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Poneromorfas Do Brasil Miolo.Indd 5 - Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae Itanna O. Fernandes Jorge L. P. de Souza Fabricio B. Baccaro SciELO Books / SciELO Livros / SciELO Libros FERNANDES, IO., SOUZA, JLP., and BACCARO, FB. Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae. In: DELABIE, JHC., et al., orgs. As formigas poneromorfas do Brasil [online]. Ilhéus, BA: Editus, 2015, pp. 43-53. ISBN 978-85-7455-441-9. Available from SciELO Books <http://books.scielo.org>. All the contents of this work, except where otherwise noted, is licensed under a Creative Commons Attribution 4.0 International license. Todo o conteúdo deste trabalho, exceto quando houver ressalva, é publicado sob a licença Creative Commons Atribição 4.0. Todo el contenido de esta obra, excepto donde se indique lo contrario, está bajo licencia de la licencia Creative Commons Reconocimento 4.0. 5 Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae Itanna O. Fernandes, Jorge L. P. de Souza, Fabricio B. Baccaro Resumo A subfamília Paraponerinae possui uma forrageando em nectários extrafl orais e carregando série de mudanças em sua classifi cação taxonômi- pequenos invertebrados ou partes deles para o ni- ca no decorrer da história. A utilização de dados nho. Estudos indicaram que o forrageamento em P. morfológicos e moleculares em fi logenias mais re- clavata é principalmente noturno, mas pode variar centes, demonstrou que a posição de Paraponeri- entre as colônias. Sua distribuição é ampla, com nae dentro do grupo de poneromorfas ainda está registros da Nicarágua, na América Central, até o indefi nida, visto que as relações fi logenéticas com Centro-Sul do Brasil. Mapas de distribuição em lar- as subfamílias Proceratiinae, Agroecomyrmecinae ga escala sugerem que a ocorrência de P. clavata está e Amblyoponinae ainda não foram esclarecidas. mais correlacionada com variáveis abióticas, como Paraponerinae possui duas espécies, pluviosidade e temperatura média, que com variá- Paraponera clavata, sua única espécie vivente, e veis bióticas, como tipo de vegetação. Paraponera dieteri, espécie fóssil oriunda de âmbar Paraponera clavata é conhecida por aplicar Dominicano. As principais características morfoló- ferroadas extremamente dolorosas, ocasionando gicas do gênero são, os escobros antenais em forma febre, tremores, sudorese fria, náuseas, vômito e de “v”, curvando-se detrás dos olhos; soquetes ante- arritmia cardíaca. A dor, no entanto, é o principal nais ocultos pelos lobos frontais, localizados longe da sintoma da picada, sendo extremamente violen- margem anterior da cabeça; garras tarsais com dentes ta e capaz de permanecer por períodos de até 24 pré-apicais; e nodo peciolar longo e sub-retangular. horas. O veneno da P. clavata é classifi cado como São formigas grandes, as operárias podem chegar a neurotóxico atuando diretamente no sistema ner- mais de 2,5 cm. voso central dos insetos, bloqueando a transmis- O tamanho das colônias pode variar entre são de sinapses e tem potencial para aplicação em 200 a 3000 operárias e o ninho está normalmente inseticidas biológicos. Os indivíduos de P. clavata localizado na base de árvores. As operárias esta- são popularmente conhecidos como “tocandiras belecem trilhas de forrageio estáveis, forrageando ou tucandeiras”, são utilizados como elementos ativamente tanto no sub-bosque como no dossel. O folclóricos. Na Região Norte do Brasil, índios Sa- hábito alimentar pode ser classifi cado como onívo- teré-Mawé utilizam as tucandeiras em rituais de ro, já que operárias são frequentemente observadas passagem para a vida adulta. FERNANDES, Itanna O.; SOUZA, Jorge L. P. de; BACCARO, Fabricio B. Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae. In: DELABIE, Jacques H. C. et al. As formigas poneromorfas do Brasil. Ilhéus: Editus, 2015. p. 43-53. Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae | 43 PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 4433 113/01/20163/01/2016 111:48:31:48:37 Abstract State of the art of the taxonomy and nectaries and carrying small invertebrates to the phylogeny of the subfamily Paraponerinae - Th e nest. Foraging studies indicate that P. clavata is Paraponerinae has experienced a series of changes primarily nocturnal, but the foraging time can vary in its taxonomic history. Indeed, morphological and among the colonies. P. clavata is widely distributed, molecular data in more recent phylogenies indicate with records from Nicaragua in Central America, that the position of Paraponerinae within the through to the center and south of Brazil. Large- poneromorph group is still undefi ned, since certain scale distribution maps in South America suggest phylogenetic relationships with Proceratiinae, that the occurrence of P. clavata is more correlated Agroecomyrmecinae and Amblyoponinae have with abiotic variables, such as rainfall and mean not been clarifi ed. Paraponerinae has two species, temperature, than with biotic variables, such as Paraponera clavata, the only living species, and type of vegetation. Paraponera dieteri, a fossil species from Dominican Paraponera clavata has an extremely amber. Th e genus diagnosis is the presence of painful sting, with symptoms of fever, chills, cold antennal scrobes in a “v” shape curving behind the sweats, nausea, vomiting and cardiac arrhythmia. eyes, antennal sockets hidden by the frontal lobes Th e pain aft er a sting is the main symptom, and located far from the anterior margin of the being extremely violent and able to remain for head, tarsal claws with pre-apical teeth and a long up to 24 hours. P. clavata venom is neurotoxic, sub-rectangular petiolar node. Ants are large, with acting directly on the central nervous system of workers reaching more than 2.5 cm. insects to block transmission at synapses, and Th e colony size can vary between 200-3000 has potential use for application in biological workers and the nest is usually located at the base insecticides. Individuals of P. clavata, locally of trees. Th e workers establish stable foraging trails, known as ‘tucandiras or tucandeiras’ are used actively foraging in both, understorey and canopy. as folklore elements. In the northern region of P. clavata has an omnivorous diet, with workers Brazil, the Sateré-Mawé Indians use tucandeiras frequently observed foraging on extrafl oral in rites of passage to adulthood. Sistemática de Paraponerinae os gêneros Paraponera e Ectatomma estariam re- lacionados, devido à presença do lobo anal na asa A subfamília Paraponerinae possui apenas posterior em ambos os gêneros. uma espécie vivente, Paraponera clavata (FABRI- Lattke (1994) revalidou a tribo Paraponeri- CIUS, 1775), e apresenta uma série de mudanças ni através de uma fi logenia baseada em caracteres em sua classifi cação taxonômica no decorrer da morfológicos, onde a ausência do processo ante- história. Em 1954, Brown apresentou a primeira roventral póspeciolar, a presença de tumosidades classifi cação para o chamado complexo Poneroi- laterais no pronoto e o escrobo antenal bem desen- de, composto pelas subfamílias Ponerinae, Cera- volvidos de P. clavata a distinguia de qualquer ou- pachyinae, Myrmicinae, Dorylinae e Leptanillinae tra Ectatommini. Neste mesmo trabalho, também (Figura 5.1). De acordo com esta classifi cação, demonstrou que a tribo Ectatommini formava um Paraponera clavata foi agrupada na subfamília agrupamento polifi lético (Figura 5.2). Ponerinae, dentro da tribo Ectatommini. Keller (2000) analisou as relações existen- Brown (1958) ao estudar as relações tes entre os membros da tribo Ectatommini e su- existentes entre os indivíduos que compunham geriu um arranjo diferente do proposto por Lattke a subfamília Ponerinae propôs uma nova classi- (1994). Keller menciona que a posição do gênero fi cação, baseada em caracteres morfológicos, que Paraponera é mal suportada para uma proposta de reduziu as três tribos da subfamília Ponerinae reclassifi cação. Ao reanalisar a matriz proposta por (Ectatommini, Paraponerini e Proceratiini) a so- Lattke, observou que a posição de Paraponera mu- mente uma: Ectatommini. Concluiu também que dava à medida que o grupo externo era modifi cado. 44 | Itanna O. Fernandes, Jorge L. P. de Souza, Fabricio B. Baccaro PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 4444 113/01/20163/01/2016 111:48:381:48:38 FIGURA 5.1 – Árvore fi logenética para as subfamílias de Formicidae proposta por Brown em 1954 (Adaptado de Keller, 2011). FIGURA 5.2 – Relações fi logenéticas dos gêneros da subfamília Ponerinae com grupo externo representado pelo gênero Myrmecia (Modifi cado de Lattke, 1984). Estado da arte sobre a Filogenia, Taxonomia e Biologia de Paraponerinae | 45 PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 4455 113/01/20163/01/2016 111:48:381:48:38 Paraponera assumia uma posição terminal quando de Ponerinae não é suportada e Ectatommini (sen- Cerapachys era utilizada como grupo externo e ba- su Lattke) continua envolvida em uma politomia. sal ao utilizar Myrmecia (Figura 5.3). Bolton (2003) desmembrou a então subfamí- Ao fi nal, Keller concluiu que Paraponera era lia Ponerinae em seis subfamílias: Amblyoponinae, grupo-irmão das demais poneríneas + Cerapachys. Ectatomminae, Heteroponerinae, Paraponerinae, De acordo com essa análise, a classifi cação tribal Ponerinae e Proceratiinae. Neste mesmo trabalho, FIGURA 5.3 – Relações fi logenéticas para a subfamília Ponerinae propostas por Keller (2000) com grupos externos representados pelos gêneros Cerapachys (A) e Myrmecia
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