Os Cicindelídeos (COLEOPTERA, CICINDELIDAE): TAXONOMIA, MORFOLOGIA, ONTOGENIA, ECOLOGIA E EVOLUÇÃO

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Os Cicindelídeos (COLEOPTERA, CICINDELIDAE): TAXONOMIA, MORFOLOGIA, ONTOGENIA, ECOLOGIA E EVOLUÇÃO Professor Germano da Fonseca Sacarrão, Museu Bocage, Lisboa, 1994, pp. 233-285. os CICINDELíDEOS (COLEOPTERA, CICINDELIDAE): TAXONOMIA, MORFOLOGIA, ONTOGENIA, ECOLOGIA E EVOLUÇÃO. por ARTUR R. M. SERRANO Departamento de Zoologia e Antropologia, Faculdade de Ciências de Lisboa, Edifício C2, 3.° piso, Campo Grande, 1700 Lisboa - Portugal. «Devemos ter sempre presente o facto de os organismos serem criaturas históricas, com um longo passado de transforma­ ções resultantes de complexas relações entre os seres vivos e os factores do ambiente». G.F. SACARRÃO (1965), ln «A Adaptação e a Invenção do Futuro». ABSTRACT The tiger heetles (Coleoptera, Cicindelidae); taxonomy, morphology, ontogeny, ecology and evolution The family of tiger beetles (Coleoptera, Cicindelidae) is an ideal group for studies in areas as taxonomy, morphology, ontogeny, physiology, eeology, behavior, evolution and biogeography. ln a time in which biodiversity is in great danger, we try in this work to present an update of our knowledge in the previously mentioned areas and therefore call attention to the importance of this group of inseets. To com pose ali these data more than 300 references were consulted fram several hundreds pertinent works. 234 Ar/ur R. M. Serrano RESUMO A família dos coleópteros cincidelídeos constitui um grupo ideal para estudos nas áreas de taxonomia, morfologia, ontogenia, fisiologia. ecologia. comportamento, evolução e biogeografia. Numa altura em que a biodiversidade se encontra em grande perigo, tentamos neste artigo apresentar os conhecimentos mais recentes naquelas áreas, chamando portanto a atenção para a importância deste grupo de insectos. Para a elaboração destes dados foram consultadas mais de 300 obras sobre o assunto. INTRODUÇÃO Desde os tempos de Lineu que foram publicados numerosos trabalhos sobre taxonomia, ou pequenas notas e estudos sobre comportamento, zoogeografia, anatomia, biologia, etc. dos coleópteros cicindelídeos, mas maioritariamente sobre os imagos. Pearson em 1988 publicou um artigo monográfico e generalista na célebre revista Annual Review of En/omology intitulado «Biology of Ti­ ger Beetles» . O objectivo deste trabalho é fazer uma actualização daquele artigo e apresentar uma súmula de conhecimentos sobre taxonomia, morfologia, fisiologia, ontogenia, citogenética, ecologia e evolução da família Cicindelidae. Salvo apontamento em contrário, as considerações de índole geral referidas neste trabalho ao género Cicindela Linnaeus, ainda no sentido lato, tiveram apenas o propósito de não fugir à regra daquilo que tem sido feito anteriormente por outros autores (e.g., Willis, 1967, Freitag, 1979, Leffler, 1979, Murray, 1980, Pearson, 1988). TAXONOMIA A família Cicindelidae está incluída nos Caraboidea, uma das superfanulias da subordem Adephaga dos Insec/a Coleop/era. O nome Cicindela apareceu pela primeira vez na 10.' edição do Sys/ema Na/urae (Linnaeus, 1758), englobando 7 espécies. Só mais tarde Latreille (1804), propôs o nome «Cicindêletae» com o mesmo significado do da farmlia (Cicindelidae). Desde essa época, a polémica sobre o nível taxonómico destes coleópteros tem-se mantido (família, subfamília, supertribo ou tribo dos Carabidae) (Ball, 1979 a). Cicindelídeos (Coleoptera, Cicindelidae) 235 Nos catálogos de Heyden et aI. (1891) e de Winkler (1924) o grupo foi considerado uma família independente. Horn (1915 e 1926) incluiu­ -o contudo, nos carabídeos, como uma subfamília. Fuente (1927), ao estudar os cicindelídeos e carabídeos da Península Ibérica, tratou-os como famílias independentes. Jeannel (1941), baseado nas estruturas esternais, no órgão copulador (genitália) e na morfologia das larvas, juntou-se aos defensores da família independente. O mesmo critério seguiu Magistreti (1965) para a fauna de Itália. Thie1e (1977) defendeu, pelo contrário, a sua inclusão nos carabídeos. Reportando-nos a vários estudos, como, por exemplo, padrões estruturais dos élitros (Shelford, 1917), microscultura cuticular (Stegemann, 1930), comparações imunu1ógicas (Basford et aI., 1968), órgãos tibiais de limpeza (Hlavac, 1971, Regenfuss, 1975), estrutura dos olhos (Kuster, 1979), das asas metatorácicas (Ward, 1979), das faneras tarsais (Stork, 1980) e das coxas (Nichols, 1985) e ainda mecanismos de determinação sexual (Serrano & Yadav, 1984), poderemos verificar a defesa de quaisquer das duas posições (vide ainda Leffler, 1979). A maioria dos especialistas do grupo têm contudo, adoptado o critério da família. Foi, por exemplo, o que ocorreu no mais recente catálogo sobre estes coleópteros (Wiesner, 1992). Horn (1926) além de ter reconhecido aproximadamente 1300 espécies, assinalou ainda centenas de formas, cuja validade taxonómica, face aos modernos conceitos da biologia populacional, tem sido gradualmente posta em causa em trabalhos de revisão. Contudo, os trabalhos de Horn (1915, 1926) tentaram explanar dentro do possível uma relação fi10genética com todos os cicindelídeos do mundo. Nessas obras foram reconhecidos 35 géneros (em 1929 Horn descreveu mais um: Pronyssiformia). O mais numeroso, o género Cicindela, foi dividido em grupos mais ou menos naturais, baseados nos padrões quetotáxicos dos adultos (não foi reconhecida a categoria subgenérica para estes grupos). Vários estudos posteriores aos trabalhos de Horn (op. cit.) vieram clarificar a taxonomia de complexos e grupos de espécies desta família. Poderemos referir para a fauna paleárctica, por exemplo, os trabalhos de Mandl (1935b, 1936, 1981 e 1982), para a neárctica Freitag (1965), WilIis (1967), Gaumer (1977), Leffler (1979), Murray (1980) e mais recentemente Spanton (1988). Freitag (1979) estudou o género Cicindela da fauna australiana. Wiesner (1986), Cassola (1987 a e b e 1991) e Naviaux (1991), reviram respectivamente, os cicindelídeos de Sumatra, da 236 Artur R. M. Serrano Nova Guiné, Ilhas Salomão e Sulawesi (Indonésia) e da Tailândia. Freitag & Barnes (1989) estudaram também o género Cicindela do continente sul-americano, com especial incidência sobre a componente brasileira. Utilizando os caracteres da genitália masculina, Rivaler (1950, 1954, 1957,1961 e 1963) dividiu o género Cicindela em 55 géneros. Pos­ teriormente, este mesmo autor (Rivalier, 1969), dividiu o género pan­ tropical Odontocheila e a finalizar redefiniu dentro da tribo Cicindelini as 5 subtribos que a compõem (Rivalier, 1971). De um modo geral, os sistematas concordaram que os géneros propostos por Rivalier (1963), são grupos naturais que têm corres­ pondência na maior parte dos casos com os de Horn (1926). Tal facto já transpareceu no recente trabalho de Wiesner (1992). Apesar de terem aceite globalmente esta divisão, os entomólogos norte-americanos preferiram considerá-los com o estatuto subgenérico (e.g., Boyd et aI., 1982). Citogenética o estudo cito genético dos cicindelídeos iniciou-se no princípio deste século (Stevens, 1906 e 1909). No entanto, só em 1952 Guenin e a se­ guir Smith & Edgar (1954) indicaram que o género Cicindela possuia um sistema de cromossomas sexuais múltiplos (X.Y/X). Somente Cylindera germanica (Linnaeus) e Cylindera paludosa (Dufour) contrariaram até agora esta homogeneidade, evidenciando sistemas sexuais simples (XY/ XX e XO/XX, respectivamente) (Giers, 1977 e Serrano et aI., 1986). O número de autossomas, no cômputo geral de todas as espécies analisadas, destaca-se pela grande estabilidade dentro do grupo (9 a 11 pares) (Serrano & Yadav, 1984). Um estudo recente sobre 11 espécies indianas do género Cylindera Westwood (Yadav & Burra, 1991) veio reforçar aquela asserção. Serrano (1986) baseado nos padrões dos complexos cromossómicos sexuais encontrados nas espécies estudadas até àquele momento, expressou a opinião da ocorrência de uma forte correlação entre cada tipo e a respectiva região biogeográfica. Assim, indicou um número básico 2n=20+X\X2Y para as espécies da Região Oriental, 2n=18+X\X2X,Y para as da Região Paleárctica e finalmente 2n=18+X\X2Y para as da Região Neárctica. Serrano et. ai (1986) adiantaram ainda que o padrão neárctico estaria na base dos outros modelos. Cicindelídeos (Coleoptera, Cicindelidae) 237 Aquela correlação foi, contudo, posteriormente contrariada com a descoberta de um sistema cromossómico sexual múl tiplo análogo ao modelo neárctico em Ceph. hispanica (Serrano & Collares-Pereira, 1989). A Península Ibérica, no contexto da Região Paleárctica, tem sido um local privilegiado no âmbito dos estudos citogenéticos (Serrano, 1980 e 1981, Serrano et ai., 1986, Serrano & Collares-Pereira,. 1989 .e 1992, Collares-Pereira & Serrano, 1990 e Galian el ai., 1990). O problema da origem dos sistemas cromossómicos sexuais múltiplos nos cicindelídeos tem sido uma questão aliciante e que continua por resolver (vide por exemplo, Smith & Edgar, 1954, Dasgupta, 1967, Serrano, 1980,1981,1986, Yadav el aI., 1985, Serrano et aI., 1986, Yadav & Burra, 1991). MORFOLOGIA GERAL o ovo Os ovos das «cicindelas» são aparentemente muito similares e dados sobre o seu tamanho foram referidos por Ponselle (1900), Shelford (1908), Huie (\915), Willis (1967) e Serrano (1990). Dados apresentados por Serrano (op. cil.) para 8 formas de POrtugal indicaram que havia maior uniformidade na forma dos ovos ·· de algumas espécies do que noutras. Os ovos apresentam uma forma ovóide mais ou menos acentuada, com um micrópilo estrelado no pólo anterior (Serrano, op. dI.). De acordo com Willis (1967) e'Serrano (1988), ocorria por vezes uma ligeira depressão no lado «ventral». O estudo da estrutura do cório realizado por Serrano (\ 990) permitiu identificar um exocório com um
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