South American Spider Mites: New Hosts and Localities

In order to contribute to taxonomic information on Tetranychid mites (Acari: Tetranychidae) in South America, surveys were conducted in Brazil (15 States and the Federal District) and Uruguay (one Department); 550 samples of 120 plant species were collected. Tetranychid mite infestations were confirmed in 204 samples, and 22 species belonging to seven genera of the Bryobiinae and Tetranychinae subfamilies were identified on 58 different host plants. Thirty-six new plant hosts were found in Brazil, South America, and worldwide for the following species: Eutetranychus banksi (McGregor); Mononychellus tanajoa (Bondar); Oligonychus anonae Paschoal; O. mangiferus (Rahman and Sapra); Tetranychus bastosi Tuttle, Baker and Sales; T. desertorum Banks, 1900, T. evansi Baker and Pritchard; T. ludeni Zacher; T. mexicanus (McGregor); T. neocaledonicus André; and T. urticae Koch. Four new localities in Brazil were reported for Eotetranychus tremae De Leon; O. anonae; Panonychus ulmi (Koch); and T. gloveri Baker and Pritchard.


Introduction
The Tetranychidae Donnadieu family includes a large number of strictly phytophagous mites; a few species are significant agricultural crop pests worldwide (Jeppson et al. 1975;Helle and Sabelis 1985).
The number of described tetranychids remained stable for 75 years and then increased when their economic incidence for agriculture became more significant (Bolland et al. 1998). In 1950, McGregor initially listed 102 species in 15 genera. Five years later, this number increased to 204 species in 18 genera (Pritchard and Baker 1955). In 1998 there were 1189 species in 71 genera registered in the World Catalogue of the Spider Mite Family (Acari: Tetranychidae) (Bolland et al. 1998), and currently there are approximately 1257 species in 76 genera listed on the Spider Mite Web, which is a comprehensive database on the Tetranychidae (Migeon and Dorkeld 2006).
In Brazil, taxonomic studies on the Tetranychidae were initially conducted between 1920 and 1930. Among the early findings was the report of Tetranychus gloveri Banks 1900 and the description of Mononychellus tanajoa (Bondar 1938) based on specimens collected in Brazil by G Bondar, a Russian researcher living in the state of Bahia (Bondar 1930(Bondar , 1938. In the 1960s numerous contributions were made by pioneers in agricultural mite studies in Brazil, such as CHW Flechtmann (Flechtmann 1967(Flechtmann , 1967a(Flechtmann , 1967b(Flechtmann , 1972(Flechtmann , 1975(Flechtmann , 1976Baker 1970, 1975) and AD Paschoal (Paschoal 1970(Paschoal , 1970a, followed by RJF Feres in subsequent decades (Feres 1986(Feres , 1992Feres and Flechtmann 1986, 1986a, 1988, 1995a, 1995b. From that time forward these and other researchers have conducted major studies on tetranychids in several fields such as taxonomy, ecology, biology, control, and resistance, bringing important advances to knowledge of Brazilian tetranychids. Currently 185 of the 1257 described tetranychid species are reported in South America, of which 104 are found in Brazil (Bolland et al. 1998;Migeon and Dorkeld 2006).
The most intensive search for Tetranychidae mites in Brazil, the largest country in South America, has been centered in the southeast region and is usually associated with economically important crops. Agriculture continues to expand, approaching biological reserves and other regions in the country. It is important to continue tetranychid research in previously unexplored areas in hopes of increasing our understanding of this family. This paper presents new information about the occurrence and distribution of Tetranychidae in five Brazilian regions and in Uruguay, on plants ranging from ornamental, fruit-bearing, vegetable, grass, oilseed, to large crops species. New hosts and localities for tetranychid mites in Brazil, South America, and the world are reported.

Materials and Methods
A legal permit to collect zoological material for scientific purposes is required by the Brazilian Institute for the Environment and Renewable Natural Resources (IBAMA). A permit was requested and permit number 231/2006 CGFAU/IBAMA was issued.  Figure 1; Appendix 1). One additional collection was conducted in Uruguay. Plants were collected when symptoms of tetranychid attacks were observed, according to Moraes and Flechtmann (2008). The geographical points of collection were logged into a Garmin 12 GPS using WGS84 (World Geodetic System 84) datum; maps were created using ARCGIS 9.0 (ESRI, www.esri.com).

Inspecting and processing plant samples
Mites were collected from the plants using a washing/sieving extraction method (Miranda et al. 2007). The resulting 70% ethanol solution was inspected under Stemi SV6 (www.zeiss.com) and Olympus SZX 122 (www.olympus.com) stereomicroscopes connected to a Highlight 3000-1 transilluminator at 50 magnification. Finally, Tetranychidae mites were directly mounted on microscopic slide preparations in Hoyer's medium. From each population, 25 females were mounted in dorso-ventral position and ten isolated males were mounted in lateral position to identify the species by morphological analysis. When only a few males were present in a sample, all of them were slide-mounted.

Identifying mite species
Microscopic slide preparations of specimens were examined under phase contrast (Leitz Wetzlar, www.leica-microsystems.com) and interference microscopes (Nikon Eclipse 80i, www.nikon.com) in 40 and 100 objectives and. Morphological identification was conducted by examining relevant taxonomic characteristics of male and female Tetranychidae systematics. The shape of the aedeagus, or male genitalia, was used to classify the species (Pritchard and Baker 1955, Meyer 1974, Baker and Tuttle 1994, Ehara, 1999, which were then compared to specimens that had been deposited in the Reference Mite Collection of the Laboratory of Plant Quarantine, Embrapa Genetic Resources and Biotechnology, Brasília, Brazil. Materials from this study were deposited in this collection as voucher specimens.

Results and Discussion
Tetranychid mites were found on 37.1% (n = 204) of the 550 plant material samples that were collected (Appendix 1). Twenty-one species of Tetranychinae and one of Bryobiinae were identified on the 204 samples from different localities and inspected hosts (Table 1).
Several samples contained more than one species of tetranychid on the same leaf. This occurred on bean (T.  Foott (1962Foott ( , 1963 reported phytophagous mite species coexisting on a single host. The fact that different species coexist on a single host requires careful sampling and collecting larger numbers of specimens in microscopic preparations. This allows for greater representation of mite fauna and increases the probability of identifying all Tetranychidae species present on a specific host and/or at a specific locality.

New hosts for Tetranychidae mites in
Brazil, South America, and the world New hosts were found for 11 tetranychid mite species: E. banksi, M. tanajoa, O. anonae, O.  The presence of E. banksi on Acacia sp. was previously reported in Colombia (Urueta 1975). The occurrence on this host from Pernambuco is the first in Brazil.
Tetranychus evansi was originally described using samples collected from tomato plants (Baker and Pritchard 1960). However, this mite had already been discovered in Brazil by Silva (1954)

who described it as T. marianae
McGregor. Since then it has been reported in the United States (Bolland et al. 1998) and more recently in Africa, Argentina, China, Spain, France, Greece, Israel, Italy, Puerto Rico, Portugal, and Taiwan (Migeon and Dorkeld 2006). Tetranychus evansi is currently reported on 93 host species (Migeon and Dorkeld 2006), most of which belong to the Solanaceae family (Bolland et al. 1998;Moraes et al. 1987). Infestations were reported in Europe of this mite across several plant families (Ferragut and Escudero 1999). However, the only reports on the biology of T. evansi are for those found on tomato plants. Carica papaya is a new host for the T. evansi. Zacher 1913 Tetranychus ludeni Zacher, 1913(Zacher 1913apud Pritchard and Baker 1955 Tetranychus ludeni was previously found on C. lanatus, P. hortorum and P. domestica in Africa, Central America, (El Salvador), Australia, and India (Meyer and Ryke 1959;Davis 1968;Andrews and Poe 1980;Gutierrez and Schicha 1983;Gupta and Gupta 1994). This is the first report of these hosts in South America.  (Flechtmann 1981). Other plant species belonging to the Ocimum and Morus genera (O. sanctum L., M. alba L., M. australis Poir, M. indica L., M. nigra L., M. sp.) were registered as hosts for T. neocaledonicus in Cuba, India, Japan, and Thailand (Livshits and Salinas-Croche 1968;Baker 1975;Gupta 1992;Gupta and Gupta 1994;Ehara and Yamaguchi 2001).

Tetranychus urticae Koch 1836
Tetranychus urticae Koch, 1836. (Koch 1836apud Pritchard and Baker 1955 Paschoal (1970b) first confirmed the presence of T. urticae on A. trifida L. in the state of São Paulo. Ambrosia polystachya is a new host for T. urticae in Brazil and in the world.
Reports of Tetranychus urticae on Parthenium sp. were made in India (Gupta and Gupta, 1994). In the Americas, the first report of its presence on Parthenium sp. occurred in Rio Grande do Sul.

Bignoniaceae
Pyrostegia venusta Miers (flame vine), Universidade Federal de Lavras, Lavras, Minas Gerais, 09.V.2006. Vargas et al. (1996) described T. urticae infestations on P. ignea C. Presl. in Costa Rica. This was the first report of T. urticae infesting P. venusta in the world. Infestations of T. urticae on P. lunatus were reported in the United States, India, and Thailand (Reeves 1963;Gupta and Gupta 1994;Ho et al. 1997) and on M. atropurpureum in Greece (Hatzinikolis 1969). Phaseolus lunatus and M. atropurpureum are new hosts for T. urticae in South America.
Tetranychus urticae on passion fruit (P. caerulea L. and P. edulis ) was reported in Venezuela (Doreste 1968) and Australia (Passiflora sp.) (Gutierrez and Schicha 1983). This is the first report of T. urticae on passion fruit (P. edulis) in Brazil.
The first T. urticae infestation of wheat crops (Triticum sp.) was registered in Greece (Hatzinikolis 1969). This wheat infestation in Brazil is the first occurrence in the Americas.

Cases
of Tetranychus urticae on Zingiberaceae (Curcuma longa L.) were reported in India (Gupta and Gupta 1994). This is the first report of T. urticae on alpinia (A. purpurata) in the world.
The single sample collected in Uruguay revealed a new host for the two-spotted spider mite (T. urticae).
Tetranychus urticae infesting L. camara was first observed in the United States and in India (Thewke and Enns 1969;Gupta and Gupta 1994); this is the first record in South America.
Reports of the European red mite (P. ulmi) on grape vines (Vitis vinifera L.) were registered for the first time in Brazil in 2008 when damages caused by these mites were first noticed on crops in Rio Grande do Sul (Ferla and Botton 2008). In this work, a second occurrence of P. ulmi infesting grape vines was reported in Pirapora, state of Minas Gerais.
Panonychus ulmi was first reported in Brazil by Flechtmann (1967a), who discovered them on apples imported from Argentina. Soon after, Bleicher (1974) reported their presence on apple trees (Malus domestica L.) growing in orchards in the south of the country. However, European countries such as France, Austria, Portugal, Italy, and Greece consider the P. ulmi a grape vine pest (Rambier 1958;Artofer 1976;Carmona and Dias 1980;Girolami and Mozzi 1983;Papaioannou-Souliotis et al. 1994). Similar reports were also registered in Morocco and the United States (Reeves 1963;Tixier et al. 2003).
Despite reports of P. ulmi on apple trees as early as 1967 (Flechtmann, 1976a), grape vines in Brazil were not affected, in contrast to the damages the P. ulmi caused to vineyards in Europe. Navia et al. (1998) hypothesized that the mites that attacked grape vines in Europe had different biotypes than those that infested apple trees. Moraes and Flechtmann (2008) emphasized the importance of avoiding introduction of European mites into Brazil, stating the reason this species was not present on grape vines in Brazil was unknown. They suggested the possibility of biological differences between Brazilian and European mites.
The spread of P. ulmi infestation on grape vines from Rio Grande do Sul (2005Sul ( /2006 to Minas Gerais (2006) state could be the result of the two states trading plant propagation material. The hypothesis that European mites were introduced into Brazil cannot be discarded considering that P. ulmi were discovered approximately 40 years ago and have only recently been causing damage to grape vines. Ferla and Botton (2008) state that the probable causes for P. ulmi dispersion in Brazil are: the proximity of apple tree plantations to vineyards in the south of Brazil; the trading of infested plant material within Brazil and abroad; and unbalanced apple orchards due to excessive application of chemical products, especially non-selective fungicides.
New occurrences of phytophagous mites in South America were recently reported in the literature. For instance, the citrus Hindu mite, Schizotetranychus hindustanicus (Hirst) was collected from citrus in the municipality of Boa Vista, State of Roraima (Navia and Marsaro Jr. 2010), and Eotetranychus smithi Pritchard & Baker 1955 from roses in Rio Branco, Acre (Mendonça et al. 2010); both states from the Northern region of the country. Tetranychus roseus Gutierrez 1969 was first observed in São Paulo in 2007 (Matioli et al. 2008) and its dispersal in the state was reported by Matioli et al (2010).
These new occurrences for mites in Brazil together with the results obtained in this study indicate the importance of intensified surveys on different host plants in the country. Doing this will broaden the understanding of the Tetranychidae family and give light to management and control of these mite species in agricultural systems.

Conclusions
Thirty-five new hosts were registered for 11 tetranychid species already known in Brazil: E. banksi (6 hosts