Phylogeny, divergence times, and evolution of Oecanthidae n. status (Insecta, Orthoptera, Grylloidea

A ordem Orthoptera, conhecida principalmente pelos grilos e gafanhotos, é distribuída mundialmente com quase 30 mil espécies válidas, sendo considerada a ordem mais diversa dentre os Polyneoptera. Em Orthoptera, os grilos verdadeiros (Grylloidea) são organismos frequentemente utilizados como organismos modelo para diversos tipos de estudo como comunicação acústica, comportamento, ecologia e até neurobiologia. No entanto, poucos desses estudos são focados no contexto evolutivo, uma vez que ainda existem poucas hipóteses filogenéticas para o grupo. Além disso, a comunicação acústica desses insetos é considerada um dos elementos mais intrigantes e presentes em sua evolução. Porém, apesar de ser um tema constantemente retratado pela comunidade científica, poucos trabalhos o abordam em um contexto filogenético. No primeiro capítulo dessa tese, apresentamos uma hipótese filogenética para uma nova família de grilos, grupo-irmão de Gryllidae n. def.: Oecanthidae n. status. As análises filogenéticas possuem como fonte de caracteres dados moleculares e morfológicos, para parcimônia e máxima verossimilhança, e dados moleculares para a análise de tempos de divergência (inferência bayesiana). Foram utilizados 107 terminais de todas as regiões biogeográficas e seis fósseis para a calibração da árvore filogenética. Todas as análises resultam em Oecanthidae n. status composta por quatro subfamílias, Euscyrtinae, Oecanthinae n. def., n. status, Podoscirtinae n. def. e Tafaliscinae n. def. n. status. Baseado nos resultados obtidos, nós revisamos a definição e a classificação interna das subfamílias, supertribos (propostas neste trabalho) e tribos. Phyllogryllini n. tribo é definida. Também atualizamos as diagnoses, listamos os gêneros de cada uma das tribos e as apomorfias dos táxons supragenéricos. Também é apresentada uma chave de identificação para todos os níveis taxonômicos de Oecanthidae n. status mais todos os gêneros de Tafaliscinae n. def. n. status. No segundo capítulo, utilizamos a filogenia de Oecanthidae, baseada em dados moleculares e morfológicos, para realizar uma reconstrução de estados ancestrais relacionados a comunicação acústica. Os caracteres são estruturas das asas anteriores, as próprias asas anteriores e tímpanos. Foram analisados seis caracteres no total. Nossos resultados demostram perdas de caracteres relacionados a comunicação acústica para vários táxons independentemente ao longo do tempo. Ao final, discutimos as possíveis causas da perda da capacidade de produção do som em Oecanthidae além das suas alternativas para uma comunicação eficiente não dependente de sinais acústicos.Orthoptera, mainly known by crickets and grasshoppers, is distributed worldwide with almost 30,000 valid species. The order is considered the most diverse order within Polyneoptera. In Orthoptera, true crickets (Grylloidea) are frequently used as models for many areas of science as acoustic communication, behavior, ecology, and neurobiology. However, only a few studies are focused on a phylogenetic context since the number of phylogenetic hypotheses for this group is low. Besides, the acoustic communication of these insects is considered an essential element of their evolution. Although it is a subject constantly debated, only a few works are in a phylogenetic frame. In the first chapter of this thesis, we present a phylogenetic hypothesis for a new family of crickets, sister-group of Gryllidae n. def.: Oecanthidae n. status. The phylogenetic analyses are based on molecular and morphological data for maximum likelihood and parsimony criteria and molecular data for divergence times analysis (Bayesian inference). Were used 107 terminals from all biogeographic regions and six fossils to calibrate the phylogenetic tree. All analyses result in Oecanthidae n. status composed of four subfamilies: Euscyrtinae, Oecanthinae n. def., Podoscirtinae n. def., and Tafaliscinae n. def. n. status. We revise the definition and internal classification of subfamilies, supertribes (proposed herein), and tribes based on our results. Phyllogryllini n. tribe is defined. We also update the diagnosis of suprageneric groups, list their apomorphies, and list the genera of each tribe. An identification key is proposed for all taxonomic levels of Oecanthidae n. status plus all Tafaliscinae n. def., n. status genera. In Chapter 2, we use the phylogeny of Oecanthidae, based on molecular and morphological characters, to reconstruct the history of characters related to acoustic communication in crickets as forewings structures, the forewings itself, and tympana. In total, the history of six characters are analyzed. Our results demonstrate the loss of characters related to acoustic communication along the time independently for many taxa in this family of crickets. Several of these taxa are not able to use forewings to stridulate. We discuss the potential causes of losing the capacity of sound-producing and hearing and their alternatives for efficient communication not exclusively based on sounds

Taxonomic and phylogenetic study of Eidmanacris Chopard, 1956 (Orthoptera; Phalangopsidae; Luzarinae)

Os ortópteros neotropicais, com exceção aos de importância econômica, de maneira geral, são muito pouco conhecidos em relação aos demais ortópteros ao redor do mundo. Isso devido à carência de trabalhos científicos para esse grupo, principalmente os estudos taxonômicos; consequentemente, o número de espécies descritas atualmente é muito subestimado. Não diferente, o gênero Eidmanacris Chopard, 1956, possui atualmente 20 espécies descritas, distribuídas nos domínios de Mata Atlântica e Cerrado, que se estendem pelas regiões sul, sudeste e centro-oeste brasileiras, além de também serem encontrados na Bolívia e Paraguai. São grilos ativos no período noturno, habitantes cavidades naturais, como tocas, barrancos, troncos de árvores mortas, fendas de rochas e cavernas. Neste estudo, o gênero foi revisado, incluindo a redescrição de espécies pouco conhecidas, descrição de sete novas espécies (E. scopula Campos, sp. nov., E. gigas Campos, sp. nov., E. neomarmorata Campos, sp. nov., E. desutterae Campos, sp. nov., E. putuhra Campos, sp. nov., E. fontanettiae Nihei & de Mello, sp. nov. e E. melloi Campos, sp. nov.), três novas combinações (E. speluncae (Mello-Leitão, 1937) comb. nov., E. minuta (de Mello, 1990) comb. nov. e E. endophallica (de Mello, 1990) comb. nov.) e uma sinonímia nova (E. lencionii Bolfarini, 2016 = E. dissimilis Desutter-Grandcolas, 1995, syn. nov.), totalizando 29 espécies para o grupo. Também foram feitos uma chave de identificação para o gênero e mapas de distribuição de suas espécies. Uma análise filogenética é apresentada incluindo 38 terminais, dos quais 12 foram considerados como grupo externo, e 98 caracteres morfológicos. Essa análise suportou a monofilia do gênero, permitiu visualizar seu relacionamento com táxons próximos de Luzarinae e sustenta a proposta de sinonímia de Endophallusia de Mello, 1990 com Eidmanacris Chopard, 1956, de modo a sua taxonomia refletir a monofilia indicada na análise filogenéticaThe Neotropical Orthoptera, excluding those with economic importance, are poorly known, in comparison with the orthopterans around the world. The main reason is the lack of scientific studies on this group, mainly taxonomic studies, resulting in a underestimated number of described species. Likewise, Eidmanacris Chopard, 1956 comprises 20 described species, and is mostly distributed on Atlantic Forest and Cerrado areas, extending from south, southeast and midwest Brazilian regions, and beyond, reaching Bolivia and Paraguay. Eidmanacris species are active at night, and inhabit natural cavities as burrows, bounds, hollow trees trunks, cavities in rocks and caves. In this study, the genus was reviewed, including the redescription of seven new species (E. scopula Campos, sp. nov., E. gigas Campos, sp. nov., E. neomarmorata Campos, sp. nov., E. desutterae Campos, sp. nov., E. putuhra Campos, sp. nov., E. fontanettiae Nihei & de Mello, sp. nov. and E. melloi Campos, sp. nov.), three new combinations (E. speluncae (Mello-Leitão, 1937) comb. nov., E. minuta (de Mello, 1990) comb. nov. and E. endophallica (de Mello, 1990) comb. nov.) and one synonym (E. lencionii Bolfarini, 2016 = E. dissimilis Desutter-Grandcolas, 1995, syn. nov.), totalizing 29 species in this genus. An identification key for the genus and a distribution map of species were made. A phylogenetic analysis is presented including 38 terminals, 12 of them as outgroup and 98 morphological characters This analysis attested the monophyly of the genus, showed its relationships with another Luzarinae taxa, and supported the proposal of synonymy of Endophallusia de Mello, 1990 with Eidmanacris Chopard, 1956, so that its taxonomy reflected the monophyly indicated in the phylogenetic analysi

Taxonomic studies on the Neotropical Landrevinae with description of new taxa (Orthoptera, Grylloidea, Gryllidae)

The understanding of the subfamily Landrevinae has been modified by different authors since its creation. In the neotropics three genera are known to the present: Odontogryllus Saussure, 1877 (one from México, the others amazonian), Brasilodontus de Mello, 1992 with two species (from Brazilian Atlantic Forest), e Valchica de Mello, 1992 with one species (from Costa Rica). De Mello (1992) erroneously created the tribe Odontogryllini for this cluster of neotropical genera, here suppressed. In the present paper we revise and add new species to Brasilodontus and describe two monotypic genera, Xulavuna n. gen. and Yarrubura, n. gen. An identification key to the genera of neotropical Landrevinae is presented as well as one for the species of Brasilodontus. The male fore wings of Xulavuna adenoptera n. sp. is remarkable regarding its shape and its glandular conditionO entendimento da subfamília Landrevinae tem sido alterada por diferentes autores desde sua criação. Até o momento, três gêneros são conhecidos da Neotropica: Odontogryllus Saussure, 1877 com 11 espécies (uma das quais do México, as demais amazônicas), Brasilodontus de Mello, 1992 com duas espécies (da Floresta Atlântica brasileira), e Valchica de Mello, 1992 monotípico (da Costa Rica). De Mello (1992) erroneamente erigiu a tribo Odontogryllini para esse grupo de gêneros neotropicais, aqui suprimida. No presente trabalho revisamos e adicionamos novas espécies ao gênero Brasilodontus e descrevemos novos gêneros monotípicos, Xulavuna gen. n. e Yarrubura, gen. n. Uma chave de identificação para os gêneros de Landrevinae neotropicais é apresentada, assim como uma para as espécies de Brasilodontus. As asas mesonotais do macho de Xulavuna adenoptera sp. n. são notáveis quanto à forma e sua condição glandula

Three new species of Amblyrhethus (Orthoptera, Grylloidea, Gryllidae, Paroecanthini) from Brazil

Amblyrhethus Kirby, 1906 is a genus of arboreal, undergrowth, and shrub crickets comprising, at present, seven species: one from Panama, one from Peru, two from Colombia, two from Brazil, and one with an unprecise locality. These crickets are seldom found in regular active collecting at night, although males produce a rather loud calling song. Unfortunately, their songs have never been recorded, and there is no ecological information for this genus so far. Here, we describe three new species from the Brazilian Atlantic Forest

Taxonomic studies on the Neotropical Landrevinae with description of new taxa (Orthoptera, Grylloidea, Gryllidae)

Campos, Lucas Denadai De, De Mello, Francisco De A. G. (2014): Taxonomic studies on the Neotropical Landrevinae with description of new taxa (Orthoptera, Grylloidea, Gryllidae). Zootaxa 3852 (2): 151-178, DOI: 10.11646/zootaxa.3852.2.

First 3D reconstruction of a forewing of a fossil Orthoptera: Interpreting the venation pattern in the smallest known cricket with a stridulatory apparatus, Picogryllus carentonensis (Orthoptera, Grylloidea, Oecanthidae)

International audienceFossil insects are valuable indicators of the evolutionary history of the clades to which they belong. According to their state of preservation, fossil insects are often partially described for key morphological characters, such as forewing venation in crickets (Orthoptera, Grylloidea). In parallel, the use of 3D microtomography is increasingly becoming common for studying some fossils, which allowed here the precise reconstruction and interpretation of the venation pattern in the smallest known cricket with a stridulatory apparatus, dagger Picogryllus carentonensis, found in opaque amber. The 3D reconstructions have revealed the general structure of the venation of the forewing and have enabled the identification of all its veins and cells, validating its similarity with that of extant crickets. Putative homologies are established according to previous studies, and some particularities are observed, such as the presence of two crossveins in the mirror, a rare feature in extant crickets that is discussed in the frame of cricket venation evolution. These findings highlight the importance of 3D microtomography as a powerful tool for examining fossil insects and also provide crucial information for taxonomic identification and evolutionary studies, offering a validated morphological basis for future phylogenetic analyses incorporating fossils

First 3D reconstruction of the male genitalia of a Cretaceous fossil cricket: Diving into the evolutionary history of the Oecanthidae family (Orthoptera: Grylloidea) with the incorporation of new fossils in its phylogeny and a total-evidence dating approach

International audienceFossils are valuable indicators of the evolutionary history of the clades to which they belong to, especially when they are incorporated as terminal taxa in a total-evidence phylogeny. According to their state of preservation, fossils are often incompletely described for key morphological characters, such as genitalic structures. Here, the internal parts of the genitalia of a male fossil cricket from Cretaceous amber, dagger Picogryllus carentonensis Josse and Desutter-Grandcolas (Oecanthidae, Podoscirtinae), together with other key morphological characters (i.e., metanotal structures and tibial spurs), were reconstructed for the first time by 3D microtomography. Total-evidence phylogeny and dating combining morphological data (fossils and extant taxa), molecular data (extant taxa only) and time calibration (fossil dates) were performed to evaluate the tempo and mode of evolution of the cricket family Oecanthidae. Divergence time estimates were thus refined and the patterns of transformation for key morphological structures contrasted through the analysis of phylogenetic morphological partitions. Our results show that Oecanthidae date back to the Upper Jurassic (Oxfordian, around 162 Ma) and attest to the presence of the Podoscirtinae in Western Europe during the Lower Cretaceous. Morphological evolution may have been driven by the conquest of new resources (as shown by leg evolution in ancestral Oecanthidae) and/or the 'conquest of silence' (as shown by repetitive and definitive losses of acoustic structures). By contrast, genitalia evolution proved more diffuse

Parasitoid flies (Diptera, Tachinidae) in true crickets (Orthoptera, Grylloidea): New host records from Brazil, identification key to parasitoids, and revision of host-parasitoid interactions

True crickets (Orthoptera, Grylloidea) are often parasitized by tachinid flies (Diptera, Tachinidae). However, the diversity of these parasitoids and their oviposition strategies remain unclear. Although some flies are specialized in locating crickets by their calling songs, such as the phonotactic fly Ormia ochracea (Bigot, 1889), a large portion of the tachinids that attack true crickets show different host search strategies and are adapted to parasitize other orthopteroid insects as well. However, these parasitoids have a complex and challenging taxonomy that precludes further improvement in the understanding of Tachinidae-Orthoptera interactions. Here, we described and illustrated seven new host records in Gryllidae and Phalangopsidae species from Brazil, including notes on the diagnostic characters of each parasitoid and host. An illustrated identification key to Tachinidae genera recorded in Grylloidea is also provided. Finally, all published records of Tachinidae parasitism in true crickets were revised and are presented in an annotated catalog in order to understand the host range and different oviposition strategies of each parasitoid lineage

Parasitoid flies (Diptera, Tachinidae) in true crickets (Orthoptera, Grylloidea): New host records from Brazil, identification key to parasitoids, and revision of host-parasitoid interactions

True crickets (Orthoptera, Grylloidea) are often parasitized by tachinid flies (Diptera, Tachinidae). However, the diversity of these parasitoids and their oviposition strategies remain unclear. Although some flies are specialized in locating crickets by their calling songs, such as the phonotactic fly Ormia ochracea (Bigot, 1889), a large portion of the tachinids that attack true crickets show different host search strategies and are adapted to parasitize other orthopteroid insects as well. However, these parasitoids have a complex and challenging taxonomy that precludes further improvement in the understanding of Tachinidae-Orthoptera interactions. Here, we described and illustrated seven new host records in Gryllidae and Phalangopsidae species from Brazil, including notes on the diagnostic characters of each parasitoid and host. An illustrated identification key to Tachinidae genera recorded in Grylloidea is also provided. Finally, all published records of Tachinidae parasitism in true crickets were revised and are presented in an annotated catalog in order to understand the host range and different oviposition strategies of each parasitoid lineage

FIGURE 5. Desutterella colombiana n. gen. n in Desutterella n. gen., a new genus of Luzarinae (Orthoptera: Grylloidea: Phalangopsidae) and the first report of the Aracambiae group Souza-Dias & Desutter-Grandcolas, 2014 in the Amazon

FIGURE 5. Desutterella colombiana n. gen. n. sp. General morphology.A, male habitus, dorsal; B, male head, pronotum and FW, dorsal; C, male head, pronotum and FW, lateral; D, male left FW, dorsal; E, male maxillary palpus; F, male head, frontal; G, male subgenital plate; H, male supra anal plate.Scale bar: A, 2mm; B–H, 1 mm