It is a nitrogen-fixing species and very drought and salt tolerant, rapidly out-competing other vegetation. Thorniness and a bushy habit enable it to quickly block paths and make whole areas impenetrable. Invasion in the native range generally involves an increase in plant density rather than an increase in its range. P. glandulosa is a declared noxious weed in Australia and South Africa, and the genus as a whole is regulated in several other countries. It is also reported as invasive in other southern African countries, notably Botswana and Namibia where it is known to hybridise with P. velutina, also in Australia, though P. glandulosa tends to dominate. In terms of ecology, uses, management and control, P. glandulosa and P. velutina can be effectively treated together, as a species complex.
Plant: T. diversifolia is 2-3 m tall with upright and sometimes ligneous stalks. It forms woody shrubs.
T. diversifolia, commonly known as the tree marigold, is a herbaceous flowering plant in the Asteraceae family. Native to Mexico and Central America, it has been introduced and is now naturalized in tropical parts of Asia and Africa. It is also naturalized in some Pacific islands, where it is found along roadsides and in disturbed areas. T. diversifolia tolerates heat and drought and can rapidly form large herbaceous shrubs. Rapid vegetative reproduction and significant production of lightweight seeds, which can be dormant in the soil for up to four months, allow T. diversifolia to quickly invade disturbed habitats. By forming dense stands it prevents the growth of young native plants. Depending on the area, T. diversifolia may be either annual or perennial. Being able to produce flowers and seeds throughout the year, coupled with the ability of seeds to be dispersed by wind, water and animals, makes it particularly easy for T. diversifolia to quickly colonize new areas. Shoot and root growth and nutrient uptake of several plants may be adversely affected by T. diversifolia.
Imeokpara and Okusanya (1994) observed that most farmers found it difficult to manage T. diversifolia infestation in most crop fields, but particularly rice and maize fields. T. diversifolia has been reported to contain some allelochemicals and therefore may be capable of posing a serious phytotoxicity threat to agricultural crops. Goffin et al. (2002) isolated tagitinin C, a known sesquiterpene lactone (Pal et al., 1977;Baruah et al., 1994), from the aerial parts of T. diversifolia. According to Ayeni et al. (1997) several studies have indicated that these allelochemicals and their derivatives are toxic and may inhibit shoot and root growth and nutrient uptake of several plants. Ilori et al. (2007) similarly observed that the radical growth of Oryza sativa was inhibited by aqueous extract of T. diversifolia.
The colonies of C. formosanus contain three primary castes: the reproductives, soldiers, and workers. The majority of the nestmates are workers that are responsible for the acquisition of nutrients, i.e. cellulose in the wood. The head width of the white soft-bodied worker is approximately 1.2-1.3 mm and the body length is approximately 4-5 mm. The thorax is narrower than head width. The alates and soldiers are most useful for identification. The alates are yellowish-brown and 12-15 mm long. There are numerous small hairs on the wings of these comparatively large swarmers. The alates are attracted to lights, so they are usually found near windows, light fixtures, windowsills and spider webs, around well-lit areas. The soldiers are approximately the same size as the workers and have an orange-brown oval-shaped head, curved mandibles and a whitish body. When disturbed, the soldiers readily attack any approaching objects and may secrete a white gluey defensive secretion from the frontal gland. There are more soldiers (10-15%) in a C. formosanus colony than in a subterranean termite colony, such as Reticulitermes spp. (1-2%).
Occasionally the foraging tubes may be observed on the wood surface or tree trunk. During the swarming season (April to June), elongated mud tubes that serve as flight exit slits may be seen. The damage by C. formosanus tends to occur in places with high moisture including the bathroom, kitchen sinks and leaky roofs. An acoustic emission device (AED) may be used to locate sites with feeding activity, but most AEDs have a limited detection range (Scheffrahn et al., 1993).
Large colonies of C. formosanus generally live underground. When these termites invade a house aboveground, the foraging tubes of approximately 0.5-1 cm in diameter may be found connecting the soil and the infested house. In severe infestations, C. formosanus hollows out the wood leaving a paper-thin surface and the hollowed wood surface may look blistered or peeled. Another characteristic of C. formosanus is carton nest material that is made from termite faeces, chewed wood and soil. The honeycomb-like carton nests can be as large as 1-1.5 m in diameter and are usually found in structure-voids such as between walls and beneath sinks.
C. formosanus is often transported by boats and shipping containers to port cities before being carried further inland via landscape materials such as railroad ties (railway sleepers). This may explain the current C. formosanus distribution in the USA with coastal areas more densely infested than inland areas (Hochmair and Scheffrahn, 2010). Temperature and humidity are primary factors affecting the establishment of C. formosanus, and it is potentially invasive to areas of high humidity approximately 35° north and south of the equator (Su and Tamashiro, 1987). Competition from native species is another limiting factor for many exotic pests, but C. formosanus is more aggressive and is known to out-compete the endemic termites such as Reticulitermes species. Another factor that has allowed the successful establishment and spread of C. formosanus in exotic areas has been the pest control industry's heavy reliance on soil termiticide barriers for subterranean termite control since the 1950s. Numerous studies, using mark-recapture methods, have revealed that a single colony of C. formosanus might contain several million termites that forage up to 100 m in the soil (Lai, 1977;Su and Scheffrahn, 1988). These agree with the results of excavation studies for C. formosanus colonies (Ehrhorn, 1934;King and Spink, 1969). Because of the large colony size, the application of soil termiticides beneath a structure does not usually have a major impact on the overall population, and the surviving colony continues to produce alates that can further infest nearby areas. Once established, C. formosanus has never been completely eradicated from an area. The dependency of soil termiticide barriers as the primary tool for subterranean termite control is probably the main reason for the establishment and spread of C. formosanus from four isolated port cities in the 1960s in the USA to all south-eastern states by 2001.
C. formosanus is an opportunistic feeder of any material containing cellulose. A large number of living plants are known to be attacked by C. formosanus, but it usually does not kill the plants unless the root system is significantly damaged (Lai et al., 1983;La Fage, 1987). Records show that living citrus, eucalyptus and sugar canes (Saccharum sp.) may be killed by C. formosanus, but in most cases damage occurs in the heartwood of a tree. The infested trees may be more easily blown over by high winds due to the loss of structural strength. The pest status of C. formosanus is most significant when it attacks wood products in a house such as structural lumbers, cabinets, etc. C. formosanus is also known to damage non-cellulose materials in search of food, including plastic, concrete and soft metal. Occasionally underground high-voltage power lines may be penetrated by C. formosanus, resulting in an area-wide power cut.
The small hairy larvae of U. lugens feed gregariously on the upper and lower epidermis, the pallisade tissue and the spongy mesophyll of the leaf but avoid the oil cells and the veins. This feeding habit results in the leaf being ‘skeletonised’, hence the common name of the insect. Larger larvae, from the fifth instar, feed individually and consume the entire leaf blade down to the mid-rib (Cobbinah, 1978). From around the fifth instar, U. lugens larvae retain their moulted head capsules on top of their head, creating a distinctive ‘head dress’. Larvae spin a camouflaged pupal cocoon incorporating their own hairs and fragments of surrounding materials. Adult moths are approximately 10 mm in length with a wingspan of 25 to 30 mm. The forewings are dark grey with several dark wavy lines connecting front and rear wing margins (Anonymous, 1979). The hindwings are pale grey-brown.
U. lugens can be detected by searching leaves. Eggs are laid in batches on the leaf surface, and young larvae feed gregariously adjacent to the egg batch after emergence. Oviposition tends to occur mainly in the lower crow of the tree (Morgan and Cobbinah, 1977). Young larvae skeletonise the leaves, making leaf damage easy to detect. Skeletonised leaves often have characteristic patches of cast skins where larvae have moulted before moving on. After the fifth instar larvae disperse and can be found singly, often in the vicinity of abandoned skeletonised leaves. When close to pupation, larvae wander in search of a suitable site. Camouflaged cocoons are formed in the bark or leaf litter and are very difficult to find. A synthetic pheromone has been developed, and can be used for detection and delimiting surveys (Suckling et al., 2005).
Early instar larvae skeletonise leaves, which then turn brown, giving the tree a scorched look when damage is heavy. Older larvae feed on the entire leaf blade down to the midrib, which can resemble defoliation.
U. lugens was first considered a serious pest of natural eucalypt forests in Western Australia in 1983 when the first severe outbreak occurred there (Strelein, 1988). Prior to that it was widely known as a pest of eucalypt forests in eastern Australia (Campbell, 1962;Harris, 1974). As these natural forests are or were managed for timber production, it is considered an economically important pest in its native range. Damage to amenity trees is also a common problem, although few trees are killed by this defoliation (Anonymous, 1979).
Morgan and Cobbinah (1977) list 149 Eucalyptus species and one Angophora species found to be oviposition hosts, out of more than 250 species surveyed in a field study in Adelaide, South Australia. Not all of the oviposition hosts proved to be suitable larval hosts. That work was part of a wider study in which over 580 tree species were surveyed (Cobbinah, 1978). Significant defoliation events have occurred in natural forests of Eucalyptus camaldulensis, E. calophylla and E. marginata in mainland Australia (Campbell, 1962;Strelein, 1988;Farr, 2002), and in plantations of E. nitens in Tasmania (Anonymous, 1994) although damage is common on a wide range of eucalypt species. E. nitens and E. fastigata are important commercial plantation and farm forestry species in New Zealand, although commercial impacts of U. lugens on these species have yet to be felt. The iconic native species Metrosideros excelsa (pohutukawa) has been recorded as a host in New Zealand, although this seems to only occur through spill-over feeding from near by eucalypts (Potter at al., 2004) and is not significantly impacting these trees. A number of other new host records have occurred in New Zealand since U. lugens arrived in that country, most notably on a range of deciduous Northern Hemisphere species. The most significant damage on these species has occurred on Betula pendula (silver birch), where some trees have been defoliated (J Bain, Scion, Rotorua, New Zealand, personal communication, 2008).
In New Zealand, U. lugens has been recorded on 58 tree species (J Bain, Scion, Rotorua, New Zealand, personal communication, 2008), mainly from the genus Eucalyptus. It is causing significant damage in New Zealand on Lophestemon confertus, which is commonly planted as a street tree in some parts of Auckland. In a laboratory study of larval suitability of 18 highly valued eucalypt species in New Zealand, Potter and Stephens (2005) found E. nitens, E. nicholii and E. fastigata were most at risk.
According to Peeters (2004) and with additional material from Hubbard (1968), P. pratense is a tall, tufted or single-stemmed, short-lived, cool-season perennial grass. The plant is robust, hairless and caespitose. Stems are erect, 20-100(-150) cm tall, often bulging at the base and forming a small bulb. The blade is rolled when young, large (3-10 mm wide), flat, slightly rough on the margin, rather long (reaching 45 cm), pale green to greyish green. The ligule is strong, obtuse and white, and there are no auricles. The spike-like panicle is cylindrical, 6-20(-30) cm long. Spikelets are 1-flowered, breaking up at maturity above the glumes. Glumes persistent, narrowly oblong, truncate, keeled, 3-nerved, the keels fringed with soft spreading hairs and produced at the tip into a rigid awn 1-2 mm long. Lemma and palea one-third to three-quarters the length of the glumes. Anthers 2 mm long. Seeds are small, 2 mm long, with a 1000-seed weight of 0.3 to 0.7 g. P. pratense produces few tillers (4000 to 10,000 tillers/m?) compared to perennial ryegrass (Lolium perenne, 6000 to 15,000 tillers/m?). However, leaves are produced at a slightly faster rate than in perennial ryegrass, and the maximum number of leaves per tiller is up to 6-7 (rarely 8) against only 3 for the ryegrass. Leaf lifetime is also longer, so that timothy is able to accumulate a lot of standing biomass before senescence commences.
Descriptions of L. trifolii refer to fresh materials. Dry specimens may be distorted due to the manner in which they have been preserved. Also, the age of the specimen, when killed, will have some effect on its preservation characteristics.
For accurate identification, examination of the leaf mine and all stages of development are crucial.
L. trifolii eggs are 0.2-0.3 mm x 0.1-0.15 mm, off white and slightly translucent.
This is a legless maggot with no separate head capsule, transparent when newly hatched but colouring up to a yellow-orange in later instars and is up to 3 mm long. L. trifolii larvae and puparia have a pair of posterior spiracles terminating in three cone-like appendages. Spencer (1973) describes distinguishing features of the larvae. Petitt (1990) describes a method of identifying the different instars of the larvae of L. sativae, which can be adapted for use with the other Liriomyza species, including L. trifolii.
This is oval and slightly flattened ventrally, 1.3-2.3 x 0.5-0.75 mm with variable colour, pale yellow-orange, darkening to golden-brown. The puparium has posterior spiracles on a pronounced conical projection, each with three distinct bulbs, two of which are elongate. Pupariation occurs outside the leaf, in the soil beneath the plant.
Menken and Ulenberg (1986) describe a method of distinguishing L. trifolii from L. bryoniae, L. huidobrensis, and L. sativae using allozyme variation patterns as revealed by gel electrophoresis.
L. trifolii is very small: 1-1.3 mm body length, up to 1.7 mm in female with wings 1.3-1.7 mm. The mesonotum is grey-black with a yellow blotch at the hind-corners. The scutellum is bright yellow;the face, frons and third antennal segment are bright yellow. Male and female L. trifolii are generally similar in appearance.
L. trifolii are not very active fliers, and in crops showing active mining, the flies may be seen walking rapidly over the leaves with only short jerky flights to adjacent leaves.
The frons, which projects very slightly above the eye, is just less than 1.5 times the width of the eye (viewed from above). There are two equal ors and two ori (the lower one weaker). Orbital setulae are sparse and reclinate. The jowls are deep (almost 0.33 times the height of the eye at the rear);the cheeks form a distinct ring below the eye. The third antennal segment is small, round and noticeably pubescent, but not excessively so (vte and vti are both on a yellow ground).
Acrostical bristles occur irregularly in 3-4 rows at the front, reducing to two rows behind. There is a conspicuous yellow patch at each hind-corner. The pleura are yellow;the meso- and sterno-pleura have variable black markings.
Length 1.3 -1.7 mm, discal cell small. The last section is M(sub)3+4 from 3-4 times the length of the penultimate one.
The shape of the distiphallus is fairly distinctive but could be mis-identified for L. sativae. Identification using the male genitalia should only be undertaken by specialists.
The head (including the antenna and face) is bright yellow. The hind margin of the eye is largely yellow, vte and vti always on yellow ground.
The mesopleura is predominantly yellow, with a variable dark area, from a slim grey bar along the base to extensive darkening reaching higher up the front margin than the back margin. The sternopleura is largely filled by a black triangle, but always with bright yellow above.
The femora and coxa are bright yellow, with the tibia and tarsi darker;brownish-yellow on the fore-legs, brownish-black on the hind legs. The abdomen is largely black but the tergites are variably yellow, particularly at the sides. The squamae are yellowish, with a dark margin and fringe.
Although individual specimens may vary considerably in colour, the basic pattern is consistent.
L. trifolii are small black and yellow flies which may be detected flying closely around host plants or moving erratically and rapidly upon the leaf surfaces. Inspection of the leaf surface will reveal punctures of the epidermis and the obvious greenish-white mines with linear grains of frass along their length. For accurate identification, examination of the leaf mine and all stages of development are crucial.
L. trifolii larvae will be found feeding at the end of the mine, or the mine will end with a small convex slit in the epidermis where the larva has left the mine to pupariate on the ground. Sometimes the puparium may be found adhering to the leaf surface, although in most cases the fully-fed larva will have found its way to the ground beneath the plant to pupariate. This is especially true in hot, dry conditions where the larva/puparia would quickly desiccate if exposed on the leaf surface. Empty puparial cases are split at the anterior end, but the head capsule is not usually separated from the rest of the case.
Mined leaves should be collected into polythene bags and transferred to a press as soon as possible. Leaves containing larvae intended for breeding should be collected into individual polythene bags, which on return to the laboratory should be slightly over-pressurized by blowing into them before sealing the end. Blowing up the bag by mouth and sealing it adds valuable carbon dioxide to the moist air mix. Constant attention is required to ensure that puparia are transferred to individual tubes until the fly emerges. If the plant material begins rotting, good material with feeding larvae must be removed to more sanitary conditions.
When puparia are observed they can be very carefully removed to tubes containing a layer of fine sand, or a small strip of blotting paper or filter paper. This should be kept damp (never wet) until the adult emerges.
On emergence, the fly should be kept for at least 24 hours to harden up. Do not allow condensation to come into contact with the fly, or it will stick to the water film and be damaged.
Field collection of the adult L. trifolii is done by netting. The use of sticky traps, especially yellow ones, placed near host plants is a very effective method of collection and estimation of infestation.
If the puparial stage is collected from the soil, care must be taken not to damage the puparial skin or death will almost certainly follow. The pupae should be stored in glass tubes on a layer of clean sand or, better still, thick filter paper. The tube must have high humidity, but be free of condensation.
When the fly emerges, it must be allowed to harden for 24 hours before killing for identification purposes. Ensure that the tube has no condensation present.
Newly emerged adult L. trifolii are generally softer than specimens aged for several days and may crinkle as drying proceeds, especially the head. The ptilinal sac may still protrude from the suture between the frons and face obliterating some important characteristics. Adults should be dried slowly in the dark in a sealed receptacle over blotting paper. If preserving wet is preferred, the live specimen should be dropped into 20-40% alcohol, and transferred to 70-90% alcohol after 2 days.
L. trifolii feeding punctures appear as white speckles between 0.13 and 0.15 mm in diameter. Oviposition punctures are usually smaller (0.05 mm) and are more uniformly round.
L. trifolii leaf mines can vary in form with the host plant, but when adequate leaf area is available they are usually long, linear, narrow and not greatly widening towards the end. They are usually greenish white.
In very small leaves the limited area for feeding results in the formation of a secondary blotch at the end of the mine, before pupariation. In Kenya, Spencer (1985) notes the growth of many L. trifolii from mines which began with a conspicuous spiral. This is not a characteristic associated with L. trifolii on other continents.
The frass is distinctive in being deposited in black strips alternately at either side of the mine (like L. sativae), but becomes more granular towards the end of the mine (unlike L. sativae) (Spencer, 1973).
Fungal destruction of the leaf may also occur as a result of infection introduced by L. trifolii from other sources during breeding activity. Wilt may occur, especially in seedlings.
The host range of L. trifolii includes over 400 species of plants in 28 families including both ornamental crops (Bogran, 2006) and vegetables (Cheri, 2012). The main host families and species include: Apiaceae (A. graveolens);Asteraceae (Aster spp., Chrysanthemum spp., Gerbera spp., Dahlia spp., Ixeris stolonifera, Lactuca sativa, Lactuca spp., Zinnia spp.);Brassicaceae (Brassica spp.);Caryophyllaceae (Gypsophila spp.);Chenopodiaceae (Spinacia oleracea, Beta vulgaris);Cucurbitaceae (Cucumis spp., Cucurbita spp.);Fabaceae (Glycine max, Medicago sativa, Phaseolus vulgaris, Pisum sativum, Pisum spp., Trifolium spp., Vicia faba);Liliaceae (A. cepa, Allium sativum) and Solanaceae (Capsicum annuum, Capsicum frutescens, Petunia spp., Solanum lycopersicum, Solanum spp.) (EFSA, 2012).
It is now considered to be the most important pest of cowpea (Vigna uniguilata), towel gourd (Luffa cylindrica), cucumber (Cucumis sativus) and many other vegetable crops in southern China (Gao, 2014). In Europe, L. trifolii is a major pest of lettuce, beans, cucumber and celery, Capsicum sp., carnations, clover, Gerbera sp., Gypsophila sp., lucerne, Senecio hybridus, potatoes and tomatoes (EFSA, 2012). It is now a major pest of the Compositae worldwide, particularly chrysanthemums (including Dendranthenum, the commercial 'Mum') in North America, Colombia, and elsewhere. It also causes severe damage to different open field crops, such as chili peppers in Mexico.
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