Phlebotomine sand flies (Diptera: Psychodidae) of the state of Minas Gerais, Brazil

PUBLIC HEALTH

Phlebotomine sand flies (Diptera: Psychodidae) of the state of Minas Gerais, Brazil

Andrey J de Andrade^I; Filipe Dantas-Torres^II

^IDepto de Parasitologia, Univ Federal de Minas Gerais, Av Presidente Antônio Carlos 6627, 31270-901 Belo Horizonte, MG, Brasil; [e-mail]

^IIDipart di Sanità Pubblica e Zootecnia, Univ degli Studi di Bari. Str. Prov. per Casamassima km 3, 70010 Valenzano (Bari), Italia; [e-mail]

ABSTRACT

The phlebotomine sand fly fauna of the State of Minas Gerais is presented based on an extensive review of the literature. The fauna, which is a rich one, is currently known to consist of at least 93 species included in 16 genera. The genus Evandromyia (16 spp.) is the most representative one, followed by Micropygomyia (13 spp.), Psathyromyia (12 spp.), Brumptomyia (11 spp.), Pintomyia (9 spp.), Lutzomyia (7 spp.), Psychodopygus (7 spp.), Martinsmyia (4 spp.), Nyssomyia (3 spp.), Deanemyia (2 spp.), Expapillata (2 spp.), Pressatia (2 spp.), and Sciopemyia (2 spp.). The genera Bichromomyia, Trichopygomyia, and Migonemyia are represented by only one species. An updated list of phlebotomine sand fly species occurring in Minas Gerais is provided. Finally, the importance of the species of public health concern is discussed.

Key words: Phlebotominae, leishmaniase

Phlebotomine sand flies play an important role in the transmission of disease agents to human beings. They are responsible for the transmission of intracellular protozoan parasites of the genus Leishmania Ross, the causative agents of a group of diseases collectively known as the leishmaniases (Desjeux 2004). Nowadays, over 460 phlebotomine sand fly species are known to occur in the Americas (Galati 2003a). In this region, approximately 40 species are involved in the transmission of Leishmania parasites (Killick-Kendrick 1990, Lainson & Shaw 2005).

In Brazil, the leishmaniases have a broad distribution and account for a great burden in terms of morbidity and mortality, which, together with other arthropod-borne diseases like malaria, represent the Achilles' heel of public health in Brazil in terms of disease control. The burden of leishmaniases is greater among the poorest segments of society, particular, but not exclusively, in the Northeastern region (Lainson & Rangel 2005, Dantas-Torres & Brandão-Filho 2006). Nowadays, the leishmaniases are also well established in other regions, including large urban areas of Southeastern Brazil, such as the metropolitan region of Belo Horizonte in the State of Minas Gerais (Passos et al 1993, Oliveira et al 2001).

Historically, the leishmaniases are endemic to Minas Gerais; the first cases were reported in the municipality of Uberaba, during the late in 1910s (Rabello 1917). Currently, the leishmaniases are widely spread in this state. Visceral leishmaniasis is a major public health problem in many municipalities of Minas Gerais (Barata et al 2004, Monteiro et al 2005), including in Belo Horizonte, the state capital (Silva et al 2001).

The first contributions to the study of phlebotomine sand flies of Minas Gerais took place at the beginning of the 1910s (Lutz & Neiva 1912), when Dr. Adolf Lutz and Dr. Arthur Neiva, working in the municipalities of Além Paraíba and Cordisburgo (Maquiné Cave), described two species that today are known as Lutzomyia longipalpis (Lutz & Neiva) and Nyssomyia intermedia (Lutz & Neiva). During the second half of the 20th century, the phlebotomine sand fly fauna of Minas Gerais was extensively studied and a number of species were recorded and/or described (Martins et al 1956, 1982, Falcão 1963, Gomes et al 1978, Mayrink et al 1979, Rangel et al 1985, Falcão et al 1988, Passos et al 1993, Genaro et al 1996, Marcondes 1996, Andrade Filho et al 1997, 1998, 1999a,b, Brazil et al 1997, Gontijo et al 2002).

According to the last lists of phlebotomine sand flies of Brazil, the fauna of Minas Gerais is known to consist of 82 species (CIPA 1999, Aguiar & Medeiros 2003, Galati 2003a). However, these lists are currently out of date. The present article intends to provide an updated list of phlebotomine sand flies occurring in Minas Gerais, based on an extensive review of the literature.

Material and Methods

The present study was based on an extensive search of the literature regarding records of phlebotomine sand flies in the State of Minas Gerais, Southeastern Brazil. All papers found, from 1912 to 2008, were carefully reviewed, including original descriptions of many species. The nomenclature of phlebotomine sand flies follows the classification system recently proposed by Galati (2003b). Abbreviations of genera and subgenera names are according to Marcondes (2007).

Data on phlebotomine sand flies occurring in Minas Gerais were compiled from several papers: Lutz & Neiva (1912), Newstead (1914), França (1920), Larrousse (1920), Lutz (1922), Newstead (1922), Brèthes (1923), Pinto (1926), Dyar (1929), Costa Lima (1932), Root (1934), Costa Lima & Antunes (1936), Mangabeira (1938, 1942), Barretto & Coutinho (1941), Barretto et al (1956), Martins et al (1957, 1961, 1962a, 1962b, 1964, 1970, 1972, 1975, 1982), Martins & Silva (1965, 1968), Ryan et al (1986), Dias et al (1987), Galati et al (1989, 1995), Andrade Filho et al (1999a, 2002).

Information on phlebotomine sand fly species occurring in Minas Gerais available at the CIPA Group website (http://cipa.snv.jussieu.fr), which is an on-line taxonomic catalogue of American sand fly species, was also used.

Additional locality records are according to information provided in the reference book American Sand Flies (Diptera: Psychodidae, Phlebotominae), by Martins et al (1978), and also based on several entomological surveys (Marcondes 1996, Andrade Filho et al 1999c, 2007, Santos de Marco et al 2002, Hamilton et al 2004).

Phlebotomine sand fly species found in Minas Gerais but not included in previous lists (Young & Duncan 1994, CIPA 1999, Aguiar & Medeiros 2003, Galati 2003a) and some synonyms (i.e., species that are included in the present checklist, but have been regarded as synonyms by some authors) are also discussed. Likewise, phlebotomine sand fly species epidemiologically involved in the transmission of Leishmania parasites are highlighted.

Results

The phlebotomine sand fly fauna of Minas Gerais is currently known to consist of 93 species included in four subtribes, 16 genera, 14 subgenera, 13 series and two species groups. Phlebotomine sand fly species described based on material collected from Minas Gerais (a), species endemic to Minas Gerais (i.e., only recorded in this state) (b) and species of public health concern (c) are highlighted.

The list include the species Brumptomyia ortizi Martins, Silva & Falcão (Caparaó), Br. travassosi (Mangabeira) (Viçosa), Evandromyia bacula (Martins, Falcão & Silva) (Sete Lagoas, AJA - personal observation), Ev. cortelezzii (Brèthes) (Intanhomi, Pedra do Indiá, Sacramento, Santa Luzia), Ev. corumbaensis (Galati, Nunes, Oshiro & Rego) (Uberlândia), Micropygomyia breviducta (Barretto) (Além Paraíba), Mi. micropyga (Mangabeira) (Parque Nacional Cavernas do Peruaçu-São João das Missões, Itacarambi and Januária); Mi. trinidadensis (Newstead) (Montes Claros, Porteirinha, Vazelândia), Pressatia choti (Floch & Abonnen) (Timóteo), Psychodopygus carrerai carrerai (Barretto) (Além Paraíba), and Ps. fairchildi (Barretto) (Parque Estadual do Rio Doce-Cava Grande). These species were not included in the previous lists either by omission or because there were recorded in Minas Gerais only recently (see Discussion).

Six phlebotomine sand fly species are endemic from Minas Gerais. For two of them, Deanemyia appendiculata (Martins et al) and De. ramirezi (Martins et al), the only bona fide records of their presence in Minas Gerais are their original descriptions.

Brumptomyiina Artemiev

Brumptomyia França & Parrot (11 spp.): Br. avellari (Costa Lima)ª, Br. brumpti (Larrousse), Br. cardosoi (Barretto & Coutinho), Br. cunhai (Mangabeira), Br. guimaraesi (Coutinho & Barretto), Br. mangabeirai (Barretto & Coutinho), Br. nitzulescui (Costa Lima), Br. ortizi Martins et al, Br. pintoi (Costa Lima)ª, Br. travassosi (Mangabeira), and Br. troglodytes (Lutz).

Lutzomyiina Abonnenc & Leger

Sciopemyia Barretto (2 spp.): Sc. microps (Mangabeira) and Sc. sordellii (Shannon & Del Ponte).

Lutzomyia França (7 spp.): Lu. alencari Martins et alª, Lu. amarali (Barretto & Coutinho), Lu. cavernicola (Costa Lima)ª, Lu. ischnacantha Martins, Souza & Falcão^b, Lu. ischyracantha Martins et al^b, Lu. longipalpis (Lutz & Neiva)^a,c, and Lu. renei (Martins, Falcão & Silva).

Migonemyia Galati (1 sp.): Mg. migonei (França)^c

Pintomyia Costa Lima (9 spp.): Pi. bianchigalatiae (Andrade-Filho et al)ª, Pi. christenseni (Young & Duncan), Pi. damascenoi (Mangabeira), Pi. fischeri (Pinto)^c, Pi. mamedei (Oliveira et al), Pi. misionensis (Castro), Pi. monticola (Costa Lima), Pi. pessoai (Coutinho & Barretto)^c, and Pi. serrana (Damasceno & Arouck).

Expapillata Galati (2 spp.): Ex. cerradincola (Galati et al) and Ex. firmatoi (Barretto et al)ª.

Pressatia Mangabeira (2 spp.): Pr. choti (Floch & Abonnenc) and Pr. equatorialis (Mangabeira).

Trichopygomyia Barretto (1 sp.): Ty. longispina (Mangabeira).

Evandromyia Mangabeira (16 spp.): Ev. bacula (Martins et al), Ev. callipyga (Martins & Silva)^b, Ev. carmelinoi (Ryan et al), Ev. cortelezzii (Brèthes), Ev. corumbaensis (Galati et al), Ev. costalimai (Mangabeira), Ev. edwardsi (Mangabeira), Ev. evandroi (Costa Lima & Antunes), Ev. lenti (Mangabeira)ª, Ev. petropolitana (Martins & Silva), Ev. sallesi (Galvão & Coutinho), Ev. rupicola (Martins et al), Ev. teratodes (Martins et al), Ev. termitophila (Martins et al)ª, Ev. tupynambai (Mangabeira), and Ev. walkeri (Newstead).

Psychodopygina Galati

Psathyromyia Mangabeira (12 spp.): Pa. aragaoi (Costa Lima)ª, Pa. barrettoi barrettoi (Mangabeira), Pa. brasiliensis (Costa Lima), Pa. dendrophyla (Mangabeira), Pa. hermanlenti (Martins et al)ª, Pa. lanei (Barretto & Coutinho), Pa. lutziana (Costa Lima) ^a, Pa. pascalei (Coutinho & Barretto), Pa. pelloni (Sherlock & Alencar), Pa. pestanai (Barretto & Coutinho), Pa. runoides (Fairchild & Hertig), and Pa. shannoni (Dyar).

Martinsmyia Galati (4 spp.): Mt. cipoensis (Martins et al)ª, Mt. gasparviannai (Martins et al), Mt. minasensis (Mangabeira)^b, and Mt. oliveirai (Martins et al)ª.

Psychodopygus Mangabeira (7 spp.): Ps. ayrozai (Barretto & Coutinho)^c, Ps. carrerai carerai (Barretto), Ps. davisi (Root), Ps. fairchildi (Barretto), Ps. hirsutus hirsutus (Mangabeira)^c, Ps. lloydi (Antunes), and Ps. matosi (Barretto & Zago).

Bichromomyia Artemiev (1 sp.): Bi. flaviscutellata (Mangabeira)^c.

Nyssomyia Barretto (3 spp.): Ny. intermedia (Lutz & Neiva)^a,c, Ny. neivai (Pinto)^c, and Ny. whitmani (Antunes & Coutinho)^c.

Sergentomyiina Artemiev

Deanemyia Galati (2 spp.): De. appendiculata (Martins et al)^b and De. ramirezi (Martins et al)^b.

Micropygomyia Barretto (13 spp.): Mi. acanthopharynx (Martins et al), Mi. borgmeieri (Martins et al)ª, Mi. breviducta (Barretto), Mi. capixaba (Dias et al), Mi. ferreirana (Barretto et al)ª, Mi. longipennis (Barretto), Mi. micropyga (Mangabeira), Mi. oswaldoi (Mangabeira), Mi. peresi (Mangabeira), Mi. quinquefer (Dyar), Mi. schreiberi (Martins et al), Mi. trinidadensis (Newstead), and Mi. villelai (Mangabeira).

Discussion

The phlebotomine sand fly fauna of Minas Gerais. In the present article, we comprehensively reviewed the literature on phlebotomine sand fly fauna of Minas Gerais, which is currently known to consist of at least 93 species in 16 genera; most of them belong to the subtribe Lutzomyiina. In comparison with other states within the Southeast region of Brazil (see Aguiar & Medeiros 2003), the phlebotomine sand fly fauna of Minas Gerais is very rich in terms of species diversity.

Checklists of phlebotomine sand fly species occurring in Minas Gerais have appeared in textbooks (Martins et al 1978, Aguiar & Medeiros 2003, Galati 2003a). These lists were not intended to treat exclusively the phlebotomine sand fly fauna of Minas Gerais, but also of other Brazilian states (Aguiar & Medeiros 2003) or other American countries (Martins et al 1978, Galati 2003a). To the best of our knowledge, this is the first comprehensive review exclusively dedicated to the study of the phlebotomine sand fly fauna of Minas Gerais.

Comments on the validity of some phlebotomine sand fly species found in Minas Gerais. Micropygomyia ferreirana (Barretto, Martins & Pellegrino) was described based on a single male specimen collected in the municipality of Jampruca (Barretto et al 1956). In their book, Martins et al (1978) mentioned Mi. ferreirana as occurring in several municipalities of Minas Gerais and stated that its female was already known, although undescribed. Dias et al (1989) described the female and redescribed the male of Mi. ferreirana. Examining the types of both species, Galati et al (2002) noticed some morphometric similarities between Mi. ferreirana and Mi. borgmeieri (Martins, Falcão & Silva) and proposed their synonym. Based on their original descriptions, males of Mi. ferreirana and Mi. borgmeieri present some differences in relation to size of the head, clypeus, antenna, number of setae on the gonocoxite, and size and form of the paramere. Thus, we follow Dias et al (1989) in maintaining both species as valid, until their synonymy status is reassessed.

Martins et al (1978) mentioned the presence of Lu. spinosa (Floch & Abonnenc) in some municipalities of Minas Gerais. According to Young & Duncan (1994), Lu. spinosa is in part a synonym of Lu. damascenoi (Mangabeira) - Pi. damascenoi, according of Galati (2003a) - and in part a synonym of Lu. christenseni Young & Duncan - Pi. christenseni, according to Galati (2003a). Aguiar & Medeiros (2003) have included both of them in their list of phlebotomine sand flies of Minas Gerais, but it is not clear whether both species occur in Minas Gerais (EAB Galati, personal communication). In fact, Young & Duncan (1994) were not clear in relation to the actual distribution of Pi. damascenoi and Pi. christenseni in Brazil. The presence of Pi. christenseni in Minas Gerais has recently been confirmed (Andrade Filho et al 2008), but the presence of Pi. damascenoi is uncertain (JD Andrade Filho, personal communication).

During the 1970s, Forattini (1973) cited Mi. goiana (Martins, Falcão & Silva) as a junior synonym of Mi. trinidadensis and Martins et al (1978) considered Mi. villelai (Mangabeira) as a junior synonym Mi. trinidadensis. Later, Williams (1988) argued that Mi. trinidadensis would be a combination of several sibling species. Curiously, Mi. villelai, Mi. goiana, and Mi. trinidadensis were subsequently been considered as valid species (Williams 1991). Contrary to Forattini (1973) and Martins et al (1978), Galati (2003a) has recently listed Mi. goiana as a junior synonym of Mi. villelai. Although very closely related, Mi. villelai, Mi. goiana, and Mi. trinidadensis appear to be distinct species, which perhaps should be grouped separately. A detailed study of the typespecimens of Mi. trinidadensis, Mi. villelai, and Mi. goiana in comparison with new specimens collected from their type localities would resolve this taxonomic question.

There has been much discussion on the synonymy of Ny. intermedia and Ny. neivai (Pinto). The two species were considered synonymous by Martins et al (1978) and Young & Duncan (1994). However, Marcondes (1996) have removed Ny. neivai from the synonymy with Ny. intermedia, based on examination of the female spermathecae. Moreover, Andrade Filho et al (2003) observed some morphological differences in the tips of genital filaments of the males. Undoubtedly, Ny. intermedia and Ny. neivai are morphologically very close (Andrade Filho et al 2006). On the other hand, the differences noticed by Marcondes (1996) and Andrade Filho et al (2003) are sufficient to separate them as distinct species.

Phlebotomine sand fly species recently recorded in Minas Gerais. The most recent list of phlebotomine sand flies found in Minas Gerais is that of Aguiar & Medeiros (2003). Galati (2003a) also provided data on the known distribution of phlebotomine sand flies occurring in all Brazilian states, which includes Minas Gerais. However, these lists were incomplete and several other species have been recorded since then. The phlebotomine sand flies that are known to occur in Minas Gerais but that were not cited in their lists include: Ev. cortelezzii, Ps. fairchildi, Mi. breviducta, and Ps. carrerai carrerai (Gomes et al 1978, Rangel et al 1985, Alexander et al 2001).

The phlebotomine sand flies that have been recorded in Minas Gerais during the last six years include: Br. ortizi, Br. travassosi, Br. troglodytes, Mi. micropyga, Mi. trinidadensis, Ev. corumbaensis (Santos & Brazil 2003, Barata et al 2004, 2005, 2008, Lemos et al 2004, Monteiro et al 2005, Dias et al 2007, Saraiva et al 2006), Ev. bacula (AJA, personal observation), and Pr. choti (Souza et al 2009).

Lemos et al (2004) recorded a single male of Ev. corumbaensis in Uberlândia. This phlebotomine sand fly species is part of the cortelezzii complex, which includes Ev. cortelezzii, Ev. sallesi, and Ev. edwardsi, all recorded in Minas Gerais. According to Carvalho et al (in press), Ev. corumbaensis occur only in Center-West Region. Therefore the record of this phlebotomine sand fly in Minas Gerais needs to be confirmed (JD Andrade Filho, personal communication).

Aguiar & Medeiros (2003) included Psathyromyia pelloni (Sherlock & Alencar) in their list of phlebotomine sand flies of Minas Gerais, but its presence in this state is doubtful (Martins et al 1978) and needs confirmation. Similarly, the presence of Ny. singularis (Costa Lima) in Minas Gerais, and even its validity, deserves further study.

The local fauna of phlebotomine sand flies can vary widely. As many municipalities of Minas Gerais are completely unexplored, new collections of phlebotomine sand flies would be greatly appreciated. Hopefully, these collections are much likely to reveal the existence of unknown species.

Phlebotomine sand fly species of public health concern found in Minas Gerais. The leishmaniases are endemic in Minas Gerais. The causative agent of visceral leishmaniasis in Minas Gerais is Leishmania (L.) chagasi Cunha & Chagas (Silva et al 2001), which is currently considered to be a junior synonym of L. (L.) infantum Nicolle (Lukeš et al 2007). In this state, cases of cutaneous leishmaniases are caused by L. (Viannia) braziliensis and rarely by L. (L.) amazonensis Lainson & Shaw (Passos et al 1999).

The phlebotomine sand flies responsible for transmission of Leishmania spp. in each disease focus in Minas Gerais are relatively poorly known. In most cases, the incrimination of a given species in the transmission cycle has been based on circumstantial evidence; that is, its presence or absence in a given area, and its relative abundance to that of other species in a given area where cases of leishmaniases have been notified. Obviously, there are other aspects that must be considered when implicating a given phlebotomine sand fly species as a vector of a given Leishmania species (for a review, see Killick-Kendrick 1990).

Similarly to what occurs in most parts of Brazil, Lu. longipalpis plays a major role in the zoonotic transmission cycle of the causative agent of visceral leishmaniasis in Minas Gerais. This phlebotomine sand fly has a broad distribution in this state usually ocuring where cases of visceral leishmaniasis have been reported (Souza et al 2004, França-Silva et al 2005, Michalsky et al 2007). Since Lu. longipalpis was originally described in 1912, it has been recorded in 49 or so municipalities of Minas Gerais (Andrade et al 2007, Carvalho et al 2008, Souza & Borges 2008). An average of 295 cases of visceral leishmaniasis is notified annually in Minas Gerais (Ministério da Saúde 2007a).

In relation to cutaneous leishmaniasis, the scenery is quite different. The role of each species in the transmission of may vary widely from municipality to municipality and even within municipalities. Among the phlebotomine sand flies that may be implicated in the transmission of L. (V.) braziliensis in Minas Gerais, Ny. intermedia and Ny. whitmani (Antunes & Coutinho) are those that deserves the most attention. Both species are suspected to be involved in the transmission of L. (V.) braziliensis in other parts of Brazil (Marcondes et al 1998, Andrade Filho et al 2007, Costa et al 2007), are highly anthropophilic, and are often present in foci of cutaneous leishmaniasis in Minas Gerais (Casanova 2001, Lemos & Lima 2005).

Both Ny. intermedia and Ny. whitmani have a broad distribution in Minas Gerais, frequently occurring in sympatry (Brazil et al 2006, Resende et al 2006, Loiola et al 2007). Since its original description in 1912, Ny. intermedia has been recorded in 74 or so municipalities of Minas Gerais. Nyssomyia whitmani is also widely spread in this state; it is known to occur in at least 76 municipalities (Hermeto et al 1998, Santos et al 2003, Margonari et al 2006). An average of 1,793 cases of cutaneous leishmaniasis has been notified in Minas Gerais (Ministério da Saúde 2007b).

It is important to highlight that Ny. intermedia occurs in sympatry with Ny. neivai in Minas Gerais (Andrade Filho et al 2007). Nyssomyia neivai has been recorded in 21 municipalities (Marcondes et al 1998, Andrade Filho et al 2007). It is a suspected vector in the municipality of Corinto, where it represents 90% of the phlebotomine sand flies caught (Andrade Filho et al 2007, Saraiva et al 2008). However, its role as a vector needs confirmation.

Other species that may eventually play a local role in the transmission of L. (V.) braziliensis Minas Gerais are: Bichromomyia flaviscutellata (Mangabeira), Migonemyia. migonei (França), Pi. pessoai (Coutinho & Barretto), Ps. ayrozai (Barretto & Coutinho), Pi. fischeri (Pinto), and Ps. hirsutus hirsutus (Mangabeira) (Rangel et al 1985, Andrade Filho et al 1997, Santos de Marco et al 2002, Souza et al 2004, Saraiva et al 2006). These species are suspected to be involved in the transmission cycle of L. (V.) braziliensis elsewhere in Brazil (Lainson 1983, Killick-Kendrick 1990, Peterson & Shaw 2003, Rangel & Lainson 2003), with the exception of Bi. flaviscutellata which is the primary vector of L. (L.) amazonensis (Lainson & Shaw 2005).

In the present article, we comprehensively reviewed the available literature about phlebotomine sand flies of Minas Gerais. Nearly a century after the first contributions to the knowledge of the phlebotomine sand fly fauna of this state, much work has been done, but it is evident that there is a lot of work to do.

From an entomological point of view, there appears to be many phlebotomine sand fly species to be known, because the fauna of many municipalities of Minas Gerais is completely unknown. From the epidemiological standpoint, it seems evident that much study is required to improve our current understanding on the distribution and behavior of phlebotomine sand flies of public health significance. A better understanding of these aspects is seminal for any vector control program to achieve success.

We hope that the present work will somehow contribute to the knowledge of distribution of phlebotomine sand flies of Minas Gerais. Finally, but not less important, we believe that our review will encourage other similar studies about the phlebotomine sand fly fauna of other Brazilian states.

Acknowledgments

This work is dedicated to the memory of Professor Paul Williams, who made outstanding contributions to the study of taxonomy and biology of phlebotomine sand flies. The authors are indebted to Dr José Dilermando Andrade Filho for his useful comments on phlebotomine sand flies of Minas Gerais and Dr Eunice A B Galati and Dr Reginaldo P Brazil for their critical review of the manuscript. A J Andrade is supported by a PhD scholarship from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq).

Received 22/II/08

Accepted 26/VIII/09

Edited by Eunice Galati - FSP/USP

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Publication Dates

• Publication in this collection
  12 Mar 2010
• Date of issue
  Feb 2010

History

• Received
  22 Feb 2008
• Accepted
  26 Aug 2009