Jose Carlos V. Rodrigues

Citrus leprosis virus vectored by Brevipalpus phoenicis (Acari: Tenuipalpidae) on citrus in Brazil.

Citrus leprosis is caused by Citrus leprosis virus (CiLV) that is transmitted by mites in the genus Brevipalpus (Acari: Tenuipalpidae). This disease directly reduces production and the life span of the citrus plant. The main symptoms of the disease include lesions on fruits, leaves, and twigs or small branches, causing premature fruit drop, defoliation, and death of the twigs or branches leading to serious tree decline. Leprosis is a highly destructive disease of citrus, wherever it occurs. The Brazilian citrus industry spends over 100 million US dollars annually on acaricides to control the vector, Brevipalpus phoenicis (Geijskes). This review contains information about the history of the etiology of citrus leprosis, its geographical distribution, host range, the role of the mite vectors, viral morphology and relationships with the infected cell, and transmissibility of the virus by the mite. In addition, data on the mite-virus-plant relationship, disease damage, and strategies for controlling disease spread are presented.

Brevipalpus-Transmitted Plant Virus and Virus-Like Diseases: Cytopathology and Some Recent Cases

An increasing number of diseases transmitted by Brevipalpus mite species (Acari: Tenuipalpidae) is being identified that affect economically important plants such as citrus, coffee, passion fruit, orchids, and several ornamentals. All of these diseases are characterized by localized lesions (chlorotic, green spots, or ringspots) on leaves, stems, and fruits. Virus or virus-like agents are considered to be the causal agents, possibly transmitted in a circulative-propagative manner by Brevipalpus mites. The virus or virus-like particles are short, rod-like, or bacilliform, that induce two characteristic types of cell alteration: (1) ‘Nuclear type’ – nuclei of parenchyma and epidermal cells in the lesions often contain a large electron lucent inclusion. Short, naked, rod-like (40–50 nm × 100–110 nm) particles may be seen in the viroplasm or nucleoplasm and in the cytoplasm. These particles are commonly arranged perpendicularly on the membranes of the nuclear envelope or endoplasmic reticulum (ER). In a very few instances, they were found to be membrane-bound, within the ER cavities. (2) ‘Cytoplasmic type’ – short bacilliform particles (60–70 nm × 110–120 nm) are present within the cisternae of the ER and often have electron dense viroplasm of varied shapes present in the cytoplasm. Bacilliform particles may be seen budding into the ER lumen near the viroplasm. These particles resemble those of members of the Rhabdoviridae, but are shorter. The only sequenced virus of this group, orchid fleck virus (OFV), has a negative sense (bipartite) type ssRNA genome, but its organization is similar to known rhabdoviruses, which are monopartite. Both types of cytopathological effects have been found associated with citrus leprosis. In orchids, OFV has a ‘nuclear type’ of cytopathology, but in some species the ‘cytoplasmic type’ has been found associated with ringspot symptoms. In Hibiscus and Clerodendron, green spot symptoms have been associated with the cytoplasmic type of cell alteration, while chlorotic spots, in the same species, are associated with the nuclear type. In a few cases, both types of cytopathological effects have been found in the same tissue and cell.

Host Plants of Brevipalpus californicus, B. obovatus, and B. phoenicis (Acari: Tenuipalpidae) and their Potential Involvement in the Spread of Viral Diseases Vectored by these Mites

The family Tenuipalpidae has over 622 species in 30 genera described worldwide. A total of 928 plant species in 513 genera within 139 families are recorded hosts of one or more of the following species: Brevipalpus californicus (Banks), B. obovatus Donnadieu, and B. phoenicis (Geijskes). B. californicus has 316 plant species reported as hosts compared with 451 and 486 host plants for B. obovatus and B. phoenicis, respectively. There are 67 genera of plants within 33 families that are reported hosts of only B. californicus, 119 genera within 55 plant families that are hosts of only B. obovatus, and 118 genera of plants within 64 families that are hosts of only B. phoenicis. There are 14 genera of plants within 12 families that are hosts to both B. californicus and B. obovatus, while there are 40 genera of host plants within 26 families that are hosts for both B. californicus and B. phoenicis. A total of 70 genera of host plants within 39 families have been reported as hosts of both B. obovatus and B. phoenicis, while 77 genera of plants within 44 families have been reported as hosts of all three Brevipalpus species. Geographical differences in the three species of Brevipalpus identified on different plant species within the same genus are common.

Brevipalpus californicus, B. obovatus, B. phoenicis, and B. lewisi (Acari: Tenuipalpidae): a Review of their Biology, Feeding Injury and Economic Importance

The genus Brevipalpus includes most of the economically important species of Tenuipalpidae. Many Brevipalpus species reproduce by theletokous parthenogenesis while other species reproduce by male fertilization of female eggs. Previous researchers have determined that Brevipalpus californicus (Banks), B. obovatus Donnadieu, and B. phoenicis (Geijskes) females were haploid with two chromosomes. The life cycle and developmental times for these three species are reviewed. Longevity of each Brevipalpus species is two to three times greater than corresponding longevities of various tetranychid mites. Brevipalpus mites inject toxic saliva into fruits, leaves, stems, twigs, and bud tissues of numerous plants including citrus. Feeding injury symptoms on selected plants include: chlorosis, blistering, bronzing, or necrotic areas on leaves by one or more Brevipalpus mites. Premature leaf drop occurred on ‘Robinson’ tangerine leaves in Florida (USA). Leaf drop was observed in several sweet orange and grapefruit orchards in Texas (USA) that were heavily infested with Brevipalpus mites feeding on the twigs, leaves, and fruit. Initial circular chlorotic areas appear on both sweet orange and grapefruit varieties in association with developing populations of Brevipalpus mites in Texas. These feeding sites become progressively necrotic, darker in color, and eventually develop into irregular scab-like lesions on affected fruit. Russeting and cracking of the fruits of other plant hosts are reported. Stunting of leaves and the development of Brevipalpus galls on terminal buds were recorded on sour orange, Citrus aurantium L., seedlings heavily infested with B. californicus in an insectary. The most significant threat posed by these mites is as vectors of a potentially invasive viral disease called citrus leprosis.

Coffee Ringspot Virus Vectored by Brevipalpus phoenicis (Acari: Tenuipalpidae) in Coffee

Coffee ringspot is characterized by conspicuous ringspot symptoms on leaves, berries, and less frequently on twigs. It is caused by coffee ringspot virus (CoRSV), a short, bacilliform virus (40 nm × 100–110 nm). The virus is not seed borne and is transmitted by Brevipalpus phoenicis (Geijskes). Transovarial transmission within the mite does not occur. CoRSV has been mechanically transmitted to Chenopodium amaranticolor Coste and Reynaud, C. quinoa Wildenow, Beta vulgaris L., and Alternanthera tenella Colla resulting in local lesions. Systemic infection within both C. amaranticolor and C. quinoa occurs. Virions are found in the nucleus or cytoplasm of infected cells, commonly associated with membranes. Occasionally, membrane bounded particles are found within the cisternae of the endoplasmic reticulum. A characteristic electron lucent, nuclear inclusion is commonly found in many infected cells. These cytopathic effects place CoRSV among the nuclear type of Brevipalpus-borne viruses. The disease has been reported in several Brazilian states (São Paulo, Paraná, Minas Gerais, and Federal District) and recently found in Costa Rica. A similar disease is known in the Philippines, but no information exists about its relationship to CoRSV. Coffee ringspot had no economical significance until recently when a large scale infection was reported in Minas Gerais that resulted in yield loss.

Citrus leprosis and its status in Florida and Texas: past and present.

According to published reports from 1906 to 1968, leprosis nearly destroyed the Florida citrus industry prior to 1925. This was supported with photographs showing typical leprosis symptoms on citrus leaves, fruit, and twigs. Support for the past occurrence of citrus leprosis in Florida includes: (1) presence of twig lesions in affected orange blocks in addition to lesions on fruits and leaves and corresponding absence of similar lesions on grapefruit; (2) yield reduction and die-back on infected trees; and (3) spread of the disease between 1906 and 1925. Transmission electron microscopy (TEM) examination of tissue samples from leprosis-like injuries to orange and grapefruit leaves from Florida in 1997, and fruits from grapefruit and sweet orange varieties from Texas in 1999 and 2000 did not contain leprosis-like viral particles or viroplasm inclusions. In contrast, leprosis viroplasm inclusions were readily identified by TEM within green non-senescent tissues surrounding leprosis lesions in two of every three orange leaf samples and half of the fruit samples obtained from Piracicaba, Brazil. Symptoms of leprosis were not seen in any of the 24,555 orange trees examined across Florida during 2001 and 2002. The authors conclude that citrus leprosis no longer exists in Florida nor occurs in Texas citrus based on: (1) lack of leprosis symptoms on leaves, fruit, and twigs of sweet orange citrus varieties surveyed in Florida; (2) failure to find virus particles or viroplasm inclusion bodies in suspect samples from both Florida and Texas examined by TEM; (3) absence of documented reports by others on the presence of characteristic leprosis symptoms in Florida; (4) lack of its documented occurrence in dooryard trees or abandoned or minimal pesticide citrus orchard sites in Florida. In view of the serious threat to citrus in the U.S., every effort must be taken to quarantine the importation of both citrus and woody ornamental plants that serve as hosts for Brevipalpus phoenicis (Geijskes), B. californicus (Banks), and B. obovatus Donnadieu (Acari: Tenuipalpidae) from countries where citrus leprosis occurs.

First report of Raoiella indica Hirst (Acari: Tenuipalpidae) and its damage to coconut palms in Puerto Rico and Culebra Island.

Abstract This is the first report of the occurrence of the red palm mite, Raoiella indica Hirst, and its damage in Puerto Rico and Culebra Island. Raoiella indica is a new invasive pest threatening ornamental palm, coconut, and banana plantations. The mite was observed in eastern Puerto Rico and on Culebra Island infesting and causing damage to coconut, Cocos nucifera L., and Veitchia merrillii L.

An annotated list of ornamentals naturally found infected by Brevipalpus mite-transmitted viruses

The first cases of ornamental plants found infected by Brevipalpus transmitted viruses (BTV) were described in the 1990s from the region of Piracicaba, State of Sao Paulo, Brazil; subsequent cases were from other regions in the country and other American countries. Currently, 37 ornamental plant species (for the sake of simplicity, orchids being considered as a single species), belonging to 18 families of dicotyledons, have been reported hosting BTV. Because of the non systemic type of infection of these viruses, the localized diseases they cause are unimportant usually, but they have the potential to cause economic losses if severe outbreaks of Brevipalpus mite populations occur. Some ornamentals may serve as reservoirs to BTV known to cause serious damage to food crops as Citrus leprosis virus- cytoplasmic type (CiLV-C), passion fruit green spot virus (PFGSV) and Coffee ringspot virus (CoRSV).

A review of the natural enemies of the red palm mite, Raoiella indica (Acari: Tenuipalpidae)

A review of all the available information about the natural enemies reported in association with the red palm mite, Raoiella indica is presented. Twenty-eight species of predatory arthropods, including mites and insects, have been reported in association with R. indica in Asia, Africa and the Neotropics. In addition, pathogenic fungi associated with R. indica in the Caribbean have been reported. The available literature indicates that each site has a different natural enemy complex with only one predator species, Amblyseius largoensis (Acari: Phytoseiidae), present in all the geographical areas. The phytoseiids, Amblyseius caudatus Berlese, Amblyseius channabasavanni Gupta and A. largoensis, were regarded as important natural enemies of R. indica, and their predatory efficiency was studied in some detail. Among the predatory insects the coccinellids Stethorus keralicus Kapur and Telsimia ephippiger Chapin were reported as major predators of R. indica. The known distribution, abundance and relative importance of each species reported in association with R. indica are discussed.

Passion Fruit Green Spot Virus Vectored by Brevipalpus phoenicis (Acari: Tenuipalpidae) on Passion Fruit in Brazil

Passion fruit green spot disease was first identified in 1997 after a severe outbreak at Vera Cruz County, state of São Paulo, Brazil. Mature yellow fruits of Passiflora edulis Simms f. flavicarpa Degener showed characteristic green spots, 2–5 mm in diameter and patches of green tissues were present on senescent leaves. The devastating effect to passion flower is caused by necrotic lesions that encircle the stems and kill the plant. In severe cases, entire orchards of a few hectares in size have been completely destroyed. The disease was always preceded by heavy infestations of Brevipalpus phoenicis (Geijskes) (Acari: Tenuipalpidae). Transmission electron microscopy of affected tissues (fruits, leaves, and stems) consistently revealed the presence of short, bacilliform particles (50–70 nm × 100–120 nm) in the cisternae of the endoplasmic reticulum, as well as the presence of a dense viroplasm in the cytoplasm. This cytopathic effect has been found in several other Brevipalpus-transmitted or associated viruses and is classified as a cytoplasmic type of disease. Experimental reproduction of the leaf and stem symptoms was achieved by transferring B. phoenicis collected from affected field passion flower plants onto healthy plants. The evidence supports a viral etiology for the disease and the agent was named passion fruit green spot virus. Its relationship with other B. phoenicis related viruses continues to be studied. The disease was also found in the Brazilian states of Bahia, Sergipe, Rondonia, Minas Gerais, Rio de Janeiro, and in the Federal District. Use of one or more of the following acaricides (hexythiazox, fenbutatin-oxide, propargite, quinomethionate, or dicofol) has significantly reduced the incidence of the disease.