LAURA JANE GISLOTI
O gênero Neosilba McAlpine (Tephritoidea: Lonchaeidae): revisão,
ocorrência e diversidade
Campinas
2014
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RESUMO
As moscas da família Lonchaeidae (Diptera), juntamente com as da família
Tephritidae formam um grupo conhecido como moscas-das-frutas, já que suas larvas utilizam como recurso alimentar frutos de interesse comercial. Espécies de Tephritidae são comumente relatadas como pragas importantes para a fruticultura e têm sua biologia e ecologia profundamente estudadas. Já as espécies de Lonchaeidae, mesmo estando associadas a diversos frutos de interesse agrícola, são pouco estudadas. Isto pode estar relacionado ao fato de que não há estudos distinguindo taxonomicamente esse grupo no nível de gênero e espécies. Paralelamente, esta inconsistência taxonômica dificulta e desencoraja estudos com o grupo, fazendo com que a história natural dessas moscas permaneça desconhecida. Em Neosilba, o gênero de Lonchaeidae alvo deste estudo, a taxonomia tradicional é baseada unicamente na análise da morfologia genital masculina e assim todos os trabalhos a respeito deste grupo excluem grande parte do material obtido: as fêmeas. Alguns estudos já foram desenvolvidos no intuito de facilitar a identificação destas moscas, envolvendo técnicas eletroforéticas de sistemas enzimáticos e análises empregando métodos morfométricos. Porém nenhuma técnica se mostrou específica o bastante para distinguir as espécies neste gênero.
Dentro desse contexto, a tese foi estruturada em três capítulos. No capítulo 1 fazemos uma revisão da biologia, ecologia e taxonomia da família Lonchaeidae. Dentro desse capítulo disponibilizamos um mapa da distribuição mundial dos oito gêneros, além de uma chave taxonômica dicotômica ilustrada, adaptada de McAlpine, para todos os gêneros de Lonchaeidae. No capítulo 2 realizamos uma revisão centrada em diversidade, distribuição e plantas hospedeiras de moscas do gênero Neotropical Neosilba. Listamos a
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distribuição e as plantas hospedeiras das 40 espécies de Neosilba. Por fim, no capítulo 3 estudamos a diversidade de espécies frugívoras do gênero Neosilba em 35 espécies de frutíferas cultivadas, de duas localidades do estado de São Paulo e concluímos que Neosilba zadolicha é a espécie que apresenta o maior número de plantas hospedeiras e é a mais abundante entre as sete espécies de Neosilba encontradas. Este trabalho acrescenta informações inéditas ao gênero Neosilba e a compilação de dados a respeito deste grupo pretende estimular o ingresso de novos pesquisadores a fim de preencher as lacunas existentes no âmbito da taxonomia, sistemática, biologia e ecologia desse táxon.
PALAVRAS-CHAVE: Acalyptratae, Diptera, moscas frugívoras, lonchaeids, taxonomia alfa.
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ABSTRACT
The flies of the Lonchaeidae family, alongside with the Tephritidae, form a group known as fruit-flies, since their larvae feed on fruits of huge economical interest in their diet. Species of Tephritidae are generally reported as important pests to the horticulture, with their biology and ecology deeply studied in detail.
However the Lonchaeidae species, despite being associated to many fruits of agricultural interest, are not as studied. This can be related to the fact that there aren’t many studies distinguishing taxonomically this group at the genus or species level. In parallel, this taxonomic inconsistency complicates and discourages studies of the group, keeping the natural history of those flies unknown. In Neosilba, the traditional taxonomy is based only on the analysis of the male genitalia morphology, and so, this method excludes great part of the material collected: the females. Some studies were done on the purpose of facilitating the identification of these flies, involving electrophoresis techniques of enzymatic systems and analysis employing morph metric methods. However, no technique demonstrated to be specific enough to distinguish the species of this genus.
Inside this context, this work was structured in four chapters. In chapter 1 a revision of the biology, ecology, and taxonomy of the Lonchaeidae family is made. Inside this chapter, we make available a map containing the global distribution of all genus, in addition to an illustrated dichotomic taxonomic key, adapted from McAlpine, to all Lonchaeidae genus. In chapter 2 we realize a revision centered in the diversity, distribution and hosting plants of the Neotropical genus Neosilba. We listed a distribution of hosting plants of 40
Neosilba species. In chapter 3, we study the diversity of frugivorous species in the Neosilba genus in 35 fruit species cultivated from 2 locations in the state of São Paulo and we ix
conclude that Neosilba zadolicha is the species that contains the biggest number of hosting plants, and is also the most abundant compared to the 7 species of Neosilba that were found.
This work adds unpublished information of the Neosilba genus and the compilation of data concerning this group aims to stimulate the entrance of new researchers for the purpose of fill the existing lacunae in the scope of taxonomy, systematics, biology, and ecology of this taxon.
KEYWORDS: Acalyptratae, alfa taxonomy, Diptera, frugivorous flies, lonchaeids.
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SUMÁRIO
RESUMO...... vii
ABSTRACT...... ix
LISTA DE ILUSTRAÇÕES...... xv
LISTA DE TABELAS...... xvii
INTRODUÇÃO GERAL...... 01
REFERÊNCIAS...... 04
CAPÍTULO 1. REVIEW OF THE TAXONOMY AND BIOLOGY OF LONCHAEIDAE (DIPTERA), FRUIT FLIES OF ECONOMIC IMPORTANCE IN THE NEOTROPICAL REGION
Abstract...... 9 Introduction...... 10 Methodology………..……………………………...... 12 Results……………………………...... 13 Discussion...... 27 References...... 44
CAPÍTULO 2 ESPÉCIES DE Neosilba (DIPTERA, LONCHAEIDAE, LONCHAEINAE): DISTRIBUIÇÃO GEOGRÁFICA E PLANTAS HOSPEDEIRAS
Resumo...... 67 Introdução...... 68 Material e Métodos ………..….....…………...... ……...... 70 Resultados e Discussão …………...... 71 Referências...... 95
CAPÍTULO 3. Neosilba McAlpine (Diptera, Lonchaeidae) Diversity on Cultivated Host Plants from Brazilian Cerrado and Atlantic Tropical Forest Biomes
Abstract...... 107 Introduction...... 108 Material and Methods………..…………….…………...... 109 Results and Discussion………………...... 111 References...... 118 xi
CONCLUSÃO………………………...... …………………………….....……………130
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Agradecimentos
"Aqueles que passam por nós, não vão sós, não nos deixam sós. Deixam um pouco de si, levam um pouco de nós (Antoine de Saint-Exupéry).
Estes 4 anos de pesquisa foram uma árdua jornada de desafio, construção e amadurecimento. De fato, nenhum empreendimento é realizado de forma fácil e sem esforço. José Saramago discorreu sobre a importância da jornada rumo à vitória: "Nem as derrotas nem as vitórias são definitivas. Isso dá uma esperança aos derrotados, e deveria dar uma lição de humildade aos vitoriosos.” Assim, neste período aprendi que uma tese ou qualquer outro trabalho é uma extensão da vida do autor. Então, para que algo de valor seja produzido, o indivíduo deve primeiramente criar algo de valor em si. Indivíduo e obra são consistentes com o resultado. Por estes motivos, agradeço sincera e profundamente todas as pessoas que me ajudaram e me incentivaram a produzir algo de valor em minha vida. Primeiramente agradeço ao meu grande mestre, Prof. Dr. Angelo Pires do Prado. É uma grande honra tê-lo tido como orientador por 6 anos (desde o mestrado). Jamais me esquecerei de seus ensinamentos, de seus preciosos conselhos e de sua inestimável confiança. Muito Obrigado, Angelo! Agradeço também a minha orientadora adotiva, Profa. Dra. Silmara Marques Allegretti. O seu apoio e a sua confiança no momento mais difícil desta jornada foram essenciais para que fosse possível finalizar essa etapa. Manifesto aqui a minha gratidão a todos os professores, funcionários e amigos do Departamento de Biologia Animal do Instituto de Biologia da Unicamp, em particular à Profa. Dra. Marlene Tiduko Ueta, à Carina Mara de Souza, à Rodrigo Labello Barbosa e à David Villas Boas Filho. Foram momentos bastante agradáveis e produtivos ao lado de vocês. Meus respeitosos agradecimentos pela contribuição da banca do Exame de Qualificação e pela participação dos membros da Banca Examinadora da Defesa.
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Meu muito obrigado à meus queridos companheiros que me encorajam em todos os aspectos da minha vida: minhas amigas Patrícia Silvana Barros, Laila Silva Leonel De Ávila Stoppa, Manuela Scarpa Silverio Pinto e meu companheiro Breno Gomes Machado da Silva. Por fim, agradeço em especial àqueles que sempre me apoiaram incondicionalmente, que apostaram em mim mais do que ninguém e que seguramente são os que mais compartilham da minha alegria: minha mãe Lucia Helena Gisloti e meu filho Lucas Gisloti Martins. Mais uma etapa vencida! Sei que ainda é só o começo.
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LISTA DE ILUSTRAÇÕES
Capítulo 1
Figure 1. Geographical distribution of the Lonchaeidae (Diptera: Tephritoidea) species around the world...... 29
Figure 2. Distribution of publications about the Lonchaeidae genera (Diptera:
Tephritoidea), since 1878 until December 2013 (n = 201) [*other = Papers about
Lonchaeidae without specified genus]……………………..……...... ………...………..…31
Figure 3. Stigmatal bristles (in red) present on the sclerite above the anterior thoracic spiracles…………………..…………..…………………………………………………….36
Figure 4. Lunule bare (see arrow in red)………………………………..………….………36
Figure 5. Lunule setulose (in red)……………………………...…………………………..37
Figure 6. Scutellum with four bristles on the edge (in red)…………………..……..……..37
Figure 7. Scutellum with more them four bristles on the edge (in red)….....…...... …...….38
Figure 8. Setae of the frons spaced and short (see arrow in red)………………...…….…..38
Figure 9. Setae of the frons long and dense (see arrow in red)……………..……………...39
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Figure 10. Scutellum with setae anterior to the marginal bristles (in red)…………………39
Figure 11. Cheek and parafacial wide (in red)………………………. ………..…………..40
Figure 12. Scutellum bare in relation to the marginal bristles (see arrow in red)……..…...40
Figure 13. Cheek and parafacial narrowed (in red)…………………..………. …………...41
Figure 14. Arista plumose (in red)……………………………………...…………...... 41
Figure 15. Surstylus with prensisetae (in red)……………………... ………………...... 42
Figure 16. Arista bare (in red)…………………………. ……………………………...... 42
Figure 17. Surstylus without prensisetae (see arrow in red)…………………...…………..43
Figure 18. Fringes of calypteres with a cluster of long blackish setae at fold (in red)…….43
Figure 19. Fringes of calypteres without cluster of long blackish setae at fold (see arrow in red)………………………………………………………...…………………………….…44
Capítulo 2
Figure 1. Distribuição geográfica das espécies de Neosilba McAlpine na Região
Neotropical...... 74 xvi
LISTA DE TABELAS
Capítulo 2
Tabela 1. Táxons de plantas hospedeiras de espécies de Neosilba (Diptera: Lonchaeidae) no mundo...... 75
Capítulo 3
Table 1. Indices of infestation and viability of pupae of Neosilba species in 35 cultivated hosts from Cerrado and Tropical Forest of Brazil (January/2010 to
March/2012)……………………………………………………………..………..………124
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INTRODUÇÃO GERAL
A família Lonchaeidae está classificada dentro da superfamília Tephritoidea, a qual apresenta características notórias como a exibição de padrão de corte em vôo, presença de asas geralmente sem pigmentação ou pigmentação difusa, além de halteres e corpo de coloração preta, sendo comum reflexos metálicos azuis, bronze ou verde no corpo. É composta pelas subfamílias Lonchaeinae e Dasiopinae, as quais apresentam ampla distribuição geográfica, sendo que na região neotropical são conhecidos representantes de ambas as subfamílias (McAlpine 1987).
No Brasil, os lonqueídeos estão representados principalmente pelo gênero Neosilba
McAlpine 1962, mas, são encontradas espécies do gênero Lonchaea Fallén 1820 e Dasiops
Rondani 1856 (McAlpine & Steyskal 1982, Strikis & Prado 2005).
As larvas de Neosilba são saprófagas, fitófagas ou predadoras e assim estão associadas a frutos, sementes, brotos e flores. Tais órgãos ou tecidos são danificados pela nutrição das larvas nesses substratos (McAlpine 1961, Fehn 1981).
Apesar de a literatura brasileira ter relatado a presença de Neosilba infestando frutos de importância econômica desde a década de 30, por um longo período, os lonqueídeos foram negligenciados e muitas vezes descartados nos levantamentos de moscas frugívoras, pela falta de conhecimento taxonômico e, principalmente, por não serem considerados de notável importância econômica. A partir da década de 70, foi crescente o interesse em estudar a família Lonchaeidae, principalmente espécies do gênero Neosilba, porque passou a ser comum o aparecimento de espécimes em frutos de importância comercial (Gatelli et al. 2008).
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Já na década posterior os estudos passaram a incluir as espécies desta família e devido a isto foram alistadas dezenas de espécies de frutos hospedeiros, em diversos estados brasileiros (Malavasi et al. 1980, Malavasi & Morgante 1980).
Posteriormente, na década de 90, o interesse em estudar Neosilba foi favorecido em razão da quantidade de pupários de lonqueídeos obtidos em inventários entomofaunísticos e em outros estudos. Assim, Silva (1993), constatou a presença de espécies de Neosilba associadas a 19 espécies de frutos, em quatro locais do Estado do Amazonas.
Nos anos seguintes, Raga et al. (1996, 1997), em um estudo de observação da incidência de moscas das frutas (Tephritidae) em frutos de café Coffea spp. e laranja Citrus sinensis, no estado de São Paulo, relataram que 17 e 4,9% dos exemplares coletados, respectivamente, pertenciam à família Lonchaeidae. Também no Estado de São Paulo,
Souza Filho (1999) constatou Neosilba associada a 40 espécies de frutos. No estado de
Mato Grosso do Sul, estas moscas foram associadas a 22 espécies de frutos (Uchôa et al.
2002). De fato, este gênero havia despertado interesse, principalmente nos pesquisadores de mosca das frutas, e isto possibilitou um aumento considerável de publicações que envolviam Lonchaeidae.
Assim as larvas de Neosilba foram registradas como tendo hospedeiros frutos de acerola, goiaba, tangerina (Araújo & Zucchi 2002), manga, mamão, sete-copas, abacate, carambola, laranja, maracujá (Uchôa et al. 2002); café ( Souza et al. 2005, Aguiar-Menezes et al. 2007); nêspera (Strikis & Prado 2009); pêssego (Striks & Prado 2005), entre diversos outros frutos, confirmando assim a importância desta família para a fruticultura tropical.
Além de frutos, imaturos destes insetos já foram notificados como tendo hospedeiros botões florais de Passiflora spp. (Norrbom & McAlpine 1997, Uchôa &
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Zucchi 1999, Causton & Peña Rangel 2002, Uchôa et al. 2002), Ipomoea spp. (Santos et al. 1992) e Manihot esculenta Crantz (Del Vecchio 1991).
Por outro lado, diversos estudos ilustram a dificuldade em distinguir as espécies das moscas deste gênero, de forma que um número elevado de trabalhos apresentam a ocorrência de espécies não identificadas de Neosilba (Malavasi & Morgante 1980, Raga et al. 2002, Uchôa et al. 2002, Souza et al. 2005, Bittencourt et al. 2006, Minzão & Uchôa
2008, Caires et al. 2009).
De fato, as espécies de Neosilba são estreitamente relacionadas e as divergências morfológicas entre os indivíduos não são detectáveis em uma análise de simples observação. Assim, a taxonomia se mostra bastante complicada para este grupo, o que dificulta os trabalhos com estes Diptera. Adicionalmente, já foi constatado um grande complexo de espécies crípticas neste gênero, o que dificulta ainda mais a identificação de suas espécies, que é baseada integralmente na análise das estruturas da genitália masculina dessas moscas (McAlpine & Steyskal 1982, De Conti 1984). Outrora, este método de identificação requeria bastante treinamento, o que muitas vezes acabava por desestimular os trabalhos com este grupo, prejudicando ainda mais a taxonomia.
As fêmeas de Neosilba são excluídas nos trabalhos, devido à ausência de ferramentas que possibilitem a sua identificação, que ainda é dificultada pelo fato de que inúmeras espécies não possuem hospedeiros específicos, e um único fruto pode apresentar diferentes espécies e até distintos gêneros de Lonchaeidae. Assim, todos os trabalhos a respeito deste gênero excluem grande parte do material obtido, e isto que pode gerar inconsistência nos dados, principalmente se o trabalho for ao âmbito de biodiversidade
Neotropical ou inventários de hospedeiros.
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Alguns estudos já foram desenvolvidos no intuito de facilitar a identificação destas moscas, alguns dos quais envolveram técnicas eletroforéticas de sistemas enzimáticos (De
Conti et al. 1984) e análises empregando métodos morfométricos (Oliveira 1992, Freitas
2008), porém nenhuma técnica se mostrou específica o bastante para distinguir espécies de
Neosilba.
Assim, no capítulo 1 revisamos aspectos biológicos, ecológicos e taxonômicos da família Lonchaeidae. No capítulo 2 demos ênfase através de uma revisão centrada em diversidade, distribuição e hospedeiros de moscas do gênero Neotropical Neosilba. E por fim, no capítulo 3 estudamos a diversidade de espécies frugívoras do gênero Neosilba em
35 espécies de frutíferas cultivadas, de duas localidades do estado de São Paulo.
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UCHÔA-FERNANDES, M. A.; OLIVEIRA, I. de, MOLINA, R. M. S.; ZUCCHI, R. A.
Species Diversity of Frugivorous Flies (Diptera: Tephritoidea) from Hosts in the
Cerrado of the State of Mato Grosso do Sul, Brazil. Neotropical Entomology. v. 31,
p. 515-524, 2002.
8
CAPÍTULO 1
REVIEW OF THE TAXONOMY AND BIOLOGY OF LONCHAEIDAE
(DIPTERA), FRUIT FLIES OF ECONOMIC IMPORTANCE IN THE
NEOTROPICAL REGION
GISLOTI, L. J.¹; PRADO, A. P.¹ ǂ & UCHÔA, M. A.² ¹Departamento de Biologia Animal, UNICAMP, Campinas-SP, Brazil. [email protected] 2Laboratório de Insetos Frugívoros, Universidade Federal da Grande Dourados (UFGD), Dourados-MS, Brazil. Awarded with research grant from CNPq-Brazil.
Abstract - Lonchaeidae consists of robust species of flies, of moderate size (from 3 to 6 mm long), body usually black, dark blue or green, with noticeable metallic reflections in some groups. Although several lonchaeid species have been associated with fruit or flowers of various species of Angiospermae of economic importance, they remain poorly studied.
This may be related to the fact that the systematics of this group presents major taxonomic difficulties. The distinction between many species and even between some genera can be quite confusing. In order to elucidate the aspects of biogeography, ecology and taxonomy of the Lonchaeidae, we searched for publications in seven large databases (Web of
Knowledge, Scopus, Elsevier, Jstor, Sciencedirect, SpringerLink, and Wiley Inter Science).
Papers that presented any aspects on the systematics, taxonomy, bioecology and/or biogeography of any species of Lonchaeidae were selected. From January 1878 to
December 2013 we found 201 papers published about Lonchaeids. In total, 538 species are described in two subfamilies and ten genera. Two monotypic genera are fossils: Morgea
9
mcalpinei Hennig and Glaesolonchaea electrica Hennig. There are eight living genera with
536 species: 127 species of Dasiops Rondani, eight of Protearomyia McAlpine, seven of
Chaetolonchaea Czerny, 23 of Earomyia Zetterstedt, 20 of Lamprolonchaea Bezzi, 213 of
Lonchaea Fallén, 98 of Silba Macquart, and 40 of Neosilba McAlpine. From the nineteenth to the twentieth century, there was a progressive increase in publications about Lonchaeids worldwide. In the XXI century, the increase of publications on Lonchaeidae reflects the growing interest of researchers in studying their systematics, ecology and biogeography.
On the other hand, the evident difficulties in the taxonomy of Lonchaeids can be a limiting factor for the entrance of beginner entomologists in the study of the lance flies. The lack of taxonomic keys and illustrations and the scarcity of research groups specialized in
Lonchaeidae hinder and discourage the production of scientific knowledge about these flies. Even though they are responsible for considerable economic losses in crops, several basic aspects of their biology, ecology and biogeography are still unknown. This review provides a map of the world distribution of the species, and a taxonomic key for identification of subfamilies and all living genera worldwide.
Key Words: Lonchaeid, Lance Flies, Hosts, Neotropics, Natural history, Taxonomy.
Introduction
The Lonchaeidae are cyclorraphan acalyptrate Diptera, belonging to the superfamily
Tephritoidea which also includes Pallopteridae, Piophilidae, Uliididae (= Otitidae =
Pterocallidae), Richardiidae, Platystomatidae, Pyrgotidae, and Tephritidae (Korneyev 1999;
Woodley et al. 2009). Lonchaeids in total sum 538 described species (two fossils) in two subfamilies and eight living genera in the world. They can be found in a wide variety of 10
habitats, from the Arctic Circle to the Equator. However, there are no reports of endemism or occurrences at the poles or in New Zealand (McAlpine 1962).
Lonchaeidae forms a peculiar group because they present a typical standard body color: black and shiny. Sizes are moderate to small (3-6 mm) and with robust body. The female of all species have long membranous ovipositor, sometimes rigid (aculeus), and when distended it can be as long as the pre-abdomen. The tip of the aculeus is usually fine and slender, but in some species (e.g., some Dasiops) this structure is rather flattened and enlarged, forming a laminated structure similar to a spear, hence the origin of the popular name “lance flies” (Korytkowski & Ojeda 1971).
Formerly all species of Lonchaeidae were included within the family Sapromyzidae, when only a single genus (Lonchaea) was recognized. Lonchaea was originally raised to genus level by Fallén (1820), who had grouped all species of Lonchaea together with the
"Ortalides".
Wiedemann (1830), Macquart (1843), Zetterstedt (1848), Walker (1860), and
Becker (1895) gradually increased the number of described species in the genus Lonchaea, but their descriptions are superficial compared to modern standards and based on flimsy or variable characters, resulting in taxonomic confusion (Korytkowski & Ojeda 1971).
Lonchaeidae have been recognized as a family since Malloch & McAtee (1924) separated the Sapromyzidae into three families: Lonchaeidae, Sapromyzidae (now
Lauxaniidae) and Pallopteridae.
Zetterstedt (1848) described the genus Earomyia; Macquart (1851), the genus Silba;
Rondani (1856), Dasiops; Bezzi (1920), Lamprolonchaea; Czerny (1934), Chaetolonchaea, and McAlpine (1962), the two genera: Neosilba and Protearomyia, thus completing the eight genera currently recognized for Lonchaeidae. 11
Currently, Lonchaeidae are divided into two subfamilies: Dasiopinae, which includes only the genus Dasiops, and Lonchaeinae, that includes the seven other genera:
Protearomyia, Lamprolonchaea, Chaetolonchaea, Earomyia, Silba, Neosilba and
Lonchaea (McAlpine 1987; Korytkowski & Ojeda 1971).
Because most Lonchaeidae species are morphologically similar to each other, their taxonomy is difficult (McAlpine & Steyskal 1982), and identification to the species level can be extremely complicated. Additionally, the lack of comprehensive revisions of most of the lance fly genera, discourages and hinders future work with Lonchaeidae. In this review, we evaluate, interpret and discuss the available literature about Lonchaeidae, with the aim that this work may stimulate advances in the study of these flies.
Methodology
In order to elucidate the biogeographical and ecological aspects of lonchaeid flies, we conducted a literature review, compiling information from seven large databases (Web of Knowledge, Scopus, Elsevier, Jstor, Sciencedirect, SpringerLink and Wiley Inter
Science). We also consulted the references cited in publications obtained in these databases.
All publications that addressed any issues that would include information regarding
Lonchaeidae were selected, reviewed, interpreted, and its content reported in this review. A map of the geographical distribution (by DIVA-GIS 7.5.0), and an illustrated taxonomic key (adapted from McAlpine 1987) with dichotomous entries for the two subfamilies, and the eight living genera of Lonchaeidae is presented, taking in account the current classification (McAlpine 1989).
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Results
Lonchaeidae was first recognized as a family in 1878 by Osten Sacken, in his catalog of species of North America, although Williston (1896) had recognized it as a subfamily of Sapromyzidae.
Coquillett (1902) described some species of Lonchaeidae in North America, and soon after, Meijere (1908, 1910, 1915) examined species from South Asia. Bezzi (1910) pioneered the description of Lonchaeidae in Brazil, when he published a work citing nine species. This author defined two genera: Palloptera Fallén and Lonchaea Fallén. In 1912,
Cameron published the first paper on the biology of these flies, having investigated some biological aspects of Lonchaea chorea (Fabricius). In the next year, Towensend (1913) added information about the family, and published the description of Dasiops saltans.
Melander (1913) decided to keep the lonchaeids within Sapromyzidae in his review of the species from North America.
In 1917, Silvestri conducted the first study assessing the damage caused by larvae of lonchaeids in host fruits. He also published a thorough morphological description for Silba adipata (as Lonchaea aristella Becker), and discussed a possible parasitoid associated with this species.
In 1919, Bezzi began to investigate the African lonchaeids in an important taxonomic and evolutionary study of the species of Lonchaea. In that paper, he established three groups of species, relating them to the habits of their larvae and the characters of the adults, such as feathering of the arista, the width of the antenna, and chaetotaxy of the body.
Bezzi (1919) defined as more primitive the Lonchaeidae species whose larvae develop in decaying organic matter or animal excrement.
13
Bezzi (1919) assumed that larvae with these habits could successively attack plant organs previously infested by other insects or pathogens, as in the cases of Lonchaea chorea (Fabricius) and Lonchaea longicornis Williston. The adults of these primitive species were characterized by having broad antennae, with micro-pubescent arista, and well defined chaetotaxy.
The second group defined by Bezzi (1919) was composed for lonchaeids with larvae galling stems of grass, whose adults were characterized by having very short antennae (not reaching a half of the face), with glabrous arista, and chaetotaxy poorly developed, represented by the species of Dasiops Rondani.
In the third group were included the carpophagous (fruit-feeding) larvae of species from the subtropical zone, in which the adults have a plumose arista, third antennal segment quite long, and well developed chaetotaxy. Bezzi (1919) indicated that the species of this group should be separated into a new taxon, and that the species Lonchaea plumosissima
Bezzi and Lonchaea pendula Bezzi should be included in it (Bezzi 1919).
The following year, based on his previous work, Bezzi (1920) recognized three groups as subgenera of Lonchaea: Subgenus Lonchaea, Subgenus Dasiops and Subgenus
Carpolonchaea. In addition, he suggested that anomalous species such as Lonchaea aurea
Macquart, whose larvae live in tomato fruit (Solanum lycopersicum Mill.) (Solanaceae), could be included in another new subgenus, Lamprolonchaea, in which the adults are characterized by having their head with a very peculiar shape, typical wing venation, and metallic body coloration. Bezzi in 1923 made a further contribution by publishing an identification key for the species of Lonchaeidae of Australia.
Malloch (1920) recognized the family Lonchaeidae as distinct from Lauxaniidae, describing some new species of the genus Lonchaea, such as L. major and L. 14
nigrocaerulea, based on material from the Amazon and Barbados, respectively. Currently, these species are classified as belonging to the genus Neosilba McAlpine.
Malloch & McAtee (1924), removed the genera Palloptera and Eurygnathomyia
(Czerny, 1904) from Lonchaeidae, and included both in Pallopteridae, thus definitely separating Lonchaeidae from Sapromyzidae. In the following year, Aldrich (1925) included a new species in Lonchaeidae: Lonchaea hirtithorax Aldrich (now Earomyia hirtithorax).
Curran (1932) added new knowledge to the taxonomy of Lonchaeidae, publishing an identification key to species of Lonchaea from Central America. In that key, Curran described Lonchaea luridipennis (= Dasiops) from Venezuela; Lonchaea trita from
Panama; Lonchaea bakeri (= Dasiops) from Nicaragua; Lonchaea fuscipennis (= Neosilba) and Lonchaea dimidiata (= Neosilba), both from Panama; Lonchaea ceres from Brazil;
Lonchaea duida from Venezuela; Lonchaea nigrocaerulea Malloch (= Neosilba) from
Barbados; Lonchaea batesi (= Neosilba) from Guatemala, and Lonchaea major Malloch (=
Neosilba) from Peru.
Czerny (1934) investigated the lonchaeids from the Palearctic Region, and Hennig
(1941) for the first time included the adult genitalia in the description of species of
Lonchaeidae, demonstrating the importance of these characters, which thereafter have generally been used in identification of these Tephritoidea.
Blanchard (1948) published the first revision of the Argentine species of
Lonchaeidae, describing some new species, and adding new information about
Carpolonchaea [You already said that]. Blanchard (1948) noted that the species of this genus are mostly Neotropical, and indicated L. plumossisima Bezzi (1919) as type species and suggested that Carpolonchaea could be elevated to the level of genus.
15
Hennig (1948), in his concept of the genus Carpolonchaea, included seven
Neotropical species, among them: Carpolonchaea dimidiata (Curran) from Panama and
Venezuela, and Carpolonchaea pendula (Bezzi), from Chile, Peru and Costa Rica. In the same paper, Hennig described Carpolonchaea magna from Peru, Carpolonchaea parva from Peru and Costa Rica, Carpolonchaea longicerata from Peru, Carpolonchaea devians from Bolivia (currently in Silba), and Carpolonchaea intermedia from Peru; this last species was later included in the genus Lonchaea.
Collin (1953) reviewed the British lonchaeids, and McAlpine (1956a) analyzed species of the Old World, transferring Lonchaea (Carpolonchaea) plumosissima Bezzi to
Silba, indicating that that species was not correctly described by Bezzi. In another publication in the same year, McAlpine (1956b) added more information about these flies, clarifying the identities of 15 species, six of which were described as new to science. In the same year, Smith (1956) added information on the biology of Lonchaeidae, describing the immature stages of some Lonchaea species.
Berry & Vaquero (1957) analyzed species of Lonchaeidae as potential pests, pointing out that this group was of little economic importance, composed by few species, and reported Carpolonchaea pendula Bezzi in El Salvador (Central America). In the following years, contributions to lonchaeid taxonomy continued, with several species described or redescribed in different geographical regions (Collin 1957; McAlpine 1958;
Sasakawa 1958; Morge 1959; McAlpine 1960).
In the 1960s, publications about the phylogeny and ecology of Lonchaeidae began to appear. McAlpine (1960) published a key based on species from southern Africa, showing how to identifying genera of Lonchaeidae. In this paper, he included
Chaetolonchaea Czerny, Lamprolonchaea Bezzi, Earomyia Zetterstedt, Dasiops Rondani, 16
Silba Macquart and Lonchaea Fallén. The following year started with publications relating the Lonchaeids as pests of agricultural importance for some cultures, such as the apricot
(Moffit & Yaruss 1961; McAlpine 1961). McAlpine (1962) published an extensive review on the Lonchaeidae worldwide, presented a phylogeny, adding new taxa, and updating to eight genera: Dasiops Rondani, Protearomyia McAlpine (new genre); Chaetolonchaea
Czerny, Earomyia Zetterstedt, Lamprolonchaea Bezzi, Silba Mcquart, Neosilba McAlpine
(= Carpolonchaea) and Lonchaea Fallén. This taxonomist recognized five genera proposed by Hennig (1948), except Spermatolonchaea Bendel, which was included within Earomyia
Zetterstedt and Carpolonchaea Bezzi, which according to the principles of priority was designated as Silba Mcquart.
McAlpine (1962) considered Lonchaea as the most evolved genus of Lonchaeidae, defining it as the sister group of Neosilba, assuming that these two genera come from a common ancestor with Silba, very similar to this, but it would have lost the setae of the prosternum. He also pointed out that some changes in morphological characters happened, including decreases in the feathering of arista. He described the genus Neosilba, designating Lonchaea batesi Curran (1932) as type species, previously assigned to the genus Carpolonchaea by Hennig (1948), and to the genus Silba by McAlpine (1960).
McAlpine (1962) recognized Silba devians as the single one species of Silba in the
Neotropical Region, and transferred all remaining species previously treated as
Carpolonchaea to the genus Neosilba.
McAlpine (1962) realized that the genus Neosilba is quite similar to Silba, except by having the third antennal segment usually three times longer than wide (usually in Silba this segment is four to five times longer than broad); arista with well developed feathering or rarely glabrous, and scutellum with setulae between the basal and apical to the scutellar 17
arrows. He mentioned that its larvae in some cases are associated with the fruit pest species, particularly with Tephritidae or frugivorous larvae of Lepidoptera, causing physiological or mechanical injury in the host fruit. McAlpine (1962) reported that Neosilba pendula
(Bezzi) had been reported as primary pest of pepper (Capsicum sp.) (Solanaceae) in Peru.
One of the most important contributions of McAlpine (1962) was the provision a key to identify species Neosilba, including N. intermedia (Hennig) (Peru), N. devians
(Hennig) (Bolivia), N. flavipennis (Morge) (Holartico), N. fuscipennis (Curran) (Panama),
N. longicerata (Hennig) (Peru), N. dimidiata (Curran) (Panama), N. magna (Hennig)
(Peru), N. glaberrima (Wiedemann) (Western India, South and Central America), N. batesi
(Curran) (Guatemala), N. nigrocaerulea (Malloch) (Barbados), N. major (Malloch)
(Brazil), N. parva (Hennig) (Peru), and N. pendula (Bezzi) from Brazil.
Morge (1963) proposed a new phylogenetic scheme based on species from
Palaearctic, in which the Lonchaeids were subdivided into three subfamilies: Dasiopinae which includes the genera Silvestrodasiops Morge and Dasiops Rondani; Earomyiinae, comprising the genera Priscoearomyia Morge; Chaetolonchaea Czerny, Earomyia
Zetterstedt and Lamprolonchaea Bezzi, and Lonchaeinae with Silba Macquart,
Setisquamalonchaea (new genre) and Lonchaea Fallén. However, this classification has never been well accepted by the experts in Lonchaeidae.
McAlpine (1964) presented a crucial paper to broaden the knowledge of
Lonchaeidae, with an extensive list of new species, as result of his elegant paper published in 1962, which describes 160 species from different regions of the world, with 20 of them from Neotropics, being 17 species of Dasiops, and three of Lonchaea.
The 1960s were marked by publications on the taxonomy and biology of
Lonchaeids, with descriptions of several new species in distinct genera, and records of 18
species occurrences from different localities worldwide (Romero & Ruppel 1973;
Sasakawa 1964; Johnsen et al. 1965; Nuorteva 1967; Hennig 1967).
Hennig (1967) did a remarkable study of fossils of Lonchaeids in amber, describing the species Morgea mcalpinei Hennig and Glaesolonchaea electrica Hennig. The age of those extinct species was estimated between 48.6 and 40.4 million years ago. He proposed a phylogenetic model for Lonchaeidae, very similar to that presented by McAlpine (1962).
In the same year, Beaver (1967) and Nuorteva (1967) added information on biology to some species Lonchaea.
Later on, McAlpine & Munroe (1968) published important information on ecology of some species of Lonchaeids, and described four new species. These authors noted a bit peculiar behavior for that flies: the swarming (formation of leks for mating). In fact, the first author continued studying these flies, producing two more publications on the biology and taxonomy of lance flies (McAlpine 1970; McAlpine & Morge 1970).
In the 70s, many articles were published on Lonchaeidae. Some authors emphasized predation by larvae of lonchaeids (Lonchaea) on larvae of other insects (Harman &
Wallace 1971; Kishi 1972). Other publications have been devoted to the economic importance of lonchaeids as agricultural pests (Romero & Ruppel 1973; Stegmaier 1973;
Brinholi et al. 1974; Boza & Waddill 1978; Steyskal 1978; Waddill & Weems 1978; Pena
& Waddill 1982). But in fact, the vast majority of publications at that time continued toward the description of new species and new habitats, such as the samplings on
Lonchaeids diversity undertaken in Russia (Kovalev 1973, 1976, 1978), Austria (Morge
1974), Sri Lanka (McAlpine 1975a), Malaysia (McAlpine 1975b), Afghanistan (Soos 1977) and Chile (McAlpine 1974).
19
Korytkowski & Ojeda (1971) reviewed the species of Lonchaeidae from Peru, based on material collected in the North of that country, and reported a list with 80 Neotropical species, distributed in three genera. In fact, many species treated in 1971 by Korytkowski &
Ojeda as Silba, currently are species of Neosilba.
In the 80s were published some fundamental papers to consolidate the knowledge about Lonchaeidae, with revisions of three genus: Chaetolonchaea (McAlpine & Steyskal
1982), Neosilba (McAlpine & Steyskal 1982), and Protearomyia (McAlpine 1983), which updated the information about these taxa. It is remarkable that the work McApine (1982) remains the main reference on Lonchaeids the Neotropics.
Another important contribution was the publication of a Lonchaeidae checklist, in which 17 species of Dasiops, one species of Lamprolonchaea, 17 species of Lonchaea, and
17 of Silba were registered to the tropical regions of Africa (McAlpine 1981).
Papers on the biology and ecology of these flies were also published in the same decade. The investigation of the predatory behavior of Lonchaea corticis Taylor upon larvae of weevils was presented by several authors (Alfaro & Borden 1980; Hulme 1989) that investigated this aspect in many laboratory experiments. The biology of Lonchaeidae was also investigated, such as some aspects of Earomyia species (Hedlin et al. 1981;
Morge & Nanu 1981), species of Lonchaea (Hedlin et al. 1981; Kovalev 1984), species of
Silba (Awadallah et al. 1980; Katsoyannos 1983a) and species of Dasiops in South
America (Peñaranda et al. 1986).
Other important biological data were added in the years 80s, such as the swarming courtship behavior of Lamprolonchaea species, and the discovery of a potent attractant to catch Silba adipata (a key pest of figs, Ficus carica L.) in traps was provided by
Katsoyannos (1983b). Guerin & Katsoyannos (1984) published a further contribution, 20
clarifying the biology of lance flies. Also, in this decade was published for the first time, the only paper one that studied the aspects of molecular biology on Lonchaeids (De Conti et al. 1984).
Regarding the damage caused by larvae of Lonchaeids in plant tissues, were published three important papers in the years 80s, involving different Lonchaeidae species, attacking different organs of different plant species. The first study was about species of
Dasiops attacking fruits and flowers of passion fruit (Passiflora sp.) (Steyskal 1978). The second paper refers to the attack of some species of Earomyia in seeds of fir (Abies amabiles) (Hedlin et al. 1981). The third paper refers to the damages caused by larvae of
Neosilba perezi in terminal buds of cassava (Manihot esculenta) (Pena 1982). Within the taxonomy of Lonchaeidae, were published keys to characterize different Lonchaeids species, including the Dasiops from Africa (Morge 1980; Shtakel'berg 1989), and
Lonchaea species from Europe (Kovalev 1981, 1984). Lonchaeidae species from Ireland were also investigated for the first time (Chandler 1986). Also in the 80s, two important book chapters were published about the Lonchaeids in the Manual of Nearctic Diptera. In
Volume II, the taxonomy and biology of that family were fully explored, with several illustrations that have enriched the work. Moreover, the author has provided a key for identification of living Lonchaeidae genera, such as be currently accepted: Chaetolonchaea
Czerny, Dasiops Rondani, Earomyia Zelterstedt, Lamprolonchaea Bezzi, Lonchaea Fallén,
Neosilba McAlpine, Protearomyia McAlpine, and Silba Macquart (McAlpine 1987 - see key adapted to the end of this review). Phylogeny and classification of Lonchaeidae were presented in Volume III of that same publication. McAlpine (1989) suggested that the aedeagus of the most ancient species of Lonchaeid was simple (unsegmented and without apical glands), membranous and subsequently were modified to form the several types of 21
aedeagus currently found in Lonchaeids. Other plesiomorphics characteristics indicated by
McAlpine are entirely black color of body, including the head and legs, the presence of the seven abdominal spiracles in males and females, and three spermathecae in females. While none of these features is unique to Lonchaeidae, no other families of Tephritoidea present all of them together. Thus, the author suggested that Lonchaeidae could be the sister group of all other Tephritoidea.
Finally, McAlpine (1989) defined the Lonchaeidae into two subfamilies: Dasiopinae
(oldest lineage) containing the genus Dasiops, and Lonchaeinae (more recent), containing the tribe Earomyiini, with the genera: Protearomyia, Chaetolonchaea, Earomyia,
Lamprolonchaea, and the tribe Lonchaeini, with the genera: Lonchaea,
Setisquantalonchaea (= now Silba), Silba, and Neosilba. In addition, he reports that
Lonchaeidae should contain about 700 species worldwide, of which about a half were undescribed species. McAlpine (op. cit.) highlights that Dasiops and Lonchaea are the most specious genera, with distribution in all zoogeographic regions; that Protearomyia,
Chaetolonchaea, and Earomyia are present throughout the Holoartic region, but
Protearomyia also occurs in Patagonia. Lamprolonchaea and Silba are distributed mainly in the Oriental region, but also occur in Ethiopia and Australia; that Neosilba is limited to
Neotropical Region, and Setisquamalonchaea (= now Silba) is endemic to the Palearctic
Region.
In the 90s, many papers about lonchaeids were published, addressing mainly on the fact that several species of Lonchaeidae are characterized as potential pests for agriculture.
Causton (1993) reported the first attack in a species of passion fruit by a species of Dasiops in Venezuela. Norrbom & McAlpine (1997) reviewed the genus Dasiops, highlighting some pest species in passion fruit (Passiflora spp.), and they suggested that other species 22
have potential to be used in biological control against invasive species of Passifloraceae in
Hawaii, and worldwide. Lourenção et al. (1996) and Bellotti et al. (1999) evaluated the infestation Neosilba perezi in shoots of cassava, reporting this species as an important pest for cassava in South America. Ahlmark & Steck (1997) pointed out the importance
Neosilba batesi as a secondary pest of avocado (Persea americana) in Florida. Finally,
Owens & Morris (1998) identified larvae Earomyia abietum causing damage in seeds of firs.
Uchôa (1999), in a pioneering study on the diversity and trophic interactions of
Tephritoidea with their host plants in southwest Brazil, found that lonchaeids in many species of native and cultivated fruits, including Citrus sinensis and Citrus reticulata
(Rutaceae) are even more abundant and economically important that Tephritidae. This research provided information that led to other Brazilian researchers to launch a different look on Lonchaeidae, whose species were previously considered secondary invaders of fruits and vegetables (Uchôa & Zucchi 1999; Uchôa et al. 2002; Uchôa et al. 2003a; Uchôa et al. 2003b; Uchôa et al. 2003c; Uchôa 2012).
From the 90s, the Lonchaeidae have been considered as potential pests of vegetables and fruit, beginning of the first studies related to control these flies. In Chile, Sánchez et al.
(1991) evaluated the effect of four different insecticides for the control of Neosilba species.
Other important studies were related to natural enemies, seeing the use of parasitoids for biological control of lance flies. Wharton et al. (1998) made associations of some species of
Eucoilidae as parasitoid in larva of Lonchaeidae that eat pulp of fruit. Gallardo et al. (2000) has described a new species of Trybliographa (Eucoilidae) in Brazil (Trybliographa infuscata Gallardo et al. 2000), associated with larvae of Neosilba pendula infesting fruit
23
of: orange, Citrus sinensis, tangerine, Citrus reticulata (Rutaceae) and pequi, Caryocar brasiliense (Caryocaraceae) in the Cerrado.
The 90s were also marked by the presence of numerous records for lonchaeids in
Europe. Werner (1995), and Maca (1999) had records of species in Germany; Maca (1997) recorded species lonchaeids the Czech Republic, and Slovakia. Chandler (1998) listed the
British species of Dasiops, and MacGowan & Rotheray (2008), reported Lonchaea species in the UK.
Dobson (1992) published a rather curious observation, pointing out that the adults of
Lonchaeid species are possible cleptoparasites of spider’s web. Since that time, this is the only paper that suggests this kind of behavior for Lonchaeidae.
The papers published during the last decade, left quite clear the need for taxonomic studies on Lonchaeids. Thus, various specimens deposited in museums were analyzed and from this material, some new species have been described. MacGowan (2004a, 2006a,
2007a, 2008) examined specimens from European museums and corrected numerous taxonomic errors in samples. He examined specimens from Malaysia deposited in the
Natural History Museum of Sweden, described a new species of Dasiops, two of Lonchaea, and nine of Silba (MacGowan 2005a). MacGowan et al. (2007) also examined specimens from a museum in Switzerland, and illustrated for the first time the male genitalia for
Lonchaea hyalipennis Zetterstedt 1848. On the other hand, some researchers explored the
Neotropical species of Neosilba, publishing papers on the ecological context (Araújo &
Zucchi 2002; Bomfim et al. 2007, Caires et al. 2009).
On taxonomy, more recently several authors described new species of lonchaeids.
Species were described in Brazil (Strikis & Prado 2005; Strikis & Prado 2006; Strikis &
Lerena 2009; Strikis & Prado 2009), Spain (MacGowan 2000; MacGowan 2009); UK 24
(MacGowan & Rotheray 2000), Andorra (MacGowan 2001a), Switzerland (MacGowan
2001b), Taiwan (MacGowan 2004b, 2007b), Israel (MacGowan & Freidberg 2008), from
Central and Southern Africa (MacGowan 2005b), China (MacGowan 2007b), and Thailand
(MacGowan 2007b).
About Lonchaeidae diversity were reported species in the Great Britain region
(MacGowan 2001c, 2006b, MacGowan & Rothery 2008), Denmark (MacGowan 2001d,
2005c), Czech Republic (Maca & Bartak 2001), London (Allen 2002, 2003), Spain (Carles-
Tolra 2004), and Brazil (Bittencourt et al. 2006).
Statement about species considered pests remained fairly addressed in this decade.
Aluja & Díaz-Fleischer (2004) found N. batesi infesting fruit avocado Persea americana in
Mexico. In Brazil, Araújo & Zucchi (2002) evaluated the levels of infestation by N. pendula on native and exotic fruits, finding quite high infestation levels for fruits in barbado's cherry (Malpighia emarginata). Carvalho & Marques (2004) and Santos et al.
(2004) reported N. zadolicha attacking umbu-caja fruit's (Spondias sp.). Still in Brazil,
Neosilba species attacking loquat's fruit (Eriobotryia japonica), and coffee (Coffea arabica) (Strikis & Prado 2005; Strikis & Prado 2006, respectively) were reported.
Fruits of fig (Ficus carica) (Moraceae) were also reported as being quite attractive to lonchaeids. Larvae of Lonchaeid species such as Silba adipata were reported (in Iraq, in
Montenegro, and in South Africa) (Abdullah & Fandy 2006; Giliomee et al. 2007; Radonjic
& Hrnci 2009), and Lonchaea aristella (in Portugal) (Gonçalves et al. 2008) as pests of fruits of Moraceae. Dasiops caustonae was also recorded infesting passion fruit (Passiflora edulis) in Colombia (Umana 2005).
The last decade was marked by a large increase in publications on the genus
Neosilba. Montes et al. (2010) investigated the infestation rates by Neosilba species in 25
varieties of peach (Prunus persica). Uchôa & Nicácio (2010) associated larvae of Neosilba species with 38 host fruit species. Gisloti & Prado (2011a, 2011b), and Strikis et al. (2012), analyzed the infestation of a Neosilba species in cassava (Manihot esculenta), and Lopes et al. (2008) analyzed the infestation by larvae of Neosilba in tangerine (Citrus reticulata) orchards. Aguiar-Menezes et al. (2007) reviewed the species Neosilba infestation in different varieties of coffee (Coffea arabica). In the same period were recorded parasitoids associated with larvae of several species of Lonchaeidae (Aguiar-Menezes et al. 2008;
Souza et al. 2012), including a new species of parasitoid (Arouca & Penteado-Dias 2011;
Gisloti & Prado 2012; Marsaro-Junior et al. 2012, Fernandes et al. 2013).
On the diversity of host fruits of Lonchaeidae in Brazil, Uchôa (2012) related species Neosilba as possible candidates for biological control of mistletoe Psittacanthus acinarius (Martius) (misidentified the Psittacanthus plagiophyllus Eichler) (Santalales:
Loranthaceae) a hemi-parasite of tropical trees; Montes et al. (2012) and Camargos et al.
(2011) investigated the infestation of these flies in Coffea arabica; Adaime et al. (2012) associated with species of Neosilba in fruits of murici (Byrsonima crassifolia), Dias et al.
(2012) found larvae Neosilba attacking fruits of Jatropha curcas, Bittencourt et al. (2013) reported five new occurrences of Neosilba species in Brazil, and Nicácio & Uchôa (2011) investigated the biodiversity of these flies in the Pantanal region.
The taxonomy of Neosilba species were well explored in the recent years. Strikis
(2011) described eleven new species in Brazil, and Galeano-Olaya & Canal (2012) described ten new species of Neosilba from Colombia. The genus Lamprolonchaea also had a major publication recently (Blacket & Malipatil 2010).
The species of Dasiops have been explored in recent papers, pointing out about their status as pests in plants of economic interest. They were considered major pests of the 26
cactus (Selenicereus megalanthus) (Medina & Kondo, 2012), and in flowers of passion fruits, (Passiflora sp). (Imbachi et al. 2012; Quintero et al. 2012; Wyckhuys et al. 2012).
In the last decade the lonchaeids remained being reported in Spain (Carles-Tolra et al. 2010), France (MacGowan 2012), Russia (MacGowan 2010), in UK (Perry 2010), the
Czech Republic (Barták & Roháček 2011), India (MacGowan et al. 2012), and Japan
(MacGowan & Okamoto 2013). Finally, MacGowan & Okamoto (2013) modified the classification of Lonchaeidae, reassessing the genus Silba, and concluding that the genus
Setisquamalonchaea Morge is a junior synonym of Silba Macquart.
Discussion
The Lonchaeid family consists of two subfamilies: Dasiopinae, containing a single genus Dasiops, and Lonchaeinae, the more biologically diverse subfamily, containing the remaining seven genera (see adapted key from McAlpine 1987, available at the end of this paper). The classification of Lonchaeids has been relatively stable. Czerny (1934) erected the genus Chaetolonchaea, and McAlpine (1962) described the genus Protearomyia, then
Setisquamalonchaea was added by Morge in 1963, but it is now regarded as a synonym of
Silba (MacGowan & Okamoto 2013).
Species of Dasiops are mainly found in the Nearctic region, though some species are found less frequently in the Palaearctic region, southern Africa, Asia, Australasia and
South America. The species of Earomyia and Chaetolonchaea are restricted to the Nearctic and Palaearctic regions. Lonchaea species are commonly found in the Palaearctic, but there are some records of endemic species in both Nearctic and Asian regions. Protearomyia species are distributed primarily by Nearctic region, but there are also some species of this genus in the Palaearctic and southern Chile. Lamprolonchaea species are confined to the 27
Palaearctic and Australasian regions. Silba species are present mainly in Africa, Asia and
Australia, having a single species in South America, but the species of Neosilba are restricted to the Neotropical region (Fig. 1).
28
Figure 1. Geographical distribution of the Lonchaeidae (Diptera: Tephritoidea) genus around the world.
29
About 201 papers on Lonchaeidae species (Fig. 2) were published from 1878 to
November 2013. A total of 538 species were described in two monotypic genera from fossils, and eight living genera, being 127 species of Dasiops (Fig. 3), 8 species of
Protearomyia (Fig. 6), 20 species of Lamprolonchaea (Fig. 8), 7 species of
Chaetolonchaea (Fig. 10, 11), 23 species of Earomyia (Figs. 12, 13), 213 species of
Lonchaea (Figs. 16, 17), 40 species of Neosilba (Fig. 18), and 98 species of Silba (Fig.
19).
Of total publications (201) about Lonchaeidae in 135 years (1878-2013), the species of Neosilba are pointed out in 60 papers, standing out in the publications. That paper mainly makes associations of larvae of Neosilba species as potential pests of several species of fruit, being many of them of commercial interest (Lourenção et al. 1996; Araújo
& Zucchi 2002; Santos et al. 2004; Uchôa 2012). The species of Neosilba appear to be the ecological equivalent for the species of Silba in the New World. Both genera are very similar in their morphological and ecological aspects. The species of Neosilba are known from Caribe, Mexico, Colombia to Brazil. Forty species were described, and at least more
60 species waits for description in the Neotropical region (McAlpine & Steyskal 1982).
30
Figure 2. Distribution of publications about the Lonchaeidae genera (Diptera:
Tephritoidea), since 1878 until December 2013 (n = 201) [*other = Papers about
Lonchaeidae without specified genus].
Although Araújo & Zucchi (2002) have speculated that the species of Neosilba probably occurs as secondary invaders of fruits, after the primary attack by species
Tephritidae, nowadays it is known that the larvae of several Neosilba species infest a wide variety of fruits and vegetables from different angiosperm families, acting as primary invaders.
The genus Chaetolonchaea has the lowest number of publications; just a revision of it by McAlpine in 1956b, beyond that, there are record of two species in Hungary
(MacGowan 2007a). This genus is restricted to the Nearctic and Palaearctic regions. Three species were recorded in Europe and two in the Nearctic. Information regarding to the natural history and ecology of Chaetolonchaea species are scarce.
The genera Protearomyia (2 papers), Lamprolonchaea (4 papers), and Earomyia (9 papers), are poorly studied. The publications on Earomyia are mainly lists of species
31
records, and some aspects of their geographical distribution worldwide (Skrzypczyńska
1984; Perry 2010). Protearomyia is poor in described species, with records of occurrence in the Nearctic and Europe (Palearctic). Earomyia is composed by species distributed in the
Palearctic and Nearctic regions.
The genus Lamprolonchaea is composed by colorful species with metallic tones, mainly distributed in the Oriental region, and in the Australasia. One species, L. smaragdi
(Walker), was found in the Mediterranean and Northern Europe (Netherlands and
Germany), associated with fruits of the following hosts: Capsicum sp. (Solanaceae), Prunus armeniaca and Prunus persica (Rosaceae), Citrus paradisi, Citrus limon, Citrus reticulata
(Rutaceae), and Persea americana (Lauraceae) (Blacket & Malipatil 2010).
The genus Lonchaea (51 publications) has been the best studied. It has the largest number of described species in Lonchaeidae, with a wide geographic distribution. There are
213 named species, present in areas of boreal forest of Palaearctic and Nearctic regions.
Currently, there are about 65 species recorded in Europe, and 72 in Nearctic region.
Researches about the species of this genus have focused on investigations of predatory potential of their larvae on larvae of wood-destroying beetles (Harman & Wallace 1971;
Kishi 1972; Alfaro & Borden 1980; Hulme 1989; Hulme 1990).
Many species of Dasiops are commonly associated as pests of fruit of some species, especially of Passifloraceae (Moffit & Yaruss 1961; Stegmaier 1973; Steyskal 1980;
Norrbom & McAlpine 1997; Causton & Rangel 2002; Uchôa et al. 2003a, 2003c, Amaya et al. 2009; Delgado et al. 2010; Quintero et al. 2012; Carrero et al. 2013). This genus is the second most species rich in Lonchaeidae, with greater species richness in Nearctic Region.
On the other hand, the genus Silba, even with a large number of described species (98), is a little-studied taxon (12 papers). Furthermore, most studies on the species of Silba focused 32
mainly on geographical distribution (Giliomee et al. 2007; MacGowan & Friedberg 2008).
This may be related to the fact that few species of Silba cause damage to fruits exploited commercially (Katsoyannos 2009; MacGowan et al. 2012; Okamoto et al. 2012).
It was found that by the 50s, the majority of publications were toward taxonomy and biogeography, with description of new species or reporting these in different geographical regions. The most studied species during this period were those of the genus Lonchaea with several studies on the taxonomy and biogeography. Two of these studies were very important: Bezzi (1919, 1920) elucidated some evolutionary aspects of the behavior of larvae, separating the species into four subgenera: Lonchaea, Dasiops, Lamprolonchaea and Carpolonchaea. Currently the species of the subgenus Carpolonchaea integrate the genus Neosilba.
The 60s, 70s and 80s were essential to the construction of knowledge about
Lonchaeidae. Important taxonomic papers were published, many of them providing keys to the correct identification of the species (McAlpine 1960, 1962; Morge 1980; Kovalev 1981,
1984).
Starting on the 90s, the investigations on Lonchaeids earn a new approach. These flies came to be seen as potential pests for many fruit species, many of which with commercial interest. There was a large increase in publications on species of Neosilba in the Neotropical region, resulting in some major publications.
In the 2000s, publications on Lonchaeidae have progressively increased, demonstrating the growing interest of researchers in the production of scientific knowledge about lonchaeids. On other hand, the taxonomic difficulties, the shortage of identification keys, and the lack of research groups specialized in Lonchaeidae, have been an obstacle in attracting entomologists beginners in the study of Lonchaeids, even being responsible for 33
considerable economic losses, the lance flies still have many aspects of their basic biology and ecology unknown.
Key to the living supraspecific taxa of Lonchaeidae (Diptera: Tephritoidea) (Modified from
McAlpine 1987)
1. Stigmatal bristles present on the sclerite above the anterior thoracic spiracles (Fig.
3)………………….………………..………DASIOPINAE...... …Dasiops Rondani
- Stigmatal bristles absent...... ………..LONCHAEINAE...... …...…...………. 2
2. Lunule bare (Fig. 4)……………………………………….………...... ……...... ……3
- Lunule setulose (Fig. 5) ..…………………LONCHAEINI ..…..………..……….....….....6
3. Scutellum with only four marginal bristles (Fig. 6)...... Protearomyia McAlpine
- Scutellum with additional bristles beyond the four marginal ones (Fig. 7)...... 4
4. Body brilliant, golden green or bronzy blue. Eye bare, hairs of frons sparse and short
(Fig. 8). Anepisternum with one to several clearly distinguishable anterodorsal bristles and two such posterodorsal bristles. Katepisternum with one clearly distinguishable bristle
...... Lamprolonchaea Bezzi
- Body predominantly black or bluish-black. Eye frequently pilose. Hairs of frons fairly dense and long (Fig. 9). Katepisternum usually with several bristles ...... 5
34
5. Scutellum with hairs anterior to the marginal bristles (Fig. 10). Anepisternum with anterodorsal bristles week or absent. Cheek and parafacial wide (Fig. 11). Calypteres with whitish margins……...... Chaetolonchaea Czerny
- Scutellum bare [i.e., without marginal bristles] (Fig. 12). Anterodorsal bristles of anepisternum long. Cheek and parafacial narrowed (Fig. 13). Calypteres always with dark margin ...... …...... Earomyia Zetterstedt
6. Usually with two bristles in the katepisternum, and plumose arista (Fig. 14). Male genitalia with surstylus presenting an organized array of prensiseta located on the inner surface, or on a basal plate along the ventral edge (Fig. 15); inner surface bare and generally with the ventral edge simple. Fringes of calypteres with or without elongated setae………………………………………………..…………...………………………...… 7
- Usually with one bristle in the katepisternum, and bare arista (Fig. 16). Male genitalia with surstylus without an organized array of prensiseta [as above] (Fig. 17); inner surface always having bristles or setae, ventral margin sometimes undulated, stretched or modified into spines or crests; Fringes of calypteres uniform in length, but if elongated setae are present, them these ones are never blackish...... Lonchaea Fallén
7. Fringes of calypteres with a cluster of long blackish setae at fold (Fig. 18). Posterior margin of anepisternum usually with a row of five to seven evenly spaced bristles
(Neotropics species)………………...... Neosilba McAlpine
- Fringes of calypteres without a cluster of long blackish setae, but if setae are present, they are never totally black (Fig. 19). Posterior margin of anepisternum usually with a discontinued row of three to five bristles...... Silba Macquart
35
Figure 3. Stigmatal bristles (in red) present on the sclerite above the anterior thoracic spiracles.
Figure 4. Lunule bare (see arrow in red).
36
Figure 5. Lunule setulose (in red).
Figure 6. Scutellum with four bristles on the edge (in red).
37
Figure 7. Scutellum with more them four bristles on the edge (in red).
Figure 8. Setae of the frons spaced and short (see arrow in red).
38
Figure 9. Setae of the frons long and dense (see arrow in red).
.
Figure 10. Scutellum with setae anterior to the marginal bristles (in red).
39
Figure 11. Cheek and parafacial wide (in red).
Figure 12. Scutellum bare in relation to the marginal bristles (see arrow in red).
40
Figure 13. Cheek and parafacial narrowed (in red).
Figure 14. Arista plumose (in red).
41
Figure 15. Surstylus with prensisetae (in red).
Figure 16. Arista bare (in red).
42
Figure 17. Surstylus without prensisetae (see arrow in red).
Figure 18. Fringes of calypteres with a cluster of long blackish setae at fold (in red).
43
Figure 19. Fringes of calypteres without cluster of long blackish setae at fold (see arrow in red).
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66
CAPÍTULO 2
ESPÉCIES DE Neosilba (DIPTERA, LONCHAEIDAE, LONCHAEINAE):
DISTRIBUIÇÃO GEOGRÁFICA E PLANTAS HOSPEDEIRAS
Gisloti, L.J¹*. & Prado, A. P.¹ ǂ
¹Departamento de Biologia Animal, UNICAMP, Campinas-SP, Brazil. [email protected]
Resumo
O gênero Neosilba McAlpine (Diptera: Lonchaeidae) é composto por espécies de moscas robustas, de tamanho moderado (variando 3 de 6 mm), geralmente de coloração negra, podendo apresentar notáveis reflexos metálicos. As espécies de Neosilba, mesmo estando associadas a diversos frutos e flores de plantas com interesse econômico, continuam negligenciadas nos estudos de moscas frugívoras. Isto pode estar relacionado ao fato de que
Neosilba apresenta grandes entraves taxonômicos, sendo a distinção entre espécies bastante confusa. Com o intuito de elucidar aspectos biogeográficos e ecológicos de Neosilba, realizou-se uma pesquisa bibliográfica em sete grandes bancos de dados (Web of
Knowledge, Scopus, Elsevier, Jstor, Sciencedirect, Springerlink e Wiley Inter Science).
Foram selecionados artigos que abordassem quaisquer aspectos bioecológico ou biogeográfico das espécies de Neosilba. Até o momento foram descritas 40 espécies distribuídas em 10 países da região Neotropical. Espécies de Neosilba obtém recurso alimentar em 107 espécies de plantas distribuídas em 35 famílias distintas. As famílias 67
botânicas que apresentaram o maior número de espécies com frutos hospedeiros de larvas de Neosilba são Fabaceae e Myrtaceae. A planta hospedeira que hospeda a maior diversidade de Neosilba é Coffea arabica. Neosilba zadolicha é a espécie mais generalista colonizando 61 espécies de frutos hospedeiros. Nove espécies de Neosilba tem o hospedeiro desconhecido, e foram registradas somente através da captura em armadilhas. A escassez de taxônomos especialistas em Neosilba é evidente, e muitos trabalhos a respeito da diversidade de moscas frugívoras apresentam dados de identificação de Neosilba apenas a nível genérico.
Palavras-chave: Acalyptratae, fruticultura, lonchaeids, moscas-das-frutas, neotrópico, história natural.
Introdução
Neosilba McAlpine (1962) é um gênero de Lonchaeidae (Diptera, Tephritoidea) com espécies de moscas cujas larvas se alimentam principalmente da polpa de diversos frutos, muito deles de interesse comercial (McAlpine 1982, Strikis & Prado 2005). Apesar de a literatura ter relatado a presença de espécies de Neosilba infestando frutos de importância econômica na região neotropical desde a década de 30, por um longo período estas moscas estiveram negligenciadas nos inventários de moscas frugívoras, sendo comum o descarte dos indivíduos provenientes desse gênero pela falta de conhecimento taxonômico e principalmente, por não representarem, naquela época, uma notável importância econômica (Costa Lima 1940, Strikis & Prado 2005, Strikis & Lerena 2009).
Na década de 70, Korytkowski & Ojeda (1971) revisaram espécies de Lonchaeidae no Peru e disponibilizaram uma chave de identificação para espécies de Neosilba 68
(classificadas equivocadamente como Silba). Nesta mesma década, uma espécie de
Neosilba (N. perezi) despertou interesse agrícola devido ao fato de suas larvas serem encontradas se alimentando de brotos de mandioca (Manihot esculenta) em plantações na
Flórida (Waddill 1978, Boza & Waddill 1978, Wadill & Weems 1978), em Porto Rico
(Romero & Ruppel 1973), na Colômbia (Steyskal 1978) e no Brasil (Brinholi et al. 1974,
Samways 1979).
A revisão elaborada por McAlpine e Steyskal (1982) foi de fundamental importância para os avanços nos estudos de Neosilba. Aquele trabalho revisou taxonomicamente 12 espécies de Neosilba previamente descritas, descreveu três novas espécies, se tornou um dos trabalhos mais relevantes até o momento, de forma que a chave para estas 15 espécies de Neosilba disponibilizada naquela publicação é utilizada amplamente por taxonomistas deste gênero.
Nos anos 90, o interesse no estudo dos Lonqueídeos foi favorecido em razão da quantidade de pupários de Neosilba obtidos em estudos de ecologia e inventários entomofaunísticos. Raga et al. (1996, 1997), em um estudo de observação da incidência de moscas das frutas (Tephritidae) em frutos de café (Coffea sp.) e laranja (Citrus sinensis) relataram que cerca de 17 e 5% dos exemplares coletados, respectivamente, pertenciam às espécies de Neosilba. Estes trabalhos geraram informações que motivaram outros pesquisadores a lançarem um olhar diferenciado sobre as espécies de Neosilba, antes consideradas invasoras secundárias de frutos (Uchôa & Zucchi 1999, Uchôa et al. 2002a,
Uchôa et al. 2003a, Uchôa et al. 2003b, Uchôa et al. 2003c, Uchôa 2012).
A partir da década de 90 as espécies de Neosilba passaram a serem consideradas potenciais pragas de hortaliças e frutíferas, com início dos primeiros estudos relacionados ao controle dessas moscas. Em um estudo pioneiro Sánchez et al. (1991) avaliaram a ação 69
de quatro diferentes inseticidas para o controle dessas moscas, no Chile. Outros estudos importantes foram relacionados aos inimigos naturais, vislumbrando o emprego de parasitóides para o controle biológico de larvas de Neosilba. Wharton et al. (1998) associaram espécies Eucoilidae parasitóides com larvas de Neosilba. Gallardo et al. (2000) decreveram uma nova espécie de Trybliographa (Eucoilidae) do Brasil, associada à moscas do gênero Neosilba infestando frutos de tangerina, Citrus reticulata (Rutaceae) e de pequi,
Caryocar brasiliense (Caryocaraceae) no Bioma Cerrado.
De fato, Lonchaeidae havia despertado interesse principalmente nos pesquisadores de Tephritidae, e isto possibilitou um aumento considerável de publicações envolvendo estas moscas, o que acabou por favorecer a compreensão de alguns aspectos biológicos e ecológicos deste grupo de Diptera (Rodrigues et al. 2006, Bonfim et al. 2007; Souza et al.
2008).
Este trabalho sintetiza e analisa os principais trabalhos realizados no âmbito do conhecimento científico sobre o gênero Neosilba na região neotropical. São apresentadas informações sobre a ocorrência, a distribuição geográfica, os hospedeiros e os inimigos naturais de todas as espécies de Neosilba. Esses aspectos são de suma importância para nortearem programas de monitoramento, manejo de risco e manejo integrado das moscas- das-frutas em fruticultura tropical.
Material e Métodos
Este trabalho apresenta uma revisão bibliográfica a partir de artigos científicos publicados entre 1830 e 2013, referentes às moscas do gênero Neosilba. Foram 70
considerados todos os trabalhos publicados durante este período que apresentasse qualquer informação referente a este gênero de Lonchaeidae.
Para a pesquisa foram utilizados sete grandes bancos de dados (Web of Knowledge,
Scopus, Elsevier, Jstor, Sciencedirect, Springerlink e Wiley Inter Science). Além disso, foram também consultadas as referências dos artigos citados nas publicações destes bancos de dados.
Finalmente, realizou-se a análise descritiva da amostra bibliográfica, acompanhada de discussão sobre os aspectos abordados por cada trabalho. Um mapa de distribuição de
Neosilba é apresentado, com uso do programa DIVA-GIS 7.5.0. A bibliografia de cada espécie foi discutida separadamente, dando ênfase à distribuição geográfica e plantas hospedeiras. As espécies encontram-se ordenadas alfabeticamente, seguida pelo local de ocorrência (país), a espécie e a família de planta hospedeira, e a respectiva referência bibliográfica.
Resultados e Discussão
O gênero Neosilba foi erigido por McAlpine (1962), representando o equivalente ecológico do gênero Silba Macquart para a região neotropical. McAlpine & Steyskal (1982) revisaram o gênero, até então composto pelas seguintes espécies: N. batesi (Curran 1932);
N. certa (Walker 1852); N. dimidiata (Curran 1932); N. fuscipennis (Curran 1932), N. glaberrima (Wiedemann 1830); N. longicerata (Hennig 1948); N. major (Malloch 1920);
N. nigrocaerulea (Malloch 1920); N. oaxacana McAlpine & Steyskal 1982; N. parva
(Hennig 1948), N. peltae McAlpine & Steyskal 1982; N. pendula (Bezzi 1919); N. perezi
(Romero & Ruppel 1973); N. pseudopendula (Korytkowski & Ojeda 1971) e N. zadolicha 71
McAlpine & Steyskal 1982. De acordo com a revisão de McAlpine & Steyskal (1982), as espécies deste gênero são amplamente negligenciadas nos estudos de Diptera, e estimaram que mais de 60 espécies novas estariam esperando descrição.
Após a primeira revisão do gênero, vinte e cinco novas espécies foram descritas: N. amphora Galeano-Olaya & Canal 2012; N. angusta Galeano-Olaya & Canal 2012; N. bella
Strikis & Prado 2008; N. bifida Strikis & Prado 2005; N. concava Galeano-Olaya & Canal
2012; N. convexa Galeano-Olaya & Canal 2012; N. cornuphallus Strikis 2011; N. delvechioi Strikis 2011; N. distospinosa Galeano-Olaya & Canal 2012; N. ilheuense Strikis
2011; N. inesperata Strikis & Prado 2009; N. laura Strikis 2011; N. mcalpiniei Strikis
2011; N. orbata Galeano-Olaya & Canal 2012; N. pantanense Strikis 2011; N. paramerolatus Strikis 2011; N. parapeltae Strikis 2011; N. piraceae Galeano-Olaya &
Canal 2012; N. plana Galeano-Olaya & Canal 2012; N. pradoi Strikis & Lerena 2009; N. pseudobifida Strikis 2011; N. pseudozadolicha Strikis 2011; N. spiculata Galeano-Olaya &
Canal 2012; N. tolimensis Galeano-Olaya & Canal 2012 e N. turgidiphallus Strikis 2011
(Strikis & Prado 2005, 2008, 2009, Strikis & Lerena 2009, Strikis 2011, Galeano-Olaya &
Canal 2012).
As espécies de Neosilba estão distribuídas em 10 países da região Neotropical:
Brasil (24 espécies), Colômbia (18 espécies), Estados Unidos da América (sul da Flórida)
(3 espécies), Guatemala (1 espécie), México (3 espécies), Panamá (2 espécies), Peru (3 espécies), Porto Rico (1 espécie), Trinidade (2 espécies) e Venezuela (1espécie) (Figura 1).
Espécies de Neosilba obtém recurso alimentar em 107 espécies de plantas distribuídas em 35 famílias distintas: Anacardiaceae, Annonaceae, Apocynaceae,
Arecaceae, Bignoniaceae, Bombacaceae, Cactaceae, Caricaceae, Chrysobalanaceae,
Combretaceae, Convolvulaceae, Euphorbiaceae, Fabaceae, Flacourtiaceae, Gnetaceae, 72
Lauraceae, Lecythidaceae, Loganiaceae, Loranthaceae, Malpighiaceae, Melastomataceae,
Moraceae, Myrtaceae, Olacaceae, Oxalidaceae, Passifloraceae, Rhamnaceae, Rosaceae,
Rubiaceae, Rutaceae, Sapotaceae, Solanaceae, Ulmaceae, Verbenaceae e Vitaceae. No centro oeste do Brasil, Uchôa et al. (2002) encontraram Neosilba spp. associadas a 22 espécies de plantas hospedeiras.
As famílias botânicas que apresentaram o maior número de espécies com frutos hospedeiros de larvas de Neosilba são Fabaceae e Myrtaceae (10 espécies de plantas em cada família). A frutífera que hospeda a maior diversidade de espécies de Neosilba é Coffea arabica (11 espécies), seguida por Inga edulis que serve de substrato alimentar e de desenvolvimento para nove espécies de Neosilba (Tabela 1).
Neosilba zadolicha é a espécie mais generalista, colonizando 61 espécies de frutos hospedeiros. N. pendula e N. glaberrima também são generalistas, colonizando 36 e 31 espécies de frutíferas, respectivamente.
Nove espécies de Neosilba: N. concava, N. convexa, N. fuscipennis, N. longicerata,
N. oaxacana, N. parapelate, N. parva, N. piraceae e N. tolimensis têm hospedeiros desconhecidos, pois foram registradas somente através da captura em armadilhas.
Bittencourt et al. (2006) encontraram N. pendula como a espécie mais comum no nordeste do Brasil, porém a utilização de armadilhas como meio de amostragem impossibilita a associação espécie-hospedeiro.
A falta de taxonomistas especializados em Neosilba é evidente em trabalhos como:
Uchôa et al. (2002a,b), Uchôa et al. (2003a,b, c); Raga et al. (2004); Rodrigues et al.
(2006); Silva et al. (2006); Bomfim et al. (2007); Minzão e Uchôa (2008) e Costa et al.
(2009), pois em tais publicações Neosilba é identificada apenas genericamente.
73
Figura 1. Distribuição geográfica do gênero Neosilba McAlpine na Região Neotropical.
74
Tabela 1. Táxons de plantas hospedeiras de espécies de Neosilba (Diptera: Lonchaeidae) no mundo.
Planta Hospedeira Espécie de Neosilba País Referências
Anacardiaceae
Anacardium humile St. Hil. N. pendula, N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Anacardium occidentale L. N. conuphallus Brasil Strikis 2011
Anacardium othonianum Rizz. N. pendula, N.zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011
Mangifera indica L. N. batesi, N.zadolicha Brasil, Colômbia Yepes & Velez 1989, Nicácio
& Uchôa 2011
Spondia dulcis Forst. N. glaberrima, N. zadolicha Brasil Uchôa & Nicácio 2010
Spondias purpurea L. N. batesi, N.certa, N.pendula, Brasil, Colômbia Yepes & Velez 1989, Araújo &
N.zadolicha Zucchi 2002, Nicácio & Uchôa
2011.
75
Spondias sp. L. N. pendula, N. zadolicha Brasil McAlpine & Steyskal 1982,
Santos et al. 2004.
Annonaceae
Annona sp. N. dimidiata, N. pendula Colômbia, Venezuela Penna & Bennetti 1995.
Annona crassiflora L. N. glaberrima, N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 201, Strikis et al.
2011.
Annona muricata L. N. batesi, N. glaberrima, Brasil, Colômbia Yepes & Velez 1989, Strikis et
N.zadolicha al. 2011.
Duguetia furfuracea (A. St.- N. certa, N. glaberrima, N. Brasil Nicácio & Uchôa 2011.
Hil.) inesperata, N.zadolicha
Guatteria discolor Fries N. cornuphallus, N. Brasil Strikis 2011.
pseudobifida
Rollinia mucosa Baill N. zadolicha Brasil Strikis et al. 2011.
76
Apocynaceae
Hancornia speciosa Gomes N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Arecaceae
Allagoptera leucocalyx (Drude) N. pantanense Brasil Strikis 2011.
Bactris gasipaes Kunth N. batesi Panamá McAlpine & Steyskal 1982.
Bignoniaceae
Crescentia cujete L. N. zadolicha Colômbia McAlpine & Steyskal 1982.
Bombacaceae
Quararibea guianensis Aubl. N. zadolicha Brasil Strikis et al. 2011.
77
Cactaceae
Selenicereus megalanthus N. batesi Colômbia Imbachi et al. 2012, Medina &
Britton & Rose Kondo 2012
Caricaceae
Caryocar brasiliense Cambess N. pendula Brasil Guimarães et al. 2003
Carica papaya L. N. batesi, N. nigrocaerulea Colômbia, Estados Unidos da McAlpine & Steyskal 1982,
América Yepes & Velez 1989.
Chrysobalanaceae
Couepia excelsa Ducke N. dimidiata Brasil Strikis et al. 2011.
Chrysobalanus icaco L. N. pendula Brasil Strikis et al. 2011.
Licania tomentosa Fritsch N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
78
Combretaceae
Buchenavia sp. Eichler N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Terminalia catappa L. N. certa, N. glaberrima, N. Brasil Uchôa & Nicácio 2010, Nicácio
inesperata, N. pendula, N. & Uchôa 2011.
zadolicha
Convolvulaceae
Operculina alata (Ham) N. certa, N. inesperata, N. Brasil Uchôa & Nicácio 2010, Nicácio
pendula, N. zadolicha & Uchôa 2011.
Operculina turpethum (Linn.) N. pendula Brasil Uchôa & Nicácio 2010.
Euphorbiaceae
Jatropha curcas L. N. bella, N. glaberrima, N. Brasil Dias et al. 2012
pendula, N. zadolicha
79
Manihot esculenta Crantz N. perezi Brasil, Estados Unidos da Romero & Ruppel 1973,
América, Porto Rico Brinholi et al. 1974, Boza &
Waddil 1978, Waddill 1978,
Waddill & Weems 1978,
Samways 1979, Peña &
Waddill 1982, Souza & Reis
1986, Lourenção et al 1996,
Arouca & Penteado-Dias 2012,
Gisloti & Prado 2011a, Gisloti
& Prado 2011b, Gisloti &
Prado 2012, Strikis et al. 2012.
Fabaceae
Andira cuyabensis Benthan N. pendula Brasil Uchôa & Nicácio 2010.
Dalbergia brasiliensis Vogel N. laura Brasil Strikis 2011.
80
Inga edulis Mart. N. bella, N. cornuphallus, N. Brasil, Colômbia Strikis et al. 2011, Strikis 2011,
dimidiata, N. distospinosa, N. Galeano-Olaya & Canal 2012.
glaberrima, N. plana, N.
spiculata, N. zadolicha
Inga feullei DC. N. glaberrima Peru McAlpine & Steyskal 1982.
Inga ingoides (Rich.) N. pseudozadolicha Brasil Strikis 2011.
Inga laurina (Sw.) N. inesperata, N. pendula, N. Brasil Strikis & Lerena 2009, Uchôa
pradoi, N. zadolicha & Nicácio 2010, Nicácio et al.
2011, Nicácio & Uchôa 2011.
Inga sp. Miller N. batesi Guatemala McAlpine & Steyskal 1982.
Inga thibaudiana DC. N. glaberrima Brasil Strikis et al. 2011.
Inga velutina Willd. N. bella, N. certa, N. Brasil Strikis et al. 2011.
glaberrrima, N. zadolicha
Inga vera Willd. N. certa, N. laura, N. pradoi Brasil McAlpine & Steyskal 1982,
Strikis 2011, Marsaro Júnior et
81
al. 2012.
Flacourtiaceae
Banara arguta Briq. N. pendula Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Gnetaceae
Gnetum sp. L. N. zadolicha Brasil Strikis et al. 2011.
Lauraceae
Persea americana Mill. N. batesi, N. zadolicha Estados Unidos da América McAlpine & Steyskal 1982,
Ahlmark & Steck 1997, Aluja
et al. 2004, Nicácio & Uchôa
2011.
82
Lecythidaceae
Eschweilera coriacea (DC.) N. mcalpiniei Brasil Strikis 2011.
Eschweilera atropetiolata Mori N. paramerolatus Brasil Strikis 2011.
Loganiaceae
Strychnos pseudoquina N. inesperata, N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
St.Hilarie & Uchôa 2011.
Loranthaceae
Psittacanthus acinarius (Mart.) N. bifida, N. certa, N. Brasil Strikis 2011, Uchôa et al. 2012.
inesperata, N. pantanense, N.
pendula, N. zadolicha
Psittacanthus plagiophyllus N. bifida, N. pendula, N. Brasil Caires et al. 2009.
Eichler zadolicha
83
Malpighiaceae
Byrsonima crassifolia (L.) N. bella, N. zadolicha Brasil Adaime et al. 2012.
Byrsonima orbignyana Jussieu N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Malpighia emarginata DC. N. bella, N. glaberrima, N. Brasil Araújo & Zucchi 2002, Silva et
pendula, N. pradoi, N. al. 1998, Strikis & Lerena
pseudozadolicha 2009, Strikis 2011, Strikis et al.
2011, Marsaro-Junior et al.
2012, Bittencourt et al. 2013.
Melastomataceae
Bellucia grossularioides (L.) N. pendula Brasil Strikis et al. 2011.
Mouriri elliptica Martius N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
84
Moraceae
Artocarpus communis Forst. N. glaberrima, N. zadolicha Brasil Strikis et al. 2011.
Artocarpus heterophyllus Lam. N. glaberrima Brasil Strikis et al. 2011.
Ficus insipida L. N. certa, N. glaberrima, N. Brasil Uchôa & Nicácio 2010, Nicácio
pendula, N. zadolicha & Uchôa 2011.
Sorocea saxicola Hassler N. bifida Brasil Uchôa & Nicacio 2010, Nicácio
& Uchôa 2011.
Myrtaceae
Campomanesia xanthocarpa N. pradoi Brasil Strikis & Lerena 2009,
Berg. Marsaro-Junior et al. 2012.
Eugenia pyriformis Cambess N. pradoi Brasil Marsaro-Junior et al. 2012.
Eugenia stipitata McVaugh N. bella, N. pseudozadolicha Brasil Strikis 2011, Bittencourt et al.
2013.
Eugenia tomentosa Berg N. pendula Brasil McAlpine & Steyskal 1982.
85
Eugenia uniflora L. N. zadolicha Brasil Nunes et al. 2012.
Psidium cattleianum Sabine N. inesperata, N. pendula, N. Brasil Strikis & Lerena 2009, Uchôa
pradoi, N. zadolicha & Nicácio 2010, Nicácio &
Uchôa 2011.
Psidium guajava L. N. amphora, N. certa, N. Brasil Araújo & Zucchi 2002, Souza-
glaberrrima, N. inesperata, N. Filho et al. 2009, Strikis &
pendula, N. pradoi, N. Lerena 2009, Strikis et al. 2011,
zadolicha Nicácio & Uchôa
2011,Galeano-Olaya & Canal
2012.
Psidium kennedyanum Morong N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Syzygium cumini (L.) N. pendula Brasil Nicácio & Uchôa 2011.
Syzygium jambos (L.) N. certa, N. glaberrima, N. Brasil Uchôa & Nicácio 2010, Nicácio
86
zadolicha & Uchôa 2011.
Olacaceae
Schoepfia sp. Schreb. N. inesperata, N. pendula, N. Brasil Uchôa & Nicácio 2010, Nicácio
zadolicha & Uchôa 2011.
Ximenia americana L. N. glaberrima, N. zadolicha Brasil Uchôa & Nicácio 2010.
Oxalidaceae
Averrhoa carambola L. N. glaberrima, N. pendula Brasil Araújo & Zucchi 2002, Strikis
et al. 2011.
Passifloraceae
Passiflora alata Curtis N. zadolicha Brasil Aguiar-Menezes et al. 2004.
Passiflora caerulea L. N. pradoi Brasil Marsaro-Junior et al. 2012.
Passiflora coccinea Aublet N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
87
& Uchôa 2011.
Passiflora edulis Sims N. batesi, N. ilheuense, N. Brasil, Colômbia Yepes & Velez 1989, Uchôa et
peltae, N. pradoi, N. zadolicha al. 2002, Strikis & Lerena
2009, Uchôa & Nicácio 2010,
Nicácio & Uchôa 2011, Strikis
2011, Strikis et al. 2011.
Passiflora elegans Mast N. pradoi Brasil Marsaro-Junior et al. 2012.
Passiflora flavicarpa Degener N. batesi Colômbia Wyckhuys et al. 2012.
Passiflora ligularis Juss. N. batesi Colômbia Wyckhuys et al. 2012.
Passiflora sp. L. N. bifida Brasil Guimarães et al. 2003.
Passiflora vitifolia Kunth N. orbata Colômbia Galeano-Olaya & Canal 2012.
Rhamnaceae
Ziziphus joazeiro Mart. N. pendula Brasil Araújo & Zucchi 2002.
88
Rosaceae
Eriobothrya japonica (Thunb.) N. bifida, N. certa, N. Brasil Strikis & Prado 2005, Souza-
cornuphallus, N. glaberrima, N. Filho et al. 2009, Strikis &
inesperata, N. pendula, N. Prado 2009, Strikis 2011.
zadolicha
Prunus avium L. N. pradoi Brasil Strikis & Lerena 2009.
Prunus persica (L.) N. certa, N. glaberrima, N. Brasil McAlpine & Steyskal 1982,
inesperata, N. pendula, N. Souza-Filho et al. 2009, Montes
zadolicha et al. 2010.
Rubiaceae
Alibertia edulis Rich N. glaberrima, N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Coffea arabica L. N. bella, N. bifida, N. certa, N. Brasil, Colômbia Portilla et al. 1994, Souza et al.
cornuphallus, N. delvechioi, N. 2005, Aguiar-Menezes et al.
89
glaberrima, N. inesperata, N. 2007, Aguiar-Menezes et al.
pendula, N. pradoi, N. 2008, Strikis & Prado 2008,
pseudopendula, N. zadolicha Camargos et al. 2011, Silva et
al. 2011, Strikis 2011, Strikis et
al. 2011, Montes et al. 2012.
Genipa americana L. N. glaberrima, N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Tocoyena formosa (Cham. & N. zadolicha Brasil Uchôa & Nicácio 2010, Nicácio
Schlechtd.) & Uchôa 2011.
Rutaceae
Citrofortunella microcarpa N. delvechioi Brasil Strikis 2011.
(Bunge.)
Citrus aurantifolia (Christm.) N. glaberrima Brasil Strikis et al. 2011.
Citrus jambhiri Lush N. inesperata, N. pendula, N. Brasil Uchôa & Nicácio 2010, Nicácio
90
zadolicha & Uchôa 2011.
Citrus reticulata Blanco N. certa, N. glaberrima, N. Brasil Gallardo et al. 2000, Araújo &
pendula, N. zadolicha Zucchi 2002, Lopes et al. 2008,
Souza et al. 2008
Citrus sinensis (L.) N. batesi, N. bella, N. bifida, N. Brasil, México McAlpine & Steyskal 1982,
certa, N. cornuphallus, N. Uchôa et al. 2003b, Silva et al.
glaberrima, N. inesperata, N. 2006, Strikis & Lerena 2009,
pendula, N. pradoi, N. Souza et al. 2008, Strikis &
zadolicha Prado 2008, Nicácio & Uchôa
2011, Strikis 2011, Strikis et al.
2011.
Fortunella sp. Swingle N. pendula Brasil Araújo & Zucchi 2002.
Metrodorea flavida Krause N. zadolicha Brasil Strikis et al. 2011.
Sapotaceae
91
Chrysophyllum soboliferum N. pendula, N. pseudobifida Brasil Uchôa & Nicácio 2010, Strikis
Rizz. 2011.
Manilkara zapota (L.) N. bella, N. dimidiata Brasil Bittencourt et al. 2013.
Pouteria caimito (Ruiz & Pav.). N. glaberrima, N. zadolicha Brasil Raga et al. 2003, Strikis et al.
2011, Fernandes et al. 2013.
Pouteria glomerata (Miq.) N. certa, N. zadolicha Brasil Uchôa & Nicácio 2010.
Pouteria macrophylla (Lam.) N. glaberrima, N. zadolicha Brasil Strikis et al. 2011
Pouteria ramiflora (Mart.) N. certa, N. glaberrima, N. Brasil Uchôa & Nicácio 2010, Nicácio
inesperata, N. pendula, N. & Uchôa 2011.
zadolicha
Pouteria sp. Aubl. N. nigrocaeruleae Brasil Strikis et al. 2011.
Pouteria torta (Mart.) N. certa, N. glaberrima, N. Brasil Uchôa & Nicácio 2010, Nicácio
inesperata, N. pendula, N. & Uchôa 2011.
zadolicha
92
Solanaceae
Duckeodendron cestroides N. zadolicha Brasil Strikis et al. 2011.
Kuhlm.
Capsicum annuum L. N. amphora, N. major, N. Colômbia Steyskal 1978, McAlpine &
pseudopendula, N. turdiphallus Steyskal 1982, Strikis 2011,
Galeano-Olaya & Canal 2012.
Solanum gilo Raddi N. zadolicha Brasil Strikis et al. 2011.
Solanum quitoense Lam. N. batesi Colômbia Yepes & Velez 1989.
Solanum sisymbriifolium Lam. N. inesperata Brasil Uchôa & Nicácio 2010, Nicácio
& Uchôa 2011.
Physalis angulata L. N. certa, N. inesperata, N. Brasil Uchôa & Nicácio 2010, Nicácio
pantanense, N. zadolicha & Uchôa 2011, Strikis 2011.
Ulmaceae
Ampelocera edentula Kuhlm N. cornuphallus Brasil Strikis 2011.
93
Celtis iguanaea (Jacq.) N. laura, N. pradoi Brasil Strikis 2011.
Verbenaceae
Cytharexyllum myrianthum N. bella, N. cornuphallus Brasil Strikis & Prado 2008, Strikis
Chamiáo 2011.
Vitaceae
Cissus sp. L. N. angusta Colômbia Galeano-Olaya & Canal 2012.
94
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CAPÍTULO 3
Neosilba McAlpine (Diptera, Lonchaeidae) Diversity on Cultivated Host Plants from
Brazilian Cerrado and Atlantic Tropical Forest Biomes
ǂ L GISLOTI, AP DO PRADO
Laboratório de Entomologia, Departamento de Biologia Animal, Instituto de Biologia,
Universidade Estadual de Campinas, Caixa Postal 6109, 13.083-862, Campinas-SP, Brazil.
Neosilba Diversity on Cerrado and Tropical Forest of Brazil
ABSTRACT - Thirty-five cultivated native fruit species (19 orders and 12 families) from
Brazil were sampled in farm fruit from two municipalities located in the state of São Paulo
(January 2010 to March 2012) in a study to evaluate the diversity of Neosilba flies. Thirty- one species of plants were infested, while five were not. Some aspects of the biology and patterns of species diversity, abundance, infestation rates, puparia viability and the interactions among species of frugivorous flies and their host plants were quantified. Seven species of Neosilba were reared from 31 host fruits: Neosilba bella Strikis & Prado (4 hosts), Neosilba certa (Walker) (4 hosts), Neosilba glaberrima (Wiedemann) (5 hosts),
Neosilba inesperata Strikis & Prado (6 hosts) Neosilba pendula (Bezzi) (15 hosts),
Neosilba pradoi Strikis & Lerena (8 hosts) and Neosilba zadolicha McAlpine (26 hosts). N. zadolicha was the specie most abundant (340 individuals), and most polyphagous occurring 107
in 26 host fruits. The association between the frugivorous flies and the fruit species is discussed.
Keywords – lance flies, frugivorous flies, biodiversity.
Introduction
The genus Neosilba (Diptera, Lonchaeidae) is comprised of Neotropical fly species whose larvae are hosted by many species of commercially important fruits (Araújo &
Zucchi 2002, Strikis & Prado 2005, Bittencourt et al 2006). The genus is restricted to the
Neotropical region being known from Caribe, Mexico and Colombia to Brazil. Forty species were described, and at least more 60 species wait for description (McAlpine &
Steyskal 1982, Strikis 2011, Galeano-Olaya & Canal 2012).
Although of the economic importance of some species of Neosilba that occur as pests on fruit and vegetables in several countries, such as Colombia (Steyskal 1978,
Peñaranda et al 1986), Peru (Korytkowski & Ojeda 1971), and Costa Rica (Sánchez et al
1991) the knowledge of these dipterous is still incipient. Also in Brazil, research on the damage in fruits and vegetables by Neosilba is very scarce (Uchôa et al 2002).
Neosilba species obtains food resource in 107 plant species distributed in 35 plant families: Anacardiaceae, Annonaceae, Apocynaceae, Arecaceae, Bignoniaceae,
Bombacaceae, Cactaceae, Caricaceae, Chrysobalanaceae, Combretaceae, Convolvulaceae ,
Euphorbiaceae, Fabaceae, Flacourtiaceae, Gnetaceae, Lauraceae, Lecythidaceae,
Loganiaceae, Loranthaceae, Malpighiaceae, Melastomataceae, Moraceae, Myrtaceae,
Olacaceae, Oxalidaceae, Passifloraceae, Rhamnaceae, Rosaceae, Rubiaceae, Rutaceae,
108
Sapotaceae, Solanaceae, Ulmaceae, Verbenaceae e Vitaceae, confirming the relevance of this genus to tropical fruit production. (McAlpine & Steyskal 1982, Raga et al 1996, Araújo
& Zucchi 2002, Uchôa et al 2002, Souza et al 2005, Bittencourt et al 2006, Strikis & Prado
2005, Strikis & Prado 2009, Uchôa & Nicácio 2010, Nicácio & Uchôa 2011, Souza et al
2012, Uchôa et al 2012).
Studies on the biology and ecology of economically important species have contributed for the management and control of agricultural pests (Carey 1993,
Papadopoulos et al 2002). Integrated pest management has been more difficult by the lack of basic studies on taxonomy, biology and ecology. In Brazil, records on the genera of
Lonchaeidae associated with fruits are scarce. Regional surveys are very important because they can provide basic information for managing insect pest populations and their natural enemies (Uchôa et al 2002).
The aim of this study was to provide an insight for the theoretical approaches to the life history and dynamic distribution of Neosilba species in different host plants, in addition to baseline information on emergence percentage, rate infestation and puparia viability for the development of prediction models of population growth and control.
Material and Methods
Collecting Host Fruits. Fruits from 35 species (Table 1) were collected, from
January 2010 to March 2012, in the fruit farm of the municipalities of Campina do Monte
Alegre (23º 53’ 37’’ S, 48º 51’ 06’’ W, 612m) (site 1), and Paraibuna (23º 27’ 94’’ S, 45º
42’ 88’’ W, 647m) (site 2). These cities are located in an important fruit producing area in 109
the state of São Paulo, Brazil. Collecting fruits, and fruit fly and lonchaeid rearings were done according to Uchôa & Zucchi (1999).
Fly Identification. The adults were identified in the Departamento de Biologia
Animal, Instituto de Biologia, Universidade Estadual de Campinas (Unicamp), Campinas-
SP, by the first author. Neosilba species were identified using keys and original descriptions (Korytkowski & Ojeda 1971, McAlpine & Steyskal 1982). Only males were used since traditional taxonomy is based on analysis of male genitalia (McAlpine &
Steyskal 1982). Plant species were identified by botanists at the Departamento de Botânica,
Universidade Estadual de Campinas (Unicamp) in Campinas. Voucher specimens of the insects were deposited at Coleção Zoológica (ZUEC), Universidade Estadual de Campinas
(Unicamp) in Campinas, SP.
Quantitative variables, infestation indices and viability of the puparia. Two quantitative variables were evaluated: number of emerged flies, and number of puparia. The fruit infestation levels were measured by 2 indexes: number of puparia / fruit, and number of puparia / mass (g) of fruit. The viability of the puparia was calculated by the equation:
%V = No. of ЕА/ PUP) X 100, where: %V = Percent of viability), No. of EA = number of emerged adults, and PUP= Total number of puparia; and the quotient was multiplied by
100.
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Results and Discussion
Host Fruits. Seven Neosilba species were reared: N. bella Strikis & Prado, N. certa
(Walker), N.glaberrima (Wiedemann), N. inesperata Strikis & Prado, N. pendula (Bezzi),
N. pradoi Strikis & Lerena and N. zadolicha McAlpine. A total of 21,496 fruit were collected weighing 335,713g, from which developed 6,910 puparia.
The highest number of individuals belong to N. zadolicha (1.289) (Table 1) which was the specie most abundant (340 individuals), and most generalist occurring in 26 host fruits. N. bella was less abundant (51 individuals).
Out of 36 fruit species sampled in 12 families, were obtained frugivorous flies from
31 plant species (Table 1). Only in four fruit species no adult Neosilba flies were collected:
Campomanesia sessiflora (Berg.), Eugenia matosii Legr., Myrciaria jaboticaba Baill.
(Myrtaceae), and Pouteria macrophylla (Lam.) (Sapotaceae) (Table 1).
Uchôa et al (2002) found Neosilba species hosted in C. sessiflora and M. jaboticaba in the cerrado of Brazil (Uchôa et al 2002). Strikis et al (2011) found N. glaberrima and N. zadolicha hosted in P. macrophylla, but in this paper, we did not record this association.
Likewise, Uchôa & Zucchi (1999) and Uchôa et al (2002) recorded Neosilba spp. in C. sessiflora, however, we did not have this record in this work.
N. bella (51 individuals in total) was the only species that occurred in only one site
(site 2), and this species had the lowest number of host fruit (4 species) from two plant families: Byrsonima crassifolia (L.), Malpighia emarginata DC. (Malphigiaceae), Eugenia stipitata McVaugh, and Eugenia uniflora L. (Myrtaceae).
Adaime et al (2012), as in this work, recorded N. bella in B. crassifolia. Bittencourt et al (2013) found N. bella hosted in E. stipitata in northeastern Brazil. 111
According Bittencourt et al (2013) N. bella has a wide geographical distribution in
Brazil ranging from Mata Atlântica, Amazon rain forest, and Cerrado. Its plasticity in occupying such different biomes and in attacking different hosts makes this species a candidate in becoming an important pest, once it is found in environments occupied by crops plantation, specially coffee. However, in this study, this species was the one with the lowest number of host fruit.
Others author’s recorded N. bella hosted in: Citrus sinensis (L.) (Rutaceae),
Cytharexyllum myrianthum Chamiáo (Verbenaceae) (Strikis & Prado 2008), Coffea arabica
L. (Rubiaceae), Inga edulis Mart., Inga velutina Willd. (Fabaceae) (Strikis et al. 2011),
Jatropha curcas L. (Euphorbiaceae) (Dias et al 2012), and Manilkara zapota (L.)
(Sapotaceae) (Bittencourt et al 2013).
N. certa (90 individuals) also presented four host fruit occurring in three plant family: Spondias mombim L. (Anacardiaceae), Inga vera Willd. (Fabaceae), Psidium cattleianum Sabine, and Psidium guajava L. (Myrtaceae). McAlpine (1982) found N. certa in I. vera and Souza-Filho et al 2009 recorded this specie in P. guajava orchards.
N. certa has already been registered in the host fruits: Citrus sinensis (L.) (Souza et al 2008), Citrus reticulata Blanco (Rutaceae) (Souza et al 2008), C. arabica (Rubiacae)
(Souza et al 2005, Strikis & Prado 2008, Aguiar-Menezes et al 2008), Duguetia furfuracea
(A. St.-Hil.) (Annonaceae), Ficus insipida Willd. (Moraceae), Operculina alata (Ham.)
(Convolvulaceae), Physalis angulata L. (Solanaceae), Pouteria glomerata (Miq.), Pouteria ramiflora (Mart.) (Sapotaceae), Pouteria torta (Mart.), Syzygium jambos (L.) (Myrtaceae),
Terminalia catappa L. (Uchôa & Nicácio 2010, Nicácio & Uchôa 2011), Eriobotrya japonica (Thunb.) (Rosaceae), P. persica, (Souza-Filho et al 2009, Strikis & Prado 2009),
(McAlpine & Steyskal1982), I. velutina (Fabaceae) (Strikis et al 2011), J. curcas 112
(Euphorbiaceae) (Dias et al 2012), and Psittacanthus acinarius (Mart.) (Loranthaceae)
(Uchôa et al 2012).
N. glaberrima (109 individuals) hosted five fruit species from five families: Rollinia sylvatica (A. St.-Hil.) (Annonaceae), Bactris gasipaes Kunth (Arecaceae), Selenicereus setaceus (Rizz) (Cactaceae), M. emarginata (Malphigiaceae), Pouteria caimito (Ruiz &
Pav.) (Sapotaceae). Strikis et al (2011) recorded N. glaberrima in M. emarginata, and
Raga et al (2003), Strikis et al (2011) and Fernandes et al (2013) found this specie in P. caimito, as we found in this work.
N. glaberrima has already been registered hosted in: Alibertia edulis Rich.
(Rubiaceae), Annona crassiflora L. (Annonaceae), D. furfuracea, F. insipida, Genipa americana L.(Rubiaceae), P. ramiflora, P. torta, Spondia dulcis Forst. (Anacardiaceae), S. jambos, T. catappa, Ximenia americana L. (Olacaceae) (Uchôa & Nicácio 2010, Nicácio &
Uchôa 2011), Annona muricata L. (Annonaceae), Artocarpus communis Forst., Artocarpus heterophyllus Lam., (Moraceae), A. carambola, Capsicum odoriferum Vell. (Solanaceae),
C. sinensis, Citrus aurantifolia (Christm.) (Rutaceae), I. edulis, Inga thibaudiana DC., I. velutina (Fabaceae), P. macrophylla, P. guajava, (Strikis et al. 2011), C. reticulata (Lopes et al 2008), J. curcas (Dias et al 2012), and Prunus persica (L.) (Rosaceae) (Souza-Filho et al 2009).
N. inesperata (287 individuals) was found in six host fruit occurring in three families: M. emarginata, Eugenia dysenterica (DC.) (Malpighiaceae), E. uniflora, P. cattleianum, P. guajava (Myrtaceae), and Rubus urticaefolius Poir (Rosaceae). All hosts found in this paper for N. inesperata are novel excepted P. guajava (Nicácio & Uchôa
2011).
113
Others author’s recorded N. inesperata hosted in: Citrus jambhiri Lush. C. sinensis
(Rutaceae), D. furfuracea, Inga laurina (Swartz) (Fabaceae), O. alata, P. angulata, P. ramiflora, P. torta, Schoepfia sp. (Olacaceae), Solanum sisymbriifolium Lam. (Solanaceae),
Strychnos pseudoquina St.Hilarie (Loganiaceae), T. catappa (Uchôa & Nicácio 2010,
Nicácio & Uchôa 2011), P. acinarius (Uchôa et al 2012), C. arabica (Camargos et al
2011, Montes et al 2012), and E. japonica (Strikis & Prado 2009).
N. pendula (340 individuals) hosted in 15 host fruit from seven plant family: S. mombim, Spondias purpurea L. (Anacardiaceae), Caryocar brasiliense Camb.
(Caryocaceae), I. vera (Fabaceae), M. emarginata (Malphigiaceae), Eugenia brasiliensis
Lam., E. dysenterica, Eugenia involucrata DC., Eugenia neonitida Sobral, E. stipitata, P. cattleianum, P. guajava, Psidium guineensis Swartz. (Myrtaceae), Ziziphus joazeiro Mart.
(Rhamnaceae), and R. urticaefolius (Rosaceae).
S. purpurea (Araújo & Zucchi 2002, Nicácio & Uchôa 2011). C. brasiliense
(Guimarães et al. 2003). M. emarginata (Araújo & Zucchi 2002, Strikis et al. 2011). P. cattleianum (Uchôa & Nicácio 2010, Nicácio & Uchôa 2011), P. guajava (Araújo &
Zucchi 2002, Souza-Filho et al. 2009, Nicácio & Uchôa 2011), and Ziziphus joazeiro Mart.
(Rhamnaceae) (Araújo & Zucchi 2002) have been reported as hosts of N. pendula.
N. pendula has already been registered hosted in: Anacardium humile St. Hil.,
Anacardium othonianum Rizz. (Anacardiaceae), Andira cuyabensis Benthan (Fabaceae),
Banara arguta Briquel (Flacourtiaceae), Chrysophyllum soboliferum Rizz. (Sapotaceae), C. jambhiri, F. insipida, I. laurina, O. alata, Operculina turpethum (Linn.) (Convolvulaceae),
P. ramiflora, P. torta, Schoepfia sp., Syzygium cumini (L.) (Myrtaceae), T. catappa (Uchôa
& Nicácio 2010, Nicácio & Uchôa 2011), A. carambola, C. reticulata, Fortunella sp.
(Rutaceae), Bellucia grossularioides (L.) (Melastomataceae), Chrysobalanus icaco L. 114
(Chrysobalanaceae) (Strikis et al. 2011), C. sinensis (Souza et al 2008), C. arabica
(Souza et al 2005, Strikis & Prado 2008, Camargos et al 2011), Eugenia tomentosa Berg.
(Myrtaceae) (McAlpine & Steyskal 1982), P. persica (Souza-Filho et al 2009), P. acinarius (Uchôa et al 2012), and J. curcas (Dias et al 2012).
N. pradoi (124 individual) hosted in eight fruits from three families: I. vera
(Fabaceae), M. emarginata (Maphigiaceae), Campomanesia aurea Berg., Campomanesia guazumaefolia (Camb.), Campomanesia phaea (Berg.) E. dysenterica, Eugenia pyriformis
Cambess and P. cattleianum (Myrtaceae).
I. vera (Marsaro-Junior et al 2012), M. emarginata, E. pyriformis, P. cattleianum
(Strikis & Lerena 2009, Marsaro-Junior et al 2012) have been reported as hosts of N. pradoi.
N. pradoi has already been registered hosted in: Campomanesia xanthocarpa Berg.,
Celtis iguanaea Cannabaceae (Jacq.), C. sinensis, I. laurina, Passiflora caerulea L.,
Passiflora edulis Sims., Passiflora elegans Mast. (Passifloraceae), Prunus avium L.
(Rosaceae), P. guajava (Strikis & Lerena 2009, Marsaro-Junior et al. 2012), and C. arabica
(Montes et al 2012).
N. zadolicha (1289 individuals) occurred in 25 fruit species from seven plant family: S. mombim, S. purpurea, Spondias tuberosa Arruda, Spondias venulosa Mart.
(Anacardiaceae), Hancornia speciosa Gomes (Apocynaceae), B. gasipaes (Arecaceae), C. brasiliense (Caryocaceae), I. vera (Fabaceae), B. crassifolia, M. emarginata
(Malpighiaceae), Acca sellowiana (Berg.), C. guazumaefolia, C. phaea, E. brasiliensis, E. dysenterica, E. involucrata, E. neonitida, Eugenia pitanga (Berg.), E. pyriformis, E. uniflora, Myrciaria dubia McVaugh, P. cattleianum, P. guajava, P. guineensis
(Myrtaceae), and P. caimito (Sapotaceae). 115
S. purpurea, H. speciosa, P. cattleianum (Uchôa & Nicácio 2010, Nicácio & Uchôa
2011), B. crassifolia (Adaime et al. 2012), E. uniflora (Nunes et al. 2012), P. guajava
(Souza-Filho et al 2009, Strikis et al 2011) and P. caimito (Fernandes et al 2013) have been reported as hosts of N. zadolicha.
N. zadolicha has already been registered in hosts: A. edulis, A. humile, A. othonianum, A. crassiflora, Buchenavia sp. (Combretaceae), Byrsonima orbignyana Jussieu
(Malphigiaceae), C. jambhiri, D. furfuracea, F. insípida, G. americana, I. laurina, Licania tomentosa Fritsch (Chrysobalanaceae), M. indica, Mouriri elliptica Martius
(Melastomataceae), O. alata, Passiflora coccinea Aublet (Passifloraceae), P. edulis, P. americana, P. angulata, P. glomerata, P. ramiflora, P. torta, Psidium kennedyanum
Morong (Myrtaceae), Schoepfia sp., S. dulcis, S. pseudoquina, S. jambos, T. catappa,
Tocoyena formosa (Cham. & Schlechtd.) (Rubiaceae), X. americana (Uchôa & Nicácio
2010, Nicácio & Uchôa 2011); A. muricata, A. communis, Duckeodendron cestroides
Kuhlm. (Solanaceae), Gnetum sp. (Gnetaceae), I. edulis, I. velutina, Metrodorea flavida
Krause (Rutaceae), P. macrophylla, Quararibea guianensis Aubl. (Bombacaceae), Rollinia mucosa Baill. (Annonaceae), Solanum gilo (Raddi) (Solanaceae) (Strikis et al 2011); C. reticulata (Lopes et al 2008, Souza et al 2008); C. sinensis (Souza et al 2008, Camargos et al 2011, Strikis et al 2011, Nicácio & Uchôa 2011, Montes et al 2012); C. arabica (Strikis
& Prado 2008); E. japonica (Souza-Filho et al 2009, Strikis & Prado 2009); J. curcas
(Dias et al 2012); P. persica (McAlpine & Steyskal 1982, Souza-Filho et al 2009); P. acinarius (Uchôa et al 2012); Passiflora alata Curtis (Passifloraceae) (Aguiar-Menezes et al 2004) and Crescentia cujete L. (Bignoniaceae) (McAlpine & Steyskal 1982).
The plant species M. emarginata was the host with the largest Neosilba diversity: six species - all species of Neosilba found in this work - excepted N. certa. Strikis et al 116
(2011) also found no association record between N. certa and Malpighia sp. when studied frugivorous flies in the Amazon rainforest. P. cattleianum was the host with the largest number of Neosilba specimens emerged (398 individuals - total of males and females).
Infestation Indices and Puparia Viability.
The mean of infestation was: 0.46 (puparia / total fruit) and 0,05 (puparia / fruit weight in grams). The mean of viability of the puparia was 67,23%.
C. brasiliensis was the plant species that showed the highest infestation indices based on the number of fruit (2.30 puparia / fruit). B. crassifolia was the plant species that presented the highest infestation indices based on mass of fruit (0.23 puparia/ mass in grams).
Adaime et al (2012) found lower infestation indices to that found in our work: 0.06 puparia/fruit to N. bella hosted in B. crassifolia. Aguiar-Menezes et al (2004) reported to N. zadolicha hosted in P. alata infestation index (2.1 puparia/fruit) higher than the all indices found in our work.
Souza et al (2012) reported infestation indexes to Neosilba species hosted in P. guajava (0.03 puparia/ g of fruit) similar than mass fruit infestation that we found in our work (0.01 puparia/ g of fruit). However to Neosilba species hosted in Z. joazeiro these authors found higher infestation indices (3.28 puparia/ g of fruit) than reported in our work
(0.09 puparia/ g of fruit).
Araújo & Zucchi (2002) found similar infestation indexes to N. pendula hosted in S. purpurea, M. emarginata and in P. guajava (0.06, 0.1 and 0.1 puparia/ g of fruit, respectively), however to Z. joazeiro (0.2 puparia/ g of fruit) these authors found infestation indices higher than reported in this paper (0.09). 117
New Records
In this work were reported following new records: N. bella in : E. uniflora.
N. certa in: P. cattleianum and S. mombim.
N. glaberrima in: B. gasipaes, R. sylvatica, and S. setaceous.
N. inesperata in: E. dysenterica, E. uniflora, M. emarginata, P. cattleianum, and R. urticaefolius.
N. pendula in: E. brasiliensis, E. dysenterica, E. involucrate, E. neonitida, E. stipitata, I. vera, P. guineensis, R. urticaefolius, and S. mombim.
N. pradoi in: C. aurea, C. guazumaefolia, C. phaea, and E. dysenterica.
N. zadolicha in: A. sellowiana, B. gasipaes, C. brasiliense, C. guazumaefolia, C. phaea, E. brasiliensis, E. dysenterica, E. involucrata, E. neonitida, E. pitanga, E. pyriformis, , I. vera, M. emarginata, M. dubia, P. guineensis, P. alata, S. mombim, S. tuberose, and S. venulosa.
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Table 1. Indices of infestation and viability of pupae of Neosilba species in 35 cultivated hosts from Cerrado and Tropical Forest of
Brazil (January/2010 to March/2012).
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Anacardiaceae
Spondias mombim L. 2 620 11160 384 289 - 48 - - 22 - 83 0.62 0.03 75.26
Spondias purpurea L. 2 460 3220 189 102 - - - - 17 - 50 0.41 0.06 53.97
Spondias tuberosa 2 1280 19200 412 302 ------127 0.32 0.02 73.30
Arruda
Spondias venulosa Mart. 1 367 5505 113 81 ------54 0.31 0.02 71.68
Annonaceae
Rollinia sylvatica (A. 1 67 5360 64 50 - - 6 - - - 12 0.96 0.01 78.13
St.-Hil.)
124
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Apocynaceae
Hancornia speciosa 1 543 10860 184 101 ------51 0.34 0.02 54.89
Gomes
Arecaceae
Bactris gasipaes Kunth 1 521 26050 101 78 - - 12 - - - 32 0.19 0.00 77.23
Cactaceae
Selenicereus setaceus 2 62 7740 37 25 - - 12 - - - - 0.60 0.00 67.57
(Rizz)
Caryocaraceae
Caryocar brasiliense 2 82 7380 189 134 - - - - 42 - 12 2.30 0.03 70.90
Camb.
Fabaceae
Inga vera Willd. 1 512 7680 201 171 - 10 - - 14 8 53 0.39 0.03 85.07
125
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Malphigiaceae
Byrsonima crassifolia 2 843 1011 234 189 30 - - - - - 71 0.28 0.23 80.77
(L.)
Malpighia emarginata 2 1384 4152 389 302 11 - 37 53 21 12 33 0.28 0.09 77.63
DC.
Myrtaceae
Acca sellowiana (Berg.) 2 100 5178 42 21 ------7 0.42 0.01 50.00
Campomanesia aurea 2 189 4725 47 19 - - - - - 8 - 0.25 0.01 40.43
Berg.
Campomanesia 1 112 2688 45 21 - - - - - 3 6 0.40 0.02 46.67 guazumaefolia (Camb.)
Campomanesia phaea 1 229 8450 79 54 - - - - - 9 12 0.34 0.01 68.35
(Berg.)
126
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Campomanesia 1 340 6667 ------sessiflora (Berg.)
Eugenia brasiliensis 2 670 1150 231 120 - - - - 12 - 31 0.34 0.20 51.95
Lam.
Eugenia dysenterica 2 578 8670 289 201 - - - 43 20 11 40 0.50 0.03 69.55
(DC.)
Eugenia involucrata 2 539 1340 211 160 - - - - 12 - 61 0.39 0.16 75.83
DC.
Eugenia matosii Legr. 1 501 8020 ------
Eugenia neonitida 1 1279 3837 467 321 - - - - 27 - 102 0.37 0.12 68.74
Sobral
Eugenia pitanga (Berg.) 1 1021 3063 312 204 ------81 0.31 0.10 65.38
Eugenia pyriformis 2 1342 8910 380 289 - - - - - 46 101 0.28 0.04 76.05
127
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Cambess
Eugenia stipitata 2 234 3880 98 69 2 - - - 31 - - 0.42 0.00 70.41
McVaugh
Eugenia uniflora L. 2 2295 4704 467 332 8 - - 119 - - 22 0.20 0.10 71.09
Myrciaria dubia 2 829 9119 231 167 ------61 0.28 0.03 72.29
McVaugh
Myrciaria jaboticaba 2 801 8604 ------
Baill.
Psidium cattleianum 1 876 5310 465 398 22 34 17 27 101 0.53 0.09 85.59
Sabine
Psidium guajava L. 1 335 33890 189 89 - 10 - 5 4 - 21 0.56 0.01 47.09
Psidium guineensis 2 378 60480 168 102 - - - - 2 - 41 0.44 0.00 60.71
Swartz.
128
Neosilba male species Indices/Rate
Plant Taxa
b
c
a
ruits
nfestation
f
erta
i
nfestation viability (%)
of fruits (g)
i
N. bellaN.
N. c N.
ruit
N. pradoiN.
Pupae (n)
f
N. pendulaN.
Nº of
N. zadolichaN.
N. inesperataN.
N. glaberrimaN.
Emerged adults
Collecting sites
Fruit
Mass
uparia
P Total Rhamnaceae
Ziziphus joazeiro Mart. 1 521 2605 232 171 - - - - 68 - - 0.45 0.09 73.71
Rosaceae
Rubus urticaefolius Poir 1 594 3564 281 188 - - - 33 31 - - 0.47 0.08 66.90
Sapotaceae
Pouteria caimito (Ruiz 2 549 16470 179 102 - - 42 - - - 24 0.33 0.01 56.98
& Pav.)
Pouteria macrophylla 2 443 15071 ------
(Lam.)
Total - 21496 6910 4852 51 90 109 287 340 124 1.289 - - -
Mean ------0.46 0.05 67.23
a- site 1: Campina do Monte Alegre.; site 2: Paraibúna.. (São Paulo, Brazil). b- number of puparia/number of fruit c- number of puparia/ weight of fruits (in grams)
129
CONCLUSÕES
1. Houve um aumento na produção científica a respeito de Lonchaeidae a partir da
segunda metade do séc. XX.
2. A maioria dos trabalhos publicados a respeito da família Lonchaeidae é sobre o
gênero Neosilba.
3. O crescimento da produção científica sobre Lonchaeidae está associado ao
aumento das publicações sobre Neosilba como praga de frutíferas de interesse
agrícola.
4. A escassez de taxonomistas dificulta os estudos com Lonchaeidae.
5. A compilação de dados e a revisão da literatura pretendeu encorajar novos
pesquisadores a iniciarem estudos com esse taxón.
6. Mais de ¼ das espécies de Neosilba descritas possuem hospedeiro
desconhecido.
7. A falta de taxonomistas especializados em Neosilba é bastante evidente.
8. Inúmeras publicações a respeito de moscas-de-frutas identificam indivíduos de
Neosilba somente em nível de gênero.
9. Não foi possível elaborar uma chave para identificação de espécies de Neosilba,
devido à inconsistência das descrições e à ausência dos holótipos em museu de
referência.
10. Espécies de Neosilba colonizaram 86% dos frutos nativos cultivados.
11. A investigação de espécies de Neosilba em frutos cultivados nativos resultou em
um grande número de associações inéditas entre essas moscas e os respectivos
frutos hospedeiros.
130
12. Há a necessidade de se revisar taxonomicamente o gênero Neosilba, além de se
desenvolver novas ferramentas taxonômicas que possibilitem a identificação das
espécies deste gênero.
131