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2 - A subfamília Amblyoponinae na Região Neotropical Flavia A. Esteves Brian L. Fisher SciELO Books / SciELO Livros / SciELO Libros ESTEVES, FA., and FISHER, BL. A subfamília Amblyoponinae na Região Neotropical. In: DELABIE, JHC., et al., orgs. As formigas poneromorfas do Brasil [online]. Ilhéus, BA: Editus, 2015, pp. 13-22. 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. 2 A subfamília Amblyoponinae na Região Neotropical Flavia A. Esteves, Brian L. Fisher Resumo A subfamília Amblyoponinae apresenta da biologia e taxonomia da subfamília na região distribuição global e na região Neotropical inclui Neotropical. Infelizmente, todos os três gêne- os gêneros Paraprionopelta Kusnezov, Prionopelta ros presentes nesta região possuem problemas Mayr e Stigmatomma Roger. com relação à delimitação de suas espécies. Neste capítulo, apresentamos uma introdu- Paraprionopelta é apenas conhecida por machos; ção à esta subfamília, destacando alguns aspectos Prionopelta amabilis, P. antillana e P. marthae de sua sistemática. Fornecemos informações sobre não podem ser claramente identifi cadas com os distribuição e biologia, e diagnoses para fêmeas e caracteres morfológicos atualmente disponíveis; machos dos gêneros de Amblyoponinae de distri- e a maioria das espécies de Stigmatomma precisa buição Neotropical. Incluímos imagens de alta re- de redescrições mais detalhadas para que carac- solução para ilustrar os gêneros que ocorrem nesta teres ambíguos, e amplamente utilizados atual- região biogeográfi ca. mente, sejam substituídos por outros de maior Finalizamos o capítulo ressaltando al- valor diagnóstico. Por fi m, apresentamos solu- guns empecilhos para um maior conhecimento ções para que estes problemas sejam superados. ESTEVES, Flavia A.; FISHER, Brian L. A subfamília Amblyoponinae na região Neotropical. In: DELABIE, Jacques H. C. et al. As formigas poneromorfas do Brasil. Ilhéus: Editus, 2015. p. 13-22. A subfamília Amblyoponinae na região Neotropical | 13 PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 1133 113/01/20163/01/2016 111:48:281:48:28 Abstract Th e subfamily Amblyoponinae in We fi nalize this chapter by highlighting the Neotropical Region - Th e subfamily some issues for a comprehensive knowledge of Amblyoponinae is widespread around the globe, the subfamily’s biology and taxonomy in the and in the Neotropical bioregion includes the Neotropical region. Unfortunately, all the three genera Paraprionopelta Kusnezov, Prionopelta genera possess problems related to the identifi cation Mayr and Stigmatomma Roger. of their species. Paraprionopelta is just known by In this chapter, we off er an overview the male caste; Prionopelta amabilis, P. antillana and of the subfamily and highlight aspects of its P. marthae cannot be clearly called apart with the systematics; and provide information on biology, morphological characters currently available; and distribution, and diagnosis for females and males the majority of Stigmatomma species need more of Amblyoponinae genera distributed in the detailed descriptions, so that ambiguous characters, Neotropics. We also included high-resolution currently broadly used, may be replaced by others images to illustrate the genera that occur in that with higher diagnostic value. Finally, we present bioregion. solutions to overcome these issues. Introdução Sistemática A subfamília Amblyoponinae se distribui A relação de parentesco entre as linhagens globalmente e atualmente agrupa os gêneros: de formigas com ramifi cação mais antiga, incluin- Adetomyrma Ward (1994: 160); Amblyopone do Amblyoponinae, ainda é incerta (BRADY et al., Erichson (1842: 260); Apomyrma Brown, Gotwald 2006; KÜCK et al., 2011; MOREAU; BELL, 2013), e Lévieux (1971: 259); Bannapone Xu (2000: limitando nossa compreensão sobre os fatores que 299); Concoctio Brown (1974: 29); Myopopone moldaram a evolução das formigas, e nossa habi- Roger (1861: 49); Mystrium Roger (1862: 245); lidade para testá-los (WILSON e HÖLLDOBLER, Onychomyrmex Emery (1895: 349); Opamyrma 2005; LUCKY et al., 2013; MOREAU; BELL, 2013). Yamane, Bui e Eguchi (2008, 56); Paraprionopelta Felizmente, o nosso conhecimento sobre a biolo- Kusnezov (1955: 270); Prionopelta Mayr (1866: gia evolutiva de formigas aumentou muito ao lon- 503); Stigmatomma Roger (1859: 250); Xymmer go da última década (BOLTON, 2003; BRADY et Santschi (1914: 311); além do gênero fóssil al., 2006; MOREAU et al., 2006; RABELING et al., Casaleia Pagliano e Scaramozzino (1990: 5). 2008; MOREAU, 2009; KÜCK et al., 2011; MORE- Atualmente são reconhecidas mais de 130 espécies AU; BELL, 2013; BRADY et al., 2014; WARD et al., para a subfamília (BOLTON, 2014). 2014), e um dos resultados é que Amblyoponinae O velho mundo possui a maior diversidade voltou a ser reconhecida como subfamília (BOL- genérica de Amblyoponinae. A região Indomalaia TON, 2003). é a mais diversa, com sete gêneros (Bannapone, A relação fi logenética entre os gêneros de Myopopone, Mystrium, Opamyrma, Prionopelta, Amblyoponinae também não está perfeitamente Stigmatomma e Xymmer), seguida pelas regiões resolvida, mas caminhamos muito nesta direção. Australiana e Afrotropical, ambas com seis A subfamília aparentemente é formada por dois gêneros (Amblyopone, Myopopone, Mystrium, clados (BRADY et al., 2006; OULLETTE et al., 2006; Onychomyrmex, Prionopelta e Stigmatomma; SAUX et al., 2004): (1) Adetomyrma + Bannapone, Apomyrma, Concoctio, Mystrium, Prionopelta, Myopopone + Mystrium + Paraprionopelta + Stigmatomma e Xymmer, respectivamente). A região Stigmatomma + Xymmer; e (2) Amblyopone Neotropical possui 3 gêneros: Paraprionopelta, + Concoctio + Onychomyrmex + Prionopelta Stigmatomma e Prionopelta (ANTWEB, 2014a; (YOSHIMURA e FISHER, 2012; FISHER, B.L., BOLTON, 2014). dados ainda não publicados); a posição de 14 | Flavia A. Esteves, Brian L. Fisher PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 1144 113/01/20163/01/2016 111:48:21:48:29 Apomyrma e Opamyrma não é conhecida. A Diagnose delimitação dos gêneros dessa subfamília também está passando por transformações – Stigmatomma Mandíbulas com um dente. Antena com 10 e Xymmer são aceitos novamente como gêneros segmentos (Figura 2-1A). Fórmula palpal 2:2 (2 depois de um longo tempo sinomizados em maxilares e dois labiais) (KUSNEZOV, 1955). Ca- Amblyopone (YOSHIMURA; FISHER, 2012), beça oblonga (Figura 2-1A). Notaulus ausente ou e dados preliminares indicam a polifi lia de fracamente desenvolvido. Pigóstilos presentes. Stigmatomma (FISHER, B.L., dados ainda não publicados). Prionopelta Mayr, 1866 Gêneros de Amblyoponinae da Prionopelta possui atualmente 21 espécies Região neotropical válidas no mundo, das quais 5 ocorrem na região Neotropical: P. am abi li s Borgmeier (1949: 203); P. Paraprionopelta Kusnezov, 1955 antillana Forel (1909: 239); P. marthae Forel (1909: 240); P. modesta Forel (1909: 241); e P. punctulata Paraprionopelta é conhecida por apenas Mayr (1866: 505). uma espécie, Paraprionopelta minima Kusnezov Este gênero de formigas geralmente ocorre (1955: 271), baseada na descrição de machos cole- em habitats de fl orestas úmidas e nidifi ca dentro tados na região de Tucumã, Argentina. de troncos podres sobre o solo, nas camadas su- Os espécimes deste gênero foram coletados perfi ciais do solo, em frutos podres e secos sobre o em diferentes ocasiões em Tucumã, Argentina, solo, e no interior do solo. através de armadilhas luminosas, e assim, nada se O conhecimento atual sobre o comporta- sabe sobre sua biologia. Desde então, machos de mento de Prionopelta é derivado basicamente de Amblyoponinae que possuem caracteres seme- três estudos baseados em observações de colônias lhantes aos de Paraprionopelta foram coletados na cativas de P. am abi li s (HOLLDOBLER; WILSON, Nicarágua, e suas imagens estão disponíveis em 1986; HOLLDOBLER et al., 1992) e P. kraepelini Fo- Antweb.org (2014b; CASENT0628950). rel (1905: 3) (ITO; BILLEN, 1998). Em P. am abi li s , Figura 2-1 – Parátipo de Paraprionopelta minima (CASENT0172797): (A) vista dorsal da cabeça e (B) lateral do - corpo. Imagens: April Nobile; disponível em AntWeb.org. A B A subfamília Amblyoponinae na região Neotropical | 15 PPoneromorfasoneromorfas ddoo BBrasil_miolo.inddrasil_miolo.indd 1155 113/01/20163/01/2016 111:48:21:48:29 as colônias são polidômicas (também descrito para P. am abi li s nunca foi observado, e aparentemente P. modesta; BROWN, 1960) e podem conter mais não existe ferormônio de alarme. de 700 operárias e uma ou duas rainhas. Aparente- mente as colônias de P. kraepelini são monogínicas. Diagnose Todas as operárias de P. am abi li s parecem ser capa- zes de botar ovos trófi cos, os quais podem ser a fon- Fêmeas (Figura 2-2): Mandíbula estreita e te principal de alimento da rainha (esta se alimenta curta, subtriangular, armada com 3 dentes (Figu- também de hemolinfa larval em P. kraepelini). ra 2-2A). Margem anterior do clípeo convexa ou Aparentemente, o recrutamento em P. moderadamente projetada, armada com pequenas amabilis é feito através de vibrações corporais e setas dentiformes (Figura 2-2A). Olhos compos- secreções de uma glândula exócrina localizada tos reduzidos nas
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  • Hymenoptera, Formicidae: Amblyoponinae, Ectatomminae, Ponerinae) of Grube Messel, Germany: High Biodiversity in the Eocene G

    Hymenoptera, Formicidae: Amblyoponinae, Ectatomminae, Ponerinae) of Grube Messel, Germany: High Biodiversity in the Eocene G

    Journal of Systematic Palaeontology, Vol. 10, Issue 4, December 2012, 725–753 The poneromorph ants (Hymenoptera, Formicidae: Amblyoponinae, Ectatomminae, Ponerinae) of Grube Messel, Germany: high biodiversity in the Eocene G. M. Dlusskya∗ and S. Wedmannb aDepartment of Evolution, Biological Faculty, M.V.Lomonosov Moscow State University, Vorobjovy gory, 119992, Moscow, Russia; bForschungsstation Grube Messel, Senckenberg Forschungsinstitut und Naturmuseum, Markstraße 35, D-64409, Messel, Germany (Received 17 September 2010; accepted 27 March 2011; printed 5 December 2012) Ants are a very successful group among insects, but the course of evolution of their biodiversity is still unclear. This study sheds light on ant diversification during the Eocene. Analysis of the ant taphocoenosis of the fossil site Grube Messel, Germany (47 Ma) yielded three poneromorph subfamilies and 22 new species in six genera, four of which are new: Pseudectatomma gen. nov., Cephalopone gen. nov., Cyrtopone gen. nov. and Messelepone gen. nov. Only one extant genus, Pachycondyla, is present in the taphocoenosis from Messel. The high diversity of poneromorph ants from Messel is very striking in comparison with middle to late Eocene European ambers. A significantly lower proportion of species in ambers can be assigned to poneromorph ants, and fewer poneromorph species are known from European ambers than from Messel. A possible gradual decline of the diversity of poneromorphs from the Eocene to the Miocene seems to be detectable worldwide. These insights are discussed in the context of the morphology and ecology of Poneromorpha and Formicomorpha. The proportion of ant castes in amber seems to indicate that already during the Eocene poneromorphs inhabited preferably litter and soil, whereas formicomorphs preferred the arboreal realm.