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Instituto Nacional de Pesquisas da Amazônia - INPA Coordenação de Biodiversidade - CBIO Divisão do Curso de Entomologia - DCEN
Taxonomia de Dasymutillini Neotropical Brothers & Lelej, 2017 com ênfase em Traumatomutilla André, 1901 (Hymenoptera: Mutillidae)
MANAUS, AMAZONAS JUNHO, 2019 ii
Pedro Reck Bartholomay
Taxonomia de Dasymutillini Neotropical Brothers & Lelej, 2017 com ênfase em Traumatomutilla André, 1901 (Hymenoptera: Mutillidae)
Orientador: Dr. Marcio Luiz de Oliveira - Instituto Nacional de Pesquisas da Amazônia (INPA)
Co-orientador: Dr. Kevin Andrew Williams - California Department of Food and Agriculture (CSCA)
Tese apresentada ao Instituto Nacional de Pesquisas da Amazônia como parte dos requisitos para obtenção do título de doutor em Ciências Biológicas (Entomologia)
MANAUS, AMAZONAS JUNHO, 2019
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Sinopse:
A taxonomia de Dasymutillini no neotrópico é abordada com ênfase no gênero Traumattomutilla resultando em mais de 10.000 espécimes examinados, cerca de 200 dos quais são espécimes tipo. São propostos dois gêneros novos na tribo, revisados os gêneros Leucospilomutilla e Cephalomutilla; são revisados ainda 12 dos 14 grupos de espécies de Traumatomutilla resultando em 68 sinonímias propostas, 61 redescrições de espécies, 13 chaves de identificações e nove espécies novas. Os resultados abrange 68 das 118 espécies atualmente válidas de Traumatomutilla tendo sido revisados 128 das mais de 180 espécies anteriormente reconhecidas para o gênero. i
“Whatever exists, he said. Whatever in creation exists without my knowledge exists without my consent. He looked about at the dark forest in which they were bivouacked. He nodded toward the specimens he'd collected. These anonymous creatures, he said, may seem little or nothing in the world. Yet the smallest crumb can devour us. Any smallest thing beneath yon rock out of men's knowing. Only nature can enslave man and only when the existence of each last entity is routed out and made to stand naked before him will he be properly suzerain of the earth.”
Cormack McCarthy, Blood Meridian. ii
Para meu avô Pedro Reck
e minhas sobrinhas
Alice e Martina.
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Agradecimentos
Agradeço primeiramente a meus pais, Maristela e Fernando, por aceitarem e entenderem o desejo de ir para tão longe, fazer algo tão estranho e com tão pouco retorno financeiro. Sou grato pelos acertos e pelos erros, pelos bons e maus exemplos, pela proteção mesmo quando excessiva, pela paciência e por acreditar que as coisas vão dar certo mesmo quando nem eu acredito mais. Vocês são um exemplo de como se deve seguir a caminhada, não importando os tropeços que damos nem as rasteiras que nos passam. Espero ser digno de tudo que vocês fizeram. Agradeço ao meu irmão Fernando e minha irmã Francisca pela paciência e por aguentarem um irmão de 33 anos de idade que “só estuda”. Ao Fernando minha admiração pela inteligência e por valores que eu admiro mesmo que os considere utópicos. Alguém sempre vai ter que lutar por essas coisas em uma sociedade saudável, fico feliz que você se sinta confiante o suficiente para levantar essas bandeiras. Sempre me sinto a vontade na sua companhia mesmo que ela seja rara. À Francisca, e seu marido Felipe, eu agradeço pelas duas lindas sobrinhas que vocês deram para a família. Fico assombrado como vocês fazem parecer simples algo que pra mim é tão difícil, construir uma vida estável e, aparentemente, nunca errar o próximo passo. Peço desculpas por ainda precisar de tanta ajuda mesmo sendo só dois anos mais novo e espero poder retribuir um pouquinho dessa bondade no futuro. É bom saber que a eficiência da dona Maristela é hereditária, talvez um dia ela apareça para mim também.
Agradeço a Paulinha, minha namorada, por tudo que passamos juntos. Não está sendo fácil, mas nos mantivemos juntos e tenho orgulho de dizer que nunca ferimos um ao outro nem mesmo nos piores momentos. Você é uma pessoa rara, bondosa, trabalhadora, guerreira e generosa, tenho muita sorte de ter te encontrado. Você é sincera e de forte personalidade, nunca tentou ser alguém que não é, nunca escondeu seu jeito de ser. Você é mais forte que eu e lhe admiro por isso. Muito obrigado, eu sei que não sou fácil, eu te amo.
Agradeço ao Minhoco, o gato do lixo, que dá muito trabalho e muito mais alegria. Um antidepressivo de quatro patas! Não foram poucas as vezes em que chegar em casa e ver ele se espreguiçando mudou o meu humor para melhor.
Aos meus amigos Williams, Kevin e Kristy, por terem me recebido durante seis meses em sua casa e permitido que eu partilhasse em igualdade do seu dia a dia. Agradeço pela oportunidade de ter conhecido o Noodle, Peixe-Boi, Wes (and his bitches), e Sauce (in memoriam). Aprendi muito com vocês e espero um dia levar uma vida tão bonita e pacífica quanto a sua. Ao Kevin em especial eu simplesmente devo quase tudo que aprendi sobre Mutillidae. Nunca conheci alguém tão inquenstionavelmente apaixonado por um grupo de insetos, com tanto conhecimento e com tanta generosidade na hora de compartilhar esse conhecimento. Parabéns por tudo que você construiu com seu esforço, dedicação e talento. Muito obrigado por tudo.
Aos colegas de longa data, Karine e Alexandre, por não terem me matado (nem matado um o outro), após 13 anos de convivência diária. Nas mesmas instituições. Na mesma sala. Todos. Os. Dias. Tenho muita admiração pelo trabalho de ambos, sei que os dois são boas pessoas e, como todos nós, estão tentando acertar mais do que errar. Espero que vocês consigam o que almejam.
Aos colegas do laboratório de Hymenoptera, Diego, Sian, Bruno, Boy, Breno e Davi. Pelos anos de convivência, intermináveis (e demasiado frequentes) pausas para o café, conversas, churrascos, bullying. Tenho muito carinho por todos vocês, não parece, mas eu tenho. iv
À turma da entomo 2012, Márlon, Sovano, Marituba, Nilba, Jejê, Patrik bb, Inaurinha, Antônio, Juci, Tati, Carol e todos aqueles que encaram a pós-graduação puramente em nome da curiosidade pela ciência, porque dinheiro nós sabemos que não dá.
À toda a equipe e amigos da California Department of Food and Agriculture - Plant Pest Diagnostics Center: Martin Hauser, Steve Gaimari, Alexey Tishechkin, Alessandra Rung, Natalia von Ellenrieder, Peter Kerr, Marc Epstein, Andy Cline, Shaun Winterton, Andrew Young, Rosser Garrison, e tantos outros que me acolheram e me ensinaram tanto. Parabéns pelo excelente trabalho que é feito em Sacramento.
Ao meu orientador Marcio Luiz de Oliveira, pelo apoio, conversas e conselhos durante os últimos sete anos. Sua ajuda foi essencial para o andamento dos trabalhos, muito obrigado.
De maneira geral a todos os amigos funcionários do INPA, professores, pesquisadores, alunos, pós-docs, faxineiras, pibics, sem os quais o instituto não funciona e que direta ou indiretamente também contribuíram com essa tese.
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Resumo
Mutillidae é uma família de vespas aculeadas cujas fêmeas são ápteras, machos geralmente alados e cujas larvas atuam como ectoparasitóides de imaturos encapsulados de outros insetos, sobretudo pupas de outros himenópteros solitários. Na região Neotropical apenas Mutillinae e Sphaeropthalminae são registradas, esta última contendo Dasymutillini que abriga cerca de um quarto das espécies reconhecidas para Mutillidae nessa região. Os gêneros neotropicais de Dasymutillini são reconhecidamente complexos e têm os limites entre si pouco definidos, com diversas espécies tendo posicionamente questionável. Muitos destes gêneros são provavelmente parafiléticos e devem ser desmembrados em múltiplos gêneros novos no futuro. Dentre tais grupos, Traumatomutilla André, responsável por mais de 180 das mais de 450 espécies de Dasymutillini, tem a taxonomia especialmente complexa e historicamente negligenciada. O dimorfismo sexual acentuado presente em Mutillidae de maneira geral faz com que Traumatomutilla tenha 136 espécies conhecidas com base apenas nas fêmeas, 48 com base apenas nos machos e duas espécies com ambos os sexos conhecidos. A descrição de mais da metade das espécies data de antes de 1920 e se baseia quase que exclusivamente em caracteres de coloração e cerdas cuja utilidade como caráter diagnóstico foi colocada em dúvida por estudos recentes. Além disso, não existem caracteres morfológicos conhecidos que possam ser usados para separar Traumatomutilla de sua contraparte norte-americana, Dasymutilla Ashmead, mesmo que ambos os gêneros formem grupos distintos filogeneticamente. O presente estudo teve como objetivo elucidar a taxonomia de Dasymutillini na região Neotropical dando ênfase à Traumatomutilla e gêneros morfologicamente próximos a este. Para tanto foram examinados mais de 10.000 espécimes de Traumatomutilla André, Cephalomutilla André, Leucospilomutilla Ashmead e Dasymutilla Ashmead, incluindo cerca de 200 espécimes-tipo. Os resultados foram: a descrição de dois gêneros novos de Dasymutillini, Atlantilla Bartholomay & Williams — contendo apenas uma espécie com base no antigo grupo de Traumatomutilla auriculata e machos da Mata Atlântica até então desconhecidos — e Quwitilla Bartholomay & Williams gen. nov. — incluindo três espécies anteriormente alocadas em Dasymutilla, Q. blattoserica (Kohl) comb. nov. (espécie-tipo), Q. peruviana (Suárez) comb. nov. e Q. bellatrix (Manley & Pitts) comb. nov.; o gênero Cephalomutilla foi revisado resultando em nove sinonímias e quatro espécies conhecidas por ambos os sexos, C. haematodes (Gerstaecker), C. zelichi Casal, C. proxima (Smith) e C. cabezona Bartholomay & Williams sp. nov.; o gênero Leucospilomutilla é revisado resultando em duas espécies conhecidas com base em ambos os sexos e uma espécie conhecida com base apenas em fêmeas; duas espécies de Dasymutilla são sinonimizadas com espécies de Traumatomutilla e uma terceira é realocada para Xystromutilla André, X. aequatorialis (André) comb. nov.; os registros sul-americanos de Dasymutilla são atualizados reduzindo a cinco o número de espécies desse gênero na região. Doze dos 14 grupos de espécie de Traumatomutilla são revisados resultando em 68 sinonímias propostas, 61 redescrições (incluindo 37 casos em que ambos os sexos foram descritos/redescritos), 38 associações sexuais, 13 chaves de identificação e nove espécies novas: T. pilkingtoni Bartholomay & Williams sp. nov., T. fratres Bartholomay & Williams sp. nov., T. anhanga Bartholomay & Williams sp. nov., T. barathra Bartholomay & Williams sp. nov., T. pantherina Bartholomay & Williams sp. nov., T. poranga Bartholomay & Williams sp. nov., T. juvenindica Bartholomay & Williams sp. nov., T. peismatara Bartholomay & Cambra sp. nov., and T. tetratrauma Bartholomay & Williams sp. nov. Três novos registros de hospedeiros são relatados: T. ocellaris (Klug) parasitando Bicyrtes sp.; e T. diopthalma parasitando Podium sp. (em condições de laboratório) e Trypoxylon sp. (em ninhos armadilha). Os resultados incluídos na presente tese abrangem 68 das 118 espécies atualmente válidas de Traumatomutilla tendo sido revisados 128 das mais de 180 espécies anteriormente reconhecidas para o gênero, com a adição de três espécies anteriormente em status de incertae sedis, Mutilla tenuis (Smith), Mutilla polita (Smith) e Mutilla impetuosa (Smith). vii
Abstract
Mutillidae is a family of aculeate wasps in which the females are always apterous, males usually fully winged, and whose larvae act as ectoparasitoids of encapsulated imatures of other insects, especially pupae of other solitary Hymenoptera. Only Mutillinae and Sphaeropthalminae are recorded for the Neotropical Region, with the latter housing the Dasymutillini which in turn encompasses one fourth of the Neotropical species of Mutillidae. The Neotropical genera of Dasymutillini are notoriously complex, with poorly defined limits, several doubtful species placements, many of which are likely paraphyletic and will probably be split into multiple new taxa in the future. One such genus is Traumatomutilla André, which holds over 180 of the approximately 450 species of the tribe, and whose taxonomy is especially complex as it is historically neglected. The sexual dimorphism typocal of Mutillidae in general resulted in numerous species of Traumatomutilla being described based on females (136) or males (48), with only two species being known from both sexes. The description of more than half of these species dates from before 1920 e it is based almost exclusively on color and setae characters whose diagnostic usefulness has been put into question by recent studies. Additionally, there are no morphological characters that can be reliably used to separate Traumatomutilla from its north american counterpart, Dasymutilla Ashmead, even though both genera form phylogenetically distinct groups. The present study aimed to elucidate the taxonomy of neotropical Dasymutillini emphasizing the Traumatomutilla species and its morphological allies. For this, more than 10.000 specimens of Traumatomutilla, Cephalomutilla André, Leucospilomutilla Ashmead and Dasymutilla were examined, including more than 200 type-specimens. The results are as follows: two new genera were described, Atlantilla Bartholomay & Williams gen. nov. — containing only one species based on the former species group of Traumatomutilla auriculata and males of the Atlantic Forest hitherto undescribed — and Quwitilla Bartholomay & Williams— to include three species formerly place within Dasymutilla, Q. blattoserica (Kohl) comb. nov. (type species), Q. peruviana (Suárez) comb. nov., and Q. bellatrix (Manley & Pitts) comb. nov.; the genus Cephalomutilla is revised resulting in nine proposed synonyms and four species known from both sexes, C. haematodes (Gerstaecker), C. zelichi Casal, C. proxima (Smith), and C. cabezona Bartholomay & Williams sp. nov.; the genus Leucospilomutilla is also revised resulting in two species known from both sexes and one species known from females only; two species of Dasymutilla are synonymized with previously known species of Traumatomutilla and a third one is transferred to Xystromutilla André, X. aequatorialis (André) comb. nov.; the South American records for Dasymutilla species are updated reduing to five the number of species of this genus to occur in the region. Twelve of the 14 species groups of Traumatomutilla are revised resulting in 68 proposed synonyms, 61 redescriptions (including 37 cases in which both sexes had to be described/redescribed), 38 sex associations, 13 identifcation keys, and nine new species: T. pilkingtoni Bartholomay & Williams sp. nov., T. fratres Bartholomay & Williams sp. nov., T. anhanga Bartholomay & Williams sp. nov., T. barathra Bartholomay & Williams sp. nov., T. pantherina Bartholomay & Williams sp. nov., T. poranga Bartholomay & Williams sp. nov., T. juvenindica Bartholomay & Williams sp. nov., T. peismatara Bartholomay & Cambra sp. nov., and T. tetratrauma Bartholomay & Williams sp. nov. Three new host records are also provided for Traumatomutilla: T. ocellaris (Klug) parasitizing Bicyrtes sp.; and T. diopthalma (Klug) parasitizing Podium sp. (in laboratory) and Trypoxylon sp. (in trap nests). The results included in the current thesis encompass 68 of the 118 currently valid species of Traumatomutilla, having revised 128 of the more than 180 species formerly recognized within the genus with the addition of three species formerly in incertae sedis status, Mutilla tenuis (Smith), Mutilla polita (Smith), and Mutilla impetuosa (Smith). vii
Sumário
1. Introdução Geral ...... 1
2. Objetivos ...... 8
2.1 Objetivo geral ...... 8
2.2 Objetivos específicos ...... 8
3. Material e Métodos ...... 9
4. Resultados e Discussões ...... 13
4.1 Capítulo 1: “Revision of Cephalomutilla André Hymenoptera: Mutillidae) including one new species, three new sex associations, and ten new synonymies” ...... 13
4.2 Capítulo 2: “Revision of Leucospilomutilla Ashmead and description of the new genus Atlantilla (Hymenoptera: Mutillidae)” ...... 33
4.3 Capítulo 3 (Publicado): “Does the genus Dasymutilla Ashmead occur in South America? Quwitilla gen. nov., a new genus, new combinations and new distribution records for Neotropical velvet ants (Hymenoptera: Mutillidae)” ...... 55
4.4 Capítulo 4 (Publicado): “Traumatomutilla André Miscellanea: Revision of the bellica, bifurca, diabolia and vitelligera species group, and a new group for the new species T. pilkingtoni (Hymenoptera: Mutillidae: Spaheropthalminae: Dasymutillini)” ...... 56
4.5 Capítulo 5 (Publicado): “New species of Traumatomutilla André in the T. tabapua and T. integella species-groups (Hymenoptera, Mutillidae)” ...... 57
4.6 Capítulo 6 (Publicado): “Revision of the Traumatomutilla americana species-group (Hymenoptera: Mutillidae)” ...... 58
4.7 Capítulo 7: “Revision of the Traumatomutilla juvenilis species-group (Hymenoptera: Mutillidae)” ...... 59
4.8 Capítulo 8: “Revision of the Traumatomutilla gemella species-group (Hymenoptera: Mutillidae)” ...... 88
4.9 Capítulo 9: “Revision of the Traumatomutilla indica species-group (Hymenoptera: Mutillidae)” 110
4.10 Capítulo 10: “Revision of the Traumatomutilla quadrinotata species-group (Hymenoptera: Mutillidae)” ...... 186 vii
4.11 Capítulo 11: “Species groups of Traumatomutilla (Mutillidae: Sphaeropthalminae: Dasymutillini): updates, keys and redescriptions to the known males”...... 227
5. Considerações finais ...... 264
6. Referências ...... 268 1
1. Introdução geral
Mutillidae é uma família de vespas solitárias reconhecida principalmente por suas fêmeas ápteras com cerdas densas e padrões de coloração complexos e chamativos (Brothers, 2006). Os machos são quase sempre alados, podendo ser ápteros ou raramente braquípteros, e geralmente possuem cerdas e coloração mais discreta que as fêmeas (Brothers, 2006). As larvas de mutilídeos atuam como ectoparasitóides de imaturos encapsulados de outros insetos, geralmente pupas ou pré-pupas de outros himenópteros solitários (Apidae, Crabronidae, Sphecidae, Pompilidae, etc.) (Brothers, 2006; Luz et al. 2016a). Entretanto, existem registros de mutilídeos parasitando outras ordens de insetos como Diptera (Tephritidae), Lepidoptera (Limacodidade), Coleoptera (Chrysomelidae) e Blattaria (Polyphagidae) (Brothers, 1989; Amini et al. 2014). Tal diversidade de hospedeiros se deve em parte ao fato de que, segundo Brothers (1989), mutilídeos não são “hospedeiro- específicos” e sim “situação-específicos”. Sendo assim, para que determinado grupo seja considerado como hospedeiro potencial, basta que os imaturos estejam encapsulados (pupa, pré-pupa, ooteca, etc) e forneçam espaço/alimento suficientes para o desenvolvimento da larva do parasitóide. Considerando-se que a lista de hospedeiros de Mutillidae é baseada em pouco mais de 3% das espécies conhecidas para a família, é presumível que a diversidade de grupos parasitados se torne maior com futuros estudos (Brothers, 2006). Espécies de Mutillidae são encontradas em todos os continentes, não ocorrendo nos pólos e sendo extremamente raras em altas latitudes (Williams, 2012). São conhecidas cerca de 5.000 espécies em todo mundo, mais de 1.500 das quais registradas para a região Neotropical em aproximadamente 50 gêneros (Brothers, 2006). Recentemente, diferentes espécies de mutilídeos foram utilizadas com sucesso para estudos biogeográficos (Wilson & Pitts, 2012; Wilson et al., 2012a), de mimetismo (Wilson et al., 2012b, 2013, 2018) e ecológicos (Boehme et al. 2012). Entretanto, sua a utilização em estudos aplicados é parcialmente inviabilizada pelos inúmeros problemas taxonômicos encontrados no grupo, derivados principalmente do dimorfismo sexual acentuado e variabilidade dos padrões de coloração e cerdas comumente utilizadas na maior parte das descrições de espécies da família (Brothers, 2006; Williams, 2012; Williams et al., 2012). Desde a filogenia recente de Brothers & Lelej (2017) o consenso aceito é de que a família pode ser dividida em oito subfamílias, Myrmosinae, Pseudophotopsidinae, Rhopalomutillinae, Ticoplinae, Sphaeropthalminae, Myrmillinae, Dasylabrinae e Mutillinae. Destas, apenas Sphaeropthalminae e Mutillinae ocorrem no Novo Mundo, representadas pelas tribos Sphaeropthalmini, Dasymutllini, Pseudomethocini para Sphaeropthalminae, assim como Mutillini (Ephutina) e Trogaspidiini para Mutillinae (Brothers & Lelej, 2017). Dasymutillini inclui espécies da Austrália, Neártico e Neotrópico, abrigando aproximadamente um quarto da fauna conhecida de Mutillidae no Novo Mundo com cerca de 450 espécies descritas (Brothers & Lelej 2017; Williams, 2012). Membros da tribo podem ser identificados pelas seguintes características: ambos os sexos com o primeiro segmento metasomal estreito e geralmente peciolado, pigídio geralmente com uma área pigidial bem definida por carenas laterais, franjas metasomais sem cerdas plumosas e machos geralmente com as axilas projetadas na forma de protuberâncias agudas ou truncadas (Williams, 2012). Além disso, quase todos os membros da tribo são diurnos e possuem coloração vívida (Williams, 2012). Os gêneros neotropicais 2
incluídos dentro de Dasymutillini são: Cephalomutilla André, 1908b; Dasymutilla Ashmead, 1899; Frigitilla Williams, 2015; Gogoltilla Williams, Brothers & Pitts, 2011a; Leucospilomutilla Ashmead, 1903; Lomachaeta Mickel, 1936; Protophotopsis Schuster, 1947; Reedomutilla Mickel, 1964, Suareztilla Casal, 1968; Tobantilla Casal, 1965; e Traumatomutilla André, 1901, este último, gênero foco do presente estudo. Com mais de 180 espécies, Traumatomutilla é o segundo gênero de Dasymutillini em diversidade e o terceiro para a família na região Neotropical. Proposto por André (1901) como um subgênero do grande grupo Mutilla Linnaeus, inicialmente com base em apenas algumas características superficiais e abrigando um pequeno número de espécies típicas da América do Sul, sem mencionar quaisquer comentários a respeito da relação do novo subgênero com outros subgêneros ou gêneros conhecidos à época. Posteriormente, André (1902) realocou o subgênero Traumatomutilla para o gênero Ephuta Say, 1836 juntamente com uma descrição mais completa do grupo e designando Mutilla indica Linnaeus, 1758 como espécie-tipo. No mesmo estudo foram listadas aproximadamente 130 espécies que, de acordo com o autor, pertenceriam a Traumatomutilla, algumas das quais hoje pertencem aos gêneros Suareztilla e Cephalomutilla. Ashmead (1904) passou a considerar Traumatomutilla como sinônimo de Sphaeropthalma, o que é contestado por André (1904) que revalida o subgênero. Os trabalhos subsequentes de André (1905, 1906, 1908a) descrevem novas espécies de Traumatomutilla ainda como subgênero de Ephuta. André (1908b), entretanto, ao descrever Traumatomutilla fissiventris (André, 1908b), eleva o então subgênero a status de gênero sem quaisquer comentários ou designação formal da mudança. O catálogo de gêneros de Mutillidae elaborado por Lelej & Brother (2008) inclui Traumatomutilla como gênero válido, mencionando a sinonimização feita por Ashmead (1904) e André (1904) como o autor que revalida o status de gênero. André (1904) claramente se refere à Traumatomutilla apenas como subgênero de Ephuta, em nenhum momento indicando estar elevando o nome à categoria de gênero. Sendo assim, o consenso adotado entre especialistas na família é de que André em algum momento entre 1904 e 1908 passou a considerar Traumatomutilla como um gênero válido, ou por suas espécies serem morfologicamente distintas dos outros gêneros existentes à época, ou por considerar que não mais pertenceria a Ephuta. Traumatomuitlla passou a ser adotado como gênero por autores subsequentes e o grupo de fato possui características morfológicas que o distingue dos demais Dasymutillini à exceção de Dasymutilla (Casal, 1969; Suárez, 1960; Nonveiller, 1990). André (1902) cita sete características como diagnósticas para Traumatomutilla: cabeça da mesma largura que o tórax ou mais estreita que o mesmo; olhos protuberantes, lisos e brilhantes; mandíbulas com ápice acuminado; tórax ovalado e alongado, estreitado posteriormente, pouco ou nada constringido medialmente, com as laterais simples ou "mais ou menos tuberculadas"; abdome peciolado e com o último segmento (sexto tergito metasomal) com uma área pigidial geralmente bem distinguível; segundo segmento abdominal com duas manchas e, mais frequentemente, quatro manchas de cor vermelho-sangue ou amarelo- avermelhado. Em relação aos machos do gênero, as características propostas no mesmo estudo são: cabeça pequena; olhos inteiros, sem emarginações, convexos e brilhantes como nas fêmeas; tórax ovóide com escutelo não saliente e epauletas pequenas; abdome "bastante" peciolado; asas providas de três células cubitais e duas veias recorrentes. A descrição de ambos os sexos é evidentemente subjetiva e as características podem 3
ser encontradas em diversos outros gêneros do Novo e Velho Mundo, o que ocasionou no posicionamento de espécies morfologicamente distintas dentro do gênero (obs. pes.). As fêmeas de Traumatomutilla atualmente são separadas dos demais gêneros de Sphaeropthalminae por uma combinação exclusiva de características que incluem principalmente a presença de uma placa pigidial definida por carenas laterais no sexto tergito, a largura da cabeça igual ou menor que a do mesosoma e o primeiro tergito subpeciolado (Brothers, 2006). Enquanto os machos possuem os ângulos axilares projetados na forma de tubérculos truncados ou agudos, mandíbulas simples, sem dentes ou emarginações, primeiro tergito metasomal peciolado e escutelo convexo (Brothers, 2006). Os caracteres utilizados por André (1901) para descrever Traumatomutilla são quase idênticos aos utilizados por Ashmead (1899) na diagnose de Dasymutilla o que torna sua separação um empecilho recorrente nos estudos com mutilídeos neotropicais. As dificuldades em se separar Traumatomutilla e Dasymutilla derivam em parte do fato destes serem os gêneros mais diversos de Dasymutillini, com mais de 180 e 200 espécies reconhecidas respectivamente (Williams 2012; Williams et al. 2017). Além disso, ambos são extremamente variáveis e notadamente difíceis de serem diferenciados morfologicamente (Manley & Pitts 2007; Williams et al. 2017). De fato, a única chave de identificação existente para gêneros neotropicais de Mutillidae inclui Traumatomutilla e Dasymutilla no mesmo terminal fazendo referência ao fato de que os dois táxons possuem características morfológicas que se confundem e acenando com a possibilidade de serem sinonimizados ou divididos em múltiplos gêneros no futuro (Brothers, 2006). Até o momento nenhuma combinação de caracteres estruturais pode ser utilizada para diferenciar Traumatomutilla de Dasymutilla com eficiência, pois os caracteres anteriormente tidos como úteis para a diagnose dos gêneros mostraram-se polimórficos dentro dos mesmos ou são compartilhados por ambos. Por exemplo, a cor do esporão das tíbias geralmente é branca ou amarelo-pálida em Traumatomutilla e preta em Dasymutilla (Quintero & Cambra, 1996). Entretanto, aproximadamente 20 espécies de Traumatomutilla possuem os esporões de cor preta enquanto que mais de 20 espécies de Dasymutilla possuem esporões brancos (obs. pes.). Outras características possuem relação semelhante entre ambos os gêneros podendo ser usadas para separar a vasta maioria das espécies, mas ineficientes para separar os gêneros como um todo. Atualmente o “caráter” que melhor pode ser utilizado para separar Traumatomutilla de Dasymutilla é sua presença na América do Sul ou do Norte (incluindo América Central e Grandes Antilhas) respectivamente. A vasta maioria das espécies de Dasymutilla ocorrem na América do Norte com cerca de dez representantes ocorrendo na América do Sul, enquanto apenas três das 186 espécies de Traumatomutilla ocorrem ao norte do istmo do Panamá (Manley & Pitts, 2007; Williams, 2012, Williams et al. 2017). Em uma tentativa de elucidar parcialmente essa questão, Williams (2012) propôs uma filogenia focada em Dasymutilla incluindo representantes de outros quatro gêneros neotropicais de Dasymutillini, entre eles 18 espécies de Traumatomutilla. As relações filogenéticas recuperadas forneceram indícios de que apesar de Dasymutilla e Traumatomutilla formarem clados distintos, ambos são parafiléticos. Os táxons de ambos os gêneros recuperados fora do clado principal dos mesmos segundo Williams (2012), possuem característcas morfológicas exclusivas e/ou distribuição atípica em relação às demais espécies. Segundo o mesmo autor, os 4
resultados indicavam a existência de dez ou mais possíveis novos gêneros dentro de Traumatomutilla e Dasymutilla algumas dessas espécies incomuns seriam realocadas a gêneros distintos em trabalhos subsequentes. Em nível de espécie, a taxonomia de Traumatomutilla sofre de problemas que atrasam o avanço no conhecimento de Mutillidae como um todo. O estudo de Williams et al. (2017) foi o primeiro a focar na taxonomia do gênero desde Casal (1969), dividindo as 135 espécies do gênero conhecidas com base nas fêmeas em grupos de espécie que patrilham características exclusivas ou conjuntos de características exclusivas. Mesmo com esse avançao significativo, a vasta maioria das espécies do gênero foi sucintamente descrita antes da década de 1920 com base em características subjetivas de coloração e/ou cerdas, ignorando quase que completamente quaisquer caracteres estruturais (e.g. Klug, 1821; Burmeister, 1845; Gerstaecker, 1874; Burmeister, 1875; Cresson, 1902). A vasta maioria das espécies descritas durante esse período jamais foi abordada em quaisquer outros estudos após sua descrição original e, até o estudo de Williams et al. (2017), seus dados de distribuição estavam restritos às localidades-tipo. Ademais, comentários a respeito de variações de coloração e ou estrutura são virtualmente inexistentes na bibliografia do gênero, muito em virtude de que quaisquer diferenças na coloração em relação a uma espécie previamente descrita tornaram-se a base para diagnose de uma nova espécie (Cresson, 1902; Casal, 1969). A dependência da utilização de caracteres de coloração e cerdas para separação de espécies em Mutillidae torna-se contestável em virtude de trabalhos recentes que demonstraram a existência de complexos miméticos envolvendo dezenas de espécies na América do Norte (Wilson et al. 2012, 2013) e África (Wilson et al. 2018). As implicações dessas novas descobertas ficaram evidenciadas em estudos como os de Williams et al. (2011a, b, 2012) nos quais foi descoberto que diversas espécies ou, em alguns casos, grupos de espécies inteiros se tratavam na verdade de uma única espécie amplamente distribuída e com variações extremas de coloração em decorrência da influência do padrão mimético predominante nas áreas em que os espécimes foram coletados. Dados preliminares a respeito da fauna Sul-Americana já dão indícios de que situações semelhantes ocorrem na região, com algumas síndromes de coloração conspicuamente predominantes em determinadas regiões e aparentemente restritas a um bioma em especial (Kevin Williams com. pes.). Outro grande empecilho nos avanços taxonômicos de Traumatomutilla existe em decorrência do dimorfismo sexual acentuado que permeia Mutillidae como um todo. Associações sexuais na família são raras e de difícil confirmação sem o auxílio de observações de casais coletados in copulo, observações de cópula em campo, dados moleculares, casais coletados no mesmo ninho de hospedeiro, ou grandes séries de espécimes que forneçam dados consistentes para associações com base em distribuição ou similaridades morfológicas (Williams et al. 2011b; Luz & Williams, 2014; Luz et al. 2016b; Lopez et al., 2019). No caso de Traumatomutilla apenas duas espécies do gênero são conhecidas com base em ambos os sexos, T. dubia (Fabricius, 1804) e T. bispiculata (André, 1906) (Williams et al. 2017). Esse dado torna-se ainda mais preocupante em virtude da enorme quantidade de machos presente em coleções e coletados em diversas regiões da América do Sul os quais podem superar as fêmeas na proporção de 20 para 1 em alguns casos (obs. pes.). Tais espécimes, virtualmente inutilizados pela ausência de meios de identificação, constituem uma fonte 5
inestimável de dados de distribuição e possíveis caracteres morfológicos com potencial para auxiliar num melhor entendimento de Traumatomutilla e sua relação com os demais gêneros de Dasymutillini. As 46 espécies de Traumatomutilla conhecidas apenas com base nos machos não foram contempladas no estudo de Williams et al. (2017) assim como os machos de T. dubia e T. bispiculata pelas mesmas razões que inviabilizam uma maior utilização de machos de Traumatomutilla na taxonomia do gênero. Primeiramente, existem muito mais espécies conhecidas com base nas fêmeas (138 incluindo as fêmeas de T. dubia e T. bispiculata), o que sugere que a maior parte da diversidade dos machos não é conhecida. Além disso, caracteres de genitália jamais foram utilizados ou até mesmo mencionados na taxonomia de machos Traumatomutilla, com as descrições focando apenas em caracteres de coloração e cerdas (Gerstaecker, 1874; Cresson 1902), com poucos casos utilizando características estruturais (Mickel, 1952), o que limita a viabilidade e confiabilidade de quaisquer identificações feitas com base na literatura. Finalmente, a existência de apenas duas espécies conhecidas com base em ambos os sexos torna impossível que seja feita uma diagnose de grupos de espécie com base em machos e fêmeas. De um modo geral, um estudo taxonômico aprofundado sobre Dasymutillini focando em uma atualização e maior organização de Traumatomutilla, justifica-se por três principais causas: 1. Sua provável parafilia indicando que algumas espécies, ou mesmo grupos de espécie, pertencem a gêneros distintos o que acarreta a necessidade de maior definição dos limites entre Traumatomutilla e os demais Dasymutillini neotropicais; 2. A defasagem na taxonomia de Traumatomutilla cujas espécies são pobremente descritas e ilustradas, virtualmente impossíveis de serem identificadas sem auxílio de especialistas, material-tipo ou material de referência; 3. A quase que ausência de conhecimento a respeito dos machos do gênero incluindo possíveis associações sexuais, seu posicionamento no contexto dos grupos de espécie e a utilidade dos caracteres de genitália. Os 11 capítulos que compõem a presente tese constituem um avanço significativo na taxonomia dos Dasymutillini neotropicais sobretudo para Traumatomutilla cuja vasta maioria das espécies passa a ser acessível para não-especialistas. A sequência dos capítulos apresentados segue uma ordem lógica para facilitar o entendimento de como as descobertas expostas em cada estágio do estudo influenciaram nos resultados e conclusões subsequentes. A ordem de publicação dos capítulos não segue a sequência apresentada aqui. Capítulo um (4.1): aborda o gênero Cephalomutilla André, cujas fêmeas distinguem-se de Traumatomutilla apenas com base na largura do vértice em relação ao corpo, enquanto os machos, até recentemente, estavam em status de incertae sedis ou alocados dentro de Traumatomutilla. Nove sinonímias são propostas, duas delas através de associações sexuais com base na distribuição de machos e fêmeas. Ambos os sexos de todas as espécies válidas são ilustrados e inseridos em uma chave de identificação incluindo uma nova espécie. Aspectos da distribuição das espécies de Cephalomutilla e sua possível participação em complexos de mimetismo Mulleriano são discutidos. Capítulo dois (4.2): o novo gênero Atlantilla Bartholomay & Williams gen. nov. é descrito com base nas fêmeas do extinto grupo de Traumatomutilla auriculata e machos recentemente coletados na Mata Atlântica. O gênero Leucospilomutilla Ashmead também é revisado por ser morfologicamente muito similar 6
ao novo gênero proposto. As relações e similaridades morfológicas entre ambos os gêneros e Traumatomutilla é discutida. Chaves e ilustrações para as três espécies de Leucospilomutilla também são incluídas. Capítulo três (4.3): aceito para publicação no periódico Zootaxa, trata das espécies de Dasymutilla Ashmead que ocorrem na América do Sul. Um novo gênero, Quwitilla Bartholomay & Williams gen. nov., é proposto com base em espécies andinas anteriormente alocadas em Dasymutilla. Duas espécies Sul- Americanas de Dasymutilla são sinonimizadas com espécies de Traumatomutilla e uma terceira espécie é transferida para o gênero Xystromutilla André. A distribuição de Dasymutilla na América do Sul também é discutida. Por questões legais referentes à política de acesso livre às publicações da Zootaxa apenas a primeira página da primeira prova do artigo é incluída na presente tese. Uma cópia do artigo original (caso publicado a tempo) ou da primeira prova na íntegra será fornecida aos membros da banca em separado. Capítulo quatro (4.4): aceito para publicação no periódico Insecta Mundi, são revisados os menores grupos de espécie de Traumatomutilla: grupo de T. vitelligera, T. bifurca, T. bellica e T. diabolica. Um novo grupo de espécies é proposto para incluir a nova espécie T. pilkingtoni Bartholomay & Williams sp. nov. Três sinonímias e duas associações sexuais com base na distribuição de ambos os sexos são propostas. A distribuição dos cinco grupos de espécie abordados é discutida assim como o próprio esquema de divisão do gênero em grupos de espécie. Capítulo cinco (4.5): publicado recentemente no periódico Zootaxa — 4433 (2): 361–385 — abordando a revisão dos grupos de espécie de T. integella e T. tabapua. Uma nova sinonímia é proposta, todas as espécies previamente conhecidas de ambos os grupos são redescritas e cinco novas espécies são descritas. A possível existência de complexos miméticos dentro da região Amazônica é discutida brevemente assim como as variações de coloração encontradas em algumas das espécies abordadas. Por questões legais referentes à política de acesso livre às publicações da Zootaxa apenas a primeira página do artigo é incluída na presente tese. Uma cópia do artigo original será fornecida aos membros da banca em separado. Capítulo seis (4.6): também recentemente publicado no periódico Zootaxa — 4608 (1): 001–034 — trata da revisão do grupo de Traumatomutilla americana incluindo 14 novas sinonímias. Todas as espécies e variantes de coloração do grupo são incluídas em chaves de identificação e ilustradas. Os machos das três espécies reconhecidas para o grupo são associados, ilustrados e incluídos em chaves de identificação. As associações sexuais se baseiam na distribuição de ambos os sexos, casais coletados in copulo e/ou observações de comportamento de cópula observado in situ. A primeira associação de hospedeiro para Traumatomutilla na América do Sul é realizada com base em observações de campo. A metodologia empregada nas medidas e descrições de Traumatomutilla é elucidada e ilustrada. São fornecidos mapas de distribuição para as três espécies válidas do grupo de T. americana e aspectos da biologia das espécies do grupo são brevemente discutidos. Assim como no capítulo cinco, apenas a primeira página do artigo original é incluída no corpo da tese e uma cópia do artigo será enviada em separado para os membros da banca pelas razões anteriormente citadas. Capítulo sete (4.7): o grupo de T. juvenilis é revisado e reduzido a cinco espécies conhecidas por ambos os sexos resultando em 10 sinonímias propostas e uma nova espécie descrita, T. juvenindica 7
Bartholomay & Williams sp. nov. Todas as espécies são redescritas, ilustradas e inseridas em uma chave de identificação para fêmeas e machos separadamente. Uma chave de identificação para as variantes de coloração das fêmeas do grupo também é fornecida. A distribuição atípica de algumas espécies do grupo de T. juvenilis e sua possível participação em complexos miméticos na América do Sul é brevemente discutida. Capítulo oito (4.8): trata da revisão do pequeno grupo de espécies de T. gemella. São reconhecidas cinco espécies válidas no grupo, incluindo uma nova espécie, T. peismatara Bartholomay & Cambra sp. nov. Quatro dessas cinco espécies têm ambos os sexos associados, descritos (ou redescritos), ilustrados e incluídos em chaves de identificação. Duas sinonímias são propostas com base em variantes de coloração encontradas no material examinado. Dois novos registros de hospedeiro são feitos com base em experimentos de ninho armadilha, constituindo os primeiros registros de espécies de Traumatomutilla parasitando espécies que não nidificam no solo. As implicações desses novos registros são brevemente discutidas. Capítulo nove (4.9): trata do grande grupo de Traumatomutilla indica resultando em 12 espécies conhecidas por ambos os sexos, quatro conhecidas com base apenas em fêmeas e duas com base apenas nos machos. São propostas 27 sinonímias e uma nova combinação baseada em um macho anteriormente em status de incertae sedis. Todas as espécies válidas são redescritas, ilustradas e incluídas em chaves de identificação para ambos os sexos (quando conhecidos). Comentários a respeito de variações morfológicas, possíveis sinonímias ou associações sexuais futuras são fornecidos para diversas espécies. A morfologia do grupo de T. indica e sua relação com os demais grupos de espécie em Traumatomutilla é discutida assim como paralelos morfológicos encontrados em ambos os sexos de algumas espécies. Capítulo dez (4.10): o grupo de Traumatomutilla quadrinotata é revisado resultando em seis espécies conhecidas por ambos os sexos, quatro conhecidas com base em fêmeas apenas e duas espécies conhecidas com base apenas nos machos. São propostas onze novas sinonímias e uma nova espécie, T. tetratrauma Bartholomay & Williams sp. nov., com base em casais coletados na Mata Atlântica. Todas as espécies válidas são redescritas, ilustradas e incluídas em chaves de identificação para ambos os sexos (quando conhecidos). Comentários a respeito de variações morfológicas, possíveis sinonímias ou associações sexuais futuras são fornecidos para diversas espécies. A morfologia do grupo de T. quadrinotata e sua relação com os demais grupos de espécie em Traumatomutilla é discutida. Capítulo onze (4.11): o restante dos machos não associados de Traumatomutilla são redescritos, ilustrados e inseridos em uma chave de identificação ilustrada para os grupos de espécie do gênero com base nos machos. As prováveis futuras associações sexuais dos machos não-associados são discutidas assim como sua relação com as fêmeas dos grupos de espécie remanescente, grupos de T. inermis e T. trochanterata. O macho de T. virginalis (Gerstaecker, 1874) é descrito e ilustrado. 8
2. Objetivo Geral
- Atualizar a taxonomia de Dasymutillini na América do Sul elucidando os limites entre gêneros morfologicamente próximos à Traumatomutilla visando tornanr este grupo mais acessível e identificável para não-especialistas.
2.1 Objetivos específicos
- Realocar espécies equivocadamente inseridas em Traumatomutilla pela ausência de um dos sexos ou por defasagem do conhecimento taxonômico em relação às mesmas, propondo novos gêneros ou revisando os gêneros aos quais elas de fato pertencem;
- Descrever e associar os machos dos grupos de espécies de Traumatomutilla mais comumente encontrados e coletados;
- Revisar as espécies dos grupos de espécie de Traumatomutilla mais comumente encontrados e coletados;
- Ilustrar e redescrever o material-tipo dos grupos de espécies de Traumatomutilla mais comumente encontrados e coletados, inserindo as espécies em chaves de identificação acessíveis para não especialistas. 9
3 Material e métodos
Mais de 10.000 espécimes das seguintes 68 instituições foram examinados ao longo do presente estudo:
AMNH - American Museum of Natural History, New York, New York, USA ANSP - Academy of Natural Sciences, Philadelphia, Pennsylvania, USA BMNH - British Museum of Natural History, London, United Kingdom BYU - Brigham Young University, Monte L. Bean Life Science Museum, Provo, Utah, USA CASC - California Academy of Sciences, San Francisco, California, USA CESC - Coleção Entomológica de Santa Cruz do Sul, Universidade de Santa Cruz do Sul, Santra Cruz do Sul, Rio Grande do Sul, Brasil CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA CISC - Essig Museum of Entomology, Department of Entomological Sciences, University of California, Berkeley, California, USA CPDC - Centro de Pesquisas do Cacau, CEPEC, CEPLAC, Divisão de Zoologia Agrícola, Itabuna, Bahia, Brazil CSCA - California State Collection of Arthropods, Sacramento, California, USA CUIC - Cornell University Insect Collection, Department of Entomology, Ithaca, New York, USA CZMA - Coleção Entomológica do Maranhã, Caxias, Maranhão, Brasil DEI - Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany DGMC - Donald G. Manley Collection, Florence, South Carolina, USA DJBC - Denis J. Brothers Collection, a ser depositada em Iziko South African Museum (SAMC), Cape Town, South Africa DZUP - Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brasil EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA FSCA - Florida State Collection of Arthropods, Gainesville, Florida, USA FMNH - Fields Museum of Natural History, Chicago, Illinois, Brasil HNHM - Hungarian Natural History Museum, Budapest, Hungary IaVH - Instituto Alexander Von Humboldt, Villa de Leyva, Boyacá, Colombia IEPA - Instituto de Pesquisas Científicas e Tecnológicas do Estado do Amapá, Macapá, Amapá, Brazil INPA - Instituto Nacional de Pesquisas da Amazônia, Coleção de Invertebrados, Manaus, Amazonas, Brasil LACM - Insect Collection, Los Angeles County Museum of Natural History, Los Angeles, California, USA 10
LRRP - Laboratório de Sistemática e Bioecologia de Parasitóides e Predadores da Agência Paulista de Teconologia dos Agronegócios (APTA), Ribeirão Preto, São Paulo, Brazil LSUK - Linnean Society United Kingdom, London, England MACN - Museo Argentino de CIenias Naturales, Buenos Aires, Buenos Aires, Argentina MCZC - Museum of Comparative Zoology, Harvard Univeristy, Cambridge, Massachusetts, USA MIUP - Museo de Invertebrados G.B. Fairchild, Universidad de Panamá, Ciudad de Panamá, Panamá MLUH - Martin Luther University Halle-Wuttenberg, Halle, Germany MLPA – Museo de La Plata, La Plata, Buenos Aires, Argentina MNCN - Museo Nacional de Ciencias Naturales, Madri, Espanha MNHN - Museum National d’Histoire Naturelle, Paris, France MNRJ* - Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brasil MPEG - Museu Paraense Emílio Goeldi, Belém, Pará, Brasil MRSN - Museo Regionale de Scienze Naturali, Torino, Italia MuBio - Museu da Biodiversidade, Universidade Federal da Grande Dourados, Dourados, Mato Grosso do Sul, Brasil MZSP - Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brasil NHMB - Natural History Museum Basel, Basel, Switzerland NHMW - Natural History Museum Viena, Viena, Austria NHRM - Natural History Museum, Stockholm, Sweden PMAE - Royal Alberta Museum, Edmonton, Alberta, Canada PMNH - Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA RBINS - Royal Belgian Institute of Natural Sciences, Brussels, Belgium RMNH - Naturalis Biodiversity Center [formerly Rijksmuseum van Natuurlijke Historie], Leiden, Netherlands SEMC - Snow Entomological Museum, Univeristy of Kansas, Lawrence, Kansas, USA TAMU - Texas A & M University, College Station, Texas, USA UCDC - The Bohart Museum of Entomology, University of California, Davis, California, USA UCRC - Entomology Research Museu, Department of Entomology, Univeristy of California, Riverside, California, USA UEFS - Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brasil UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brasil UFMG - Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brasil UFMT - Universidade Federal do Mato Grosso, Cuiabá, Mato Grosso, Brasil UFRN - Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte, Brasil UFRR - Universidade Federal de Roraima, Boa Vista, Roraima, Brasil UFT - Universidae Federal do Tocantins, Palmas, Tocantins, Brasil UMMZ - University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA 11
UMSP - University of Minnesota, Saint Paul, Minnesota, USA UNEMAT - Universidade do Estado de Mato Grosso, Tangará da Serra, Mato Grosso, Brazil USNM - United States Natural History Museum, Smithsonian Institution, Washington D.C., USA UNSM - University of Nebraska State Museum, Lincoln, Nebraska, USA ZIN - Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia ZMB - Zoological Museum Berlin, Berlin, Germany ZMUC - Zoological Museum, University of Copenhagen, Copenhagen, Denmark *Material destruído no incêndio de 02/09/2018. Dados provenientes da planilha elaborada pelo Dr. Kevin Williams em visita ao MNRJ em 2013.
A terminologia morfológica e medidas utilizadas foram baseadas em três principais estudos: Harris (1979), Bartholomay et al. (2018) (Capítulo 5), Cambra et al. (2018), com a adição das correções de termos e medidas propostas no capítulo 6. As abreviações T1–6 e S1–6 nas fêmeas e T1–7 e S1–7 nos machos referem-se os tergitos e esternitos metasomais respectivamente. Tais abreviações estão em concordância com a terminologia utilizada em estudos com Mutillidae mesmo estando em discordância com o site Hymenoptera Anatomy Ontology Portal (HAO, Yoder et al. 2010) onde as mesmas são utilizadas em referências aos tergitos e esternitos abdominais. Diversos termos e medidas utilizadas por Bartholomay et al. (2018) (Capítulo 5) divergem dos demais capítulo apresentados em decorrência do artigo em questão haver sido publicado antes que se houvesse chegado a um consenso quanto à terminologia e medidas a serem utilizadas entre especialistas da família. Todos as alteraçãoes feitas são mencionadas em detalhes na seção “Material and methods” de Bartholomay et al. (2019a) (Capítulo 6). Os seguintes termos e conceitos de esculturação foram propostos por Harris (1979) e utilizados no presente: “areolate”: dividido em pequenos espaços irregulares; “punctate”: com marcas pontuais finas, perfurações semelhantes às feitas com um alfinete; “foveolate”: com pequenos poços profundos; “micropunctate”: com minúsculas perfurações na superfície visíveis apenas em grandes aumentos; “puncticulate”: esparsamente perfurado com pontos finos e amplamente espaçados; “interval”: o espaço entre duas estruturas ou esculturações, isto é, entre duas ‘unidades de escultura’ (e.g. espaço entre duas puncturas); “interstice”: o espaço entre duas linhas, sejam elas estrias ou linhas compostas por pontuações; “confused”: marcas sem contornos definidos ou econtrando-se desordenadamente, sem padrão definido; “granulate” ou “granulose”: aparentemente coberto ou composto por pequenos grânulos; “rugose”: enrugado, com dobras ou elevações irregulares; “scabrous”: irregular, possuindo projeções e/ou dobras curtas e agudas; “reticulate”: semelhante a uma rede ou composto por uma rede de linhas; 12
“carinate”: elevado, com uma ou muitas cristas longitudinais estreitas (ecarinate = sem tais estruturas). Combinações de tais termos (e.g. “areolate-punctate”) foram utilizadas para referir-se a esculturações que apresentem elementos de ambas as definições. Salvo raras exceções, fêmeas de Mutillidae possuem todas as placas mesosomais completamente fusionadas dorsalmente, de modo que o mesonoto, mesoescutelo e metaescutelo não podem ser distinguidos externamente entre si. Além disso, segundo Huber & Goulet (1993) e o site Hymenoptera Anatomy Ontology Portal (HAO, Yoder et al. 2010), as famílias da subordem Apocrita possuem apenas o mesoescutelo evidente, sendo chamado apenas de escutelo. Sendo assim, utilizamos o termo “scutelar area” para a região aparentemente composta por tecidos meso- e metaescutelares localizada entre os espiráculos propodeais no dorso do mesosoma. Do mesmo modo, o termo “scutelar scale” se refere a uma carena ou lamela transversa encontrada anteromedialmente nessa mesma região. Três novos termos no estudo de Mutillidae foram introduzidos em um ou mais capítulos da presente tese visando a utilização de estruturas importantes na diagnose e separação de algumas espécies examinadas: 1. “post-spiracular area" se refere uma área de integumento bem definida imediatamente posterior aos espiráculos propodeias que pode ser completamente lisa e brilhante ou com esculturação conspicuamente distinta em relação às áreas adjacentes; 2. “post-mesonotal tubercles” são pequenos tubérculos ou dentículos subagudos no terço posterior das margens laterais do mesonoto, aproximadamente a meio caminho entre o meio do mesonoto e os espiráculos propodeais; 3. “scabrous intervals” — mencionado como “cariniform reticulations” no Capítulo 5 — se refere aos intervalos entre areolações ou foveolações da esculturação na área escutelar de alguns grupos que pode projetados grosseiramente e de maneira irregular. O comprimento dos olhos foi medido pela distância da margem dorsal à margem ventral do mesmo em vista frontal em relação à distância da margem ventral dos olhos até o côndilo mandibular anterior também em vista frontal. A largura do vértice foi medida imediatamente posterior aos olhos e dividida pela largura máxima do pronoto, ambos em vista dorsal (exceto no Capítulo 5). As medidas dos flagelômeros foram tomadas em vista frontal. Para comparação entre comprimento e largura do mesosoma foi medida a distância entre a margem anteromedial do pronoto (excluindo o colar pronotal) e uma linha imaginária entre a margem anterior dos espíráculos propodeais, que por sua vez foi dividida pela largura máxima do mesosoma em vista dorsal. A comparação entre as larguras de T1 em relação a T2 foi feita dividindo a largura máxima de ambos os segmentos em vista dorsal. O comprimento de T2 foi medido em vista dorsal da margem anterior exposta (não encaixada sob o ápice de T1) até a margem posterior do mesmo. Na seção “Material examined” de todos os capítulos os seguintes padrões foram adotados: meses são inseridos em números romanos maiúsculos; abreviações, acrônimos e dados corrigidos e/ou adicionados pelos autores são colocados entre colchetes. Nomes de estados/províncias são destacados em itálico. Capítulos formatados para submissão à Insecta Mundi divergem em alguns detalhes na seção de material examinado por normas específicas do periódico.
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4 Resultados e discussões
CAPÍTULO 4.1 (em formato de submissão à revista JOURNAL OF NATURAL HISTORY)
Revision of Cephalomutilla André (Hymenoptera: Mutillidae) including one new species, three new sex associations, and nine new synonymies
Kevin A. Williams1*, Pedro R. Bartholomay2, Roberto A. Cambra3, James P. Pitts4, & Márcio L. Oliveira2
1 Plant Pest Diagnostics Center, California Department of Food and Agriculture, Sacramento, California, USA. 2 Instituto de Pesquisas da Amazônia, Manaus, Amazonas, Amazonas, Brazil. 3Museo de Invertebrados G. B. Fairchild, Estafeta Universitaria Apartado 00017, Universidad de Panamá, Panamá 0824, República de Panamá 4 Department of Biology, Utah State University, Logan, Utah, USA.
* Corresponding author. E-mail: [email protected]
Abstract
Cephalomutilla André, 1908, composed of eleven species previously recognized from females only, is here revised. A new species from Colombia, C. cabezona, is recognized from both sexes and newly described. The previously unknown male of C. zelichi Casal, 1963 is associated and described. Mutilla vulnerifera André, 1908, syn. nov., is recognized as the male of C. haematodes (Gerstaecker, 1874) and the following species are recognized as synonymous color variants: C. albicalcaris Mickel, 1960, syn. nov.; C. argyrosticta (Burmeister, 1875), syn. nov.; and C. flavigastra Mickel, 1960, syn. nov. Cephalomutilla proxima (Smith, 1879), comb. nov. is recognized as the male and senior synonym of the following species previously recognized from females only: C. confluenta Mickel, 1960, syn. nov.; C. distincta Mickel, 1960, syn. nov.; C. fasciata Mickel, 1960, syn. nov.; C. transversa Mickel, 1960, syn. nov.; and C. vivata (Cresson, 1902), syn. nov. Keys and illustrations are provided for the species of Cephalomutilla. Key words: Sphaeropthalminae, Dasymutillini, Dasymutilla, Traumatomutilla, velvet ants, Neotropics, Brazil
Introduction
Cephalomutilla André, 1908 was described to include four South American species, known from females only. André considered these females intermediate between Traumatomutilla André, 1902 and Pseudomethoca Ashmead, 1899 (as Sphinctomutilla Andre, 1908), having the broad head of Pseudomethoca and slender metasoma of Traumatomutilla (Fig. 1). Although Andre suggested Traumatomutilla diabolica (Gerstaecker, 1874) as the type for Cephalomutilla, this was based on a misidentification. Mickel (1957), therefore, proposed a case to ICZN (1959) to change the type species designation to C. graviceps (Andre, 1902). After this type designation, Mickel (1960) revised the genus, treating 10 species known from females only. Later, Casal (1963) described an 11th species for the genus. For over 50 years, no taxonomic changes were proposed, until Brothers and Lelej (2017) transferred the first male into the genus, C. vulnerifera (Andre, 1908). Now, 60 years after Mickel’s revision, we present a revision of the genus, including one new species, one new combination, three new sex associations, and nine new synonymies.
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Materials and Methods
The following acronyms are used for institutions housing the material discussed in the current study:
AMNH – American Museum of Natural History, New York, New York, USA. BMNH – British Museum of Natural History, London, England. CMNH – Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA. CASC – Department of Entomology, California Academy of Sciences, San Francisco, California, USA. CSCA – California State Collection of Arthropods, Sacramento, California, USA. DGMC – Donald G. Manley Collection, Florence, South Carolina, USA. DJBC – Collection of Denis J. Brothers, to be deposited in Iziko South African Museum (SAMC), Cape Town, South Africa. DZUP – Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. EMUS – Department of Biology Insect Collection, Utah State University, Logan, Utah, USA. FSCA – Florida State Collection of Arthropods, Gainesville, Florida, USA. IAVH – Instituto Alexander von Humboldt, Villa de Leyva, Boyaca, Colombia. MIUP – Museo de Invertebrados G.B. Fairchild, Universidad de Panamá, Panama City, Panama. MNRJ – Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil. MPEG – Museu Paraense Emílio Goeldi, Belém, Pará, Brazil. MZSP – Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo, Brazil. SEMC – Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA. UCDC – The Bohart Museum of Entomology, Universisty of California, Davis, California, USA. UEFS – Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil. UNSM – Univeristy of Nebraska State Museum, Lincoln, Nebraska, USA.
Transcription of label data as well as morphological terminology and measurements followed those adopted by Bartholomay et al. (2018, 2019a), Cambra et al. (2018), and Williams et al. (2011). For C. haematodes and C. proxima, which had their color variants formerly named as discrete species, these former names are provided for each observed female.
Cephalomutilla André, 1908
Cephalomutilla André, 1908: 194. Cenhalomutilla Quintero & Cambra, 1996: 13. Incorrect subsequent spelling.
Type species. Ephuta (Traumatomutilla) graviceps André, 1903 (female), designated by ICZN, 1959: 103 according to application by Mickel, 1957: 25.
Diagnosis. FEMALE: Females of Cephalomutilla can be separated from other Dasymutillini by the following combination of characters: vertex broader than mesosoma; scutellar scale present (sometimes rudimentary); T2 marked with two to four integumental spots; T6 with defined pygidial plate. MALE: Males can be differentiated from other Dasymutillini by the following combination of characters: axilla armed with posteriorly directed tooth or lobe; mesopleuron not projected into conspicuous tubercle in dorsal half, at most simply swollen; S2 medially flattened and lacking seta-filled pit; and metasoma at least partly reddish or orange in color. The genitalia also have useful diagnostic features: the paramere is slightly downcurved medially and upcurved apically and often somewhat dorsoventrally compressed; the cuspis has an elongate brush of posteriorly directed setae; and the penis valve has two simple teeth posteroventrally. 15
Description (Hitherto unknown). MALE. Integument mostly blackish with strong dense punctuation, lacking any reflection, except metasoma partly or entirely reddish to orange-brown. Setae mostly simple, sometimes with few brachyplumose setae on head or plumose setae in genitalia. Head. Round to transverse, usually narrower than mesosoma. Occipital carina distinct dorsally. Vertex often conspicuously swollen posteriorly. Eye broadly semispherical, strongly protruding, surface shining, ommatidia discernible. Ocelli small. Antennal scrobe deep, reaching eye margin, with a transverse lamellate carina on dorsal margin. Clypeus depressed immediately below insertion of antennae, overall slightly convex, anterior/ventral margin with sub-truncate lamella. Malar space shorter than basal height of mandible. Genal carina absent. Hypostomal carina evident, evenly high throughout, extending in a smooth curve to posterior mandibular condyle. Scape bicarinate. Pedicel virtually as wide as long, shorter than F1, which is generally shorter than F2. Mandible in dorsal/anterior view evenly curved, bidentate apically, lacking any ventral tooth or notch. Maxillary palp 6-segmented; labial palp 4-segmented; basal palpomeres strongly flattened and broadened, densely covered with short setae. Mesosoma. Surface finely and densely foveolate to punctate except smooth or nearly so on metapleuron, posteroventrally on mesopleuron, and anteriorly on lateral face of propodeum; mesoscutellum and most of propodeum areolate. Epaulet weak, reduced to a row or tuft of short setae; in anterior view humeral angle rounded. Tegula broadly subcircular, convex, almost entirely smooth, except along punctate anterior and inner margins. Mesoscutum with notaulus and parapsids practically obliterated. Mesoscutellum virtually flat; axilla generally flat dorsally, narrowly connected to scutellum apicointernally, terminating in a truncate or dentate lobe; dorsal surface of axilla coarsely and sparsely punctate except along smooth impunctate posterior margin. Metanotum simple, transverse, dorsellum poorly delimited laterally, usually concealed by dense setae. Propodeum evenly convex, dorsal and lateral faces indistinct. Metasternal processes convergent to unidentate apex, shorter than metacoxal length. Wings. Fore wing with elongate sclerotized pterostigma; marginal cell elongate, rounded to truncate apically; three closed submarginal cells, apical veins of third often inconspicuous or obliterated. Legs. Tibial spur formula 1–2–2. Mid and hind tibiae with two weak rows spines; apical spurs densely clothed with microsetae. Metacoxa with weak inner carina. Metasoma. Punctation generally dense and fine; T1 longer than wide, about 0.5 × width of T2, anterior and dorsal faces indistinct, slightly constricted apically. T2 with lateral felt line narrow and long. S1 with vestigial longitudinal carina. S2 in lateral view flattened postero-medially, lacking felt line. Hypopygium generally longer than wide, surface virtually flat, posterior margin medially unidentate or narrow bidentate. Genitalia. Paramere long, free length as long or longer than length from base of genital capsule to apex of parapenial lobe, sub-cylindrical to dorso-ventrally flattened, acute apically, apex weakly or clearly curved dorsad in lateral view. Parapenial lobe well-developed. Cuspis sub-cylindrical, paracuspis a rounded lobe. Digitus narrow digitiform, slightly wider basad. Penis valve without any conspicuous concave or convex areas; with two sharp apical teeth on ventral margin; apical margin of penis valve with long distinct setae. Species included. Five species: one known from females only, C. graviceps (André, 1903), and four known from both sexes: C. cabezona, sp. nov.; C. haematodes (Gerstaecker, 1874); C. proxima (Smith, 1879), comb. nov.; and C. zelichi Casal, 1963. Distribution. Widespread in South America (Argentina, Bolivia, Brazil, Colombia, Paraguay, Uruguay). Remarks. This genus has been intertwined with Traumatomutilla since its initial description, wherein Andre stated that it was intermediate between Pseudomethoca and Traumatomutilla. Three of the females, including the type, were formerly included in Traumatomutilla (André 1902, 1903) and the first associated male, T. vulnerifera, was most recently included in that genus. Williams (2012) found that Cephalomutilla was indeed closely related to Traumatomutilla but should be treated as a distinct genus. The first recognized sex association was suggested, but not formalized, by Williams (2012), based on identical ITS1 sequences shared by a male of C. proxima and a female C. vivata (Cresson, 1902), both collected in Minas Gerais, Brazil. The remaining sex associations presented here are based on overlapping distribution and morphological similarities; they are further explained in the Remarks section for each species.
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Key to species of Cephalomutilla (females) 1. Vertex armed with tubercles (Figs. 30, 33); scutellar scale distinct, usually surrounded by transverse carinae … 2 -- Vertex unarmed (Figs. 35, 36); scutellar scale indistinct … 3 2. Vertex with small sub-triangular tubercles, distance between tubercles less than distance between epaulets (Figs 29, 30) … C. proxima (Smith, 1879) -- Vertex with strong rectangular tubercles, distance between tubercles greater than distance between epaulets (Figs 32, 33) … C. zelichi Casal, 1963 3. Pygidium dense microreticulate (Fig. 37); so far known only from Colombia … C. cabezona, sp. nov. -- Pygidium rugose or striate (e.g. Figs 31, 34); predominantly found southern South America … 4 4. Vertex punctures medium-sized, coarse punctate, some confluent; dorsal face of T1 convex; pleura entirely pale pubescent; frons, vertex, femora, and tibiae usually with extensive white setae (Figs 9–12); widespread in southern South America … C. haematodes (Gerstaecker, 1874) -- Vertex punctures mostly small, not coarse or confluent; dorsal face of T1 flat; pleura black pubescent; frons, vertex, femora, and tibiae mostly with black setae (Figs 4–6); predominantly known from arid regions of Argentina and Paraguay … C. graviceps André, 1903
Key to species of Cephalomutilla (males) Unknown in C. graviceps (André, 1903) 1. Axilla flat with truncate posterior face (Figs 38–41) … 2 -- Axilla posterior face angular or sub-dentate (Figs 42–45) … 3 2. Paramere downcurved medially and upcurved at apex, dorsoventrally flattened (Figs 46–48); tegula usually with black setae (Figs 23–25), except in northern Brazil (Fig. 22); widespread in South America … C. proxima (Smith, 1879) -- Paramere basically straight, sub-cylindrical (Figs 51–53); tegula with silvery setae (Fig. 28); Argentina, Bolivia, SW Brazil, Paraguay … C. zelichi Casal, 1963 3. Axilla contorted and sharply dentate posteriorly (Figs 44, 45); T7 microreticulate; Colombia … C. cabezona, sp. nov. -- Axilla more or less flat, posterior face angular (Figs 42, 43); T7 rugose; southern South America … C. haematodes (Gerstaecker, 1874)
Cephalomutilla cabezona Williams & Bartholomay sp. nov. (Figs 1–3, 35–37, 44–45, 61–65)
Diagnosis. FEMALE. This species is unique in the pygidial sculpture, which is uniform microreticulate (Fig. 37). The unarmed vertex and poorly defined angular scutellar scale are also useful for diagnosis. Body length 7.5–13 mm. MALE. The male of this species can be recognized by the T7 sculpture, which is uniform micro- reticulate. Additional useful characters include the posteriorly incurved sharp dentate axilla; the cuspis, which is twice as long as the digitus, slender sub-cylindrical, and has sparse elongate setae directed posteriorly and ventrally; and the scarcely recurved scarcely flattened paramere with a ventral setal brush. Body length 8.5–12 mm. Description. Holotype FEMALE. Body length 11.5 mm. Head. Quadrate, head width 1.15 × pronotal width. Head, except ventral surface, irregularly finely and densely foveolate-punctate. Mandible oblique, robust, with sharp subapical tooth. Dorsal scrobal carina well-defined, continuing onto antennal scrobe as small tubercle; lateral scrobal carina reduced to longitudinal impunctate area. Flagellomere 1 2.5 × pedicel length; flagellomere 2 1.9 × pedicel length. Genal carina present, long, broadly separated from gular carina and occipital carina. Occipital carina distinct, without any connected tubercles. Mesosoma. Mesosoma 0.8 × as long as wide. Mesosomal dorsum densely and coarsely areolate-punctate, areolations denser and smaller 17
mediad, with some intervals scabrous. Anterior face of pronotum well-defined, mostly smooth with some parallel striae and punctures. Humeral carina present, extending to rounded epaulet, antero-lateral corner of pronotum forming right angle in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Sculpture of lateral face of pronotum and mesosomal pleurae obscured by dense setae, except dorsal fourth of metapleuron smooth. Lateral face of propodeum virtually impunctate. Ratios of dorsal width of mesosoma at humeral angles, pronotal spiracles, widest point of mesonotum, propodeal spiracles, and propodeum posterolateral angles, 76:83:85:79:50. Lateral margin of mesosoma scarcely emarginated anterior to propodeal spiracle, smoothly diverging anterad and converging slightly posterior to pronotal spiracles. Propodeal spiracle slightly swollen, sides scarcely diverging between constriction and spiracle opening. Scutellar scale weak, angular, with interrupted longitudinal carina anterior and posterior to scale. Dorsal and posterior faces scarcely differentiated. Metasoma. T1 width 0.45 × T2 width. T2 length 1.05 × T2 width, with maximum width at midlength. Disc of T2 dense uniform foveolate-punctate with scabrous intervals; sculpture of integumental spots sparse shallow foveolate-punctate. S1 with short longitudinal carina. S2 coarse foveolate-punctate, medially flattened. Pygidium as broad as long, defined by lateral carinae, basal third ecarinate; surface uniform micro-reticulate. MALE. Body length 9 mm. Head. Sub-quadrate; head width subequal to pronotal width. Eye almost circular. Ocelli minute; OOD 7.3 × DLO, IOD 2.5 × DLO. Occipital carina distinct. Frons, vertex, and gena densely foveolate-punctate. Gena ecarinate. Antennal scrobe with prominent transverse dorsal carina. Clypeus shallow convex medially, concave laterally below antennal insertion; densely punctate; with smooth transverse sub- truncate lobe apically. Scape bicarinate. F1 length 1.9 × pedicel length; F2 length 1.9 × pedicel length. Mandible obliquely bidentate apically, inner tooth distinct; lacking dorsal or ventral projections. Mesosoma. Epaulet sub-linear, not connected with humeral carina. Anterior face of pronotum punctate dorsally and laterally, smooth ventro-medially without longitudinal medial furrow. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum dense foveolate-areolate without longitudinal carina medially; notaulus and parapsis mostly obliterated. Mesoscutellum convex, areolate-foveolate. Axilla produced posterolaterally as incurved dentiform projection with sharp posterior face; dorsal face areolate-foveolate basally, smooth laterally and posteriorly. Area between mesoscutum and mesoscutellum smooth. Metanotum virtually equally wide throughout, punctation obscured by dense setae. Propodeum dorsal and posterior faces scarcely differentiated, areolate, dorsally obscured by dense setae; lateral face mostly impunctate. Mesopleuron with punctation obscured by dense setae, evenly convex without tubercle on dorsal half. Metapleuron virtually smooth, with micropunctures and sub-appressed microsetae. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, rounded apically; three submarginal cells, third cell scarcely defined. Legs. Setae simple, meso- and metatibia each with two rows of inconspicuous spines dorsally; spurs densely microsetose. Metasoma. T1 width 0.5 × T2 width. T2 length 0.95 × T2 width. T2 disc sculpture dense foveolate-punctate. T7 dense microreticulate, lateral carinae distinct. S1 with longitudinal medial carina, highest posteriorly. S2 medially flattened, with interspersed small and large foveolations. S7 longer than broad, punctured area narrower basally, apical margin unidentate medially. Genitalia. Paramere long, free length 1.2 × length from base of genital capsule to apex of parapenial lobe; sub-cylindrical, acute apically, apex weakly curved dorsad in lateral view; with ventral brush of long setae along basal 0.5 × of free paramere length. Cuspis 0.45 × free paramere length, subcylindrical, with sparse long setae ventrally and fanned apical brush of posterior to ventrally directed setae 0.52 × cuspis length. Paracuspis short, lobe-like, rounded apically, densely setose. Digitus 0.17 × free paramere length, narrow digitiform, slightly wider basad. Penis valve scarcely extending beyond parapenial lobe, ventral margin with two apical teeth; apical distance between teeth 0.13 × length of valve; apical margin with row of distinct setae. Coloration and variations. FEMALES. Body and appendages vary from dark reddish-brown to black, legs and metasoma usually lighter than head and mesosoma; scape, pedicel, flagellomeres, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs whitish; T2 with four separated orange or 18
yellow integumental spots. Appendages and lateral and pleural body regions generally with whitish setae; dorsum of body with erect and appressed dense black setae except with sparse erect whitish setae on T2 integumental spots and denser patches of appressed whitish setae as follows: transverse curved band on vertex, lateral longitudinal mesosomal stripes extending from propodeum anterior to just past pronotal spiracle-level, small apicomedial patch on T2 (sometimes obliterated), and broader medial patches on T3–5. The specimens from Zambrano have the T2 spots yellow and smaller than the holotype; the specimen from Manaure has the orange T2 spots larger than the holotype. MALES. No significant variations in color and setae were observed in males. Head, mesosoma, and appendages dark reddish-brown to black, except scape, pedicel, flagellomeres, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs whitish; metasoma orange, except pygidium pale yellow-brown. Head, mesosoma, and appendages with mostly dense erect and appressed whitish setae, except mesoscutum, tegula, and axilla with setae blackish, and mesoscutellum setae erect interspersed black and whitish setae; and metapleuron and lateral propodeal face mostly bare. Metasoma setae predominantly whitish laterally, ventrally, and dorsally on T1 and T2 basally; T2–6 mostly with orange setae. Wings light brown, with darker cloud just beyond cells; veins dark brown. Distribution. Colombia. Etymology. From the Spanish “cabezona”, meaning “big headed”, noun in apposition. Material examined. Holotype: female, COLOMBIA, [La Guajira], Cerrejon, Esteril, Hanval, T2E5 (EMUS). Paratypes (5 females and 48 males). COLOMBIA: Bolivar, Zambrano: HOA Monterey: 7.IV.1994 (20 males, IAvH); 15–70m, 21.IV.1994 (8 males, IAvH); malaise 4, 14.IV.1994 (7 males, IAvH); malaise 10, chile borde, 14.III.1994 (2 males, IAvH); rutiuol ano Nellina 6N Pa-payo, parime 1, 24.IV.1994 (9 males, IAvH); Hd. Monterey, 120 m, 9°37′48"N 74°54′44"W, bosque seco: malaise 4, bosque de chile, 11.V.1994 (1 female, AMNH); malaise 10, Casa Neuva, 30.VIII.1993 (1 male, AMNH); site and date unknown (1 female, AMNH); La Guajira: Cerrejon, Esteril: Hanval, T2E5 (1 female, EMUS?); pitfall, T1E10, 14.VII.2007 (1 female, EMUS); Manaure, 20.IX.1968, B. Malkin (1 female, AMNH); Valle del Cauca, Farrajones, 1730m, 24– 31.III.1998 (1 male, IAvH). Remarks. This is now the northernmost distributed species known in Cephalomutilla. It was initially surprising to find this genus in Colombia, as Cephalomutilla seemed to be concentrated in southern South America. Some other mutillid genera, like Tallium André, 1902 and Tobantilla Casal, 1965, were previously known from Argentina and southern South America (Nonveiller 1990) but were subsequently discovered in Colombia (Cambra and Quintero 2003, Williams et al. 2011b). Additionally, a new species from the Traumatomutilla juvenilis species-group, also predominantly from Argentina and southern South America, has been recently found in similar areas in Colombia and sharing the same color pattern as C. cabezona sp. nov. (KAW & PRB pers. obs.). Both sexes of C. cabezona sp. nov. have the pygidium microreticulate. This differentiates them from all other Cephalomutilla and further supports a sex association that was already obvious based on their co- occurrence and geographic isolation from other congeners.
Cephalomutilla graviceps (André, 1903) (Figs 4–6)
Ephuta (Traumatomutilla) graviceps André, 1903: 453, holotype [by monotypy], f#, Argentina, Santiago del Estero (MNHN). Traumatomutilla graviceps: André, 1904: 40. Cephalomutilla graviceps: André, 1908 [as a synonym of C. diabolica]: 194. Cephalomutilla diabolica: André, 1908 (nec. Gerstaecker, 1874): 194. (synonymized in Mickel 1960). Cephalomutilla graviceps: Mickel, 1960: 163.
19
Diagnosis. FEMALE. The following combination of characters is needed for diagnosis: the vertex is unarmed; the vertex has small, dense but not confluent, punctures; the scutellar scale is rounded posteriorly and surrounded by raised transverse intervals anteriorly and anterolaterally; T1 has the anterior face flattened; the pygidium sculpture is rugose; and the frons, vertex, mesopleuron, femora, and tibiae are covered with black setae only. Body length 9.1–13.3 mm. MALE. Unknown. Extended diagnosis. FEMALE. See Andre (1903). MALE. Unknown. Coloration and variations. FEMALES. The lateral longitudinal mesosomal whitish setal stripes vary in length, in some individuals they extend anteriorly to just beyond the pronotal spiracles and in others they are limited to the dorsum of the propodeum and posterior portion of the mesonotum. Distribution. Arid regions of Argentina (Catamarca, Cordoba, La Rioja, Mendoza, San Luis, Santiago del Estero, Tucuman) and Paraguay (Chaco). Material examined (11 females). ARGENTINA: Cordoba: Balnearia, III.1971, M.A. Fritz (1 female, EMUS); Guanaco Muerto, II.1980, Viana (1 female, EMUS); La Rioja: Mascasin, II.1958, F. Walz (9 females, MIUP); Patquia, XII.1932–I.1933, K.J. Hayward, B.M. 1933–187 (1 female, BMNH); Salta, La Vina, III.1985, M.A. Fritz (1 female, AMNH); San Luis, San Geronimo, II.1980, Fritz (1 female, CSCA). PARAGUAY, Chaco, Filadelfia, I.1995, Arriagagda (1 female, EMUS). Remarks. This species is similar to C. haematodes but differs in head punctation and T1 shape. Additionally, this arid distributed species has darker coloration overall, with the head, mesopleuron, and legs with black setae only. The male is not yet recognized but will likely be similar to C. haematodes.
Cephalomutilla haematodes (Gerstaecker, 1874) (Fig. 7–12, 42–43, 56–60)
Mutilla haematodes Gerstaecker, 1874: 63, lectotype [designated by Mickel (1960)], f#, Uruguay, Montevideo, Sello (ZMB). Mutilla argyrosticta Burmeister, 1875: 477, lectotype [designated by Mickel (1960)], f#, Argentina, Mendoza (MACN), new synonym. Ephuta (Traumatomutilla) argyrosticta: André, 1902: 54. Ephuta (Ephuta) haematodes: André, 1902: 60. Traumatomutilla argyrosticta: André, 1904: 40. Traumatomutilla haematodes: André, 1904: 40. Cephalomutilla argyrosticta: André, 1908: 163. Traumatomutilla vulnerifera André, 1908: 207, lectotype [designated here], m#, [no indication of locality] (MNHN), examined, new synonym. Cephalomutilla albicalcaris Mickel, 1960: 161, holotype, female, Argentina, Santiago del Estero, Chaco de Santiago del Estero, env. d’Icano [outskirts of Icaño], Bords du Rio Salado [banks of the Salado River], E.R. Wagner, 1904 (MNHN), new synonym. Cephalomutilla flavigastra Mickel, 1960: 160, , holotype, female, Argentina, Santiago del Estero, Chaco de Santiago del Estero, env. d’Icano [outskirts of Icaño], Bords du Rio Salado [banks of the Salado River], E.R. Wagner, 1909 (MNHN), new synonym. Cephalomutilla vulnerifera: Brother & Lelej, 2017: 62.
Diagnosis. FEMALE. The following combination of characters is needed for diagnosis: the vertex is unarmed; the vertex punctures are small to moderate, mostly confluent with some sub-scabrous intervals; the scutellar scale is rounded posteriorly and surrounded by raised transverse intervals anteriorly and anterolaterally; T1 has the anterior face convex; the pygidium sculpture is rugose; and the body has more extensive whitish setae, usually with whitish setae on the frons, vertex, mesopleuron, femora, and tibiae. Body 20
length 8–12 mm. MALE. The male of this species can be recognized by the cuspis, which is scarcely longer than the digitus, thick sub-cylindrical, and has dense elongate apical setae directed postero-ventrally. additional useful characters include the posteriorly sub-dentate to sub-truncate axilla, the rugose T7 sculpture, and the relatively straight sub-cylindrical paramere with a ventral setal brush basally. Body length 7.5–13.5 mm. Extended diagnosis. FEMALE. For morphology, see Mickel’s (1960) descriptions of C. flavigastra and C. albicalcaris. For coloration, see Coloration and Variations section below. MALE. Head. Subquadrate; posteromedially expanded; head width 0.9 × pronotal width. Eye almost circular. Ocelli minute; OOD 6.7 × DLO, IOD 4.0 × DLO. Occipital carina distinct. Frons, vertex, and gena densely foveolate-punctate. Gena ecarinate. Antennal scrobe with prominent transverse dorsal carina. Clypeus shallow convex medially, concave laterally below antennal insertion; densely punctate; with smooth transverse sub- truncate lobe apically. Scape bicarinate. F1 length 2.1 × pedicel length; F2 length 2.7 × pedicel length. Mandible obliquely bidentate apically, inner tooth distinct; lacking dorsal or ventral projections. Mesosoma. Epaulet small, not connected with humeral carina. Anterior face of pronotum punctate dorsally and laterally, smooth ventro-medially without longitudinal medial furrow. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum dense foveolate-areolate without longitudinal carina medially; notaulus and parapsis mostly obliterated. Mesoscutellum flat, areolate-foveolate. Axilla produced posterolaterally as sub-angular to sub-truncate projection with sharp posterior face; dorsal face areolate-foveolate, except lateral and posterior margins smooth. Area between mesoscutum and mesoscutellum smooth. Metanotum virtually equally wide throughout, medially areolate-foveolate. Propodeum dorsal and posterior faces scarcely differentiated, areolate; lateral face mostly impunctate. Mesopleuron foveolate-punctate, evenly convex without tubercle on dorsal half. Metapleuron virtually smooth, with micropunctures and sub-appressed microsetae. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate or rounded apically; three submarginal cells, third cell scarcely defined. Legs. Setae simple, meso- and metatibia each with two rows of inconspicuous spines dorsally; spurs densely microsetose. Metasoma. T1 width 0.4 × T2 width. T2 length 0.9 × T2 width. T2 disc sculpture foveolate-punctate, denser medially and laterally, sparser on sublateral cuticular spots. T7 rugose, lateral carinae distinct. S1 with scarcely defined longitudinal medial carina. S2 medially flattened, foveolate. S7 longer than broad, punctured area not narrowed posteriorly, laterally defined by carinae, apical margin unidentate medially. Genitalia. Paramere long, free length 1.1 × length from base of genital capsule to apex of parapenial lobe; sub-cylindrical, acute apically, apex weakly curved dorsad in lateral view; with ventral brush of long setae along basal 0.35 × of free paramere length. Cuspis 0.22 × free paramere length, scarcely laterally compressed, with scattered short setae throughout and apical postero-ventrally directed brush of setae 0.8 × cuspis length. Paracuspis lobe-like, rounded apically, setose. Digitus 0.15 × free paramere length, narrow digitiform, slightly wider basad. Penis valve scarcely extending beyond parapenial lobe, ventral margin with two apical teeth; apical distance between teeth 0.13 × length of valve; apical margin with row of distinct setae. Coloration and variations. FEMALES. Body and appendages vary from reddish-brown to black, head or S2 often red-orange; antenna, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs dark-brown or whitish; T2 with two or four separated or confluent yellow, orange, or reddish integumental spots. Appendages and lateral and pleural body regions generally with whitish setae; dorsum of body with erect and appressed dense black setae except with sparse erect whitish setae on T2 integumental spots and denser patches of appressed whitish setae as follows: transverse curved band on vertex (head sometimes with setae entirely black or entirely whitish), lateral longitudinal mesosomal stripes extending from propodeum anterior to just past pronotal spiracle-level (stripes sometimes terminating on mesonotum or extending to anterior pronotal margin), small apicomedial patch on T2 (sometimes obliterated, sometimes broad), medial patches on T3–5 (sometimes missing or reduced on T3, sometimes with entire tergites clothed with whitish- yellow setae). MALES. Head, mesosoma, T1, T3–7, and appendages dark reddish-brown to black, except scape, pedicel, flagellomeres, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs 21
whitish; T2 black with large-orange red integumental spots or orange-red with large orange yellow integumental spots (sometimes confluent) and the posterior and lateral margins black to orange-brown; S2 black or orange. In specimens with T2 black, body setae predominantly blackish with variably extensive sparse whitish setae in the following areas: frons, gena, mesosomal pleurae, propodeum, tibiae, T1, T2 posterolaterally, T3–4 laterally and medially, and S2–4 apical fringes. In specimens with T2 orange, body setae mostly dense erect and appressed whitish, except with blackish setae in the following areas: mesoscutum, tegula, axilla, mesoscutellum, T2 posterior and lateral margins, and T6–7. Wings dark or light brown, lighter in basal portion among wing cells; veins dark brown. Distribution. Argentina, Bolivia, Paraguay, and Uruguay. Material examined (119 females and 16 males). ARGENTINA: Colon, Liebig, II.1997, L. Caire (1 female, FSCA, C. haematodes); Cordoba: Freyre, IV.1992, Williner (4 females, EMUS, C. haematodes); La Cumbre, 25.XII.1989, M.A. Fritz (1 female, EMUS, C. albicalcaris); Villa Soto, II.2003, A. Ugarte (1 female, FSCA, C. albicalcaris); Entre Ríos, Liebig: 25.XII.1996–15.I.1997, L. Caire (4 females 3 males, EMUS MIUP, C. haematodes); II.1997, L. Caire, (18 females 1 male, MIUP, C. haematodes); II.1997, M. Zelich (9 females, MIUP, C. haematodes); 7.II.2003, L. Caire (5 females, MIUP, C. haematodes); Pronunciamiento, II.1959, F. Walz (2 females, MIUP, C. haematodes); Formosa, Gran Guardia, II.1953 (9 females, MIUP); Jujuy, Bahia Blanca (1 female, UNSM, C. haematodes); La Pampa, Reserva Parque Luro, Santa Rosa, 12.XII.1997, D. Rojas (3 females, MIUP, C. haematodes); Mendoza, Desaguadero, I.1979, Williner (1 female, EMUS, C. albicalcaris); Rio Negro: Choele Choel, XII.1997, M.A. Fritz (1 male, AMNH); Lamarque, I.1974, M.A. Fritz (4 males, AMNH); Salta: La Vina, I.1983, M.A. Fritz (3 females, EMUS, C. albicalcaris and C. argyrosticta); 22–25.XII.1983, M. Wasbauer (1 female, UCDC, C. argyrosticta); II.1984, M.A. Fritz (1 male, AMNH); Sumalao, II.1991, M.A. Fritz (2 females, EMUS, C. argyrosticta); San Luis: Arizona, II.1980, M.A. Fritz (5 females, EMUS FSCA, C. albicalcaris and C. argyrosticta); San Geronimo, I.1979, Fritz (4 females, CSCA FSCA, C. argyrosticta); Santa Fe, La Rubia, I.1978, Williner (1 female, EMUS, C. haematodes); Santiago del Estero, col. MacBrinlay, 13.II.1962, O.H. Casal (1 female, FSCA, C. haematodes); Monte Quemado, 15.XII.1971, D.J. Brothers (4 males, DJBC, SEMC); Ruta 9, 70 km S Sgo. del Estero, 20.I.2002, F. & P. Cassola (5 females, DGMC, C. albicalcaris); Tucuman, 12 km N El Cadillal, 8–9.XI.1989, J.G. Rozzzen & A. Roig (1 female, AMNH, C. argyrosticta). BOLIVIA, Santa Cruz, Campo Guairay, 24 km S Camiri, 4.III.1999, LA Stange (1 male, FSCA). PARAGUAY: Alto Paraguay, La Victoria, Puerto La Esperanza (Puerto Sastre), 79 msnm, 22° 01’ S 58° 01’ W, 29.XI–4.XII.2006, B. Garcete & C. Aguilar (17 females, MIUP); Boqueron: en camino hacia Filadelfia, 6.XII.1993, B. Garcete (6 females, MIUP); P.N. Tnte. Enciso, zona administracion, 213 msnm, 21° 12’ S 61° 39’ W, 3–4.II.2007, B. Garcete (1 female, MIUP); Chaco, Filadelfia, I.1995, Arriagagda (1 female, EMUS, C. albicalcaris and C. argyrosticta); Presidente Hayes: Cruce Loma Plata, 14.I.2003, U. Dreschel (1 female, FSCA, C. haematodes); Cruce Los Pioneros, 20– 21.I.1996, B. Garcete (4 females, MIUP); Lolita, Yaragui: 7.II.2003, U. Dreschel (1 female, FSCA, C. flavigastra); 5.III.2003, U. Dreschel (2 females, FSCA, C. haematodes); 10.I.2002, U. Dreschel (1 female, FSCA, C. haematodes); Ruta Transchaco km 252, 20.I.1996, B. Garcete (1 female, MIUP); Cruce Los Pioneros, Estac. Experim. del Ministerio de Agricultura, 22° 50’ S 59° 45’W, 9–10.XII.2001, C. Aguilar (2 females, MIUP). URUGUAY, Rio Negro, Carbonell (1 male, AMNH). Remarks. In this species and C. proxima, males and females each show significant color variation across their distribution. Southwestern specimens (Figs 7, 9) of C. haematodes from drier areas like the Chaco generally tend to have darker overall coloration than Eastern specimens (Figs 8, 11) from more humid areas like the Pampas. Surprisingly, this widespread species has not been found in Brazil, even though the authors have examined nearly 200 Cephalomutilla from that country. Males can sometimes have the axilla apparently truncate. In this case, they would key out to C. proxima, but can be immediately recognized by the paramere and cuspis shape (Figs 56–60). The sex association is based on overlapping distribution, including multiple examples of males and females found in the same localities. 22
Even though the type series of Traumatomutilla vulnerifera has no locality labels, André (1908) mentioned that the specimens are likely to be from Brazil or Argentina. This was later confirmed by numerous specimens of T. vulnerifera identified from Argentina and a peculiar color syndrome found in males of Southern Brazil and northern Argentina (KAW & PRB pers. obs.). This particular syndrome involves three South American Dasymutillini males, all of which are predominantly black with orange to reddish spots on T2 and were included in Traumatomutilla at one time or another (Nonveiller, 1990). These three males have their specific epithets beginning with vulner-: Cephalomutilla vulnerifera, syn. nov., Suareztilla vulneriventris (André, 1903), and Traumatomutilla vulnerata (Gerstaecker, 1874), likely due to their color similarities. In the AMNH collection, three specimens of Traumatomutilla vulnerata (Gerstaecker, 1874) were labeled by M.A. Fritz in 1997 as Cephalomutilla vulnerata. Examination of the genitalia and external structures of these specimens and the type revealed that T. vulnerata truly belongs to Traumatomutilla, and not Cephalomutilla (PRB pers. obs.). This species overlaps in distribution and coloration with males of C. haematodes (formerly recognized as C. vulnerifera). It is not clear whether Fritz properly recognized C. vulnerifera as C. haematodes and then assumed that T. vulnerata was a congeneric relative, whether he confused T. vulnerata with C. vulnerifera in the field, or whether the attribution of T. vulnerata to Cephalomutilla was just an unlucky guess. Either way, with the synonymy of C. vulnerifera under C. haematodes, all mutillid workers should be happy to be dealing with a bit less confusion around Argentinean vulner- names amongst the Dasymutillini males.
Cephalomutilla proxima (Smith, 1879), comb. nov. (Figs 13–25, 29–31, 38–39, 46–50)
Mutilla proxima Smith, 1879: 220, holotype [by monotypy], male, Brazil, Para [sic] (BMNH). Mutilla vivata Cresson, 1902: 51, lectotype [designated by Cresson (1902)], female, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, new synonym. Mutilla proxima André, 1902: 74 (incertae sedis). Cephalomutilla confluenta Mickel, 1960: 164, holotype, female, Argentina, Reimoser (BMNH), new synonym. Cephalomutilla distincta Mickel, 1960: 165, holotype, female, Paraguay, [Paraguari], Sapucay, February, W.T. Foster, Catalogue number 6228 (USNM), examined, new synonym. Cephalomutilla transversa Mickel, 1960: 166, holotype, female, Brazil, [Pará], Santarém (UMSP), examined, new synonym. Cephalomutilla fasciata Mickel, 1960: 167, holotype, female, Brazil (BMNHW), new synonym.
Diagnosis. FEMALE. This species can be recognized by the shape of the vertex tubercles, which are subtriangular and are separated by less than the inter-epaulet distance. Additionally, scutellar scale is apically truncate with transverse carinae anteriorly and anterolaterally; also, the pygidium is striate to longitudinally rugose. Body length 9–13 mm. MALE. The male of this species can be recognized by the paramere, which is dorsoventrally recurved with a dense brush of setae ventro-laterally. Additional useful characters include the posteriorly truncate axilla, the rugose T7 sculpture, and the cuspis that is over twice as long as the digitus, sub-cylindrical, and having elongate posteriorly directed setae apically. Body length 8.5–15 mm. Extended Diagnosis. FEMALE. For morphology, see Mickel’s (1960) descriptions of C. confluenta, C. distincta, C. fasciata, and C. transversa. For coloration, see Coloration and Variations section below. MALE. Head. subquadrate, posteromedially expanded; head width 0.9 × pronotal width. Eye almost circular. Ocelli minute; OOD 9.7 × DLO, IOD 3.0 × DLO. Occipital carina distinct. Frons, vertex, and gena densely foveolate-punctate, nearly scabrous in some areas. Gena ecarinate. Antennal scrobe with prominent transverse dorsal carina. Clypeus shallow convex medially, concave laterally below antennal insertion; densely punctate 23
with smooth transverse sub-truncate lobe apically, outlined above by transverse carina. Scape bicarinate. F1 length 1.8 × pedicel length; F2 length 2.2 × pedicel length. Mandible obliquely bidentate apically, lacking dorsal or ventral projections. Mesosoma. Epaulet small, not connected with humeral carina. Anterior face of pronotum punctate laterally, with broad longitudinal smooth furrow. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum dense foveolate-punctate with scarcely defined longitudinal carina anteriorly; notaulus and parapsis mostly obliterated. Mesoscutellum flat, areolate-foveolate. Axilla produced posterolaterally as truncate projection with flat posterior face; dorsal face areolate-foveolate anteriorly, smooth posteriorly. Area between mesoscutum and mesoscutellum smooth. Metanotum virtually equally wide throughout, medially areolate-foveolate. Propodeum dorsal and posterior faces scarcely differentiated, coarsely areolate with dense setae basally; lateral face mostly impunctate. Mesopleuron punctation obscured by dense setae, evenly convex without tubercle on dorsal half. Metapleuron virtually smooth, with micropunctures and sub-appressed microsetae. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate or rounded apically; three submarginal cells, third cell scarcely defined. Legs. Setae simple, meso- and metatibia each with two rows of inconspicuous spines dorsally; spurs densely microsetose. Metasoma. T1 width 0.5 × T2 width. T2 length 0.9 × T2 width. T2 disc sculpture dense foveolate. T7 rugose, lateral carinae distinct. S1 with scarcely defined longitudinal medial carina. S2 medially flattened, foveolate. S7 longer than broad, punctured area narrower posteriorly than medially, laterally defined by carinae, apical margin narrow bidentate medially. Genitalia. Paramere long, free length 1.3 × length from base of genital capsule to apex of parapenial lobe; dorso- ventrally flattened, acute apically, medially curved ventrad with apex recurved dorsad in lateral view; with distinct ventral brush of long setae along basal 0.75 × of free paramere length. Cuspis 0.35 × free paramere length, subcylindrical, with scattered short setae throughout and apical posteriorly directed brush of setae 0.75 × cuspis length. Paracuspis short, lobe-like, rounded apically, densely setose. Digitus 0.1 × free paramere length, narrow digitiform, slightly wider basad. Penis valve nearly as long as digitus, ventral margin with two apical teeth; apical distance between teeth 0.13 × length of valve; apical margin with row of distinct setae. Coloration and variations. FEMALES. Body and appendages vary from reddish-brown to black; antenna, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs whitish; T2 with two or four separated or confluent yellow, orange, or reddish integumental spots. Appendages and lateral and pleural body regions generally with whitish setae; dorsum of body with erect and appressed dense black setae except with sparse erect whitish setae on T2 integumental spots and denser patches of appressed whitish setae as follows: transverse curved band on vertex (head usually with setae entirely black), lateral longitudinal mesosomal stripes extending from propodeum anterior to just past pronotal spiracle-level (stripes sometimes terminating on mesonotum or mostly obliterated), and apicomedial patches on T2–5 (sometimes obliterated on T2 or reduced on T3). MALES. Head, mesosoma, and appendages dark reddish-brown to black, except antenna, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs whitish; metasoma variably black and orange, at least T1 and T6 largely blackened (Fig. 22), at least T2 and S2 orange basally (Fig. 25). Head and mesosoma setae mostly dense erect and appressed whitish, except with blackish setae in the following areas: mesoscutum, tegula, axilla, mesoscutellum (some individuals with head and pronotum setae mostly black, Fig. 25); metasoma setae interspersed black and whitish, orange and silvery, or predominantly black. Wings dark brown, often lighter in basal portion among cells; veins dark brown. Distribution. Argentina, Bolivia, Brazil, and Paraguay. Material examined (194 females and 52 males). ARGENTINA, Salta, La Vina, 22–25.XII.1983, M. Wasbauer (1 female, UCDC, C. distincta). BOLIVIA: Santa Cruz: Prov. Chiquitos: Santiago, 700 msnm, XII.1959 (3 females, MIUP, C. vivata); San Jose, 14.III.1990, E. Orihuela (2 females, MIUP, C. distincta and C. vivata); 3 km N Brazilio, 27.II–8.III.1999, Irwin & Parker (1 female 13 males, EMUS, C. distincta); 5 km N Pedro Lorenzo, 2.III.1999, L.A. Stange (3 females, FSCA, C. distincta); Santa Cruz de la Sierra, XI.1910, J. Steinbach (1 female, CMNH, C. vivata). BRAZIL: Amapá, Calçoene, 29.X.1978, M.F. Torres (1 female, MPEG, C. transversa); Bahia, Arembepe, Sítio Mingú, 15.VI.2001, L. Patricia (1 female, UEFS, C. 24
transversa); Feira de Santana, Campus UEFS, 12°13'S 38°58'W, 4.VIII.2000, G. Melo (1 female, UEFS, C. transversa); Santa Rita de Cassia, Riacho Veredão 13, 18.VII.1991, C.R.F. Brandão (1 female, MZSP, C. fasciata); Ceará: Carquejo, Moumbo, V.1968, Dirings (1 female, MZSP, C. cf. fasciata); various dates and collectors (4 females, MZSP, C. cf. fasciata); Goias: Aragarças, 30.III.1953, M. Alvarenga (3 females, MNRJ, C. vivata); Santa Isabel, Ilha de Bananal, Rio Araguaia, 15–29.VII.1957, B. Malkin (1 female, CASC, C. vivata); Maranhao: Carolina, 8.VIII.1953, M. Alvarenga (1 female, MNRJ, C. transversa); Chapadinha, 17.III–19.V.2012, J.R. Souza (9 females, DZUP, C. cf. fasciata); Grajaú, 15.VI.1966 (1 female, MZSP, C. cf. transversa); Minas Gerais, Pirapora, XI.1975, Seabra et al. (1 female, MNRJ, C. fasciata); Mato Grosso do Sul: Três Lagôas, various dates and collectors (20 females, MZSP, C. distincta and C. vivata); Fazenda Nhumirim Pantanal, 16.IX.2011, R. Aranda (3 females, DZUP, C. distincta and C. vivata); Mato Grosso, Barra do Tapirape: 15.VII–15.VIII.1962, B. Malkin (1 female, MIUP, C. vivata); 11.VIII.1962, B. Malkin (1 female, MPEG, C. transversa); various dates and collectors (3 females, MZSP, C. transversa); XII.1960, B. Malkin (1 female, CASC, C. transversa); 30.XII.1962, B. Malkin (1 female, CASC, C. transversa); 1– 2.II.1964, B. Malkin (1 female, MZSP, C. transversa); Cáceres, 11.II–27III.1985, C. Elias (3 females, DZUP, C. vivata); Chapada dos Guimarães, 26.III.2001, W.O. Sousa (1 female, DZUP, C. vivata); Confluencia Xingu Kuluene, 6–47, date unknown, J.C.M. Carvalho (1 female, MNRJ, C. vivata); Diamantino, Alto Rio Arinos, X.1983, B. Silva (1 female, MNRJ, C. vivata); Fazenda Dr. Jose Mendes, Tres Lagoas, 27.V.1964, Exp. Dep. zool. (1 female, MPEG, C. vivata); Fazenda Joao, Pinheiros, Rio Tapirape (1 male, AMNH); Nova Xavantina, Campus of UNEMAT, 12–17.XII.2001, Packer, Mateus, & Coelho (2 females, EMUS, C. vivata); Rosario Oeste, II.1972, Butterfly Co. (1 female, FSCA, C. vivata); various dates and collectors (21 females, MZSP, C. distincta and C. vivata); Utiariti, various dates and collectors (20 females, MZSP, C. distincta and C. vivata); Para, Campus Universitario, UFPA Bragança, 23.XI.2005, R. Pereira, 1 female, MPEG, C. fasciata); Gorotire, Rio Fresco: 18.IV.1983, W.L. Overal (3 females, MPEG, C. transversa); X.1985, W.L. Overal (3 females, MPEG, C. transversa); Mocajuba, Mangabeira, O. M. Rego (8 females, MNRJ, C. transversa); Pedras, Rio Cuminá, 1.X.1969, Exp. Perm. Amaz. (3 females, MPEG, C. transversa); Tumukumake, Aldeia - Araibá, 5.II.1981, E. Oliveira (1 female, MPEG, C. transversa); Sao Paulo: various dates and collectors (5 females, MZSP, C. distincta); Province Unknown, Prainha, 10.V.1905, Ducke, 1 female, MPEG, C. transversa). PARAGUAY: Alto Paraguay: P.N. Defensores del Chaco, mojon 59, 20° 30’ S 59° 47’ W, 15.I.2001 (1 female, MIUP, C. distincta); P.N. Chovoreca, 19° 32’ S 59° 29’ W, 17.V.2002, B. Garcete (1 female, MIUP, C. distincta); Amambay: P.N. Cerro Corá, zona historica, 1–6.II.2006, B. Garcete (4 females, MIUP, C. distincta and C. vivata); P.N. Cerro Corá, en la zona del cerro Muralla, 280 msnm, 22° 39’ S 55° 59’ W, 12.II.2007, B. Garcete (1 female, MIUP, C. vivata); Boqueron: Ganadera Don Daniel, establecimiento Maria Auxiliadora, 171 msnm, 23° 04’ S 61° 01’ W, 12.XII.2003, B. Garcete (1 female, MIUP, C. distincta); Chaco, Filadelfia, I.1995, Arriagagda (1 female, EMUS, C. distincta); Concepción: Cororo: 25.II–1.III.1997, B. Garcete (1 female, FSCA, C. vivata); 25.II–1.III.1997, B. Garcete (7 females, MIUP, C. distincta and C. vivata); Estancia Don Carlos, 160 msnm, 23°24’S 56°30’ W, 16–20.XII.2004, B. Garcete (4 females, MIUP, C. vivata); Estancia San Luis, 22° 24’ S 57° 28’ W, 19–27.X.1999, B. Garcete (1 female 1 male, MIUP, C. distincta); 25.II.1999, B. Garcete (1 female, MIUP, C. distincta); Estancia Santa Herminia, 173 msnm, 21.IV.2004, B. Garcete (1 female, MIUP, C. vivata); San Carlos del Apa, 22° 14’ S 57° 17’ W, 30.X.2002, B. Garcete (1 female 1 male, MIUP, C. distincta); 25 mi SE San Carlos, 27.X.2002, U. Dreschel (3 females, FSCA, C. distincta). Cordillera: Naranjo, barrio San Jose, 290 msnm, 25 32’ S, 57° 03’ W, 5–7.XI.2004, B. Garcete (1 female, MIUP, C. distincta); Paraguari: Compañía Naranjo, 3–8.II.1996, Garcete & Aguilar (3 females, MIUP, C. distincta); La Colmena, Caatymi, 175–320 msnm, 25° 55’S, 56° 47’W, 26–31.I.2005, B. Garcete (3 females 1 male, MIUP, C. distincta); P.N. Ybycui, 129–336 msnm, 26° 04’S 56° 52 W, 20– 25.I.2006, B. Garcete (6 females, MIUP, C. distincta); 14.I.1996 (1 male, MIUP); Presidente Hayes: Cruce Los Pioneros, Estac. Experim. del Ministerio de Agricultura, 22° 50’ S 59° 45’W, 9–10.XII.2001, C. Aguilar (1 female, MIUP, C. distincta); Lolita, Yaragui, 5.III.2003, U. Dreschel (4 females, FSCA, C. distincta); San Pedro, Rio Ypane, Cororo: II.1979, M.A. Fritz (2 males, AMNH); X.1979, Viana (1 female, AMNH, C. 25
vivata); XI.1979, M.A. Fritz (1 male and 2 females, AMNH EMUS, C. vivata); III.1991, Pena (1 female, EMUS, C. vivata); II.1991, Arriagagda (1 female, EMUS, C. vivata); 24.XI–9.XII.1983, malaise trap, M. Wasbauer (20 males, UCDC CSCA); XII.1983, M.A. Fritz (11 males, AMNH). Remarks. The sexes of C. proxima were associated based on identical ITS1 sequences of a male C. proxima and female C. vivata from Minas Gerais, Brazil (Williams, 2012). Subsequently, males have been examined from the same localities as C. distincta and C. vivata in Bolivia and Paraguay. The types of C. proxima and C. transversa are both from Para State in Brazil. Males circumstantially associated with these different female forms have identical genitalia and differ only in minor color differences (Figs 29–31). Females of these formerly recognized species are identical in structurally diagnostic features, being separated by their color patterns only. These color patterns are approximated by various Traumatomutilla species where they co-occur. In fact, each of the color differences that separate two of these previously recognized species has been revealed to be polymorphic within a single Traumatomutilla species in the T. americana species-group (Bartholomay et al. 2019a). For example, in some forms (C. confluenta, Fig. 13, and C. distincta, Fig. 16) the T2 spots are reddish, while in others (C. vivata, Fig. 17, and C. transversa, Fig. 19) the spots are yellow. Every one of the three species in the T. americana species-group, T. americana (Linnaeus, 1758), T. ocellaris (Klug, 1821), and T. quadrum (Klug, 1821), is polymorphic for this trait (Bartholomay et al. 2019a). Intermediate forms are commonly observed between these apparent species, multiple forms are often found in the same locality, and some forms that clearly belong to C. proxima based on structure, possess combinations of color features that preclude them from fitting any of the previously established names (e.g. Figs 14, 15). Cresson (1916) provided a list of specimens with their respective catalog number in his collection, designating them as “types” and stating that those were the specimens used in the descriptions of all his previous studies. Amongst these specimens is the lectotype of Mutilla vivata, which, though never referred to nor labeled as lectotype by Cresson, is considered herein as the lectotype of this species and Cresson (1916) as the paper that established it as such.
Cephalomutilla zelichi Casal, 1963 (Figs 26–28, 32–34, 40–41, 51–55)
Cephalomutilla zelichi Casal, 1963: 74, holotype, femal, Argentina, Salta, Pocitos, XII.1956, M.A. Fritz (AMNH), examined.
Diagnosis. FEMALE. This species can be recognized by the shape of the vertex tubercles, which are rectangular and are separated by greater than the inter-epaulet distance. Additionally, scutellar scale is apically truncate with transverse carinae anteriorly, anterolaterally, and posterolaterally; also, the pygidium is longitudinally rugose. Body length 6–9.5 mm. MALE. The male of this species can be recognized by the cuspis, which is scarcely longer than the digitus, dilated subapically, and has elongate plumose apical setae directed straight backward. Additional useful characters include the posteriorly truncate axilla, the rugose T7 sculpture, and the relatively-straight sparsely-setose sub-cylindrical paramere. Body length 6–10 mm. Description. FEMALE. See Casal (1963). MALE (Hitherto unknown). Head. Rounded, posteromedially expanded; head width 0.9 × pronotal width. Eye almost circular. Ocelli small; OOD 5.7 × DLO, IOD 2.2 × DLO. Occipital carina distinct. Frons, vertex, and gena densely foveolate-punctate, nearly scabrous in some areas. Gena ecarinate. Antennal scrobe with prominent transverse dorsal carina. Clypeus shallow convex medially, concave laterally below antennal insertion; densely punctate with smooth transverse sub-truncate lobe apically. Scape bicarinate. F1 length 1.5 × pedicel length; F2 length 2.1 × pedicel length. Mandible obliquely bidentate apically, lacking dorsal or ventral projections. Mesosoma. Epaulet sub-linear, not connected with humeral carina. Anterior face of pronotum punctate laterally, with broad longitudinal smooth furrow. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum irregularly foveolate without longitudinal carina; notaulus and parapsis mostly obliterated. 26
Mesoscutellum flat, areolate-foveolate. Axilla produced posterolaterally as truncate projection with conspicuous flat posterior face; dorsal face areolate-foveolate anteriorly, smooth posteriorly. Area between mesoscutum and mesoscutellum smooth. Metanotum virtually equally wide throughout, medially areolate- foveolate. Propodeum dorsal and posterior faces scarcely differentiated, coarsely areolate with dense setae basolaterally; lateral face mostly impunctate. Mesopleuron punctation obscured by dense setae, evenly convex without tubercle on dorsal half. Metapleuron virtually smooth, with micropunctures and sub-appressed microsetae. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate or rounded apically; three submarginal cells, third cell scarcely defined or partly obliterated. Legs. Setae simple, meso- and metatibia each with two rows of inconspicuous spines dorsally; spurs densely microsetose. Metasoma. T1 width 0.45 × T2 width. T2 length 0.85 × T2 width. T2 disc sculpture foveolate. T7 rugose, lateral carinae distinct. S1 with scarcely defined longitudinal medial carina. S2 medially flattened, foveolate. S7 longer than broad, punctured area slightly narrowed posteriorly, laterally defined by weak carina, apical margin unidentate medially. Genitalia. Paramere long, free length 1.05 × length from base of genital capsule to apex of parapenial lobe; sub-cylindrical, acute apically, apex weakly dorsad in lateral view; with obscure ventral brush of long setae scattered throughout free paramere length. Cuspis 0.4 × free paramere length, laterally flattened and thickened subapically, with scattered short setae throughout and apical posteriorly directed brush of branched setae 0.74 × cuspis length. Paracuspis short, lobe-like, rounded apically, setose. Digitus 0.32 × free paramere length, narrow digitiform, slightly wider basad. Penis valve scarcely extending beyond parapenial lobe, ventral margin with two apical teeth; apical distance between teeth 0.15 × length of valve; apical margin with row of distinct setae. Coloration and variations. FEMALES. Body and appendages vary from reddish-brown to nearly blackish, metasoma usually darker than head and mesosoma; antenna, mandible, and tarsomeres often partly yellow- brown; tibial spurs whitish; T2 with four separated yellow, orange, or reddish integumental spots. Appendages and lateral and pleural body regions generally with whitish setae; head and mesosoma dorsum mostly dense appressed and erect whitish except longitudinal stripe of black setae medially from frons to medial portion of propodeum. Dorsal metasomal setae mostly black, except T2 integumental spots sparse erect whitish and dense appressed and erect whitish patch on T2 apicomedially (sometimes obliterated), T3 medially (sometimes obliterated or extended to lateral margin, T4–5 medially (sometimes expanded to lateral margin). The specimens from Paraguay, Estancia Don Carlos have the integumental maculae of T2 yellow; Estancia San Luis females have the maculae orange; and in other specimens from Boquerón, Presidente Hayes, and Bolivia, the T2 maculae are red. Two specimens from Presidente Hayes have the anterior maculae of T2 fully fused with the posterior ones; the specimen from Boquerón has them almost in contact; and other specimens have the four maculae on T2 completely separated. Color of setae on T2–5 variable, with a median longitudinal band and lateral bands of white pubescence, the bands separated by black setae; or T3–5 almost totally white pubescence with few black setae. MALES. No significant variations in color and setae were observed in males. Entire body dark reddish-brown to black, except antenna, mandible, and tarsomeres often partly red-brown to yellow-brown; tibial spurs whitish; and T2 and S2 mostly orange except their blackish lateral and posterior margins. Entire body with mostly dense erect and appressed whitish setae, except mesoscutum, axilla, mesoscutellum, and T1 with some erect blackish setae, T2 and T5 posterior half setae blackish; and T6–7 setae mostly blackish. Wings light brown, veins dark brown. Material examined (26 females and 8 males). ARGENTINA, Salta, Pocitos: 5.II.1971, M.A. Fritz (1 male, AMNH); XII.1971, M.A. Fritz (1 male, AMNH); Tartagal, XI.1971, M.A. Fritz (5 males, AMNH). BOLIVIA, Santa Cruz, 3 km N Brazilio, 8.III.1999, Irwin & Parker (1 male, EMUS); II.1971, Fritz (1 female, MIUP, det. Fritz). BRAZIL: Mato Grosso, Rosario Oeste, various dates and collectors (13 females, MZSP). PARAGUAY: Boquerón, Picada Siracua, 3 km, NE del límite Parque Nacional Teniente Enciso, 20° 57’ S, 61° 34’ W, 21.III.2004, B. Garcete (1 female, MIUP); Concepcion: Estancia Don Carlos, 160 m, 23° 24’ S, 56° 30’ W, 16–20.XII.2004, B. Garcete (3 females, MIUP); Estancia San Luis, 22° 24’ S, 57° 28’ W, 25.II.1999 (2 females, MIUP); Parque Nacional Paso Bravo, 22° 27’ S, 57° 13’ W, 27.X.2002, B. Garcete (1 27
female, MIUP); Parque Nacional Teniente Enciso, 400–590 m, pit fall trap, 4–6.XI.2001, M. Leponce (1 female, MIUP); Presidente Hayes: Cruce Los Pioneros: 20.i.1996, B. Garcete (1 female, MIUP); 9– 10.XII.2001, C. Aguilar (1 female, MIUP); Lolita, Yaragui, 5.III.2003, U. Dreschel (2 females, FSCA). Distribution. Argentina, Bolivia, Brazil, and Paraguay. Remarks. Like C. haematodes and C. proxima, the maculae on T2 of females are variable in shape and color. This is the smallest species in the genus. The sex association is based on discovery of males in Pocitos and Tartagal of Salta, Argentina, which were found in the same localities as some of the original paratype females (Casal, 1963).
Discussion
The Mullerian mimicry complexes involving Dasymutilla in North and Central America revealed that the limits between species described by early taxonomists were unclear and that most color and setae characters seemed unreliable for differentiating species (Pilgrim et al. 2009, Wilson et al. 2012). Structural and morphological data coupled with molecular analyses confirmed the synonymy of many species and, in some cases, entire species-groups were in fact a single widespread and highly variable species (Williams et al. 2011a). Recent taxonomic revisions in Traumatomutilla suggest that similar mimetic complexes may be occurring in South America as well, resulting in the synonymy of numerous species previously differentiated by color and setae characters only (Bartholomay et al. 2019a). Cephalomutilla appears to be a third Dasymutillini genus involved in a similar issue, with nine of its previously recognized species being synonymized in the current study. In fact, both Cephalomutilla and Traumatomutilla appear to be influenced by widespread and relatively well-established Mullerian mimicry complexes throughout South America involving numerous genera (KAW pers. obs). Mimicry studies on South American mutillids could confirm this hypothesis and elucidate how each color syndrome is distributed. Cephalomutilla has parallel distribution with certain Traumatomutilla species, especially those of the T. juvenilis species-group (Bartholomay et al. in prep.). In both cases, species are predominantly found in southern South America — T. duplicata (Gerstaecker, 1874), T. miniata (Gerstaecker, 1874), C. zelichi, C. haematodes, and C. graviceps — with other species more commonly found in Brazilian Cerrado and, in some cases, even reaching certain areas of the eastern Amazon forest, such as T. bivittata (Gerstaecker, 1874), T. juvenilis (Gerstaecker, 1874), and C. proxima. More interesting, though, is the fact that both seem to “disappear” in central, western, and northern Amazonian Forest and “reappear” with disjunct Colombian species northeast of the Andes — i.e. T. juvenindica Bartholomay & Williams sp. nov. and C. cabezona Williams sp. nov. These areas West of the Andes in Colombia, particularly the Cesar, Magdalena, and Bolívar departments, seem to be fertile ground for research on the biogeography and dispersal of Dasymutillini. Not only are there unique Dasymutillini species from South American genera in this region, but this is the only area in South America with members of the predominantly North American genus Dasymutilla (Bartholomay et al. 2019b). Further studies with the Northern South American fauna west of the Colombian and Venezuelan Andes will undoubtedly provide valuable insight into the complex relations between Dasymutillini genera.
Acknowledgements
We are grateful for the subject editor and two anonymous reviewers for their help in editing and publishing this paper. We would also like to thank the collection managers and curators that provided for this study, including: Christine LeBeau (AMNH), John Rawlins (CMNH), Robert Zuparko (CASC), Denis Brothers (DJBC), Donald Manley (DGMC), Gabriel Melo (DZUP), Felipe Vivalo (MNRJ), Orlando Tobias Silveira (MPEG), Carlos Brandão (MZSP), and Lynn Kimsey (UCDC). KAW was supported in part by CNPq’s Sciences without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, 28
Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013–0). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9). VML was supported by CAPES.
References
André, E. (1902 [“1903”]) Fam. Mutillidae. In: Wytsman, P., Genera Insectorum, Fasc. 11. Bruxelles, 77 pp. + 3 pls. André, E. (1903) Mutillides nouveaux ou imparfaitemente connus de divers pays. Annales de la Societé Entomologique de France, 72, 417–459. André, E. (1904) Examen critique d’une nouvelle classification proposée par M. le Dr. W. H. Ashmead pour le famille des Mutillides. Revue d’Entomologie. 23(1): 27–41. André, E. (1908) Étude sur les Mutillides du Musée National d'Histoire naturelle de Buénos Aires. Anales del Museo Nacional de Buenos Aires (Ser. 3a), 10, 169–214. Ashmead, W.H. (1899) Superfamilies in the Hymenoptera and generic synopses of the families Thynnidae, Myrmosidae, and Mutillidae. Journal of the New York Entomological Society, 7, 45–60. Bartholomay, P.R., Williams, K.A. & Oliveira, M.L. (2018) New species of Traumatomutilla André in the T. tabapua and T. integella species-groups (Hymenoptera, Mutillidae). Zootaxa, 4433 (2): 361–385. Bartholomay, P.R., Williams, K.A., Lopez, V.M. & Oliveira, M.L. (2019a) Revision of the Traumatomutilla americana species-group (Hymenoptera, Mutillidae). Zootaxa, 4608 (1), 001–034. Bartholomay, P.R., Williams, K.A., Cambra, R.A. & Oliveira, M.L. (2019b) Does the genus Dasymutilla Ashmead occur in South America? The new genus Quwitilla, new combinations, and new distribution records for Neotropical velvet ants (Hymenoptera: Mutillidae). Zootaxa, 4623 (2): 261–282. Brothers, D.J. & Lelej, A.L. (2017) Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research, 60, 1–97. https://doi.org/10.3897/jhr.60.20091 Burmeister, H.C.C. (1875) Mutillae Argentinae. Boletín de la Academia Nacional de Ciencias Exactas existente en la Universidad de Cordova, 1, 461–502. Cambra, R.A. and Quintero A., D. (2003) Tallium lattkei n. sp. (Hymenoptera: Mutillidae) from Venezuela and Colombia, the northernmost distribution record for this neotropical genus. Transactions of the American Entomological Society, 129, 497–502. Casal, O.H. (1963 [“1961”]) Mutillidae neotropicales XVIII. (Hymenoptera). Dos nuevas especies de Argentina y Brasil. Revista de la Sociedad Entomológica Argentina, 24, 71–76. Casal, O.H. (1965a) Tobantilla montonera, género y especie nuevos de Sphaeropthalminae (Hymenoptera, Mutillidae). Revista de la Sociedad Mexicana de Historia Natural, 25, 209–215. Cresson, E.T. (1902) Descriptions of some Brasilian Mutilla. Transactions of the American Entomological Society, 28, 1–82. Cresson, E.T. (1916) The Cresson types of Hymenoptera. Memoirs of the American Entomological Society 1: 79–85. Gerstaecker, A. (1874) Mutillarum Americae meridionalis indigenarum synopsis systematica et synonymica. Archiv fur Naturgeschichte, 40, 41-77, 299-328. International Commission on Zoological Nomenclature (ICZN). (1959) Opinion 542. Designation under the Plenary Powers of a type species in harmony with accustomed usage for the nominal genus 29
"Cephalomutilla" André, (1908) (Class Insecta, Order Hymenoptera). Opinions and Declarations rendered by the International Commission on Zoological Nomenclature, 20(2), 103–110. Klug, J.C.F. (1821) Entomologie brasilianae specimen. Nova Acta Academica Caesareae Leopoldino- Carolinae Germanicae Nature Curiosorum, 10 (2), 305–324. Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Pars II. Editio Duodecimae Reformata. Laurentii Salvii, Holmiae, 795 pp. [pp. 533–1327] Mickel, C.E. (1957) Request for the designation under the plenary powers of a type species for the genus "Cephalomutilla" André, 1908 (Class Insecta, Order Hymenoptera) in harmony with accustomed usage. Bulletin of Zoological Nomenclature, 13(1), 22–25. Mickel, C.E. (1960) Review of the Mutillid Genus Cephalomutilla Andre (Hymenoptera: Mutillidae). Revista Brasileira entomologica de Sao Paulo, 9, 157–168. Nonveiller, G. (1990) Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta, Hymenoptera). Hymenopterorum Catalogus (Nova Editio), 18. Den Haag, SPB Academic Publishing. 1–150. Pilgrim, E.M., Williams, K.A., Manley, D.G. & Pitts, J.P. (2009) Addressing the Dasymutilla quadriguttata Species-Group and Species-Complex (Hymenoptera: Mutillidae): Several Distinc Species or a Single, Morphologically Variable Species? Journal of the Kansas Entomological Society, 82 (3), 231–249. Quintero A., D. & Cambra T., R.A. (1996) Mutillidae of Paraguay. Sphecos, 30, 11–14. Smith, F. (1879) Descriptions of New Species of Hymenoptera in the Collection of the British Museum. Taylor and Fransis, London, 240 pp. Williams, K.A. (2012) Systematics of Mutillidae (Hymenoptera) with special emphasis on Dasymutilla and their Alles. All Graduate Theses and Dissertations. Paper 1200. Utah State Univeristy Press: Logan, UT, USA. 327 p. (Available at ~http://digitalcommons.usu.edu/etd/1200/. Las accessed November 2018.) Williams, K.A., Manley, D.G., Pilgrim, E.M., von Dohlen, C.D. & Pitts, J.P. (2011a) Multifaceted assessment of species validity in the Dasymutilla bioculata species group (Hymenoptera: Mutillida). Systematic Entomology, 36, 180–191. Williams, K.A., Brothers, D.J. & Pitts, J.P. (2011b) New species of Tobantilla Casal, 1965 and a new genus and species, Gogoltilla chichikovi gen. et sp. nov., from Argentina (Hymenoptera: Mutillidae). Zootaxa, 3064, 41–68. Wilson, J.S., Williams, K.A., Forister, M.L., von Dohlen, C.D. & Pitts, J.P. (2012) Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications, 3: 1272.
Figure legends.
Figures 1–3. Cephalomutilla cabezona, sp. nov. 1. Dorsal habitus, female. 2. Lateral habitus, female. 3. Lateral habitus, male.
Figures 4–6. Cephalomutilla graviceps (André, 1903), females. 4, 5. Dorsal habitus. 6. Lateral habitus. 4. San Luis Province, Argentina. 5, 6. La Rioja Province, Argentina.
Figures 7–12. Cephalomutilla haematodes (Gerstaecker, 1874). 7, 8. Lateral habitus, male. 9-12. Dorsal habitus, female. 7. C. vulnerifera, syn. nov. 8. Previously unnamed male. 9. C. argyrosticta, syn. nov. 10. C. albicalcaris, syn. nov. 11. C. haematodes, nominotypical form. 12. C. flavigastra, syn. nov.
Figures 13–21. Cephalomutilla proxima (Smith, 1879), females. 13-19. Dorsal habitus. 20, 21. Lateral habitus. 13. C. confluenta, syn. nov. 14, 21. C. cf. distincta, syn. nov. 15. C. cf. transversa, syn. nov. 16. C. 30
distincta, syn. nov. 17. C. vivata, syn. nov. 18. C. vivata, syn. nov. 19. C. transversa, syn. nov. 19, 20. C. fasciata, syn. nov.
Figures 22–25. Cephalomutilla proxima (Smith, 1879), males, lateral habitus. 13. Holotype: Pará, Brazil. 23, 24. San Pedro, Paraguay. 25. Santa Cruz, Bolivia.
Figures 26–28. Cephalomutilla zelichi Casal, 1963. 26. Dorsal habitus, female. 27. Lateral habitus, female. 28. Lateral habitus, male
Figures 29–37. Cephalomutilla, females. 29-31. C. proxima. 32-34. C. zelichi. 35-37. C. cabezona, sp. nov. 29, 32, 35. Vertex and pronotum, photographs. 30, 33, 36. Vertex and pronotum, line drawings. 31, 34, 37. Pygidial plate, photographs.
Figures 38–45. Cephalomutilla, males. 38, 39. C. proxima. 40, 41. C. zelichi. 42, 43. C. haematodes. 44, 45. C. cabezona, sp. nov. 38, 40, 42, 44. Head and mesosoma, photographs. 39, 41, 43, 45. Mesosoma, line drawings.
Figures 46–65. Male genitalia: dorsal view, ventral view, lateral view, penis valve lateral view, cuspis lateral view. 46-50. C. proxima. 51-55. C. zelichi. 56-60. C. haematodes. 61-65. C. cabezona, sp. nov.
31
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33
CAPÍTULO 4.2 – (em formato de submissão à revista NEOTROPICAL ENTOMOLOGY)
Review of Leucospilomutilla Ashmead and description of the new genus Atlantilla Williams & Bartholomay
(Hymenoptera: Mutillidae: Dasymutillini)
Pedro R. Bartholomay 1*, Kevin A. Williams 2, Roberto A. Cambra3, & Márcio L. Oliveira1
1 Instituto de Pesquisas da Amazônia, Manaus, Amazonas, Brazil.
2 Plant Pest Diagnostics Center, California Department of Food and Agriculture, Sacramento, California, USA.
3Museo de Invertebrados G. B. Fairchild, Estafeta Universitaria Apartado 00017, Universidad de Panamá, Panamá
0824, República de Panamá.
* Corresponding author. E-mail: [email protected]
Abstract
Atlantilla gen. nov., is proposed based on the combination of previously undescribed males from the Atlantic Forest and females of Traumatomutilla auriculata (Gerstaecker, 1874). The previously unknown male of Leucospilomutilla staurogastra Suárez, 1973 is described. Keys and illustrations are provided for each of the three known
Leucospilomutilla species.
Key words: Sphaeropthalminae, Dasymutillini, velvet ants, Neotropics, Brazil
Introduction
Williams et al. (2017) established 14 species-groups based on females in the highly diverse Tramatomutilla André,
1902 using exclusive combinations of structural characters. They highlighted the need for future studies on each group to further organize this species-rich genus. Perhaps the most unusual Traumatomutilla belonged to the monospecific
Traumatomutilla auriculata species-group. This insect has strange thickened bristle-like setae covering most of the body, unlike other Traumatomutilla. Additionally, this species has T2 marked with setal spots only, rather than the 34 cuticular spots that mark ~95% of Traumatomutilla species (KAW pers. obs.). In these and other features mentioned below, Traumatomutilla auriculata (Gerstaecker, 1874) is apparently closer to Leucospilomutilla Ashmead, 1903 than to other Traumatomutilla. Although they recognized this issue, Williams et al. (2017) kept T. auriculata inside
Traumatomutilla, waiting for additional evidence that might support altering its genus placement.
Recent sampling in southeastern and northeastern Atlantic Forest areas of Brazil, the only known distribution for T. auriculata, yielded several males that appear morphologically intermediate between Traumatomutilla and
Leucospilomutilla. In this study we describe these males and associate them with T. auriculata based on distribution and morphology. Based on derived unique combinations of characters in both sexes and apparent autapomorphies of the male axilla and genitalia, we describe Atlantilla gen. nov. to contain this species.
The closely related genus Leucospilomutilla includes three species, but only one of them, Leucospilomutilla cerbera (Klug, 1821), is known from both sexes. During this study, males of Leucospilomutilla staurogastra (Suárez,
1973) were identified and are described below. Habitus photographs and keys are provided for all known members of the genus.
Materials and Methods
The following acronyms are used for institutions housing the material discussed in the current study:
AMNH - American Museum of Natural History, New York, New York, USA.
CASC - Department of Entomology, California Academy of Sciences, San Francisco, California, USA.
CPDC - Centro de Pesquisas do Cacau, CEPEC, CPDC, Divisão de Zoologia Agrícola, Itabuna, Bahia, Brazil.
CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA.
DZUP - Coleção Entomológica Padre Jesus Santiago Moure, Departamento de Zoologia, Universidade
Federal do Paraná, Curitiba, Paraná, Brazil.
EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA.
MIUP - Museo de Invertebrados G.B. Fairchild, Universidad de Panamá, Panamá, Panamá.
MNHN - Muséum nationale d’Histoire naturelle, Paris, France.
MNRJ - Museu Nacional, Universidade Federal do Rio de Janeiro, São Cristovão, Rio de Janeiro, Brazil.
UEFS - Museu de Zoologia da Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil. 35
UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brazil.
USNM - United States National Museum of Natural History, Smithsonian Institution, Washington D.C., USA.
ZMB - Museum für Naturkunde Berlin, Berlin, Germany
ZMUC - Københavns Universitet Zoologisk Museum, Entomologi Afdeling, Copenhagen, Denmark
Transcription of label data as well as morphological terminology and measurements for females followed those adopted by Bartholomay et al. (2018) with the addition of a new term, “bristle setae”, used to refer to setae that are conspicuously thicker and easily distinguishable from one another in comparison to simple, brachyplumose, or plumose setae. Measurements for male ocellar and ocular distances and acronyms followed those used in Cambra et al. (2018). Genitalic features and measurements follow Williams et al. (2011).
Atlantilla Williams & Bartholomay gen. nov.
Type species. Mutilla auriculata Gerstaecker, 1874, designated here.
Diagnosis. FEMALE: Bristle-like setae present at least on T2; antennal scrobe with strong sinuate dorsal carina almost reaching eye; mandible apically unidentate; T1 shape sub-nodose, much narrower than T2, smoothly rounded, dorsal and anterior faces indistinct; Lateral felt line wide and short; T6 slightly convex, with subcircular pygidium coarsely and irregularly rugose; S6 with bilobate projection apically. MALE: Simple setae present only; scrobal carinae absent laterally, vestigial dorsally; clypeus medially weakly elevated, ventral margin with a pair of short blunt projections; mandible unidentate; notaulus and parapsis vestigial, restricted to posterior third or half of mesoscutum; axilla with distinct flattened dorsal surface, connected to scutellum apically by its internal margin and produced as a hook-like projection on its external margin apically; fore wing marginal cell acute apically; T1 shape subpetiolate, slightly constricted posteriorly and much narrower than T2; T2 with narrow lateral felt line; hypopygium simply rounded medioapically; parapenial lobe well developed, blunt apically, digit-like; penis valve with strong apical tooth and greatly expanded subapical tooth ventrally.
Description. FEMALE: Integument usually black, mostly densely and finely punctate to reticulate. Setae simple to bristle-like. Head. Strongly transverse, slightly produced behind eyes with posterolateral angle armed with smooth tubercle, entirely finely and densely punctate to reticulate. Occipital carina conspicuous. Eye sub-spherical, strongly 36 protruding, surface shining, ommatidia scarcely discernible. Antennal scrobe broad, horizontal, with strong transverse dorsal carina reaching over antennal tubercle, lacking lateral carina. Clypeus with slightly protruded basomedial triangular area and strongly concave apicomedial transverse area, anteroventral margin virtually straight. Malar space greater than basal height of mandible. Hypostomal carina strong, extending sinuously to posterior mandibular condyle, conspicuously elevated medially. Scape simple. Pedicel virtually as long as wide, shorter than F1; F1 longer than wide and longer than F2. Mandible weakly curved basally, slender, tapering, unidentate apically, no ventral basal tooth, notch, or flange. Maxillary palp 6-segmented; labial palp 4-segmented; basal palpomeres strongly flattened and broadened, densely covered with short erect setae. Mesosoma. Longer than wide, sub-rectangular, constricted anterior to propodeal spiracles, evenly narrowed posterior to propodeal spiracles, dorsum convex; propodeum weakly rounded posteriorly, disk and declivity indistinct; surface densely and coarsely reticulate, coarsest and densest dorsally, weakly and sparsely areolate to impunctate laterally. Pronotal dorsal face (excluding anterior collar) with anterior margin slightly convex and sharp in profile; epaulet variable, poorly distinguished from anterior margin of pronotum and reduced to a minute tuft of short fine setae on slight tubercle, to strongly produced from anterior margin of pronotum, slightly sharp; humeral angle acute; posterior margin vestigially defined by aligned intervals. Scutellar scale absent.
Pronotal-mesopleural suture entirely distinct. Mesopleuron almost smooth in small anterior area; vertical mesopleural ridge present throughout, swollen but not angulate, rounded. Metapleuron sculptured throughout. Metapleural- propodeal suture entirely evident but sometimes obliterated anteriorly, concealed by dense setae; endophragmal pit small, inconspicuous, obscured by setae, surrounding area virtually flat. Lateral face of propodeum coarsely and sparsely reticulate to unsculptured anterad. Mesosternum with strong oblique carina and conspicuous median pit between mesocoxae. Metasternal process subtriangular, slightly less than a third of metacoxal length. Legs. Tibial spur formula 1–2–2. Foreleg with tarsal comb. Meso- and metatibiae each with two rows of prominent stout spines, few spines in each row, spines smaller basad; each subapically with inconspicuous secretory pore near base of inner apical spur; apical spurs narrow, finely serrate laterally, outer spur shortened. Metacoxa with strong slightly arcuate carina on inner margin, ending abruptly before coxal apex. Metasoma. Surface entirely finely and densely punctate, sparser and coarser on sterna and sparser anterolaterally on T2. T1 shorter than wide, about 1.8 × width of T2, strongly constricted on anterior third and evenly expanded from constriction apicad, weakly convex in lateral view, anterior and dorsal faces indistinct; anterior auricle prominent, vertically lamellate but apparently dentate from above. T2 with a pair of large sub-rounded patches of appressed bristle-like setae; lateral felt line wide. T6 with conspicuous pygidium, slightly convex, large, rounded, defined laterally by carinae and with coarse irregular rugosities throughout. S2 evenly convex, 37 with conspicuous medial longitudinal crest-fold on anterior third, without posterior transverse groove, lacking felt line.
S6 posterior margin with strong bilobate projection.
MALE: Integument black with strong dense punctuation, lacking any reflection. Setae simple throughout. Head.
Roundly transverse, conspicuously narrower than mesosoma, entirely finely and densely punctate. Occipital carina distinct dorsally. Vertex conspicuously swollen posteriorly, swelling almost carinate posterolaterally. Eye broadly semispherical, strongly protruding, surface shining, ommatidia discernible. Ocelli small. Antennal tubercles slightly widely separated. Antennal scrobe deep, reaching eye margin, with a weak inconspicuous blunt tubercle on its dorsal margin, lacking lateral carina. Clypeus depressed immediately below insertion of antennae, overall slightly concave, anterior/ventral margin with a pair of short blunt tubercles medially. Malar space shorter than basal height of mandible. Genal carina absent. Hypostomal carina evident, evenly high throughout, extending in a smooth curve to posterior mandibular condyle. Oral fossa longer than smooth postgenal bridge. Scape with single sharp longitudinal carina anteroventrally. Pedicel slightly virtually as wide as long, shorter than F1, which shorter than F2. Mandible in dorsal/anterior view strongly and evenly curved, tapering apically, with vestigial longitudinal carina dorsally on basal half, unidentate apically, lacking any ventral tooth or notch. Maxillary palp 6-segmented; labial palp 4-segmented; basal palpomeres strongly flattened and broadened, densely covered with short setae. Mesosoma. Surface finely and densely punctate to reticulate except smooth or nearly so on metapleuron, posteroventrally on mesopleuron, and anteriorly on lateral face of propodeum; scutellum and most of propodeum reticulate. Pronotal dorsal face (excluding anterior collar) with anterior margin slightly convex, rounded in profile, epaulet indistinguishable from anterior margin of pronotum in dorsal view, reduced to a narrow tuft of short setae in anterior view humeral angle rounded; posterodorsal margin broadly V-shaped. Tegula broadly subcircular, strongly convex, almost entirely smooth and glabrous, except along punctate anterior margin. Mesoscutum with notaulus and parapsidal lines vestigial, restricted to posterior third of scutum, posterolateral corner slightly forming a rounded projection discontinuous with axilla.
Scutellum virtually flat; axilla virtually as long as wide, flat dorsally, narrowly connected to scutellum apicointernally, terminating on a hook-like projection apicoexternally; projection curved inwardly in posterior view and posteriorly in lateral view; dorsal surface of axilla coarsely and sparsely punctate except along smooth impunctate posterior margin.
Metanotum simple, transverse, dorsellum poorly delimited laterally, concealed by dense setae. Propodeum evenly convex, dorsal and lateral faces indistinct. Metasternal process bluntly bidentate, apex obtuse, much shorter than metacoxal height, ending well before metacoxae. Wings: Forewing elongate sclerotized pterostigma nearly 4 × longer than wide; marginal cell elongate its maximum length more than 3 × its maximum width, acute apically; three closed 38 submarginal cells, apical veins of third inconspicuous. Legs: Tibial spur formula 1–2–2. Meso- and metatibiae without dorsal spines; apical spurs with microserrated margins, densely clothed with microsetae. Metacoxa ecarinate.
Metasoma. Punctation overall dense and fine except much sparser on S2 and anteriorly on T1; T1 much longer than wide, about 0.5 × width of T2, more or less evenly expanded from base but conspicuously swollen on posterior half, anterior and dorsal faces indistinct, slightly constricted apically; conspicuously angulate dorsolaterally on anterior third anterior auricle poorly developed, roundly lamellate vertically, scarcely discernible from above. T2 with lateral felt line narrow and long. S1 with vestigial longitudinal carina, posterior half of carina flat, smooth and shinning. S2 in lateral view evenly convex throughout, lacking felt line. Hypopygium wider than long, transversely sub-elliptical, surface virtually flat, posterior margin simply rounded, lacking any medial projections or notches. Genitalia. Paramere moderately long, almost as long as length from base of genital capsule to apex of parapenial lobe, almost cylindrical, subacute apically, apex weakly curved dorsad in lateral view, virtually straight in dorsal view. Cuspis scarcely laterally compressed, basal half shallow incurved, apical half parallel, setae denser ventrally, apical half with some ventral setae sinuous; paracuspis a short truncate lobe; digitus slightly elongated, slightly wider basad, prominent.
Penis valve without any conspicuous concave or convex areas; with two apical sharp teeth on ventral margin, basal tooth greatly expanded; basal tooth vestigially serrated on ventral margin; apical margin with sparse short inconspicuous setae.
Species included. Type species only.
Distribution. Southeastern and northeastern Atlantic Forest regions of Brazil (Espírito Santo and Bahia).
Etymology. From the Latin Atlanticus “of or connected with Mount Atlas or the adjacent ocean, Atlantic”, in reference to the Atlantic Forest biome to which this genus distribution is restricted.
Remarks. Williams et al. (2017), highlighted the presence of bristle setae and the resemblance of the females of A. auriculata (then Traumatomutilla auriculata) to the closely related Leucospilomutilla Ashmead. Though females of A. auriculata are similar to Leucospilomutilla in overall setae and shape of the pygidium, they lack the diagnostic mesosomal projections of Leucospilomutilla. Additionally, females of A. auriculata have a conspicuous transverse and quadrate head and a stouter body shape, as opposed to the comparatively slender females of Leucospilomutilla that also have the body sculpture coarser and more irregular than Atlantilla. Unlike females, the male of A. auriculata only resembles Leucospilomutilla in genitalia, especially the penis valve. Externally, the males closely resemble those of
Traumatomutilla differing mainly by having the axillar projections connected to the scutellum apico-internally and the posterior margin of the hypopygium simply rounded, lacking any projections or invaginations. 39
Atlantilla auriculata (Gerstaecker, 1874) comb. nov.
(Figs 1–13, 33–37)
Mutilla auriculata Gerstaecker, 1874. Archiv Für Naturgesichte 40: 67. Holotype female, Brazil (ZMB), examined.
Ephuta (Traumatomutilla) auriculata: André, 1902, p. 54.
Traumatomutilla auriculata: André, 1904, p. 40.
Diagnosis. FEMALE and MALE. This is the only known species in the genus.
Redescription. FEMALE. Body length 19 mm. Coloration. Head, mesosoma, and metasoma reddish-black; mandibles reddish-brown basally and black apically; antennae reddish-black, except F2–10 ventrally reddish-brown; legs reddish-brown. Tibial spurs yellowish-white. Head clothed with interspersed dense decumbent black setae and erect reddish-black setae, except frons with medial patch of sparse thick silvery-golden setae; spots of frons bifurcated and narrower dorsad and expanded ventrad, reaching internal margin of eyes; ventral surface of head with sparse erect silvery-golden setae; clypeus with dense silvery-golden setae on apical margin. Lateral face of pronotum with fine, dense, appressed and sericeous silvery-golden setae. Mesopleuron with fine, dense, appressed and sericeous silvery- golden setae throughout, except mesopleural ridge with dense erect silvery-golden setae and ventrally above mesocoxa with dense appressed silvery-golden setae. Metapleuron with dense appressed silvery-golden setae. Lateral propodeal face with sparse erect silvery-golden setae. Mesosomal dorsum with dense decumbent lanceolate and sparse erect simple reddish-black, except mesonotum posterior margin with sparse appressed thick silvery-golden setae extending antero-laterally nearly to pronotal spiracle; propodeal dorsum clothed with dense appressed and sparse erect silvery- golden setae, except narrow longitudinal area medially with dense appressed black setae. Legs with silvery-golden setae, except tarsi ventral surface setae dense golden-brown. T1 with sparse erect and appressed silvery-golden setae laterally; dense appressed and sparse erect silvery-golden setae sublaterally, dense appressed black setae medially; T1 virtually asetose antero-medially. T2, except orange setae spots, with dense decumbent lanceolate black setae and sparse erect reddish-black setae, except antero-laterally with sparse decumbent and erect silvery-golden setae, lateral margins and felt line with dense appressed silvery-golden setae. T2–5 fringes with dense appressed silvery-golden thick setae, except medially with dense appressed black setae; medial areas of black setae narrower posterad. T6, except pygidial plate, with dense decumbent silvery-golden thick setae. S1–5 clothed with sparse erect and decumbent 40 silvery-golden setae. S2–5 fringes with dense appressed silvery-golden setae. S6 with dense decumbent silvery-golden setae. Head. Posterior margin concave. Occipital carina virtually straight dorsally in posterior view, evenly equally wide throughout, except for lateral tubercles of vertex. Tubercles of vertex “comma-shaped”, wider at apex, 2.0 × as long as apical width. Head width 0.8 × pronotal width. Eye almost circular, length in frontal view slightly longer than distance from ventral margin to mandibular condyle. Front, vertex, and gena densely, finely and sharply areolate; areolations larger on gena and malar space. Genal carina present, greatly reduced, irregular and interrupted by punctures. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, separated from eyes, reaching and advancing over antennal tubercles as conspicuous ridge; lateral scrobal present carinae, reaching eyes, nearly receiving dorsal carina subapically. Antennal tubercles irregularly rugose, with longitudinal ridge medially, slightly conical in dorsal view. Scape densely punctured. F1 2.0 × pedicel length; F2 1.75 × pedicel length. Mesosoma. Mesosoma 0.8 × as long as wide. Pronotum virtually as wide as mesothorax. Mesosomal dorsum densely sharply finely areolate, areolations slightly larger postero-laterally and over propodeum. Anterior face of pronotum conspicuously longitudinally striate. Humeral carina present, disconnected from epaulet, not produced apically; antero-lateral corners of pronotum slightly angulate in dorsal view. Epaulets strongly produced from anterior margin of pronotum, slightly sharp. Pronotal spiracle virtually flat against lateral of pronotum. Lateral face of pronotum sparsely and shallowly punctured with interspersed dense micropunctures, except low micropunctate swelling antero-ventral in relation to pronotal spiracle. Mesopleuron micropunctate anteriorly, dense and coarsely reticulate along mesopleural ridge, and sparsely punctate basally above mesocoxa. Metapleuron sparsely and finely punctate. Lateral face of propodeum sparsely punctate to densely and coarsely reticulate posterad.
Post-spiracular area undefined. In dorsal view, mesosoma widest at middle of mesonotum. Post mesonotal tubercle absent. Lateral margin of mesosoma simply diverging anterior to propodeal spiracle. Propodeal spiracle virtually flat against side of mesosoma. Scutellar scale present, divided sub-medially into two separated unequal scales; wide, separated from lateral margins by little more than combined width. Anterolateral carinae absent. Scabrous intervals on scutellar area present but inconspicuous. Propodeum narrowed posterior to spiracle, convex in lateral view, posterior face significantly longer than dorsal face. Legs. Meso- and metafemora conspicuously rounded apico-externally; metatarsi without any conspicuous expansion apically, with a pair of conspicuously modified setae laterally.
Metasoma. T1 shape sub-nodose 0.55 × as wide as T2. T2 0.9 × as long as wide, with maximum width at midlength.
Disc of T2 densely and coarsely punctate; punctures sparser and larger laterally. T3–6, except pygidial plate, densely and finely punctate. S1 with low rounded longitudinal carina; carina notched posteriorly and slightly higher anteriorly. 41
S2 densely and shallowly punctate, punctures conspicuously denser and smaller antero-medially; antero-medial crest- fold conspicuous, impunctate; subapical slope anterior to fringe absent. S3–6 densely punctate; punctures smaller and denser on S5–6. Pygidium broadly subcircular, defined by lateral carinae, except at plate base; surface longitudinally irregularly striate throughout; striae conspicuously transverse at apical fifth of plate.
MALE (Hitherto undescribed). Body length 8.0 –12.0 mm. Coloration. Head, mesosoma, metasoma and appendages black. Tibial spurs white. Wings dark fuscous throughput, veins brown; hindwing overall slightly lighter basad. Head clothed with dense decumbent and erect black setae, except for few dark-golden setae on ventral surface and base of mandibles. Mesosoma clothed with dense decumbent and erect black setae, except for mesopleuron with scattered inconspicuous appressed dark-golden setae, metanotum with sparse decumbent and erect dark-golden setae and propodeum clothed with sparse erect dark-golden setae. Legs clothed with sparse erect and decumbent dark-golden setae, except femora with few decumbent dark-brown setae apico-dorsally; meso- and metatibiae with setae lighter dorsally. Metasomal terga, including fringes, clothed with dense decumbent and erect black setae except T1with dense appressed and erect dark-golden setae, sparser anterad; T2 with sparse decumbent dark-golden setae latero-basally; lateral felt lines on T2 with dense appressed brownish-black setae; lateral margins of T2 below felt lines completely asetose. Metasomal sterna clothed with dense decumbent erect black setae, except S1 and S2 with sparse erect golden setae; S2 with brownish decumbent interspersed with golden setae; S3–4 with sparse golden setae interspersed with black setae. Head. Rounded sub-quadrate, posterolateral angles rounded, conspicuously swollen postero-medially, swelling markedly angulate laterally. Head width 0.85 × pronotal width. Eye almost circular. Ocelli small; OOD 4.25
× DLO, IOD 0.8 × DLO. Occipital carina distinct. Vertex, front and gena densely and finely punctate. Gena ecarinate.
Antennal scrobe concave to eye margin, virtually devoid of any carinae, with inconspicuous transverse prominence meso-dorsally. Clypeus weakly convex medially, concave laterally immediately below antennal insertion; finely and densely punctate; with a pair of short tooth-like projections medially on apical margin. Scape unicarinate.
Flagellomere 1 1.7 × pedicel length; flagellomere 2 2.0 × pedicel length. Mandible oblique unidentate, lacking dorsal or ventral projections. Maxilla and labium longer than broad. Maxillary palp 6-segmented, third and fourth segments slightly flattened and apically expanded, other segments almost cylindrical. Labial palp 4-segmented, second and third segments slightly flattened and apically expanded, other segments almost cylindrical. Mesosoma. Epaulets inconspicuous, poorly defined, reduced to a narrow micropunctate area starting at anterior face of pronotum and ending on small rounded area on anterior margin of dorsal face of pronotum, disconnected from humeral carina.
Anterior face of pronotum virtually impunctate, smooth, with medial longitudinal concave are starting at pronotal 42 collar and ending in an arcuate carinae on anterior margin of dorsal face of pronotum. Dorsal face of pronotum coarsely and densely punctate to reticulate. Tegula convex, mostly smooth, except antero-inner margins coarsely punctate with decumbent setae. Mesoscutum densely and coarsely punctate; notaulus and parapsis vestigial, reduced to posterior third of mesoscutum. Scutellum virtually flat, sloping posteriorly, densely reticulate. Axilla produced postero-laterally as truncate projections, connected to scutellum apico-internally, terminating in a postero-ventrally produced hook-like projection apico-externally; axillar projections flat dorsally, coarsely and punctate. Metanotum equally wide throughout, with dense short silvery-golden setae, dorsellum obscured by dense setae. Propodeum roundly convex, densely and finely reticulate throughout; reticulae smaller and denser laterad; dorsal face smoothly convex throughout; posterolateral and dorsolateral corners rounded; dorsal and posterior indistinguishable.
Mesopleuron smoothly rounded, slightly swollen on dorsal half; micropunctate anteroventrally and postero-ventrally; coarsely, shallowly and sparsely punctate meso-ventrally, punctures denser and deeper dorsad along mesopleural ridge; coarsely and densely punctate to reticulate dorso-medially; mesopleural ridge not at all pronounced, slightly swollen, rounded. Metapleuron virtually smooth, shinning, except dorsal and basal fourths micropunctate. Wings. Fore wing pterostigma sclerotized, elongated, 3 × as long as wide; marginal cell elongated, acute apically; three submarginal cells, apical veins of third cell weak. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.85 × its width. T1 densely and coarsely reticulate; T2–7, except pygidium, densely and coarsely punctate; pygidial area mostly unsculptured basally, sparsely and finely punctate apically; not defined by lateral carinae. S1 sparsely and coarsely punctate, longitudinal carina vestigial, equally high throughout, slightly pronounced basally. S2 sparsely and finely punctate, weakly swollen longitudinally at base, lacking any pits. S3–7 (hypopygium) densely and coarsely punctate; punctures distinctly sparser medially and on S7. Hypopygium transversely ovate, broader than long, apical margin simply rounded, lacking any medial or lateral projections or invaginations. Genitalia. Parapenial lobe produced to incurved digitiform process.
Paramere subcylindrical, basically straight laterally, downcurving to shallow upcurved apex, ventro-laterally with long dense setae. Cuspis 0.7 × free paramere length, scarcely laterally compressed, basal half shallow incurved, apical half parallel, setae denser ventrally, apical half with some ventral setae sinuous. Paracuspis sessile, lobe-like, rounded apically, densely setose. Digitus 0.3 × free paramere length, incurved in dorsal view, slightly upcurved in lateral view.
Penis valve scarcely extending beyond parapenial lobe, ventrally with large apical tooth and larger subapical tooth with comb of short sharp teeth along basal slope of subapical tooth; apical distance between teeth 0.25 × length of valve; row of minute setae along apical margin; outer margin with large projecting tubercle. 43
Distribution. Brazil (Bahia and Espírito Santo).
Material examined. Type material. Holotype. f#, BRAZIL, v. Olfers, 21031 (ZMB). Additional material. BRAZIL,
Bahia, 1f#, Coll[ection] E. André 1912 (MNHN); 1f#, same label data except (ZMUC); Espírito Santo, Pinheiros,
Res[erva] Bio[lógica] Córrego do Veado, Trilha Jaboti, 18°21’S 40°10’’W, 17m#, 27.xi–06.xii.2011, [Armadilha de]
Malaise 7, M.T. Tavares & eq[uipe] col[etores], UFES nº[número] 96600–96613, 96615, 96598, 96599 (UFES); 3m#, same label data except, [Armadilha de] Malaise 7, UFES nº[número] 96237–96238 (UFES); 3m#, same label data except, [Armadilha de] Malaise 8, UFES nº[número] 96426, 96429, 96432 (UFES); 2m#, same label data except,
[Armadilha de] Malaise 6, UFES nº[número] 98205, 98026 (UFES); 1m#, same label data except, [Armadilha de]
Malaise 4, UFES nº[número] 97458 (UFES); 1m#, same label data except, [Armadilha de] Malaise 5, UFES nº[número] 98079 (UFES); Sooretama, Res[erva] Bio[lógica] de Sooretama, Trilha Ana, 19°03’18’’S 40°08’43’’W,
2m#, 06–14.XII.2011, [Armadilha de] Malaise 6, M.T. Tavares & eq[uipe] col[etores], UFES n°[número] 78117,
78114 (UFES); Itapemirim, Faz[enda] Usina Paineiras, P2[sic], 20°57’02’’S 41°03’20’’W, 1m#, 19–26.XI.2010,
[Armadilha de] Malaise B13[sic], M.T. Tavares & eq[uipe], UFES n°[número] 66673 (UFES).
Remarks. Female setal color varies considerably. Specimens have been observed with the head completely black, mesonotal silvery-golden setae virtually absent, medial area of black setae on propodeum and medial areas of black setae on metasoma conspicuously wider, silvery-golden setae of T6 interrupted medially with black setae. The setae of the holotype’s metasomal sterna have severely deteriorated but are still observable and this was confirmed by examining other recent specimens of A. auriculata. The tubercles on the vertex are conspicuously larger on the holotype than in other examined specimens. Minor variations on the sculpture of the head were observed in certain specimens wherein the areolations where sparser and shallower on the malar space and ventral half of the gena.
Variations regarding the carina on S1 were also observed, with some specimens having the carina much more developed than the type.
Leucospilomutilla Ashmead, 1903
Type species. Mutilla cerbera Klug, 1821 (designated by Ashmead, 1903)
Male description. Casal, 1963.
44
Diagnosis. FEMALE: Females can be recognized by having abundant bristle-setae on the body, a smooth tubercle postero-laterally on the vertex, a bi-lobed scutellar scale, the mesonotum armed laterally with a tooth-like projection, the felt line setae black and the metasoma marked only with black and whitish setal patterns. MALE: Males can be recognized by the presence of bristle-setae, the elevated truncate axillar process, the mesopleuron with a lateral dentiform projection, the dorsoventrally flattened and subapically contorted paramere, and the large triangular subapical ventral penis valve tooth.
Species included. Three species: L. cerbera (Klug, 1821); L. chemea Casal, 1961; and L. staurogastra Suárez, 1973.
Distribution. Widespread in South America (Bolivia, Brazil, and Paraguay).
Remarks. In South America, this genus is immediately recognizable by the numerous sharp tubercles around the lateral margins of the mesosomal dorsum and the contrasting black and white setal markings. The genus
Bothriomutilla Ashmead in Australia, which also belongs to tribe Dasymutillini, is superficially similar to
Leucospilomutilla. Both genera have tooth-like projections on the pronotum, mesonotum, and propodeum, but
Leucospilomutilla have denser setae and T2 convex, while Bothriomutilla have sparser setae, especially on the mesosoma, and T2 with a medial depression. Males are less obviously similar: Bothriomutilla have the axillae unarmed, while Leucospilomutilla have a distinct truncate axillar projection posteriorly.
Key to females of Leucospilomutilla species
1. Apex of hypopygium with four teeth (see Casal 1963, Fig. 10); head entirely with whitish setae (Figs 14, 21); T2 laterally with entire longitudinal band of whitish setae (Figs 14, 17); predominantly from Caatinga areas of Brazil
(Fig. 48) … L. cerbera (Klug, 1821)
— Apex of hypopygium with two teeth; head with variably extensive black setal markings (Figs 15–16); T2 laterally anterior and posterior white setal markings, widely interrupted by black setae above felt line (Figs 15–16); known from farther South or East in Bolivia, Brazil, and Paraguay (Fig. 48) … 2
2(1). Head setae mostly whitish, with white setal mark on front reaching inner eye margin (Fig. 16); T2 sub-apically with medial and lateral white setal marks connected by transverse whitish setal line (Fig. 16); known from Cerrado and Chaco areas of Brazil and Paraguay (Fig. 48) … L. staurogastra Suárez, 1973
— Head setae mostly blackish, with medial white setal mark vertex (Fig. 15); T2 sub-apically with medial and lateral white setal marks widely separated by black setae (Fig. 15); known only from Bolivia (Fig. 48) … L. chemea Casal,
1961 45
Key to the males of Leucospilomutilla species (male of L. chemea unknown)
1. Head and mesosoma with setae predominantly whitish (Fig. 23); S1 elevated medially, forming pronounced tooth
(see Casal 1963, Fig. 8); cuspis basically straight with spatulate apex (Fig. 42); predominantly from Caatinga areas of
Brazil (Fig. 48) … L. cerbera (Klug, 1821)
— Head and mesosoma with setae predominantly blackish, whitish setae restricted to mesoscutellum, metanotum, and propodeum (Fig. 28); S1 medially not forming pronounced tooth; cuspis incurved basally with slender apices parallel
(Fig. 47); known from Cerrado and Chaco areas of Brazil and Paraguay (Fig. 48) … L. staurogastra Suárez, 1973
Leucospilomutilla cerbera (Klug, 1821)
(Figs 14, 17, 20, 21, 23–27, 38–42)
Mutilla cerbera, Klug, 1821. p. 312, Holotype, female, Brazil, Bahia (ZMB).
Leucospilomutilla cerbera: Ashmead, 1903, p. 310.
Leucospilomutilla cerbera: Casal, 1963 (male description).
Diagnosis. FEMALE. This is the only Leucospilomutilla species with the head setae entirely white (Fig. 21) and the apex of hypopygium with four teeth. The T2 color pattern is also diagnostic: there is an entire sub-lateral longitudinal band of whitish setae and sub-apically the medial and lateral white setal marks are widely separated by black setae
(Fig. 14). Additionally, the dentiform projections of the epaulet and pronotal spiracle are each nearly as large and sharp as the mesonotal projection, and the pygidium has dense irregular rugae (Fig. 20). Body length 14–20 mm.
MALE. The male of this species has the head and mesosomal setae mostly whitish (Fig. 23) and the cuspis basically straight with a spatulate apex (Fig. 42). Additionally, white setae entirely cover the apical portions of T1, T2, and T3 and a large patch on the T2 disc; the wings are transparent pale brown; and the paramere is strongly contorted subapically. Body length 15–18 mm.
Material examined (136 females and 6 males). BRAZIL: Bahia: Baixa Grande, 8.V.1999, C. Lopes (1f#, UEFS);
Barreiras, X.1952, E.M. Barbosa (1f#, MNRJ); Castro Alves, 1994, C.A. Carvalho (1f#, MIUP); Feira de Santana,
Campus da UEFS, I.1999, J.M.F. Soares (3f#, CPDC); Juazeiro, XI.1907, E.Garza (1f#, CASC); Vitoria da Conquista,
Poço Escuro, 145028S 0405016W, 12.I.2001, J.R. Maia (1f#, CPDC); Maracas, II.1965, F.M. Oliveira (6f#, MNRJ); 46
Vitoria da Conquista, I.1963, F.M. Oliveira (1f#, MNRJ); Lençois, 4 km NW de Lençois, 22–24.X.1999, Freddy,
Mazzarolo, (2f#, UEFS); Lençois, Pai Inácio, 22–23.X.1999, Freddy; Mazzarolo, (2f#, UEFS); Lençois, 29–
30.IV.1999, A. Pales, 1m#, UEFS); Feira de Santana, Matinha, 22.IV.2000, Marcia (1f#, UEFS); Maracás, Maracás,
19.XI.2004, (1f#, UEFS); Lençois, 28–29.IV.1999, Ereira-Dias (1f#, UEFS); Serra do Orobo, 8.IV.2005, (1f#, UEFS);
Salvador, IV.1999, André (1f#, UEFS); Feira de Santana, 9.XI.1999, Cristiano (1f#, UEFS); Costa do Sauípe,
14.XI.2004, I. Castro (1f#, UEFS); Pedra Branca, X.2004, Balbino (1f#, UEFS); [locality unknown], G. Bondar (1f#,
MNRJ); Ceara: Aracati, VI.1952, F.S. Silva (1f#, MNRJ); Barbalha, V.1969, A. Alvarenga (4m#, EMUS); Ceara
1931, Dias Da Roche (3f#, USNM); São Benedito, Sao do Meu, 28.XI.1957, J. Souza (1f#, MNRJ); São Benedito,
São Barra, 27.II.1957, Finburo (1f#, MNRJ); Maranhão: Sao Luis, Foresta Sacavem, R. Cambra & D. Quintero (4f#,
MIUP); Vitoria do Mearim, 25–26 sep 1992, R. Cambra & D. Quintero (12f# 1m#, MIUP); Minas Gerais: 15 km sul de Jaíbe, 9.IV.1998, G.A.R. Melo (3f#, DZUP); 12 km a SE de Jaí 25.I.2008, G.A.R. Melo, (2f#, DZUP); Barro Alto,
XI.1931, J. Blaser (1f#, EMUS); Aguas Vermelhas, XII.1983, M. Alvarenga (2f#, MNRJ); Para, Óbidos, 1955 (1f#,
MNRJ); Paraiba: Juazeirinho: III.1956, A.G.A. Silva (2f#, MNRJ); V.1956, A.G.A. Silva (3f#, MNRJ); VI.1956,
A.G.A. Silva (45f#, DZUP, EMUS, MNRJ); IX.1956, C.R. Gonçalves, (2f#, MNRJ); Patos, UFCG-CSTR,
15.VI.2009, D.A.A. Lucena (1f#, DZUP); Santa Luzia, Brandão: 28.VI.1956, A.G.A. Silva (1f#, MNRJ); VII.1956,
A.G.A. Silva (2f#, MNRJ); Santa Luzia, VIII.1956, Cincinato (2f#, MNRJ); Itapora, IV.1999, Alexandre (1f#, UEFS);
Pernambuco, Base de S. Negra Floresta, V.1960, A.B.M. Machado (1f#, CASC); Rio Grande do Norte: Ceara-Mirim,
W.M. Mann (1f#, compared with type of Mutilla cerbera Klug by C.E. Mickel 1962, USNM); Iguatu (1f#, USNM);
Natal (1f#, USNM); Natal, X.1951, M. Alvarenga (1f#, MNRJ); Sergipe, Villa Nova: n.o proc. 52/354 (3f#, MNRJ); n.o proc. 52/356 (3f#, MNRJ); n.o proc. 52/349 (1f#, MNRJ); n.o proc. 52/352 (1f#, MNRJ); n.o proc. 52/342 (1f#,
MNRJ); Villa Nova, n.o proc. 52/355 (1f#, MNRJ); n.o proc. 52/351 (1f#, MNRJ).
Distribution. This species is found in the Caatinga ecoregion of Northeastern Brazil.
Remarks. This is the most common species of Leucospilomutilla in collections, and one of the commonest species in the Brazilian Caatinga ecoregion. The black and white contrasting color pattern is shared by numerous velvet ants in the Caatinga, including Atillum bucephalum (Perty, 1833); Hoplomutilla gigantea (Perty, 1833); and Traumatomutilla bifurca (Klug, 1821).
Leucospilomutilla chemea Casal, 1961
(Figs 15, 18) 47
Leucospilomutilla chemea Casal, 1961. p. 43, Holotype f#, Bolivia, Santa Cruz, Ichilo, 20km OSO [West by
Southwest of] Buenavista (AMNH).
Diagnosis. FEMALE. This species has the head setae mostly black, with only a medial whitish patch on the frons and vertex (Fig. 15). The T2 color pattern is also diagnostic: the sub-lateral whitish setal marks are widely separated by black setae and sub-apically the medial and lateral white setal marks are widely separated by black setae (Fig. 15).
Additionally, the dentiform projection of the epaulet is much smaller than the sharp projections of the pronotal spiracle and mesonotum, and the pygidium has longitudinal rugae. Body length 18–20 mm. MALE. Unknown.
Material examined (2 females). BOLIVIA: [Cochabamba], Chapare, Prosen (1f#, AMNH); Santa Cruz, Prov.
Florida, Pampagrande, 26.X.1994, A. Langer (1f#, MIUP).
Distribution. Known from only three localities in Bolivia, each near the confluence of the Bolivan Yungas and
Southwest Amazon moist forest ecoregions.
Remarks. This apparently rare species represents the westernmost records for Leucospilomutilla and seems to occur in wetter habitats than L. cerbera or L. staurogastra. No good candidate for the male has been found.
Leucospilomutilla staurogastra Suárez, 1973
(Figs 16, 19, 22, 28–32, 43–47)
Leucospilomutilla staurogastra Suárez, 1973. p. 141, Holotype f#, Brazil, Mato Grosso, Três Lagoas, Fazenda Canaã,
Rio Sucuriú (MZSP).
Diagnosis. FEMALE. This species has the head setae mostly whitish with interspersed erect black setae and a small patch of sub-appressed black setae on the frons (Fig. 16). The T2 color pattern is also diagnostic: the sub-lateral whitish setal marks are widely separated by black setae and sub-apically the medial and lateral white setal marks are connected by a transverse band of whitish setae (Fig. 16). Additionally, the dentiform projections of the epaulet and pronotal spiracle are much smaller than the sharp projection of the mesonotum, and the pygidium has sparse longitudinally oriented rugae (Fig. 22). Body length 17–22 mm. MALE. The male of this species has the head and mesosomal setae mostly black (Fig. 28) and the cuspis basically straight with a spatulate apex (Fig. 47). Additionally, 48 white setae entirely cover the apical portions of T1, T2, and T3 and a large patch on the T2 disc; the wings are transparent pale brown; and the paramere is strongly contorted subapically. Body length 18–20 mm.
Description. MALE (Hitherto undescribed). Body length 20 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs black. Wings dark fuscous throughout, veins blackish. Setae entirely black except with erect whitish setae on posterior surface of mesoscutellum, metanotum, posterolateral stripe on propodeum, dorsum of metatibia, lateral and medial stripes on T1 and base of T2, and medial and lateral spots of T2 apex and T3. Head.
Rounded, posteromedially expanded. Head width 0.7 × pronotal width. Eye almost circular. Ocelli small; OOD 6.4 ×
DLO, IOD 2.0 × DLO. Occipital carina distinct. Frons, vertex, and gena densely foveolate-punctate. Gena ecarinate.
Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before torulus. Clypeus shallow convex medially, concave laterally below antennal insertion; densely punctate; with rounded apical margin. Scape bicarinate. Flagellomere 1 1.9 × pedicel length; flagellomere 2 2.8 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely punctate with complete median longitudinal carina; notaulus and parapsis present. Mesoscutellum flat to concave, foveolate-punctate. Axilla produced posterolaterally as high truncate projection with conspicuous flat posterior face, dense foveolate-punctate dorsally, except narrow posterior margin impunctate. Area between mesoscutum and mesoscutellum deeply concave, smooth. Metanotum virtually equally wide throughout, its surface obscured by dense setae. Propodeum angular, densely areolate dorsally with conspicuous dorsolateral tubercle posterior to propodeal spiracle; lateral face mostly impunctate; dorsal, lateral, and posterior faces relatively flat, dorso-posterolateral angle sharp, sub-dentate. Mesoscutum with distinct flat subtriangular process or tubercle anterior to mesocoxa. Mesopleuron areolate-punctate with large sharp tubercle on dorsal half. Metapleuron virtually smooth, with micropunctures and erect microsetae. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, meso- and metatibia each with one row of inconspicuous spines dorsally; spurs densely microsetose. Metasoma. T1 width 0.5 ×
T2 width. T2 length 0.9 × T2 width. Dorsal metasomal sculpture predominantly foveolate-punctate. T7 finely rugose posteromedially with transverse arcuate smooth area subapically, lateral carinae obscure. S1 longitudinally elevated medially, not forming pronounced carina or teeth. S2 with interspersed small and large punctures. S3–6 with dense small punctures. S7 as long as broad, basally hollowed-out beneath S7, with obscure diagonal carinae from basolateral 49 to apicomedial margins, apical margin rounded. Genitalia. Parapenial lobe produced into thick incurved digitiform process. Paramere thick, slightly compressed laterally, downcurving to shallow upcurved apex, laterally converging in basal two thirds, diverging in apical third, ventro-laterally with long dense setae. Cuspis 0.7 × free paramere length, laterally compressed and contorted, basal half incurved, apical half parallel, setae denser ventrally, short and bristle- like in apical half. Paracuspis short, lobe-like, rounded apically, densely setose. Digitus 0.35 × free paramere length, incurved in dorsal view, slightly upcurved in lateral view. Penis valve scarcely extending beyond parapenial lobe, ventrally with large apical tooth and large subapical tooth with comb of short sharp teeth along basal slope of subapical tooth; apical distance between teeth 0.25 × length of valve; apical margin with row of minute setae.
Material examined (9 females and 3 males). BRAZIL: Minas Gerais: Cáceres, XI.1984, A. Alvarenga (1m#,
EMUS); Mato Grosso, Chapada dos Guimarães: 12–18.XI.2013, D.R. Luz, G.A.R. Melo, and K.A. Williams (4f#
1m#, DZUP); Areia Vermelha, margem direita, 23.III.2002, T. Mott (1f#, DZUP); Parana, Jaguariahyva, VII.1947,
Justus (1m#, DZUP). PARAGUAY: Caaguazu, Caaguazu, 1977, Fritz (2f#, AMNH); Cordillera, Naranjo,
24.XII.2001, R. Barrera (2f#, EMUS).
Distribution. This species is found in Brazilian Cerrado and surrounding ecoregions; the specimens from Paraguay were found in the Humid Chaco ecoregion.
Remarks. The male was easily recognized by its overall darker coloration than L. cerbera and overlapping distribution with females of L. staurogastra. During an expedition to Chapada dos Guimarães in 2013, four females and one male were collected by KAW and David R. Luz. One of these females stung DRL, who has also experienced the sting of various mutillid species. While the pain from a mutillid sting often lasts for less than 30 seconds, the sting of L. staurogastra remained intense for more than 10 minutes (DRL pers. obs.). Pain in velvet ant stings is often correlated with large body size (KAW pers. obs.) and L. staurogastra is one of the largest velvet ants in Brazil. The high duration of pain from this sting, however, may be indicative of different components in the venom rather than increased volume of venom. Studies of chemical composition of velvet ant venoms, especially if accompanied by
“comparative field trials”, would be most welcome in this group of insects.
Discussion
The recently recognized tribe Dasymutillini Brothers and Lelej, 2017 included 21 genera, eight from the Australian region and 13 from the New World (Brothers and Lelej 2017). Of the New World genera, nine seem to form a natural 50 group in being generally robust-bodied brightly-colored diurnal wasps that have males with armed axillae. These genera were informally referred to as the “dasymutilline genera” by Williams (2012) and seven of the nine genera were included in an unpublished molecular phylogeny by Williams (2012). Atlantilla gen. nov. is the 22nd genus recognized in the Dasymutillini and the 10th member of the “dasymutilline genera”. The genus shares structural similarities with Leucospilomutilla in both sexes, as mentioned in the remarks above and the sole included species, A. auriculata, was formerly included within Traumatomutilla. Sadly, Leucospilomutilla was one of only two genera not available for the phylogenetic reconstruction by Williams (2012) and molecular data are yet unavailable for Atlantilla gen. nov. We hypothesize, however, that Leucospilomutilla + Atlantilla could form a sister relationship with the species rich Traumatomutilla, and hope that the diagnoses and illustrations provided here will permit recognition of these taxa by researchers interested in future phylogenetic reconstructions, who could test the hypothesized genus limits proposed here.
Acknowledgements
We thank David R. Luz for collecting some of the specimens and for his comments about L. staurogastra. We are grateful for the subject editor and two anonymous reviewers for their help in editing and publishing this paper. We would also like to thank the collection managers and curators that provided for this study, including: Christine LeBeau
(AMNH), John Rawlins (CMNH), James Pitts and David Wahl (EMUS), Agniéle Touret-Alby (MNHN), Gabriel
Melo (DZUP), Orlando Tobias Silveira (MPEG), and Carlos Brandão (MZSP). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10
Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae
(Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9).
References
51
André, E. (1902) Fam. Mutillidae. In: Wytsman, P., Genera Insectorum, Fasc. 11. Bruxelles, 77 pp. + 3 pls.
Ashmead, W.H. (1903) Classification of the fossorial, predaceous and parasitic wasps, or the superfamily Vespoidea.
The Canadian Entomologist, 35, 303– 310.
Bartholomay, P.R., Williams, K.A., Luz, D.R., and Oliveira, M.L. (2018) New species of Traumatomutilla André in the T. tabapua and T. integella species-groups (Hymenoptera, Mutillidae). Zootaxa, 4433, 361–385.
Cambra, R.A., Williams, K.A., Quintero, D., Windsor, D.M., Pickering, J., and Saavedra, D. (2018) Dasymutilla
Ashmead (Hymenoptera, Mutillidae) in Panama: new species, sex associations and seasonal flight activity. Insecta
Mundi, 608, 1–17.
Casal, O.H. (1961) Mutillidae Neotropicales (III parte). Neotropica, 7 (23), 43–46.
Casal, O.H. (1963) Mutillidae Neotropicales. XXII (Hymenoptera). Comentarios a propósito de Leucospilomutilla
Ashmead con la descripción de los caracteres genéricos del macho. Physis, 24, 149–157.
Cresson, E.T. (1902) Descriptions of some Mutilla from Brazil. Transactions of the American Entomological Society,
28, 1–82.
Gerstaecker, A. (1874) Mutillarum Americae meridionalis indigenarum synopsis systematica et synonymica. Archiv für Naturgeschichte, 40, 41–77, 299–328.
Klug, J.C.F. (1821) Entomologiae brasilianae specimen. Nova Acta Academica Caesareae Leopoldino-Carolinae, naturae Curosiorum 10 (2), 305–324.
Nonveiller, G. (1990) Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta, Hymenoptera). Hymenopterorum Catalogus (Nova Editio), 18. Den Haag, SPB Academic
Publishing. 150 pp.
Perty, M. (1833) Delectus animalium articulatum quaie in itinere per Brasiliam, anais 1817 – 1820, digesit, descripsit et pingenda curavit Dr. M. Perty, 137–138.
Suárez, F.J. (1973) Datos sobre Mutilidos neotropicales VIII. Una nueva especie de Leucospilomutilla Ashmead
(Hymenoptera). Archivos del Instituto de Aclimatación, Almería, 18, 139–146, plate 1.
Williams, K.A., Brothers, D.J. & Pitts, J.P. (2011) New species of Tobantilla Casal, 1965 and a new genus and species, Gogoltilla chichikovi gen. et sp. nov., from Argentina (Hymenoptera: Mutillidae). Zootaxa, 3064, 41–68.
Williams, K.A., Bartholomay, P.R. & Oliveira, M.L. (2017) Species groups of Traumatomutilla André (Hymenoptera:
Mutillidae). Insecta Mundi, 0533, 1–33.
52
Figure legends.
Figures 1–2. Atlantilla auriculata (Gerstaecker, 1874), female. 1. Dorsal habitus. 2. Lateral habitus. Scale bar = 2 mm.
Figures 3–5. Atlantilla auriculata (Gerstaecker, 1874), female. 3. Face. 4. Pygidium. 5. Labels.
Figures 6–7. Atlantilla auriculata (Gerstaecker, 1874), male. 6. Lateral habitus. 7. Dorsal habitus. Scale bar = 2 mm.
Figures 8–13. Atlantilla auriculata (Gerstaecker, 1874), male. 8. Face. 9. Mesosomal dorsum. 10. Axilla, posterior view. 11. Axilla, lateral view. 12. T7. 13. S8.
Figures 14–16. Leucospilomutilla females, dorsal habitus. 14. Leucospilomutilla cerbera (Klug, 1821). 15. L. chemea
Casal, 1961. 16. L. staurogastra Suárez, 1973. Scale bar = 5 mm.
Figures 17–22. Leucospilomutilla females. 17, 20, 21. Leucospilomutilla cerbera (Klug, 1821). 18. L. chemea Casal,
1961. 19, 22. L. staurogastra Suárez, 1973. 17-19. Habitus, lateral view. 20, 22. Pygidium. 21. Head, lateral view.
Scale bar = 5 mm.
Figures 23–27. Leucospilomutilla cerbera (Klug, 1821), male. 23. Habitus, lateral view. 24. Mesosomal dorsum. 25.
Face. 26. S8. 27. T7. Scale bar = 5 mm.
Figures 28–32. Leucospilomutilla staurogastra Suárez, 1973, male. 28. Habitus, lateral view. 28. Mesosomal dorsum.
30. Face. 31. S8. 32. T7. Scale bar = 5 mm.
Figures 33–47. Male genitalia: dorsal view, ventral view, lateral view, penis valve lateral view, cuspis lateral view.
33–37. Atlantilla auriculata (Gerstaecker, 1874). 38–42. Leucospilomutilla cerbera (Klug, 1821). 43–47. L. staurogastra Suárez, 1973.
Figures 48. Atlantilla and Leucospilomutilla distributions.
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CAPÍTULO 4.7 - [formatado para submissão à revista ZOOTAXA]
Review of the Traumatomutilla juvenilis species group (Hymenoptera, Mutillidae)
PEDRO R. BARTHOLOMAY1’*, KEVIN A. WILLIAMS2 ROBERTO A. CAMBRA3 & MARCIO L. OLIVEIRA1 1Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, Laboratório de Hymenoptera, Av. André Araújo, 2936, Manaus, Amazonas, Brazil. 2Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento CA 95832, USA. 3Museo de Invertebrados G. B. Fairchild, Universidad de Panama, Panamá 0824, Panamá *Corresponding author, e-mail: [email protected]
Abstract
Traumatomutilla bivittata bivittata (Gerstaecker, 1874), T. duplicata feia Casal, 1969, T. bruchi André, 1908, T. cristata (Gerstaecker, 1874), T. aterrima (Gerstaecker, 1874) and T. lorena (Cresson, 1902) are proposed as junior synonyms of Traumatomutilla duplicata (Gerstaecker, 1874). Traumatomutilla immaculiceps André, 1901, T. bivittata rubroguttata André, 1901 and T. estrella (Cresson, 1902) are proposed as junior synonyms of T. juvenilis (Gerstaecker, 1874). Traumatomutilla bispiculata (André, 1907) is proposed as a junior synonym of T. miniata (Gerstaecker, 1874). The previously unknown males of T. guarata Casal, 1969 and T. juvenilis are described and illustrated. All species of the T. juvenilis species groups are redescribed and illustrated. A new species Traumatomutilla juvenindica Bartholomay & Williams sp. nov., is described based on males and females from northern South America. Additionally, identification keys for the species and known color forms of the T. juvenilis species-group are provided.
Key words: Spheropthalminae, Dasymutillini, Velvet ants, Neotropical, Taxonomy
Introduction
Mutillidae are aculeate wasps in which the females are wingless and the males are fully winged, brachypterous or rarely apterous (Brothers, 2006). Traumatomutilla André, 1901, is one of the most diverse genera of Mutillidae in the Neotropical region, with over 180 species ranging from Mexico to Argentina (Nonveiller, 1990). The extreme sexual dimorphism in Mutillidae caused most species in the genus to be described based only on females (140) or males (46) and sexual associations to be rare and difficult without in situ observations, molecular data, or large numbers of distribution data (Williams et al. 2011a, Luz et al. 2014, Luz et al. 2016). Also, most species of Traumatomutilla were described prior to the establishment of the genus and based almost exclusively on highly variable color and setae characters (Gerstaecker, 1874; Cresson, 1902; Casal, 1969). Recent works dealing with the Müllerian mimicry rings of closely related Dasymutilla Ashmead, 1899 have indicated that taxonomic revisions based on structural characters can reveal that several species and indeed entire species-groups might be in fact individual widespread and highly variable species (Williams et al. 2011, 2012; Wilson et al. 2012, 2013, 2018). The recent establishment of species-groups in Traumatomutilla, allowed the genus to be revised in smaller and more manageable units resulting in a series of recent taxonomic treatments that include this current revision of the T. juvenilis species-group. The T. juvenilis group is one of two species-groups in the genus with a species known from both sexes, T. bispiculata André, 1907, associated by André (1908b). Prioritizing the revision of such species-groups is important due to the rarity of sex associations for velvet ants in general and because knowing the characters of one male provides countless morphological clues to associate the remaining species within the group (Bartholomay et al. 2019).
Materials and terminology 60
For this study, approximately 1,050 specimens from 34 different collections were examined. The following codens are used for institutions housing the material discussed in the current study:
AMNH - American Museum of Natural History, New York, New York, USA ANSP - Academy of Natural Sciences, Philadelphia, Pennsylvania, USA BMNH - British Museum of Natural History, London, England CASC - Department of Entomology, California Academy of Sciences, San Francisco, California, USA CPDC - Comissão Executiva do Plano da Lavoura Cacaueira, Ilhéus, Bahia, Brazil CESC - Coleção Entomológica da Universidade de Santa Cruz do Sul, Santa Cruz do Sul, Rio Grande do Sul, Brazil CISC - Essig Museum of Entomology, Department of Entomological Sciences, University of California, Berkeley, California, USA CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA CSCA - California State Collection of Arthropods, Sacramento, California, USA CUIC - Cornell University Insect Collection, Department of Entomology, Ithaca, New York, USA DJBC - Collection of Denis J. Brothers, to be deposited in Izilo South Africa Museum (SAM), Cape Town, South Africa DEI - Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany DGMC - Donald G. Manley Collection, Florence, South Carolina, USA DZUP - Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA FMNH - Field Museum of Natural History, Chicago, Illinois, USA FSCA - Florida State Collection of Arthropods, Gainesville, Florida, USA LACM - Insect Collection, Los Angeles County Museum of Natural History, Los Angeles, California, USA MNCN - Museo Nacional de Ciencias Naturales, Madrid, Spain MNHN - Muséum Nationale d’Histoire Naturelle, Paris, France MNRJ - Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil MLUH - Martin Luther Universität Halle-Wittenberg, Halle, Germany MCZC - Museum of Comparative Zoology, Harvard Univeristy, Cambridge, Massachusetts, USA MuBio-UFGD - Museu da Biodiversidade, Universidade Federal da Grande Dourados, Dourados, Mato Grosso do Sul, Brazil MZSP - Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo, Brazil NHRM - Naturhistoriska Riksmuseet, Stockholm, Sweden PMNH - Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA RBINS - Royal Belgian Institute of Natural History, Brussels, Belgium UMSP - University of Minnesota Insect Collection, St. Paul, Minnesota, USA USNM - United States National Museum of Natural History, Simthsonian Institution, Washington D.C., USA UFMG - Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil UCDC - The Bohart Museum of Entomology, Universisty of California, Davis, California, USA ZMB - Museum für Naturkunde Berlin, Berlin, Germany ZMUC - Zoological Museum University of Copenhagen, Copenhagen, Denmark
General morphological terminology, definitions, abbreviations, and measurements followed that of Cambra et al. (2016) and Bartholomay et al. (2018, 2019). Additionally, sculpture terminology follows Harris (1979). 61
In the description and redescription sections we refrain from mentioning coloration and setal patterns due to the highly variable nature of these characters in Traumatomutilla. Instead, we provide a separate section titled Coloration and variations, in which we provide an overall description of the known color and setae patterns for a particular species. In the material examined section abbreviations, acronyms and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of the material examined section. The types of T. duplicata (Gerstaecker, 1874), T. aterrima (Gerstaecker, 1874), and T. miniata (Gerstaecker, 1874), although selected and labeled as lectotypes by the specialist Clarence E. Mickel, were never published as such by the author. In this paper we maintain and officially designate those specimens selected by C.E. Mickel as the lectotypes for their respective species. Likewise, many of the species described by Cresson (1902) have more than one specimen in their type series which is why Cresson (1916) provided a list of specimens with their respective catalog number in his collection, designating them as “types” and stating that those were the specimens used in the descriptions of all his previous studies. Though these specimense were not refered to nor labeled as lectotypes by the author, we consider them as the lectotypes of his species and Cresson (1916) as the paper that established them as such. Taxonomy
Traumatomutilla juvenilis species-group
Diagnosis. The females of this species group can be recognized by their mesosomal sculpture: the scutellar scale is broad and separated from the well-developed and medially emarginate anterior transverse carina. Other important features include: head unarmed posterolaterally, at most having the occipital carina transversely swollen dorsolaterally; femora rounded apically; and pygidial plate subpyriform ovate, with the lateral carinae converging basally, but weak and obscured by dense setae. The males of this species group can be recognized by their broad trapezoidal hypopygium and penis valve with closely spaced apicoventral teeth. Other important features include: axillar projection always truncate, mesopleuron with strongly projected tubercle on dorsal half, S2 with narrow longitudinal pit densely filled with setae. Included taxa. Traumatomutilla juvenilis (Gerstaecker), 1874; Traumatomutilla duplicata (Gerstaecker), 1874; Traumatomutilla miniata (Gerstaecker), 1874; T. guarata Casal, 1969 and T. juvenindica Bartholomay &Williams, sp. nov. Distribution. Colombia, Brazil, Bolivia, Paraguay, Argentina and Uruguay Remarks. Williams et al. (2017) mentioned that the anterolateral carinae on the scutellar area of this group is bilobate. Closer examination of specimens revealed that in fact these carinae are emarginated medially and the lateral areas are projected beyond the length of the setae that obscure most of the carinae thus appearing to be bilobate. Though the overall body shape and size of these species is similar in many aspects to the species of the T. indica species-groups, Williams (2012) provided evidence of a close relation between the T. juvenilis species-group and T. bifurca (Klug, 1821). The penis valve of the males of this species-group and T. bifurca are the only ones within the genus with closely spaced apicoventral teeth. Additionally, the genal carina of the females of the T. juvenilis species-group is greatly reduced whilst the genal carina of T. bifurca is ecarinate.
Key to females of the recognized species of the T. juvenilis species-group.
1. Metasoma strongly constricted anterior to propodeal spiracles; lateral margins of mesonotum strongly divergent anterad; propodeal spiracles projected from lateral margins of mesosoma. (Figs. 1–6) … 2 Metasoma not so constricted anterior to propodeal spiracles; lateral margins of mesosoma slightly divergent anterad; propodeal spiracles virtually flat against lateral margins of mesosoma. (Figs. 29, 31, 33, 35) … 3
2. Lateral propodeal face sculpture sparse foveolate-punctate anteriorly, coarsely and densely foveolate-punctate along posterior margin with coarse vestigial longitudinally oblique rugosities; silvery-white setae of head absent or 62 present as longitudinal stripes from posterior margin of vertex to antennal tubercles. (Figs. 1–6, 11–16) - predominantly found in Cerrado, Pampa and Chaco areas of South America - … T. duplicata (Gerstaecker) Lateral propodeal face sculpture dense and coarse foveolate-punctate throughout, vestigial rugosities absent; head with front completely covered with silvery-golden setae, vertex black. (Figs. 70–71) - predominantly found in Atlantic Forest areas in Brazil - … T. guarata Casal
3. Metatibia sparsely setose dorsally, surface mostly smooth and shinning; head with transverse stripe of silvery- white setae on vertex. (Figs. 49–50) - predominantly found in xeric areas and dry forests northwest of the Andes in Venezuela and Colombia - … T. juvenindica Williams & Bartholomay sp. nov. Metatibia densely setose dorsally, surface mostly concealed by dense setae, never shinning where visible; head setae completely black or with longitudinal stripes of silvery-white setae laterally. … 4
4. Pronotal spiracle strongly projected from lateral margins of pronotum, subacute, frequently forming a vestigial crest-fold dorsolaterally; scabrous intervals present on posterior third of mesonotum; T2 disc conspicuously angulate laterally, thus forming distinct dorsal and lateral faces of T2; head setae completely black. (Figs. 57–58) - predominantly found in Chaco and Pampa areas of Argentina - … T. miniata (Gerstaecker) Pronotal spiracle usually not strongly projected, if strongly projetcted then always rounded and never with vestigial crest-fold dorsolaterally; scabrous intervals absent on mesonotum; T2 disc smoothly and evenly convex; head setae variable, completely black or with longitudinal silvery-white stripes. (Figs. 29–36) - predominantly found in Cerrado and Chaco areas of South America - … T. juvenilis (Gerstaecker)
Key to males of the recognized species of the T. juvenilis species-group.
1. Cuspis slender, narrow, slightly broadened apically, virtually asetose except for sparse conspicuous setae ventrally at apex (Figs. 24–27, 52–55, 74–77) … 2 Cuspis broad, wide, slightly tapered basally and apically, densely setose throughout (Figs. 44–47, 64–67) … 4
2. Pygidial plate restricted to apical third of T7, defined by basal carinae as well as lateral carinae; basal carinae converging medially and extending as a single longitudinal medial carina towards base of T7; head and pronotum completely clothed with silvery-white setae. (Figs. 51–56) - predominantly found in xeric areas and dry forests northwest of the Andes in Venezuela and Colombia - … T. juvenindica Williams & Bartholomay sp. nov. Pygidial plate extending through at least apical half of T7, defined only by lateral carinae; head and pronotum completely clothed with black setae, at most with sparse silvery-white setae in certain areas (Figs. 17–20, 40–41, 60, 62, 73). … 3
3. Paracuspis stout; penis valve wider basally than apically (excluding anteroventral teeth); setae of lateral face of propodeum virtually all black. (Figs. 40–48) - predominantly found in Cerrado, Pampa and Chaco areas of South America - … T. duplicata (Gerstaecker) Paracuspis elongate; penis valve as wide basally as dorsally (excluding posteroventral teeth); setae of lateral face of propodeum predominantly silvery-white. (Figs. 73–78) - predominantly found in Atlantic Forest areas in Brazil - … T. guarata Casal
4. Lateral margins of propodeum sharply angulate, posterolateral corners of propodeum strongly projected in lateral view; T2 disc conspicuously angulate laterally, thus forming distinct dorsal and lateral faces of T2. (Figs. 60–68) - predominantly found in Chaco and Pampa areas of Argentina - … T. miniata (Gerstaecker) 63
Lateral margins of propodeum rounded, posterolateral corners of propodeum rounded in lateral view; T2 disc smoothly and evenly convex throughout, at most slightly angle laterally. (Figs. 17–28) - predominantly found in Cerrado and Chaco areas of South America - … T. juvenilis (Gerstaecker)
Key to the known color forms of females in the T. juvenilis species-group:
1. Head clothed with black setae only … 2 Head with well-defined areas of dense silvery-white setae … 6
2. Mesosoma completely black, without any silvery-white setae dorsally, on lateral face of pronotum, pleurae and lateral face of propodeum (Figs. 6, 16) … T. duplicata in part [specimens from high altitude areas in Bolivia, 1200-2500m a.s.l.] Mesosoma with conspicuous patterns of silvery-white setae at least on propodeal dorsum … 3
3. Mesosoma laterally constricted anterior to propodeal spiracles, conspicuously diverging anterad, sinuous; silvery- white setae stripes restricted to propodeum or mesonotum; integumental spots of T2 reddish, semi-arid Argentina, Bolivia, and Paraguay … 4 Mesosoma not at all constricted anterior to propodeal spiracles, slightly divergent anterad, virtually straight; silvery-white stripes reaching pronotal spiracles; integumental spots of T2 reddish to yellowish; more widely distributed, reaching semi-arid Brazil … 5
4. Silvery-white setae stripes of mesosoma ending on mesonotum; integumental spots of T2 broad, reddish to orange (Figs. 5, 15) … T. duplicata [formerly known as T. bispiculata (females sensu André, (1908b)) Silvery-white setae stripes of mesosoma restricted to propodeum; integumental spots of T2 broad, reddish, anterior pair broadly connected to posterior pair (Figs. 3, 13) … T. duplicata [formerly known as T. bruchi]
5. Integumental spots of T2 yellowish, posterior pair transversely linear to subrectangular (Figs. 29–30) … T. juvenilis in part Integumental spots of T2 reddish, posterior pair rounded (Figs. 31–32) … T. rubroguttata [formerly known as T. immaculiceps]
6. Silvery-white setae of head forming rounded spot on front or transverse band on vertex … 7 Silvery-white setae of head forming lateral longitudinal stripes along inner eye margin … 8
7. Silvery-white setae of head restricted to front; silvery-white stripes of mesosoma restricted to propodeum; integumental spots of T2 reddish to orange, posterior spots subrectangular; distribution restricted to the Atlantic Forest of Brazil (Figs. 70–71) … T. guarata Silvery-white setae of head restricted to vertex; silvery-white stripes of mesosoma reaching prontoal spiracle; integumental spots of T2 yellowish, posterior spots linear, Venezuela and Colombia (Figs. 49–50) … T. juvenindica sp. n.
8. Silvery-white stripes of mesosoma divided, appearing to from four spots (Figs. 1, 11) … T. duplicata in part [formerly known as T. duplicata duplicata] Silvery-white stripes of mesosoma continuous, uninterrupted or at least narrowly connected … 9
9. Silvery-white stripes of mesosoma almost interrupted, integumental spots of T2 reddish, small, posterior pair separated by at least the diameter of one spot (Figs. 4, 14) … T. duplicata [formerly known as T. duplicata feia] 64
Silvery-white stripes of mesosoma conspicuously connected, integumental spots of T2 reddish or yellowish, posterior pair usually larger, separated by less than diameter of one spot … 10
10. Integumental spots of T2 reddish (Figs. 33–34) … T. juvenilis in part [formerly known as T. bivittata rubroguttata] Integumental spots of T2 yellowish … 11
11. Mesosoma laterally constricted anterior to propodeal spiracles, conspicuously diverging anterad, sinuous; posterior pair of integumental spots of T2 transversely subrectangular (Figs. 2, 12) … T. duplicata in part [formerly known as T. bivittata bivittata] Mesosoma not at all constricted anterior to propodeal spiracles, slightly divergent anterad, virtually straight; posterior pair of integumental spots of T2 rounded (Figs. 35–36) … T. juvenilis in part [formerly known as T. estrella]
Traumatomutilla bivittata (Gerstaecker, 1874) (Figs. 1–28)
Mutilla bivittata Gerstaecker, 1874: 72, holotype [by monotypy], f#, Brazil, [Minas Gerais], Lagoa Santa (ZMB) examined. Mutilla duplicata Gerstaecker, 1874: 72, lectotype [designated here], f#, Brazil, [Rio Grande do Sul], Alegrete (ZMB) examined, syn. nov. Mutilla cristata Gerstaecker, 1874: 317, holotype [by monotypy], m#, Brazil, [Rio Grande do Sul], Santa Cruz [do Sul] (ZMB) examined, syn. nov. Mutilla aterrima Gerstaecker, 1874: 318, lectotype [designated here], m#, Brazil, [São Paulo], Salto Grande (ZMB) examined, syn. nov. Mutilla duplicata: Burmeister, 1875, p. 469 (sex association) Mutilla (Traumatomutilla) bivittata bivittata: André, 1901, p. 257. Mutilla lorena Cresson, 1902: 72, holotype [by monotypy], m#, Brazil, [Mato Grosso do Sul], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) aterrima: André, 1902, 54. Ephuta (Traumatomutilla) bivittata: André, 1902, 54. Ephuta (Traumatomutilla) cristata: André, 1902, 55. Ephuta (Traumatomutilla) duplicata: André, 1902, 55. Ephuta (Traumatomutilla) lorena: André, 1902, 56. Ephuta (Traumatomutilla) miniata: André, 1902, 55. Traumatomutilla aterrima: André, 1904, 40. Traumatomutilla bivittata: André, 1904, 40. Traumatomutilla cristata: André, 1904, 40. Traumatomutilla duplicata: André, 1904, 40. Traumatomutilla lorena: André, 1904, 40. Traumatomutilla miniata: André, 1904, 40. Traumatomutilla bruchi André, 1908a: 230, lectotype [designated here], f#, Argentina, Catamarca (MNHN) examined, syn. nov. Traumatomutilla bispiculata: André, 1908b, 201 (female descrpition) Traumatomutilla duplicata: André, 1908b, 206 (sex disassociation) Traumatomutilla duplicata feia Casal, 1969: 291, holotype f#, Brazil, São Paulo, Barueri (AMNH) examined, syn. nov.
65
Diagnosis. FEMALE. Mesosoma strongly constricted anterior to propodeal spiracles, lateral face of propodeum with vestigial longitudinally oblique rugosities posteriorly; setae of lateral face of propodeum virtually entirely black. MALE. Cuspis slender, predominantly asetose; setae on lateral face of propodeum and mesopleuron mostly or completely black. Description. FEMALE. Body length 9–19 mm. Head. Posterior margin virtually straight. Occipital carina conspicuously swollen and smoothyl curved dorsolaterally. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate- punctate to areolate-punctate, slightly sparser on gena and malar space. Mandible with small subapical tooth. Dorsal scrobal carina present defined, separated from antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercle finely and irregularly rugose. Flagellomere 1 2.25 × pedicel length; flagellomere 2 1.7 × pedicel length. Mesosoma. Dorsal thoracic length slightly shorter than width. Mesosomal dorsum well differentiated from pleurae, lateral margins rounded, not sharp or angulate; sculpture predominantly obscured by dense setation, dense and coarse areolate-punctate to foveolate-punctate where visible, denser and smaller mediad. Anterior face of propodeum defined, coarsely striated longitudinally ventrad, confused punctured dorsad; dorsal face roundly angulate into anterior face in lateral view. Humeral carina present broadly separated from well-defined low rounded epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum, subacute. Sculpture of lateral face of pronotum sparsely punctate with dense micropunctures; mesopleuron densely micropunctate anteriorly, dense coarse and confused areolate-punctate to foveolate-punctate along mesopleural ridge, concealed by dense setation elsewhere; metapleuron sculpture predominantly concealed by dense setation, except dorsal half unsculptured and asetose, smooth. Lateral face of propodeum sparsely foveolate-punctate anteriorly, densely coarsely and confusedly areolate-punctate to foveolate punctae posteriorly. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 69:99:97:70:71. Lateral margin of mesonotum strongly emarginated anterior to propodeal spiracle, strongly diverging anterad. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular absent. Scutellar scale present, well-developed, slightly narrower than and separated from well developed anterolateral carinae which are connected thus forming a single transverse carina with shallow emargination medially; scabrous intervals present on scutellar area. Posterior face of propodeum longer than and poorly distinguished from dorsal face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 39:89:86. Disc of T2 densely and coarsely foveolate-punctate to punctate, denser and smaller mediad; punctures sparser and shallower over integumental spots. T3–6, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate with interspersed micropunctures where visible. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, equally high throughout. S2 sparsely foveolate-punctate, punctures conspicuously sparser and smaller posteromedially, denser and smaller anteromedially and with a conspicuous anteromedial longitudinal unsculptured area; anteromedial crest-fold present, well-defined. S3–6 densely and coarsely foveolate-punctate. Pygidial plate broadly subpyriform, defined by lateral carinae at apical half of plate; surface mostly irregularly longitudinally rugose, rugae vestigial laterally. MALE. Body length 11–18 mm. Head. Rounded subrectangular in dorsal view, posterolateral angles rounded. Vertex width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 5.6 × DLO, IOD 1.4 × DLO. Occipital carina distinct. Head surface densely and finely foveolate- punctate to punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina narrowly separated from internal eye margin. Clypeus slightly concave laterally immediately below antennal insertion, virtually flat elsewhere; coarsely and densely foveolate-punctate to punctate; with a pair of short, sessile, blunt tubercles on apical margin. Scape bicarinate. Flagellomere 1 2 × pedicel length; flagellomere 2.6 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, rounded, broadly separated from humeral carina, anterolateral angles of pronotum rounded. Anterior face of propodeum vestigially and longitudinally striate mesoventrad, coarsely punctate laterodorsad, with a conspicuous longitudinal slightly concave medial area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures anteriorly and along inner margin. Mesoscutum densely and coarsely foveolate-punctate to punctate, notaulus and parapsis indistinguishable. Scutellum convex, densely and coarsely areolate-punctate; with longitudinal irregular carina medially formed by aligned intervals. 66
Axilla produced posterolaterally as truncate projection, with conspicuous flat posterior face, coarsely and densely foveolate-punctate dorsally; posterior face of axillar projection arched. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeum dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face predominantly densely areolate, areolations less defined anterad; posterolateral margins rounded, posterodorsal corners rounded in lateral view; dorsal and posterior faces indistinguishable. Lateral face of pronotum densely coarsely and confusedly foveolate-punctate to punctate with interspersed micropunctures; mesopleura with strong subacute tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures anteriorly. Metapleuron densely and coarsely areolate punctate on ventral fourth, coarsely and confusedly rugose to punctate on dorsal fourth, micropunctate to unsculptured elsewhere. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate where visible, except pygidial plate on T7 irregularly rugose and weakly defined by parallel carinae apicolaterally, rugosities predominantly transverse. S1 longitudinally elevated medially, forming pronounced carina slightly higher posteriorly. S2 foveolate-punctate, foveolations conspicuously sparser and smaller mediad; with well- developed longitudinal anteromedial crest-fold ending on elongate and narrow longitudinal pit densely filled with setae. S3–7 sparsely foveolate-punctate to punctate. S7 slightly broader than long, with a pair of subacute closely spaced tooth- like projections on posterior margin. Genitalia. Parapenial lobe slightly pronounced apically. Ratios of free length of paramere, cuspis and digitus, 70:63:8; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense long setae ventrally except at apically with setae conspicuously reduced; cuspis narrow, slender, in dorsal view slightly curved inward at basal half and curved outward in apical half; conspicuously and smoothly widened apically; predominantly asetose with sparse conspicuous setae ventrally at apex; paracuspis well-developed, not sessile, slightly elongate vertically, roundly subtriangular at apical margin, densely setose, setae predominantly shorter than paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose dorsally; penis valve strongly concave on internal surface, with very closely spaced pair of teeth apicoventrally; apical tooth more acute and longer; subapical tooth acute, externolateral vestigial; apical distance between teeth 0.03 × length of valve; dense setae present along apical margin and at base of teeth on external surface. Coloration and variations. FEMALE. Head integument always black, at most brownish-black, setation can be completely black or with a pair of longitudinal silvery-white stripes along inner eye margin starting at posterior margin of vertex and ending at antennal tubercles; stripes sometimes reduced to a small longitudinal spot dorsally on inner eye margin. Mesosoma integument is always completely black, at most brownish-black; setation can be completely black or with longitudinal silvery-white stripes along lateral margins that can be restricted to propodeal dorsum or extend as far as into anterior margin of pronotum; stripes that extend beyond propodeum can be complete or interrupted near scutelar area. Integument of metasoma is always predominantly black, at most brownish-black, except for disc of T2 which is always marked with integumental spots that can be reddish, orange, or yellowish; integumental spots vary greatly in size and can be subquadrate to subrectangular, the anterior pair can be fused with the posterior pair thus forming a pair of large longitudinal integumental spots. Metasomal setae predominantly black, except for silvery-white setae varying in density on the following: T1 laterally, T2 lateral margins and T2 lateral felt lines, T2 fringe laterally, T3–5 medially and laterally, T6 medially, S1–5 (except fringe of S5), and over integumental spots of T2. Appendages color predominantly black, except mandibles and flagellomeres partially reddish-brown. MALE. Integument always completely black, at most brownish-black, with mandibles partially reddish-brown. Body setae is predominantly black with silvery-white setae varying in density and extension in the following: metanotum, dorsum of propodeum, T1, base of T2, lateral margins of T2, lateral felt line of T2, fringes of T2–4 laterally, and S1–4; tibiae and tarsomeri. Distribution. Brazil, Bolivia, Paraguay, Argentina and Uruguay Material examined. (392 f#, 146m#) Type material. Lectotype: Traumatomutilla duplicata, f#, BRAZIL, [Rio Grande do Sul], Allegrette [Alegrete], Sello S. (ZMB); Holotype: Traumatomutilla bivittata, f#, BRAZIL, [Goiás], Lagoa Santa, Burm. S. (ZMB); Holotype: Traumatomutilla cristata, m#, BRAZIL, [Rio Grande do Sul], Santa Cruz [do Sul], Hensel S. 67
(ZMB); Lectotype: Traumatomutilla aterrima, m#, BRAZIL, [São Paulo], Salto Grande, Sello S. (ZMB); Holotype: Traumatomutilla lorena, m#, BRAZIL, [Mato Grosso do Sul], Chapada [dos Guimarães] (CMNH); Lectotype: Traumatomutilla bruchi, f#, ARGENTINA, Catamarca (MNHN); Holotype: Traumatomutilla duplicata feia, f#, São Paulo, Barueri, III.1958, K. Lenko (AMNH). Additional material. BRAZIL: Mato Grosso: 1f#, XII.1947, De Castelnau (MNHN); Utiariti, 5f# (MZSP); Tapirape Indian Village, at confluence of Rio Tapirape and Rio Araguaia, 1f#, 01– 11.XII.1960, B. Malkin (FMNH); Parque Nacional Chapada dos Guimarães, 3f#, 12.XI.2013, Melo G.A.R., Luz D.R., Williams K.A. (DZUP); Rio Caraguatá, 1m#, III.1953, F. Plaumann (DJBC); Goiás: Jataí: 15f# (MZSP); Faz. [Fazenda] Cachoeirinha, 1f#, X.1962, Exp. [Expedição] Dep. [Departamento] Zool. [Zoologia] (MNCN); Distrito Federal, Brasília, 1f# (ZMUC); Minas Gerais: Lavras, 1f#, XII.2002, A. Foucart (EMUS); P. [Parque] N. [Nacional] da Serra do Cipó, 1f#, 09–13.XII.2011, Cavichioli R.R. (DZUP); Ouro Preto, Cachoeira do Abacaxi, 1f#, 07.XI.2001, Paprocki, Blahnik & Amarante (UMSP); Uberlândia, 1f#, X.1962, EXp. [Expedição] Dep. [Departamento] Zool. [Zoologia] (MNCN); Lassance, 1f#, 09–19.XI.1919, Cornell Univ. [University] Expedition (CUIC); Uberaba, 1f# (RBINS); Sete Lagoas, 1f#, XII.1962, A. Martinez (AMNH); nr. Timoteo, 1m#, 13–20.X.1997, Eurico R De Paula (EMUS); Mato Grosso do Sul: [Porto] Murtinho, 1f#, XI.1929, R. Spitz (MZSP); Campo Grande, 1f#, X.1953, Bastos Filho (DZUP); Três Lagôas, 5f#, (MZSP); São Paulo: Barueri, 3f# (MNRJ); 1f#, III.1958, K. Lenko (AMNH); Jundiaí, 1f#, 13.XII.1960 (MNRJ); Pompéia, urban área, 1f#, 09.IV.1993, Silva R. (EMUS); Pereira Barreto, 1f#, 09.VI.1993, Lima M.A. (EMUS);São José dos Campos, 1f#, 01–04.XI.1960, D.L. Tiemann (EMUS); Ribeirão Preto, 1f#, 21.X.1962, Exp. [Expedição] Dep. [Departamento] Zool. [Zoologia] (MNCN); Campinas, 1f#, XI.1962, Himri Gridi-Papp (MZSP); Ibitinga, Faz.[enda] Itaquerê, 2f#, 25.I.1964, K. Lenko (MZSP); Nova Europa, Faz.[enda] Itaquerê, 1f#, XI.1963, K. Lenko (MZSP); Franca, 1f#, I.1911, E. Gazbe (MZSP); Rio Claro, Horto Florestal Navarro de Andrade, 1m#, 14.xi.1988, R.P. Martins (UFMG); Rio de Janeiro, [Rio de Janeiro]: Corcovado, 1f#, 3.IV.1955, Gruman (DZUP); 1m#, II.1912, E.P. Reed (CASC); Paraná: Ponta Grossa, 1m#, II.1948, Justus (DZUP); 1f#, II.1945, Justus (DZUP); Vila Velha, 1m#, 09.XI.1974, Moure J.S. (DZUP); 1m#, 03.II.1974, Rozen, Thomson & Moure (AMNH); Santa Catarina, [Florianópolis], Morro das Pedras, 1f#, Pe.[Padre] Buck (CESC); 9 km E de Aranaguá, área de dunas, 1m#, 20.XI.2007, Melo, G.A.R., Parizotto, D. (DZUP); Nova Teutonia, 1m#, XII.1969, F. Plaumann (DJBC); Rio Grande do Sul: Alegrete, 1f#, 19.I.2009, P. Bartholomay (CESC); Arroio do Sal: 2f#, 29.I.2008, J.T. Klein (CESC); 1f# 1m#, 29.I.2008, P. Bartholomay (CESC); Pelotas: 1f#, XII.1954 (DEI); 1m#, 23.XII.1962, C.M. Biezanko (PMNH); 1f#, 20.III.1962 (DEI); 1f#, 02.III.1956 (DEI); Porto Alegre: 1f#, 27.XII.1928, Pe. [Padre] (CESC); 1f#, I.1942 (CESC); 1f#, I.1968, Pio [sic!] Buck (MNCN); São Leopoldo: 3f#, J.W. Stahl (NHRM); 2f#, J.W. Stahl (MNCN); Xangri-lá: 2f#, 28.I.2008, J.T. Klein (CESC); 1f#, 28.I.2008, P. Bartholomay (CESC); São João do M. [Monte] N. [Negro], 1f# (CMNH); BOLIVIA: La Paz, La Paz, 1f#, 20.XII.1908, Jorgensen (ZMUC); Santa Cruz: Cordillera, Rio Seco, 1f#, I.1959, A. Martinez (AMNH); Buena Vista, 17°27.68’S 63°39.63’W, 2m#, 23.II.1999, F.D. Parker (EMUS); Potrerillos de Guenda, Pres. Nat. 40km w. Santa Cruz de la Sierra, 370m, 17°40’S 63°27’W, 1m#, 04.X.2007, Wappes & Morris (UCDC); ARGENTINA: Cordoba: Cordoba: 3f#, L. Bahamondes (USNM); 4m#, 1945, L Bahamondes (CISC); Dep.[Departamiento] de Calamuchita: 1m#, I.1974, Martinez (AMNH); El Sauce 3f#, XII.1938, M.J. Viana (USNM); Miramar, 1f#, III.1992 Fritz (EMUS); Dolores, ca.[circa] La Cumbre, 1f#, 14.III-07.IV.1996, C. Porter (FSCA); Balnearia, 2f#, II.1971, Fritz (AMNH); Tanti, 4f#, Fritz (AMNH); Quilpo, 2f#, II.1980, Fritz (AMNH); Punilla, Valle Hermoso, 1f#, I.1943, M.J. Viana (MNCN); Alta Garcia, 2m#, 28.II.1922, C. Bruch (UMSP); Cosquin, Sierra de Cordoba, 1m#, 1.III.1920, Cornell Expedition (CUIC); Corrientes, Ytuzaingo, 1f#, X.1979, Fritz (EMUS); Entre Rios: Lazo, 1f#, XII.1940 (LACM); 1f#, XII.1940 (EMUS); [Pueblo] Liebig: 1f#, III.1999 (FSCA); 1m#, II.1997, Zelich (AMNH); Dpto.[Departamientio] Colon, Parque Nacional, 2f# (AMNH); 1f#, 1974, Fritz (AMNH); Mercedes, 2f#, XII.1974 (AMNH); Palmar Colon: 1m#, Fritz (AMNH); 2m#, XII.1974, Fritz (EMUS); Santa Fé: Las Garzas: Rio Las Garzas, 25km [kilometers] O. [West] d’Ocampo [of Ocampo], Chaco, 2f#, 1903, E.R. Wagner (MNHN); Bord du R.[io] Las Garzas, 25km O d’Ocampo [of Ocampo], Chaco, 2m#, 1903, E.R. Wagner (MHNH); Huerta Gr.[ande], 9m#, I.1910 (UMSP); Estancia La Noria, Rio San Javier, 7m#, 22.XII.1911, G.E. Bryant (BMNH); Catamarca: Coneta, 16km [kilometers] S [South] Catamarca, 554m [meters], 28°34.07'S 65°52.74'W, 8f#, 25.X–12.XI.2003, M.E. Irwin F.D. Parker (EMUS); Palo Labrado, 23km [kilometers] S [South] La Merced, 734m [meters], 28°19.94'S 65°37.13'W, 1f#, 24.X–12.XI.2003, M.E. Irwin F.D. Parker (EMUS); 68
Trampasacha, 8km [kilometers] W [west], Chumbicha, 3f#, 25.XI–12.XII.2003, M.E. Irwin F.D. Parker (EMUS); Punta [de] Balasto, 1f#, I.1994, Fritz (EMUS); 1f#, 03.I.1991, Fritz (FSCA); Santa Maria: 1f#, 19.I.1991, Scaramozzino (EMUS); 1m#, 4.I.1991, Fritz (AMNH); 1f#, I.1993, M.A. Fritz (AMNH); Villa Vil, 1f#, 24.I.1972, JCS (CASC); Londres, 1f#, 06.XII.2002, L.A. Stange (FSCA); Capillitas, 1f#, II.1986, Fritz (AMNH); Punto Esperanza , 33m#, I.1992, Fritz (AMNH); Tucumán: Aimacha del Valle, 1f#, 28.XII.1965, H & M Townes (EMUS); 4km S Capitan Cáceres, 430m, 27°13.54'S 65°38.34'W, 1f# 1m#, 24.X-12.XI.2003, M.E. Irwin F.D. Parker (EMUS); San Pedro de Colalao, 1f#, Foerster (AMNH); Trancas, Las Tacanas, 1f#, II.1951, J.M. Arnau (MNCN); Rt.[Ruta] 40, Ruinas Indigenas de Quilmes, 1f#, 28.III.1992, DeVries, di Lorio, Quinter, Yeates (AMNH); La Rioja: Iliar, 1f#, II.1934, M. Gomez (MNCN); Mendoza: Estancia Pedregal, 5f#, 05.I.1907, Jorgensen (ZMUC); Salta: Cafayate, 1f#, 19.II.1965, Hayward (FSCA); 10–20 km N Amblayo, 1f#, 22.III.1990, J.G. Rozen & A. Roig (AMNH); La Vina, 2f#, II.1993, M.A. Fritz (AMNH); Alemania, 1f#, I.1983, Fritz (EMUS); Sumalao, 1f#, III.1995, Fritz (EMUS); Rosario de Lerma, 1f#, I.1983, Fritz (EMUS); Los Medanos, 8km NE Cafayate, 1f#, 02–09.II.1972, H.E. Evans (EMUS); Cachi, 1f#, 20–22.I.1966, C.C. Porter, (DGMC); San Carlos, 1f#, 23.I.1950, F. Monrós & A. Willink (MNCN); El Maray, 1m#, II.1996, Fritz (AMNH); San Luís: Arizona, 2f#, II.1980, Fritz (AMNH); Alto Pencoso, 1m#, 20.XII.1908, Jorgensen (ZMUC); Santiago del Estero: Bord du R. Salado River [Banks of the Salador River], environ d’Icano [outskirsts of Icaño], 2f#, 1910, E.R. Wagner (MNHN); Missiones, Missiones, 1m#, 23.I.1986, L.E. Pena (AMNH); URUGUAY: Colônia, Colônia, 2f#, 06.III.1944, P.A. Berry (USNM); Rocha, Santa Teresa, 1f#, III.1965, Achaval (EMUS); Tacuarembó, Tacuarembó, 1f#, II.1954, Carbonell (AMNH); San Jose, San Jose, 1f#, Zorron (AMNH); Paysandu, Paysandu, 1m#, 28.XII.1862–14.II.1863, F.Amor - Expdeción al Pacífico (1862-1865) (MNCN); Rio Negro, Arroyo Negro, 15km S Paysandu, 4m#, 27.XII.1962, R.G. VanGelder (AMNH); An additional 40f# from Brazil, 19f# from Paraguay and 167f#/57m# from Argentina were also examined (ANSP, AMNH, MNHN, EMUS, EMUS, CMNH, MNCN, MNRJ, CUIC, ZMUC, DEI, DZUP, USNM, LACM, MCZC, MZSP, FSCA, BMNH, CSCA). Remarks. The first attempted sex association for this widespread and variable species was by Burmeister (1875), who described and associated a male from Argentina with T. duplicata females. Later, André (1908b) determined those males to be T. cristata and, due to the lack of further data, “disassociated” them from T. duplicata. In the same paper, André (1908b) associated and described the female of T. bispiculata based on their similarities in size and distribution. When PRB examined André’s series of T. bispiculata in MNHN, he found that the males represented a mixed series of two structurally distinct species with identical color patterns. The holotype of T. bispiculata, described by Andre in 1907, is actually associated with females of T. miniata (Gerstaecker, 1874) and is discussed further below. The other males in the series are structurally identical to T. cristata but have a color pattern typical of Traumatomutilla species in arid regions of Argentina (Fig. 20). Likewise, the females associated with T. bispiculata by Andre are structurally identical to T. duplicata (Fig. 1) from more humid Atlantic areas in Argentina, Brazil, and Uruguay, but have coloration typical to arid Argentina (Fig. 5). The males and females associated by André (1908b) as T. bispiculata are, therefore, correctly associated, but they were not correctly named. Both sexes of T. bispiculata sensu Andre 1908 are actually synonymous color variants of T. duplicata. The types of T. atterima, T. cristata, and T. lorena, each known from males only, were examined and were found to be structurally identical to T. bispiculata sensu Andre 1908, differing only in the distribution and density of whitish setae. Traumatomutilla atterima presents one extreme, having the body setae entirely black (Fig. 18), while T. bispiculata sensu Andre 1908 with its dense white setae on the propodeum, base of T2, and lateral margins of T2-4 presents the other extreme (Fig. 20). The other species (Figs. 17, 19), and numerous unplaceable males from across the range of T. duplicata, fall along a continuum between these forms. Likewise, the types of T. bivittata (Fig. 2, 12) and T. bruchi (Fig. 3, 13), described from females, are identical to T. duplicata and differ only in color features across a geographic continuum. With these synonyms, T. duplicata is the most widespread, abundant, and variable species in the T. juvenilis species-group. André (1908a) described Traumatomutilla bruchi based on “multiple specimens from the La Plata museum”. The specimen at MNHN was labelled as lectotype by the staff upon dispatching the loan to PRB and bear a handwritten label reading “Museo La Plata”. We assume that this was one of the specimens in the series used by André (1910) for 69 describing T. bruchi, likely donated by the museum to the author, and hereby officially designate this specimen as the lectotype for this species. A remarkable series of four female specimens examined revealed an extreme color variant for this species-group, wherein the integument and setae of the head and mesosoma are completely black; these specimens are from high altitude xeric areas on the eastern slope of the Andes in Bolivia (1200–2500m a.s.l.). Although these females are very different from the main pattern for the T. juvenilis group and their distribution appears to be isolated, we refrain from describing them as a distinct species based on their structural similarity with other specimens of T. duplicata. Additionally, males formerly known as T. cristata and T. atterima have been collected in the same areas as these females. Females of T. duplicata vary greatly in length and consequently in certain structural characters, including a variably distinct subapical tooth on the mandibles, presence of rugosities on the metapleuron anterior to the propodeal spiracles, scutellar scale sometimes wider than its anterolateral carinae, and the propodeum strongly broadened behind the propodeal spiracles in dorsal view. In the case of the males slight structural variations were also observed, such as the parapsis and notaulus being vestigial but still observable, intervals of the scutellar sculpture rudimentary aligned so as to form an irregular longitudinal medial carina, sculpture variably dense on sterna, and size/length of the sternal pit varying in certain specimens.
Traumatomutilla juvenilis (Gerstaecker, 1874) (Figs. 29–48)
Mutilla juvenilis Gerstaecker, 1874: 75, holotype, f#, Brazil, [São Paulo], Salto Grande (ZMB), examined. Mutilla (Traumatomutilla) bivittata rubroguttata André, 1901: 257. Holotype [by monotypy], f#, Paraguay, [Alto Paraguay], Porto Casado [Puerto Casado/Puerto La Victoria] (location unknown) syn. nov. Mutilla (Traumatomutilla) bivittata immaculiceps André, 1901: 257, lectotype [designated here], f#. Paraguay, Rio Monday [sic] (MNHN) examined, syn. nov. Mutilla estrella (Cresson, 1902): 47, lectotype [designated by Cresson (1916)], f#, Brazil, Corumbá (CMNH) examined, syn. nov. Ephuta (Traumatomutilla) bivittata rubroguttata: André, 1902, 54. Ephuta (Traumatomutilla) bivittata immaculiceps: André, 1902, 54. Ephuta (Traumatomutilla) estrella: André, 1902, 55. Ephuta (Traumatomutilla) juvenilis: André, 1902, 55. Traumatomutilla bivittata rubroguttata: André, 1904, 40. Traumatomutilla bivittata immaculiceps: André, 1904, 40. Traumatomutilla estrella: André, 1904, 40. Traumatomutilla juvenilis: André, 1904, 40.
Diagnosis. FEMALE. In addition to the structural characters referenced in the species groups diagnosis, T. juvenilis females can be defined by the metasoma with lateral margins anterior to propodeal spiracle simply divergent anterad, not constricted; dorsal surface of hindtibia densely setose; pronotal spiracle projected; and T2 disc evenly convex. MALE. In addition to the structural characters referenced in the species groups diagnosis, T. juvenilis males can be defined by having the cuspis broad, club-like, densely setose throughout; lateral margins of propodeum simply rounded, not angulate; and T2 evenly convex. Description. Body length 14–17 mm. Head. Posterior margin virtually straight. Vertex width 0.8 × pronotal width. Eye length in frontal view virtually as long distance from its ventral margin to mandibular condyle. Head densely, coarsely and confusedly foveolate-punctate, sparser and finer on gena and malar space. Mandible with small subapical tooth. Dorsal scrobal carina present defined, separated from antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercle densely finely and confusedly rugulose. Flagellomere 1 2.2 × pedicel length; flagellomere 2 1.6 × pedicel length. Genal carina present, well defined, short, broadly separated from gular carina and hypostomal carina. Occipital carina well 70 defined, slightly swollen and curved in straight angle dorsolaterally. Mesosoma. Dorsal thoracic length 0.9 × its width. Mesosomal dorsum well differentiated from pleurae, lateral margins rounded, not angulate or sharp; sculpture predominantly obscured by dense setation, dense confused and coarse areolate-punctate to foveolate-punctate where visible. Anterior face of propodeum defined, superficially and longitudinally striate mesoventrad, coarsely punctate laterodorsad; dorsal face roundly angulate into anterior face in lateral view. Humeral carina present broadly separated from well-defined low rounded epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Sculpture of lateral face of pronotum densely micropunctate with sparse coarse punctures anteriorly; mesopleuron densely micropunctate anteriorly, sparse and coarse foveolate-punctate along mesopleural ridge, concealed by dense setation elsewhere; metapleuron sculpture concealed by dense setation on ventral half, unsculptured and smooth on dorsal half. Lateral face of propodeum sparsely and shallowly foveolate-punctate anteriorly, with dense coarse confused horizontally oblique rugosities along posterior margin. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 52:61:58:48:45. Lateral margin of mesonotum not emarginated anterior to propodeal spiracle, smoothly and slightly diverging anterad. Propodeal spiracle virtually flat against lateral margin of mesosoma; post- spiracular area absent. Scutellar scale present, well-developed, slightly narrower than and separated from well-developed anterolateral carinae which are connected thus forming a single transverse carina with shallow emargination medially; scabrous intervals present on scutellar area. Posterior face of propodeum longer than and poorly distinguished from dorsal face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 32:77:75. T2 slightly longer than wide, with maximum width posterior to midlength; Disc of T2 dense and coarse foveolate-punctate, sculpture denser and smaller mediad; sculpture of integumental spots sparse and shallow foveolate-punctate. Sculpture of T3–6, except pygidial plate, predominnalty concealed by dense setation, dense and coarse foveolate-punctate with sparse interspersed micropunctures where visible. S1 surface cuneiform, longitudinally elevated medially, equally high throughout. S2 sparse foveolate- punctate, foveolations sparser and smaller mediad; subapical transverse slope virtually absent medially, conspicuous laterally; anteromedial crest-fold well-developed. S3–6 dense foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third; surface with irregular, mostly longitudinal coarse rugosities; interstice apparently granulose. MALE. Body length 11–18 mm. Head. Rounded subrectangular in dorsal view, posterolateral angles rounded. Vertex width 0.75 × pronotal width. Eye almost circular. Ocelli small; OOD 6.1 × DLO, IOD 1.75 × DLO. Occipital carina distinct. Head surface densely and coarsely foveolate-punctate to punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting narrowly separated from internal eye margin. Clypeus convex medially, concave laterally immediately below antennal insertion; coarsely and densely foveolate-punctate; with a pair of short, sessile, blunt tubercles on apical margin. Scape bicarinate. Flagellomere 1 1.9 × pedicel length; flagellomere 2.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, low against anterior margin of pronotum, rounded, broadly disconnected from humeral carina; anterolateral angles of pronotum subangulate. Anterior face of propodeum coarsely and densely punctate laterally, impunctate medially, with a conspicuous longitudinal concave medial area. Tegula convex, mostly glabrous and impunctate except for dense punctures anterointernally. Mesoscutum densely and coarsely foveolate- punctate, notaulus and parapsis indistinguishable. Scutellum convex, densely and coarsely areolate-punctate. Axilla produced posterolaterally as truncate projection, with conspicuous flat posterior face, coarsely and densely foveolate- punctate dorsally; posterior face of axillar projection arched. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeum dorsum convex, predominantly concealed by dense setation, densely areolate where visible; lateral face predominantly densely areolate, areolations vestigial anterad; posterolateral margins rounded, posterodorsal corners rounded in lateral view; dorsal and posterior faces indistinguishable. Lateral face of pronotum with dense coarse punctures and interspersed micropunctures; mesopleura with strong subacute projected tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures annteriorly. Metapleuron predominantly unsculptured, smooth, except ventral fourth, densely and coarsely areolate- punctate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. 71
T1 0.5 × as wide as T2. T2 slightly longer than broad. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely foveolate-punctate to simply punctate where visible, except pygidial plate on T7 irregularly rugose and weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, forming pronounced carina slightly higher posteriorly. S2 densely foveolate-punctate, foveolations conspicuously sparser and smaller mediad; with well- developed longitudinal anteromedial crest-fold ending on small and narrow longitudinal pit densely filled with setae. S3–7 sparsely and dinely foveolate-punctate to punctate. S7 as long as broad, with single subacute tooth-like projection on posterior margin. Genitalia. Parapenial lobe slightly pronounced apically. Ratios of free length of paramere, cuspis and digitus, 51:39:7; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense long setae ventrally, setae sparser apicad; cuspis moderately thick, club-like, at its widest point half the width of paramere in dorsal view, virtually straight and equally wide throughout in dorsal view, densely covered in thick setae throughout; paracuspis well-developed, not sessile, amorphous, lobe-like, densely setose, setae as long as paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose dorsally; penis valve strongly concave on internal surface, with very closely spaced pair of teeth apicoventrally; apical tooth more acute and longer; subapical tooth subrounded, externolateral pocket vestigial; apical distance between teeth 0.05 × length of valve; dense setae present along apical margin and at base of teeth on external surface. Coloration and variations. Head integument always black, at most brownish-black; head setation can be completely black or with a pair of narrow longitudinal stripes of silvery-white setae along extending from the posterior margin of vertex along the inner eye margin to antennal tubercles; in certain specimens these stripes are reduced to a pair of lateral longitudinal spots of silvery-white setae on vertex only; specimens with silvery-white setae on vertex and/or front usually have silvery-white setae on malar space and ventral surface of head. Mesosoma integument is always completely black, at most brownish-black; setation is predominantly black dorsally with all forms having lateral longitudinal stripes of silvery- white setae usually extending from posterior margin of propodeum to posterior margin of pronotum; stripes might end before pronotal spiracles or beyond posterior margin of pronotum in certain specimes; pleural setae is predominantly black with silvery-white setae ventrally on mesopleuron and metapleuron in all forms. Integument of metasoma is always predominantly black, at most brownish-black, except for disc of T2 which is always marked with integumental spots that can be reddish or yellowish; integumental spots vary greatly in size and can be subquadrate, subrounded or linear/subrectangular. Metasomal setae are predominantly black, except for silvery-white setae varying in density on the following: T1 laterally; T2 lateral margins, lateral felt lines, fringe laterally, and over integumental spots; T3–5 medially and laterally; T6 medially; S1–5 (except fringe of S5). Appendages color predominantly black, except mandibles and flagellomeres partially reddish-brown. MALE. Integument always completely black, at most brownish-black, with mandibles and most flagellomeres partially reddish-brown. Body setae is predominantly black with silvery-white setae varying in density and extension in the following: antennal tubercles; metanotum; propodeum; T1, base of T2, lateral margins of T2, lateral felt line of T2, and fringes of T2–4 laterally; S1–S5 except fringe of S5; legs. Distribution. Brazil, Bolivia, Paraguay and Argentina. Material examined. (324 f#, 28m#) Type material. Lectotype: Traumatomutilla juvenilis, f#, BRAZIL, [São Paulo], Salto Grande, Sello S. (ZMB); Syntype: Traumatomutilla immaculiceps, f#, PARAGUAY, [Distrito Capital], Asunción, 1891 (MNHN); Holotype: Traumatomutilla estrella, f#, BRAZIL, [Mato Grosso do Sul], Chapada [dos Guimarães] (CMNH). Additional material: BRAZIL: Mato Grosso do Sul: Corumbá, 2f#, IV (CMNH); Chapada [dos Guimarães], Campo, 5f#, IX (CMNH); Mato Grosso: Confluência Xingú-Kuluene, 1f#, VI.1947 (MNRJ); P.N. [Parque Nacional] do Xingú, Jacaré, 1f#, Alvarenga & Werner (DZUP); Barra do Tapirapé, 5f# (MZSP); Diamantino, Alto Rio Arinos, 4f#, X.1983 (MNRJ); Alto Xingú, P.I. [Posto Indígena] Col. [ Coronel] Vasconçelos, 2f#, XI.1958 (MNRJ); Aldeia Camariura, 1f#, 24.XI.1960 (MNRJ); marg. esq. [margem esquerda] rio Sucuriú, Faz. [Fazenda] Canaã, Três Lagoas, 1f#, 10.XII.1967, F. Lane (MZSP); Rio Papagaios, Utiariti, 22–31.X.1966, Lenko & Pereira (MZSP); Rosário Oeste, 5f# (MZSP); Santa Isabel, 1f#, 18.I.1944, P.A. Berry (CISC); Tapirapé Indian Village, at confluence of Rio Tapirapé and Rio Araguaia, 1f#, 11.XII.1960, B. Malkin (FMNH); Poconé, Pouso Alegre, 1f#, 28.I.2001, A. Köhler (CESC); Mato Grosso do Sul: Anastácio, Fazenda Boa Esperança, 1f#, 16–26.II.2008, Bossi R. Bervian C. (MZSP); Bodoquena, Cara da Onça, 310m [above sea level], 20°44’26’’S 56°44’04’’W, 1f#, XII.2012, T.H. Auko e eq. [equipe] (UFGD); Morra D. Sul [sic], 72
Califórnia, 250m [above sea level], 20°42’07’’S 56°52’47’’W, 1f#, II.2007, V. Carbonari (UFGD); Três Lagoas, 5f# (MZSP); Pará: Santarém, 1f#, VII.1919, S.M. Klages (CMNH); Conceição do Araguaia, 1f#, 18.XI.1979, I.S. Gorayeb (MPEG); Gorotire, Rio Fresco, 1f#, X.1985, W.L. Overal (MPEG); Mocajuba, Mangabeira, 9f# (MNRJ); Rondônia, MZ [sic] Polo Noroeste, Pimenteiras, 1f#, 15–27.XI.1985 (MZSP); São Paulo: Ilha Solteira, Campos Nativos, 1f#, 05.V.1990, Borrmann (EMUS); Reserva de Jataí, Luis Antônio, 16.X.1999, Melo G.A.R. (DZUP); Cotia, 1f# (DEI); Faz. [Fazenda] Itaquerê, Nova Europa, 1f#, 04.III.1964, K. Lenko (MZSP); N. [Novo] Horizonte, 1f#, XI.1944 (MNRJ); Angatuba, 1f#, III.1917 (MNRJ); Andradina, 1f#, 10.II.1959 (MNRJ); Anhembia, Faz. [Fazenda], Barreiro Rico, 1f#, 15.XI.1965, W.D. Edmonds (MNCN); Araraquara, 1f#, 17–18.I.1941, M. Carrera (MNCN); Franca, 1f#, X.1910, E. Garbe (MNCN); Jundiaí, 1f#, 17.XI.1895, Schrottky (MNHN); Ribeirão Preto, 1f#, 21.X.1962, Exp. Dep. Zool. [Expedição Departamento Zoologia] (MNCN); São Paulo, 1f#, 06.I.1925 (DEI); Rio Claro, 2f#, Braulio Dias (AMNH); Avaré, 1f#, E. Amante (AMNH); Minas Gerais: Passos, 1f#, 23–31.I.1963, Elias C. (DZUP); Araxá, 1f#, 06.XI.1965, Elias C. & Elias T. (DZUP); Ituiutaba, 1f#, XI.1963 (MNRJ); Paraopeba, 1f#, n.o proc. [número de procedência?] 16/53 (MNRJ); Araguari, 1f#, X.1931, R. Spitz (MNCN); Lassance, 1f#, 09–19.XI.1919, Cornell University Expedition (CUIC); Sertão de Diamantina, Fazenda das Melâncias, 1f#, 10.XI.1902, E. Gounelle (MNCN); Uberlândia, 1f#, X.1962, Exp. Dep. Zool. [Expedição Departamento Zoologia] (MNCN); Goiás: Campinas, 1f#, X.1937 (MNCN); Fazenda Cachoeirinha, Jataí, 1f#, X.1962, Exp. Dep. Zool. [Expedição Departamento Zoologia] (EMUS); Jataí, Faz. [Fazenda] Aceiro, 1f#, X.1962, Exp. Dep. Zool. [Expedição Departamento Zoologia] (MNCN); Piranhas, 1f#, 02.XI.1961, Werner (MNCN); Viannópolis, 1f#, XI.1931, R. Spitz (MNCN); Distrito Federal, Brasília, 1f#, Braulio Dias (AMNH); BOLIVIA: Santa Cruz: El Carmen, 1f#, 25–27.II.1954, C. Gans & F. Pereira (MNCN); Buena Vista, 2f#, 1928, J. Steinbach (CUIC); Santa Cruz de la Sierra, 450m [above sea level], 2f#, XI.1910, J. Steinbach (CMNH); Puerto Suárez, 150m [above sea level], 1f#, XII.1908, J. Steinbach (CMNH); Pacajes, Corocoro, 1f#, 19.II.1995, L. Peña (AMNH); Beni: Cavinas, 1f#, I.1922, W.M. Mann (USNM); Guayaramerín, 1f#, XI.1956, Fritz (AMNH); Reyes, 1#, X.1921, W.M. Mann (USNM); PARAGUAY: San Pedro: Cororo, Rio Ypane: 1f#, XII.1983, M.A. Fritz (EMUS); 7m#, II.1979, Fritz (AMNH); Pto. [Puerto?] Pablo, 8f# 3m#, XI–XII.1936 (RBINS); Kanindeyu, Tava Yopoi, 24°22’S 55°53’W, 26.X–04.XI.2007, U. Dreschel (EMUS); Guairá, Villa Rica, 1f#, 1900, Burgdorf (MNHN); Distrito Capital, Asunción, 1f#, 06.XII.1905 (MNHN); Paraguari: E [East] of Ybicui, 1f#, 02.IV.2006, U. Dreschel (FSCA); Sapucay, 1f# (USNM); La Rosada, 26°06’S 56°50’W, 1f#, 27–30.XII.2008, U. Dreschel (FSCA); Presidente Hayes, Puerto Galileo, 3f#, 02–05.III.2006, U. Dreschel (FSCA); Lolita, Yaragui, 23°06’S 59°38’W, 1f#, 05.III.2003, U. Dreschel (EMUS); Cordillera, San Bernardino, 1f#, K. Fiebrig (USNM); Caaguazu: Zudañez, 1f#, I.1949 (UMMZ); Caaguazu, 2m#, XII.1977, Fritz (AMNH); Caazapa, Colonia Neufield, 390’[sic], 26°28’S 55°55’W, 1f#, 24.IX–02.XI.2008, U. Dreschel (EMUS); Concepción: 25 mi [Miles] SE [southeast] San Carlos, 1f#, 27.X.2002, U. Dreschel (FSCA); Chaparral, 1f#, 27.X.2002, U. Dreschel (FSCA); Cororo, 2f#, II.1993, Arriagada (AMNH); Amambay, 2km [kilometers] S [south] Cerro Cora, 1f#, 28.XI.1973, O.S. Flint (DGMC); Alto Paraguay, Cuaacupe, 1f#, 14.XII.1944, P.A. Berry (CISC); Chaco, Est. [Estación] Hermosa, Rio Siete Puntas, 1f#, 27.X.1946, C. Olrog (NHRM); ARGENTINA: Chaco: Ciervo Petizo, 1f#, 25.XI.1949, Duret (MNHN); San Bernardo, 1f#, II.1990, Di Lorio (AMNH); Formosa, Gran Guardia, 2f#, J. Foerster (AMNH); Mendoza, Estancia Pedregal, 4f#, 21.XII.1906, Jorgensen (ZMUC); Entre Rios, San Jose, 1m#, X.1925, Lindner D. Chaco Exp. [Departamiento Chaco Expedition] (ZMB). An additional 149f# from Brazil, 5f# from argentina, 2f# from Bolivia, and 41f# as well as 15m# from Paraguay were also examined (AMNH, ANSP, CASC, CMNH, DGMC, DZUP, EMUS, FEIS, FSCA, LACM, MNHN, MNRJ, MPEG, MZSP, RBINS, USNM, ZMB, ZMUC) Remarks. The discovery of the male of T. miniata coupled with a large series of males and females from Puerto Pablo and Cororo in Paraguay (RBINS and EMUS) facilitated the sex association for T. juvenilis. The females of that series are all clearly specimens of T. juvenilis (the T. immaculiceps color form) whilst the males have nearly identical genitalia to those of T. miniata but lack the sharply angulate propodeum and T2. Females of T. juvenilis are similar in structure to females of T. miniata apart from inconspicuous sculpture characters and the conspicuously pronounced, almost tuberculate, pronotal spiracles of T. miniata. Therefore, the sexes of T. juvenilis were associated based on repeated geographical co-ocurrence and morphological “clues” provided by both sexes of T. miniata. 73
André (1901) provided few comments regarding T. bivittata rubrogutttata and referred to two specimens “reported by M. Boggiani” from Puerto Casado in Paraguay which, we assume, the author considered as the type series for T. bivittata rubroguttata. Unfortunately, there is no indication of where the types were deposited at the time and we were unable to retrieve any information on such specimens. In the description of T. bivittata rubroguttata, however, André mentioned an additional specimen collected in Brazil, which belonged to his personal collection and agreed in every aspect with the Boggiani specimens. We have examined every specimen of Traumatomutilla in André’s collection at MNHN and found four specimens labeled as T. bivittata rubroguttata, all from Mato Grosso state in Midwestern Brazil. It’s not clear, however, which of the specimens is the one André (1901) referred to. Although we haven’t seen the actual types of T. bivittata rubroguttata (Boggiani specimens), all four specimens from André’s collection agree with each other in every aspect. Additionally, André (1901) reported that the Boggiani specimens were in agreement with Gerstaecker’s description of T. bivittata except for the T2 integumental spot color. The fact that Gerstaecker’s descriptions focused almost exclusively on color and setae characters could explain why such structurally distinct species were considered to be so closely related. Traumatomutilla immaculiceps was also originally described as a subspecies of T. bivittata by André (1901) and elevated to species level by André (1910) based on minor setae differences of the head and metasomal segments. According to André (1910) the type series from T. immaculiceps consisted of four females from Asunción, Paraguay, with collecting dates, December 30th 1904, March 14th 1906, and April 11th 1906. We examined a series of four specimens labeled by André as T. immaculiceps deposited at MNHN, two of which were labeled SYNTYPE upon preparation of the loan. One of the SYNTYPES, though identified as T. immaculiceps by André himself, is actually Traumatomutilla tristis (Gerstaecker)from the T. indica species-group. None of the specimens in the series matches the label data provided by André (1910) and according to the collection managers at MNHN there are no other specimens in André’s collection labeled as T. immaculiceps by André himself. The correctly identified syntype is from Asunción, apparently collected in 1891, whilst the other two specimens, though both from Paraguay, have different collecting locality and/or collecting date. It is unclear at this moment why are there such differences in the label data between André (1910) specimens and the specimens deposited at MNHN. Based on the fact that the correctly identified syntype was labeled by André and is from the correct locality provided by the author upon description, we designate this specimen as the lectotype for T. immaculiceps. We refrain, however, from designating the other two correctly identified specimens of T. immaculiceps as paralectotypes since, though from Paraguay, they are not from the same locality mentioned by André (1910). Casal labeled numerous specimens in different collections as “T. guarania sp. nov.” without, however, publishing this putative new species or mentioning it in his 1969 paper on Traumatomutilla which are all identical to T. estrella. Slight structural variations were observed in the specimens examined: mesosoma as wide as, narrower, or wider than distance between pronotal spiracles; presence of obvious to vestigial to absent oblique rugae on lateral face of propodeum; and medial intervals at mesosomal dorsum vestigially aligned so as to form an irregular longitudinal medial carina.
Traumatomutilla juvenindica Williams & Bartholomay, sp. nov. (Figs. 49–56)
Diagnosis. FEMALE. Head with transversal stripe of dense silvery-white setae; genal carina vestigial, porrly defined; scabrous intervals present on posterior third of mesosoma. MALE. Cuspis slender and predominantly asetose, pygidial plate restricted to apical third of T7, head and pronotum clothed with dense silvery-white setae. Description. FEMALE. Body length 12–14 mm. Head. Posterior margin virtually straight. Vertex width 0.8 × pronotal width. Eye length in frontal view 1.1 × distance from its ventral margin to mandibular condyle. Front and vertex densely, finely and confusedly areolate-punctate to foveolate-punctate; gena and malar space sparsely foveolate-punctate, with well-defined unsculptured and smooth areas. Mandible with small subapical tooth. Dorsal scrobal carina present defined, separated from antennal tubercles and from vestigial lateral scrobal carina; lateral scrobal carina reduced to longitudinal impunctate area. Antennal tubercle virtually unsculptured, smooth. Flagellomere 1 2.4 × pedicel length; flagellomere 2 1.8 74
× pedicel length. Genal carina vestigial, to a narrow vertical, mostly unsculptured, smooth area, broadly separated from gular carina and hypostomal carina. Occipital carina well defined, conspicuously swollen and curved in a straight angle dorsolaterally. Mesosoma. Dorsal thoracic length 0.8 × its width. Mesosomal dorsum predominantly dense confused and coarse areolate-punctate to foveolate-punctate, areolations denser and smaller mediad; scabrous intervals present on posterior third of mesonotum. Anterior face of propodeum defined, conspicuously and finely striated longitudinally basad, confusedly and densely punctate dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina present broadly separated from slightly pronounced blunt epaulet, antero-lateral corners of pronotum subrounded in dorsal view. Pronotal spiracle strongly projected from lateral margin of pronotum, rounded, not at all acute. Sculpture of lateral face of pronotum confusedly punctate with interspersed micropunctures; mesopleuron densely micropunctate anteriorly, sparsely and coarsely foveolate-punctate along mesopleural ridge, elsewhere obscured by dense setae; metapleuron sculpture on dorsa half unsculptured, smooth, ventral half concealed by dense setation. Lateral face of propodeum mostly unsculptured on anterior two thirds, smooth, sparsely and shallowly foveolate-punctate; posterior third densely, confusedly foveolate- punctate with interspersed vestigial oblique rugosities. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 54:70:55:53:47. Lateral margin of mesosoma not emarginated anterior to propodeal spiracle, smoothly and slightlu diverging anterad and converging slightly posterior to pronotal spiracles. Propodeal spiracle virtually flat against lateral margin of mesosoma; post-spiracular absent. Scutellar scale present, well-developed, slightly narrower than and separated from well developed anterolateral carinae which are connected thus forming a single transverse carina almost reaching lateral margin of mesosoma; scabrous intervals present on scutellar area. Posterior face of propodeum longer than and poorly distinguished from dorsal face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 39:82:87. T2 slightly longer than wide, with maximum width posterior to midlength. Disc of T2 dense and fine foveolate-punctate with sparse interspersed micropunctures to dense and fine punctate mediad; sculpture of integumental spots sparse shallow foveolate-punctate. Sculpture T3–6, except pygidial plate, dense and fine foveolate-punctate with sparse interspersed micropunctures. S1 surface cuneiform, longitudinally elevated medially, equally high throughout. S2 sparse foveolate-punctate, foveolations sparser mediad; subapical transverse slope virtually absent, vestigial laterally; antero-medial crest-fold well-developed. S3–6 dense fine foveolate-punctate. Pygidial plate broad, subpyriform, defined by lateral carinae at apical third; surface with irregular, mostly longitudinal coarse rugosities; interstice apparently sparsely granulose. MALE. Body length 13–15 mm. Head. Rounded subrectangular in dorsal view, posterolateral angles rounded. Vertex width 0.75 × pronotal width. Eye almost circular. Ocelli small; OOD 2.9 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface mostly concealed by dense setation, except ventral surface densely and shallowly punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina narrowly separated from internal eye margin and slightly extending into lateral margin of antennal scrobe. Clypeus slightly concave laterally immediately below antennal insertion, convex medially; coarsely and densely foveolate-punctate to punctate; with a pair of short, sessile, blunt tubercles on apical margin. Scape bicarinate. F1 2.2 × pedicel length; F2 2.6 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, rounded, broadly separated from poorly defined humeral carina, anterolateral angles of pronotum rounded. Anterior face of propodeum densely micropunctate laterally, unsculptured medially, with a conspicuous longitudinal slightly concave medial area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures anteriorly and along inner margin. Mesoscutum densely and finely foveolate-punctate, notaulus and parapsis indistinguishable. Scutellum convex, densely and coarsely foveolate-punctate. Axilla produced posterolaterally as truncate projection, with conspicuous flat posterior face, coarsely and densely foveolate-punctate dorsally; posterior face of axillar projection arched. Metanotum virtually equally wide throughout, surface obscured by dense setation. Propodeum dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face predominantly densely and coarsely areolate, areolations less defined anterad; posterolateral margins rounded, posterodorsal corners rounded in lateral view; dorsal and posterior faces indistinguishable. Lateral face of pronotum densely and coarsely foveolate-punctate to punctate with interspersed micropunctures; mesopleura with strong subacute tubercle on dorsal half; mesopleural sculpture densely and coarsely 75 areolate-punctate to foveolate-punctate with interspersed micropunctures anteriorly. Metapleuron virtually unsculptured and asetose throughout. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate where visible; T7, except pygidial plate, foveolate punctate; Pygidial plate restricted to apical third of T7, conspicuously wider than longer, defined by lateral carinae laterally, with conspicuous carina laterobasally converging into longitudinal medial carina extending basad on T7; pygidial plate surface irregularly rugulose. S1 longitudinally elevated medially, forming pronounced carina slightly lower medially. S2 saprsely foveolate- punctate, foveolations conspicuously denser laterad; with well-developed longitudinal anteromedial crest-fold ending on elongate patch of silvery-white setae. S3–7 sparsely foveolate-punctate to punctate. S7 slightly broader than long, with a pair of subacute closely spaced tooth-like projections on posterior margin. Genitalia. Parapenial lobe slightly pronounced apically. Ratios of free length of paramere, cuspis and digitus, 93:70:13; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense setae ventrally except apically with setae conspicuously reduced; cuspis narrow, slender, slightly curved inward at basal half and curved outward in apical half in dorsal view; conspicuously and smoothly widened apically; predominantly asetose with sparse conspicuous setae ventrally at apex; paracuspis well- developed, not sessile, longer than wide, subrounded at apical margin, densely setose, setae predominantly shorter than paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose basodorsally; penis valve strongly concave on internal surface, with very closely spaced pair of teeth apicoventrally; apical tooth more acute and slightly longer; subapical tooth acute, externolateral pocket virtually absent, vestigial; apical distance between teeth 0.03 × length of valve; dense setae present along apical margin and at base of teeth on external surface; outer surface with conspicuous medial projection on ventral margin. Coloration and variations. FEMALE. Body and appendages black to brownish-black, except most antennal flagellomeres and mandibles partially reddish-brown. Body setae predominantly black, except the following areas with silvery-white setae varying in density: transverse stripe on vertex of head; lateral longitudinal stripes from posterior margin of propodeum to anterior margin of mesonotum; ventral third of mesosomal pleurae; most of legs; T1 laterally; integumental spots, lateral area, lateral fringes and lateral margins of T2; T3–5 laterally and medially; most of T6; S1–6. No significant coloration or setae variations were observed in the specimens examined. MALE. Body and appendages black, at most mandibles and/or most flagellomeres partially reddish-black. Body setae predominantly silvery-white varying in density except for the following areas with black setae varying in density: scutum, axillae, anterior half of scutellum, posteiror third of T2, T6–7, most of S6, and S7 (hypopygium). Certain specimens might have a longitudinal narrow strip of silvery-white setae extending medially from anterior half to fringe of T2. Distribution. Colombia and Venezuela. Etymology. A merging of the epithets of T. juvenilis and T. indica (Linnaeus). An allusion to the fact that this species has the structural traits of T. juvenilis and the overall color and setal patterns of T. indica. Material examined. (2f#, 4m#) Type material. Holotype: f#, VENEZUELA, Lara, Guamacire [sic!], Hda. [Hacienda?] Sigata, 26.VIII.1976, J.M. Osorio (FSCA); Paratype: f#, COLOMBIA, Cesar, Becerril, 24.VII.1968, B. Malkin (AMNH). Allotypes: m#, COLOMBIA, [Cesar], Lake Sapatoza region, Chiriguana district, VIII–IX.1924, C. Allen (BMNH); m#, same label data as previous specimen (BMNH); m#, COLOMBIA, [Bolivar], Magdalena Valley, El Banco, C. Allen (BMNH); #m, COLOMBIA, Bolívar, Zambrano, Hda. [Hacienda] Monterrey, 09°45’N 74°49’W, Malaise 5, Andaluz 14, 10m [above the ground], 08.VII.1993, F. Fernández (UNAL). Remarks. Before the discovery of T. juvenindica, the northernmost distribution record for a species of the T. juvenilis species-group was a single female collected in Santarém at the Brazilian Amazon, nearly 2.200 km away from the closest record for T. juvenindica (Guamacire, Lara, Venezuela) with males and females collected even further away in Becerril, César, Colombia (~5.000 km from Santarém). This apparent isolation of T. juvenindica from the main bulk of the T. juvenilis species-group leaves us with almost no doubt about this sex association. It is peculiar, however, that the males of T. juvenindica have genitalic characters like males of T. duplicata and T. guarata whilst the females are clearly morphologically closer to those of T. juvenilis and T. miniata. 76
Minor structural variations were observed between the female specimens in the type series, namely 1) the sculpture of the lateral face of the propodeum, which varies in density and 2) a conspicuous longitudinal carina medial carina on the mesosomal dorsum with its adjacent sculpture simply punctate and contrasting with the areolate-punctate sculpture on the margins of the mesosomal dorsum. The medial longitudinal carina on the mesosoma is present in all species of the T. indica group, but vestigial forms of such a carina can also be found in specimens of the T. quadrinotata and T. gemella species-groups. Females of T. juvenindica have the legs remarkably asetose, smooth and shinning in comparison with other females in the group and indeed in the genus as a whole. All but one specimen in the type series were collected northwest of the Andes in Colombia, which has a peculiar fauna including the only South American records for the mainly North and Central American genus Dasymutilla Ashmead (Bartholomay et al. 2019c). It is also noteworthy that both sexes of this species bear the conspicuous Amazonian color pattern that can be observed in other species such as T. indica (Linnaeus), T. guayaca Casal, and T. selligera (Gerstaecker).
Traumatomutilla miniata (Gerstaecker, 1874) (Figs. 57–68)
Mutilla miniata Gerstaecker, 1874: 75, lectotype [designated here], f#, Argentina, Catamarca (MLUH), examined. Ephuta (Traumatomutilla) miniata: André, 1902, 55. Traumatomutilla miniata: André, 1904, 40. Ephuta (Traumatomutilla) bispiculata André, 1907: 350, holotype [by monotypy], m#, Argentina, Santiago del Estero, Icaño, Laguna Mamaita (MNHN), examined, syn. nov. Traumatomutilla bispiculata: Nonveiller, 1990, 201.
Diagnosis. FEMALE. Head entirely clothed with black setae; pronotal spiracles strongly projected from lateral margins of mesosoma, subacute; mesosomal silvery-white setae stripes restricted to propodeum. MALE. Mesopleural projections sharp, acute; lateral margins of pronotum sharply angulate, projected; propodeal dorsum concave, posterolateral corners angulate; T2 conspicuously flattened, roundly angulate laterally, nearly forming distinct dorsal and lateral faces; cuspis club-like, densely setose throughout. Description. Body length 14–17 mm. Head. Posterior margin virtually straight. Vertex width 0.7 × pronotal width. Eye length in frontal view 0.6 × distance from its ventral margin to mandibular condyle. Front and vertex densely, coarsely and confusedly areolate-punctate, more densely so on vertex; gena and malar space sparsely and coarsely foveolate- punctate, with well-defined unsculptured and smooth areas; sculpture virtually absent near base of mandibles. Mandible worn-out subapical tooth not observable. Dorsal scrobal carina present defined, separated from antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercle densely and confusedly rugulose. Flagellomere 1 2.4 × pedicel length; flagellomere 2 1.4 × pedicel length. Genal carina present, well defined, short, broadly separated from gular carina and hypostomal carina. Occipital carina well defined, slightly swollen and smoothly curved dorsolaterally. Mesosoma. Dorsal thoracic length 0.9 × its width. Mesosomal dorsum sharply differentiated from pleurae, lateral margins angulate; sculpture predominantly obscured by dense setation, dense confused and coarse areolate-punctate where visible. Anterior face of propodeum defined, sculpture mostly concealed by dense short setation, vestigially striated longitudinally where visible; dorsal face roundly angulate into anterior face in lateral view. Humeral carina present broadly separated from strongly pronounced rounded epaulet, anterolateral corners of pronotum subangulate in dorsal view. Pronotal spiracle strongly projected from lateral margin of pronotum, subacute. Sculpture of lateral face of pronotum sparsely punctate with interspersed micropunctures; mesopleuron densely micropunctate anteriorly, dense coarse and confused areolate-punctate to foveolate-punctate along mesopleural ridge and basally above mesocoxa; metapleuron sculpture densely micropunctate, except on dorsa third mostly impunctate with vestigial oblique rugosities anterior to propodeal spiracle. Lateral face of propodeum sparsely punctate anteriorly, with dense coarse confused horizontally oblique rugosities along posterior margin interspersed with coarse areolations. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 72:91:81:67:62. Lateral 77 margin of mesonotum not emarginated anterior to propodeal spiracle, smoothly and slightly diverging anterad. Propodeal spiracle virtually flat against lateral margin of mesosoma; post-spiracular absent. Scutellar scale present, well-developed, slightly narrower than and separated from well-developed anterolateral carinae which are connected thus forming a single transverse carina with shallow emargination medially; scabrous intervals present on scutellar area. Posterior face of propodeum longer than and poorly distinguished from dorsal face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 42:94:93. T2 slightly wider than long, with maximum width posterior to midlength; roundly angled laterally, with apparent dorsal and lateral faces. Disc of T2 dense, coarse and confused foveolate-punctate, denser and coarser mediad; sculpture of integumental spots sparse coarse shallow foveolate-punctate. Sculpture T3–6, except pygidial plate, dense and coarse foveolate-punctate with sparse interspersed micropunctures. S1 surface cuneiform, longitudinally elevated medially, equally high throughout. S2 sparse foveolate-punctate, foveolations sparser and smaller mediad; subapical transverse slope virtually absent medially, conspicuous laterally; anteromedial crest-fold well-developed. S3–6 dense foveolate-punctate with interspersed sparse micropunctures. Pygidial plate broad, subpyriform, defined by lateral carinae at apical third; surface with irregular, mostly longitudinal coarse rugosities; interstice apparently granulose. MALE. Body length 11–16 mm. Head. Rounded subrectangular in dorsal view, posterolateral angles rounded. Vertex width 0.7 × pronotal width. Eye almost circular. Ocelli small; OOD 5.0 × DLO, IOD 3.1 × DLO. Occipital carina distinct. Head surface densely and coarsely foveolate-punctate punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting narrowly separated from internal eye margin. Clypeus convex medially, concave laterally immediately below antennal insertion; coarsely and densely foveolate-punctate; with a pair of short, sessile, blunt tubercles on apical margin. Scape bicarinate. Flagellomere 1 2.2 × pedicel length; flagellomere 2 5 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, strongly projected from anterior margin of pronotum, rounded, broadly disconnected from humeral carina; humeral carina projected dorsally, anterolateral angles of pronotum sharp. Anterior face of propodeum coarsely and densely areolate-punctate laterally, impunctate medially, with a conspicuous longitudinal concave medial area. Tegula convex, mostly glabrous and impunctate except for dense punctures anterointernally. Mesoscutum densely and coarsely foveolate-punctate, intervals aligned so as to form vestigial longitudinal carinae; notaulus and parapsis indistinguishable. Scutellum convex, densely and coarsely areolate-punctate; with longitudinal irregular carina medially. Axilla produced posterolaterally as truncate projection, with conspicuous flat posterior face, coarsely and densely foveolate-punctate dorsally; posterior face of axillar projection arched. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeum dorsum concave, predominantly concealed by dense setation, densely areolate where visible; lateral face predominantly densely areolate, areolations vestigial anterad; posterolateral margins sharply angulate, posterodorsal corners in a straight angle in lateral view; dorsal and posterior faces indistinguishable. Lateral face of pronotum with dense coarse and confused longitudinally oblique rugosities; mesopleura with strongly projected sharply acute tubercle on dorsal half, tubercle slightly curved posteriorly in dorsal view; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate. Metapleuron densely and coarsely areolate punctate on ventral fourth, virtually unsculptured and smooth on dorsal fourth, micropunctate elsewhere. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 length 0.8 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate where visible, except pygidial plate on T7 irregularly rugose and weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, forming pronounced carina slightly higher posteriorly. S2 sparsely foveolate-punctate, foveolations conspicuously sparser and smaller mediad; with well-developed longitudinal anteromedial crest-fold ending on small and narrow longitudinal pit densely filled with setae. S3–7 sparsely and shallowly foveolate-punctate to punctate. S7 longer than broad, with a single subacute tooth-like projection on posterior margin. Genitalia. Parapenial lobe slightly pronounced apically. Ratios of free length of paramere, cuspis and digitus, 119:98:16; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense long setae ventrally except at apical third; cuspis thick, club-like, at its widest point slightly narrower than paramere in dorsal view, virtually straight throughout, slightly narrower apically and basally, densely covered in thick setae throughout; paracuspis well-developed, 78 not sessile, roundly subtriangular, densely setose, setae as long as paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose dorsally; penis valve strongly concave on internal surface, with very closely spaced pair of teeth apicoventrally; apical tooth more acute and longer; subapical tooth subrounded, externolateral pocket present, very reduced; apical distance between teeth 0.05 × length of valve; dense setae present along apical margin and at base of teeth on external surface. Coloration and variations. FEMALE. Body and appendages predominantly black, except mandibles and most flagellomeres partially reddish-brown, and T2 with four usually subrounded reddish spots, varying in size in certain specimens; certain specimens might have very large spots which cause the anterior pair to be conspicuously or narrowly converging with the posterior pair. Body setae predominantly black except for the following areas with silvery-white setae varying in density: propodeal dorsum along lateral margins; coxae and ventral surface of meso and metafemora; T1 laterally; lateral areas of T2, lateral magins of T2, lateral felt lines of T2, and over posterior integumental spots of T2; fringes of T2–4 medially and laterally; fringe of T5 medially; T6, except pygidial plate, medially; S1–3. MALE. Body and appendages black, at most mandibles and most flagellomeres partially reddish-brown. Body setae predominantly black except the following areas with silvery-white setae varying in density: metanotum and propodeum dorsum; legs partially; T1, anterior third of T2, lateral felt lines of T2, lateral margins of T2, and fringes of T2–5 laterally; S1-5 except fringe of S5. No conspicuous variations were observed in the specimens examined. Distribution. Bolivia, Paraguay and Argentina Material examined. (101f# 32m#) Type material. Lectotype: Traumatomutilla miniata, f#, ARGENTINA, Catamarca (MLUH); Holotype: Traumatomutilla bispiculata, m# (nec f#), ARGENTINA, Santiago del Estero, La Balisa del Bracho, 20km N.N.O. d’Icaño [kilometers north-northwest of Icaño], Laguna Mamaita, E.R. Wagner, 1904 (MNHN). Additional material. BOLIVIA: Chugulasca [sic], 30km SE [kilometers southeast of] Carandaity, 1f#, 25.I.1957, Stephen C. Bremley (EMUS); Cordillera, Aimiri, 1f#, 16.II.1971, Fritz (AMNH); PARAGUAY: Presidente Hayes, Yaragui, 23°06'S 59°38'W, 1f#, 05.III.2003, U. Dreschel (FSCA); Chaco, Loma Plata, 2f#, I.1956, Gerlach (AMNH); Filadelfia, 1f# 1m#, I.1995, Arriagada (EMUS, AMNH); Agua Dulce, 2f#, X.1979, Fritz (AMNH); ARGENTINA: La Rioja: La Riojas, 1m#, XI.1967 (DGMC); Patquia, 1m#, XII.1933, K.J. Hayward (BMNH); Mendoza, Mendoza, 1m#, 4.I.1908, Jorgensen (ZMUC); Estancia Pedregal, 4f#, 04.I.1907, Jorgensen (ZMUC); Estación Santa Rosa, 3f#, Jensen-Haarup (ZMUC); Desaguadero, 1f#, 13.II.2005, F. Stipo (EMUS); Salta: La Viña, 1m#, 22–25.XII.1983, M. Wasbauer (UCDC); Pocitos, 1f#, Martinez (AMNH); Campo de Cuervo, 1f#, II.1968, O.H. Casal (AMNH); Cobos, 1f#, I.1993, Fritz (EMUS); Tartagal, 1m#, XI.1971, Fritz (AMNH); Santiago del Estero: Icaño: bords du [banks of the] Rio Salado, environs d’Icaño, 1m#, XI.1909, E.R. Wagner (MNHN); Rio Salado, 15f#, 1916 (MNCN); bañados du [marshes of] Rio Dulce, 60 km O d’Icaño [west of Icaño], 1f#, 1909, E.R. Wagner (MNHN); Guarda Esolta, environs d’Icaño [outskirts of Icaño], 1f#, 1909, E.R. Wagner (MNHN); Ruta 9, 70 km S Sgo. del Estero [70 kilometers south of Santiago del Estero], 1f#, 20.I.2002, F. & P. Cassola (DGMC); Sumampa, 1f#, IV.1955, Duret (MNHN); Los Telares, 1m#, 15.VII–13.VIII.1976, J.S. Noyes (BMNH); 37–47 k. s.e. [37 to 47 kilometers southeast of] Añatuya, 1f#, 20.XI.1979, C. & M. Vardy (BMNH); Thermas de Rio Hondo, 1f#, 27–28.XI.1979, C. & M. Vardy (BMNH); 13k s.e. [kilometers southeast of] Colonia Dora, 1f#, 17–19.XI.1979, C. & M. Vardy (BMNH); Chaco: El Cabure, 1f#, XII.1983, Fritz (AMNH); Miraflores, 1f#, II.1992, González (AMNH); Fuerte Esperanza, 2f#, XI.1978, Fritz (AMNH); Rio Tapenago, 1f#, fevrier [february] (MNCN); Cordoba: Cordoba, 3f#, III.1971, Fritz y Martinez (AMNH); Copacabana, 3f#, II.1980, Fritz (AMNH); Guanaco Muerto, 4f#, II.1971, Fritz (AMNH); Jujuy: San Pedro, 1f#, 25.XI.1947, Duret (MNHN); La Rioja: Iliar, 1f#, 1929, M. Gomez (MNCN); Olta, 1f#, 08.XII.1959, Libof [sic] (LACM); Patquia, 2f#, 04.XII.1960, Yivoff (USNM); Villa Mazan, 1m#, 15.I.2005, L.A. Stange (FSCA); Santa Fé: Las Garzas, 1f#, E. Le Moult (MNCN); San Luis, San Jeronimo, 2f#, I.1979, Williner (AMNH); Tucuman, 4km S [kilometers south of] Capitan Caceres, 2m#, 24.X.2003, Irwin & Parker (EMUS); Catamarca: Coneta, 16km S Catamarca, 11m#, 25.X.2003, Irwin & Parker (EMUS); Palo Labrado, 23 km S [kilometers south of] La Merced, 2m#, 24.X.2003, Irwin & Parker (EMUS); Trampasacha, 8 km W [kilometers west of] Chumbicha, 3m#, 25.X.2003, Irwin & Parker (EMUS). An additional 37 females and six males from Argentina as well as four females from Paraguay were also examined (AMNH, DEI, EMUS, FSCA, MNHN, ZMUC). 79
Remarks. There are conspicuous structural parallels between males previously known as T. bispiculata and females of T. miniata that permitted these species to be associated. Females of T. miniata have various body segments remarkably angulate, especially the limits between the mesosomal dorsum and mesosomal sides, to the extent that the sides of the mesosoma are virtually flat to concave in some specimens. Additionally, T2 is angulate to the point of having a somewhat distinct lateral face. The T2 character can also be observed in males of T. bispiculata, which have a sharply angulate propodeum that can be viewed as parallel to the overall angulate female mesosoma. Certain male specimens may have the propodeum almost flat, but never convex, and in these cases, they are also slightly depressed sublaterally on the dorsal face. Females can have the front with vestigial longitudinal carina or conspicuous smooth unsculptured medial area. Additionally, females can have a variably conspicuous anteroventral tubercle on the lateral face of the pronotum, anteroventral in relation to the pronotal spiracles.
Traumatomutilla guarata Casal, 1969 (Figs. 70–78)
Traumatomutilla guarata Casal, 1969: 291, holotype, f#, Brazil, Distrito Federal [Rio de Janeiro], [Rio de Janeiro], Guaratiba, 10.III.1952, A.G.A. Silva (AMNH) examined.
Diagnosis. FEMALE. Mesosoma strongly constricted anterior to propodeal spiracle in dorsal view, lateral face of propodeum evenly foveolate-punctate, frons with golden setae. MALE. Cuspis slender and predominantly asetose, pygidial plate through apical half of T7, lateral face of propodeum with golden setae. Description. FEMALE. Body length 12–17 mm. Head. Posterior margin virtually straight. Occipital carina slightly swollen and smoothly curved dorsolaterally. Vertex width 0.55 × pronotal width. Eye almost circular, its length in frontal view 0.8 × to the distance from its ventral margin to mandibular condyle. Head sculpture concealed by dense setation, except gena and malar space densely and coarsely foveolate-punctate. Mandible with small subapical tooth. Dorsal scrobal carina present, well-defined, reaching but not extending over antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercles virtually unsculptured, smooth and shinning. Flagellomere 1 2.4 × pedicel length; flagellomere 2 1.8 × pedicel length. Mesosoma. Dorsal thoracic length 0.8 × its width. Mesosomal dorsum well differentiated from pleurae, lateral margins rounded, not sharp or angulate; sculpture predominantly obscured by dense setation, dense and coarse areolate-punctate where visible, denser and smaller mediad; intervals vestigially scabrous. Anterior face of propodeum well-defined, coarsely striated longitudinally medially, sparsely punctate laterally; dorsal face roundly angulate into anterior face in lateral view. Humeral carina present, broadly separated from well-defined, slightly projected, rounded epaulet; anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle projected from lateral margin of pronotum, rounded. Sculpture of lateral face of pronotum sparsely punctate with dense micropunctures; mesopleuron densely micropunctate anteriorly, dense coarse and confused foveolate-punctate along mesopleural ridge, concealed by dense setation elsewhere; metapleuron sculpture predominantly concealed by dense setation, except dorsal fourth unsculptured and asetose, smooth. Lateral face of propodeum densely and coarsely foveolate punctate with interspersed micropunctures on posteroventral half. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 52:67:65:55:53. Lateral margin of mesonotum slightly emarginated anterior to propodeal spiracle, smoothly diverging anterad. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale present, well-developed, slightly narrower than and separated from well developed anterolateral carinae which are connected thus forming a single transverse carina with shallow emargination medially; scabrous intervals present on scutellar area. Posterior face of propodeum slightly longer than and poorly distinguished from dorsal face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 38:88:86. Disc of T2 densely and coarsely foveolate-punctate to punctate, sculpture denser and smaller mediad, sparser and shallower over integumental spots. T3–6, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate where visible. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, equally high throughout. S2 densely and coarsely foveolate-punctate, more sparserly so 80 posteromedially; anteromedial crest-fold vestigial. S3–6 densely and coarsely foveolate-punctate. Pygidial plate broadly subovate, defined by lateral carinae at apical half of plate; surface mostly irregularly longitudinally rugose, rugae vestigial laterally. MALE. Body length 15 mm. Head. Rounded subrectangular in dorsal view, posterolateral angles rounded. Vertex width 0.75 × pronotal width. Eye almost circular. Ocelli small; OOD 3.5 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface densely and coarsely foveolate-punctate to punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina narrowly separated from internal eye margin. Clypeus slightly concave laterally immediately below antennal insertion, convex medially; coarsely and densely foveolate-punctate to punctate; with a pair of short, sessile, blunt tubercles on apical margin. Scape bicarinate. Flagellomere 1.75 × pedicel length; flagellomere 2.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, rounded, broadly separated from humeral carina, anterolateral angles of pronotum subangulate. Anterior face of propodeum coarsely foveolate-punctate to punctate latearlly, unsculptured medially, with a conspicuous longitudinal slightly concave medial area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures anteriorly and along inner margin. Mesoscutum densely and coarsely foveolate- punctate, notaulus and parapsis indistinguishable. Scutellum convex, densely and coarsely foveolate-punctate. Axilla produced posterolaterally as truncate projection, with conspicuous flat posterior face, coarsely and densely foveolate- punctate dorsally; posterior face of axillar projection arched. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeum dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face predominantly densely areolate, areolations less defined anterad; posterolateral margins rounded, posterodorsal corners rounded in lateral view; dorsal and posterior faces indistinguishable. Lateral face of pronotum densely coarsely and confusedly foveolate-punctate to punctate with interspersed micropunctures; mesopleura with strong blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures anteriorly. Metapleuron sculpture concealed by dense setation. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate where visible, except pygidial plate on T7 irregularly rugose and weakly defined by parallel carinae apicolaterally, rugosities predominantly transverse. S1 longitudinally elevated medially, forming pronounced carina slightly higher posteriorly. S2 foveolate-punctate, foveolations conspicuously sparser and smaller mediad; with well-developed longitudinal anteromedial crest-fold ending on elongate and narrow longitudinal pit densely filled with setae. S3–7 sparsely foveolate-punctate to punctate. S7 slightly broader than long, with a pair of subacute closely spaced tooth-like projections on posterior margin. Genitalia. Parapenial lobe slightly pronounced apically. Ratios of free length of paramere, cuspis and digitus, 64:55:7; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense long setae ventrally except at apically with setae conspicuously reduced; cuspis narrow, slender, in dorsal view slightly curved inward at basal half and curved outward in apical half; conspicuously and smoothly widened apically; predominantly asetose with sparse conspicuous setae ventrally at apex; paracuspis well-developed, not sessile, twice as long as wide, roundly subtriangular at apical margin, densely setose, setae predominantly shorter than paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose dorsally; penis valve strongly concave on internal surface, with very closely spaced pair of teeth apicoventrally; apical tooth more acute and longer; subapical tooth acute, externolateral vestigial; apical distance between teeth 0.03 × length of valve; dense setae present along apical margin and at base of teeth on external surface. Coloration and variations. FEMALE. Integument predominantly black to reddish-black except mandibles basally and most flagellomeres ventrally reddish-brown. Setae predominantly silvery-white to silvery-golden varying in density except the following areas with black setae varying in density: gena, malar space, posterior margin of vertex, ventral surface of head, pronotum, mesonotal dorsum, propodeal dorsum medially, T1–2 medially, fringe of T2 medially, and fringe of T3–5 sublaterally; no conspicuous color or setae variations were observed in the specimens examined. MALE. Integument entirely black except most flagellomeres reddish-brown ventrally. Setae predominantly silvery-white varying 81 in density except the following areas with black setae varying in density: head predominantly, pronotum, mesopleuron above tubercle, mesonotum, axillae, scutellum, T2 posteromedially, fringe of T2–5 medially and T6–7; no conspicuous color or setae variations were observed in the specimens examined. Distribution. Brazil. Material examined. (3m# 15f#) Type material. Holotype of Traumatomutilla guarata, f#, BRAZIL, D.[istrito?] Federal [Rio de Janeiro], [Rio de Janeiro], Guaratiba, 10.III.1952, A.G.A. Silva (AMNH); Additional material. BRAZIL, 1f#, (BMNH); Bahia, Freitas, 1f#, 30.III.2004, Santos J.R.M. (CPDC); Espírito Santo, Linhares. Parque Sooretama, 1f#, V.1953, P.A. Teles (MNCN); Parque Sooretama, 3f#, XI.1967, Oliveira F.M. (DZUP); 2f#, 02.XI.1964, Seabra, Werner & Oliveira (DZUP); Linhares, 1f#, 18.I.1971, Alvarenga (DZUP); 1f#, XI.1965, Alvarenga (DZUP); 1f#, 20.XI.1971, Domingos A.C. (DZUP); Pinheiros, 1f#, 06.XI.1971, Lima J.M. (DZUP); Minas Gerais, nr[near] Timóteo, 1m#, 13– 20.X.1997, Eurico R. DePaula (EMUS); 1m#, 30.IX.1999, Eurico R. DePaula (EMUS); Nova Resende, 1f#, VII.1961, Elias C. (DZUP); Rio de Janeiro, 1m#, II.1912, (CASC); 1f#, II.1960, Silva A. (DZUP). Remarks. Mickel identified numerous specimens of this species in different collections as “T. speciosa sp. n.” and even labeled certain specimens as paratypes. This new species, however, was never published by the author and no mention of the name was ever made in published literature. Males herein associated with T. guarata, though almost structurally identical to the other males of the T. juvenilis species-group, can be reliably associated with the females based on distribution and their distinct setae pattern, which can be observed in both sexes of species typical of the Atlantic Forest such as Hoplocrates cephalotes (Swederus), Hoplomutilla spinosa (Swederus), Pappognatha patruelis (André), and Ephuta chrysodora (Perty) (KAW, pers. obs.).
Discussion
The geographical distribution of this species-group differs from the main bulk of Traumatomutilla, because there are no typical Amazonian species even with ample sampling in different parts of the region (PRB pers. obs.), even though there are a few records of T. juvenilis in the northeastern and southern fringes of the Amazonian forest, and T. juvenindica occurs in the Colombian Amazon. Furthermore, this entire species group seems to be significantly more abundant in the savannah-like dry areas of Southern and Midwestern South America such as the Cerrado, Chaco and Pampa. There is a peculiar similarity between males of the T. juvenilis group and males of the T. bifurca group regarding the posteroventral teeth of the penis valve, which are closely spaced in all males of both groups, and a character that appears to be exclusive to these groups within Traumatomutilla. The females, however, differ in many aspects, except for the genal carina which is absent to vestigial in T. bifurca (Klug, 1821), T. miniata and T. juvenindica. The sex associations of T. juvenindica and T. guarata are good examples of how color patterns and apparent mimetic syndromes can be a helpful tool to associate sexes when there are enough samples and when we know where to look. Traumatomutilla guarata was associated with the only males of the juvenilis group that fit the predominant pattern for the Atlantic Forest and we haven’t observed any intermediate specimens between the females of T. guarata and other species in the group. Indeed, we have found reliable structural characters to separate them effectively. Likewise, matching the sexes for T. juvenindica was logical because, not only are they the only male and female with the defining characteristics of the T. juvenilis group in Colombia, they match perfectly the color syndrome for each sex of most species in the area. Having a large genus such as Traumatomutilla divided into species-groups based on exclusive characters or combinations of characters has proven to be immensely useful for sex associations. Having one reliable sex association for the T. juvenilis group allowed us to have a morphological “ground plan” for the males thus drastically reducing the number of possible associations in a given area, especially when adding genitalia characters. That coupled with large series of males and females repeatedly collected in the same area allowed us to confidently deduce the remaining sex associations. Additionally, reducing the number of possible matches for a given male or female evidenced that there are structural parallels to be traced between sexes and these structures served as another important clue for sex associations. 82
We are confident that such characters will prove useful in associating sexes in future revisions within Traumatomutilla and indeed in other Mutillidae genera.
Acknowledgements
We are grateful for the collection managers and curators that provided specimens for this study, including: Christine LeBeau (AMNH), Robert Zuparko (CASC), Andreas Köhler (CESC), John Rawlins (CMNH), Stephan Blank (DEI), Gabriel Melo (DZUP), James Pitts (EMUS), Mercedes Paris (MNCN), Agniéle Touret-Alby (MNHN), Felipe Vivallo (MNRJ), Orlando Silveira (MPEG), Kelli Ramos (MZSP), Fernando Silvestre (MuBio-UFGD), Gavin Broad (BMNH), Lynn Kimsey (UCDC), Fernando Silveira (UFMG), Robin Thomson (UMSP), Brian Harris (USNM), Michael Ohl, Viola Richter and Lukas Kirscher (ZMB), Lars Vilhelmsen (ZMUC), Denis Brothers (DJBC), Donald Manley (DGMC), Wouter Dekoninck (RBINS) and Karla Schneider (MLUH). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9).
References
André. E. (1901) Descriptions de quelques espèces et variétés nouvelles de Mutilles d’Amerique appartenant au Museé Civique de Gênes. Zeitschrift für Hymenopterologie und Dipterologie. 1: 257–264.André, 1902 André, E. (1902) Fam. Mutillidae. In: Wytsman, P. (Ed.), Genera Insectorum, Bruxelles, Fasc. 11, pp. 1–77. André, E. (1904) Examen critique d’une nouvelle classification proposée par M. le Dr. W. H. Ashmead pour le famille des Mutillides. Revue d’Entomologie. 23(1): 27–41. André, E. (1908a) Une nouvelle espèce de Mutillide de la Republique Argentine [Hym.]. Bulletin de la Société Entomologique de France, 230–231. André, E. (1908b) Descriptions de quelques nouveaux Mutillides du Museé National d’Hongrie. – Annales Musei Nationalis Hungarici, 6, 378–383. André, E. (1910) Liste des Mutillides recueillis par M. le Prof. J. D. Anisits au Paraguay. Zoologische Jahrbücher. Abteilung für Systematik, 29, 229–230. Bartholomay, P.R. Williams, K.A. Luz, D.R. & Oliveira, M.L. (2018) New species of Traumatomutilla André in the T. tabapua and T. integella species-groups (Hymenoptera, Mutillidae). Zootaxa 4433 (2): 361–385. Bartholomay, P.R., Williams, K.A., Lopez, V.M. & Oliveira, M.L. (2019) Revision of the Traumatomutilla americana species-group (Hymenoptera: Mutillidae). Zootaxa, 4608 (1): 001–034. Brothers, D.J. (2006) Capítulo 14.4. Familia Mutillidae. In: Hanson, P.E. e Gauld, I.D. (Eds), Hymenoptera de la Región Neotropical. The American Entomological Institute, Gainesville, FL, pp. 586–593. Burmeister, H.C.C. (1875) Mutillae Argentinae. Boletín de la Academia Nacional de Ciencias Exactas existente en la Universidad de Cordova 1: 461–502. Casal, O.H. (1969) Sobre Traumatomutilla André (Hymenoptera, Mutillidae). – Physis, 28 (77), 279–298. Cresson, E.T. (1902) Descriptions of some Brazilian Mutilla. – Transactions of the American Entomological Society, 28, 1–82. Cresson, E.T. (1916) The Cresson types of Hymenoptera. Memoirs of the American Entomological Society, 1: 79–85. 83
Gerstaecker, A. (1874) Mutillarum Americae meridionalis indigenarum synopsis systematica et synonymica. Archiv für Naturgeschichte, 40, 41-77, 299–328. Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in Entomology 28: 1–31. Klug, J.C.F. (1821) Entomologiae brasilianae specimen. – Nova Acta Academica Caesareae Leopoldino-Carolinae, 10 (2), 305–324. Luz, D.R. & Williams, K.A. (2014) The first sexual associations in the genus Darditilla Casal, 1964 (Hymenoptera, Mutillidae). Zookeys 454: 41–68. Luz D.R., Williams, K.A. & Bartholomay, P.R. (2016) The mutillid wasps of the Dasymutilla paradoxa species-group (Hymenoptera: Mutillidae). Zootaxa, 4193 (2): 361–372. Nonveiller G. (1990) Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta, Hymenoptera). Hymenopterorum Catalogus (Nova Editio), 18. Den Haag, SPB Academic Publishing. 1–150. Williams, K.A., Manley, D.G., Pilgrim, E.K., von Dohlen, C.D. & Pitts, J.P. (2011a) Multifaceted assessment of species validity in the Dasymutilla bioculata species group (Hymenoptera: Mutillidae). – Systematic Entomology, 36, 180– 191. Williams, K.A., Brothers, D.J. & Pitts, J.P. (2011b) New species of Tobantilla Casal, 1965 and a new genus and species, Gogoltilla chichikovi gen. et sp. nov., from Argentina (Hymenoptera: Mutillidae). – Zootaxa 3064, 41–68. Williams, K.A., Manley, D.G., Deyrup, M., von Dohlen, C. & Pitts, J.P. (2012) Systematic review of the Dasymutilla monticola species-group (Hymenoptera: Mutillidae): using phylogenetics to address species-group placement and sex associations. – Zootaxa 3554, 1–29. Williams, K.A., Bartholomay, P.R. & Oliveira, M.L. (2017) Species groups of Traumatomutilla André (Hymenoptera: Mutillidae). Insecta Mundi, 0533: 1–33. Wilson, J.S., Williams, K.A., Forister, M.L., von Dohlen, C.D. & Pitts, J.P. (2012) Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications, 3: 1272. Wilson, J.S., Jahner, J.P., Williams, K.A. & Forister, M.L. (2013) Ecological and Evolutionary Processes Drive the Origin and Maintenance of Imperfect Mimicry. PlosONE, 8 (4), 1–7. Wilson, J.S., Pan, A.D, Limb, E.S. & Williams, K.A. (2018) Comparison of African and North American velvet ant mimicry complexes: Another example of Africa as the ‘odd man ou’. 13(1), 1–15.
Figure legends.
Figure 1–10. 1–6. Traumatomutilla duplicata, females, dorsal habitus, lines 2mm; 1. Traumatomutilla duplicata (Gerstaecker, 1874), lectotype; 2. Traumatomutilla bivittata (Gerstaecker, 1874), holotype; 3. Traumatomutilla bruchi André, 1908, lectotype; 4. Traumatomutilla duplicate feia Casal, 1969, holotype; 5. Traumatomutilla bispiculata (nec #m), allotype; 6. Traumatomutilla duplicate, eastern slope of Bolivian Andes color form; 7–10. Type labels.
Figures 11–16. Traumatomutilla duplicata (Gerstaecker, 1874), females, lateral habitus, lines 2mm; 11. Traumatomutilla duplicata (Gerstaecker, 1874), lectotype; 12. Traumatomutilla bivittata (Gerstaecker, 1874), holotype; 13. Traumatomutilla bruchi André, 1908, lectotype; 14. Traumatomutilla duplicate feia Casal, 1969, holotype; 15. Traumatomutilla bispiculata (nec #m), allotype; 16. Traumatomutilla duplicate, eastern slope of Bolivian Andes color form.
Figures 17–28. 17–20. Traumatomutilla duplicata (Gerstaecker, 1874), males, lateral habitus; 17. Traumatomutilla cristata (Gerstaecker, 1874), holotype, line 5mm; 18. Traumatomutilla aterrima (Gerstaecker, 1874), lectotype, line 5mm; 19. Traumatomutilla lorena (Cresson, 1902), holotype, line 2mm; 20. Traumatomutilla duplicata (Gerstaecker, 1874), color form, Argentinian xeric áreas, line 5mm; 21–23. Type labels; 24–28. Traumatomutilla cristata (Gerstaecker, 1874), holotype, genitalia; 24. Dorsal view (halved); 25. Ventral view (halved); 26. Lateral/inner view (halved, penis valve 84 removed); 27. Cuspis and paracuspis, lateral/inner view (removed, not to scale); 28. Penis valve, lateral/outer view (removed, not to scale).
Figures 29–39. 29–30. Traumatomutilla juvenilis (Gerstacker, 1874), female, lectotype, dorsal and lateral habitus, line 2mm; 31– 32. Traumatomutilla immaculiceps (André, 1901), female, lectotype, dorsal and lateral habitus, line 2mm; 33– 34. Traumatomutilla rubroguttata (André, 1901), female, reference material from André collection (MNHN), line 2mm; 35–36. Traumatomutilla Estrella (Cresson, 1902), female, lectotype, line 2mm; 37–39. Type labels.
Figures 40–48. Traumatomutilla juvenilis (Gerstaecker, 1874), male; 40. 41. Lateral habitus, line 5mm; 42. Mesosoma, dorsal view; T7, posterodorsal view; 43. Hypopygium, ventral view; 44–48. Genitalia; 44. Dorsal view (halved); 45. Ventral view (halved); 46. Lateral/inner view (halve, penis valve removed); 47. Cuspis and paracuspis, lateral/inner view (removed, not to scale); 48. Penis valve, lateral/outer view (removed, not to scale).
Figures 49–50. Traumatomutilla juvenindica Bartholomay & Williams sp. nov., female, holotype, lines 2mm; 49. Dorsal habitus; 50. Lateral habitus.
Figures 51–56. Traumatomutilla juvenindica Bartholomay & Williams sp. nov., male, allotype, lines 5mm; 51. Lateral habitus; 52–56. Genitalia; 52. Dorsal view (halved); 53. Ventral view (halved); 54. Lateral/inner view (halve, penis valve removed); 55. Cuspis and paracuspis, lateral/inner view (removed, not to scale); 56. Penis valve, lateral/outer view (removed, not to scale).
Figures 57–59. Traumatomutilla miniata (Gerstaecker, 1874), female, lectotype; 57–58. Dorsal and lateral habits, line 2mm; 59. Type labels.
Figures 60–68. Traumatomutilla bispiculata (André, 1907), holotype; 60. Lateral habitus, line 2mm; 61. Type labels; 62. Mesosoma, dorsal view; 63. T7, posterodorsal view; 64–68. Genitalia; 64. Dorsal view (halved); 65. Ventral view (halved); 66. Lateral/inner view (halve, penis valve removed); 67. Cuspis and paracuspis, lateral/inner view (removed, not to scale); 68. Penis valve, lateral/outer view (removed, not to scale).
Figures 70–72. Traumatomutilla guarata Casal, 1969, female, holotype; 70–71. Dorsal and lateral habitus, lines 2mm; 72. Type labels.
Figures 73–78. Traumatomutilla guarata Casal, 1969, male;73. Lateral habitus, line 4mm; 74–78. Genitalia; 74. Dorsal view (halved); 75. Ventral view (halved); 76. Lateral/inner view (halve, penis valve removed); 77. Cuspis and paracuspis, lateral/inner view (removed, not to scale); 78. Penis valve, lateral/outer view (removed, not to scale).
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CAPÍTULO 4.8 - (formatado para submissão à ZOOTAXA)
The Traumatomutilla gemella species group (Hymenoptera, Mutillidae)
PEDRO R. BARTHOLOMAY1’*, KEVIN A. WILLIAMS2, ROBERTO A. CAMBRA3 & MARCIO L. OLIVEIRA1 1Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, Laboratório de Hymenoptera, Av. André Araújo, 2936, Manaus, Amazonas, Brazil. 2Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento CA 95832, USA. 3Museo de Invertebrados G. B. Fairchild, Universidad de Panamá, Panamá 0824, Panamá *Corresponding author, e-mail: [email protected]
Abstract
Traumatomutilla angustata André, 1906 and T. rastra Casal, 1969 are proposed as junior synonyms of T. diophthalma (Klug, 1821). The males of T. diopthalma, T. chuza and T. gemella are described and illustrated. All females are redescribed and a new species T. peismatara Bartholomay & Cambra, sp. nov., is described based on females from Peru and males from Acre state in Brazil. Additionally, two host records are provided for T. diophthalma based on a laboratory experiment and trap nests.
Key words: Sphaeropthalminae, Dasymutillini, velvet ants, Neotropics, South America
Introduction
Velvet ants (Mutillidae) are solitary wasps whose larvae act as parasitoids of encapsulated immature of other insects, especially other solitary Hymenoptera (Brothers, 2006). Females of this family are always wingless whilst the males are usually fully winged, rarely brachypterous or apterous (Brothers, 2006). This remarkable sexual dimorphism has resulted in many species being described based only on males or females and usually differentiated solely on color and setae characters (Gerstaecker, 1874; Cresson, 1902; Casal, 1969). The reliability of such characters alone for differentiating species in Traumatomutilla André, 1901 was brought into question by research on the closely related Dasymutilla Ashmead, which demonstrated the existence of Müllerian mimicry complexes within these wasps (Williams et al. 2011, 2012; Wilson et al. 2012, 2013). The influence of such mimicry complexes in the coloration of these wasps is such that several species once thought to be distinct have been proven to be a single widespread and highly variable species (Williams et al. 2011, 2012). Similar results have already been found in recent revisions of other Traumatomutilla species-groups (Bartholomay et al. 2019). These revisions were facilitated by Williams et al. (2017), which organized the 135 species of Traumatomutilla based on females into 14 species-groups based on shared structural characters or shared combinations of structural characters. With only six known species, the Traumatomutilla gemella species-groups is one of the smallest, rarest, and most morphologically distinct species-groups within Traumatomutilla (Williams et al. 2017). In this study we review and provide the first sex associations for the T. gemella species-group, as well as two novel host records for the genus.
Materials and terminology
Approximately 100 specimens were studied in the current work and the following codens are used for institutions housing the material discussed:
AMNH - American Museum of Natural History, New York, New York, USA. 89
CESC - Coleção Entomológica da Universidade de Santa Cruz do Sul, Santa Cruz do Sul, Rio Grande do Sul, Brazil. EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA. IAvH - Insituto Alexander Von Humboldt, Villa de Leyva, Boyacá, Colombia INPA - Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil. HBMNH - Hungarian Natural History Museum, Budapest, Hungary. MIUP - Museo de Invertebrados G.B. Fairchild, Panamá, Panamá MNCN - Museo Nacional de Ciencias Naturales, Madrid, España MNHN - Museum Nationale d’Histoire Naturelle, Paris, France MPEG - Museu Paraense Emílio Goeldi, Belém, Pará, Brasil MZSP - Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo, Brazil UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brazil UMMZ - University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA
General morphological terminology, definitions, abbreviations, and measurements followed that of Cambra et al. (2016) and Bartholomay et al. (2018, 2019). Additionally, sculpture terminology follows Harris (1979). In the description and redescription sections we refrain from mentioning coloration and setal patterns due to the highly variable nature of these characters in Traumatomutilla. Instead, we provide a separate section titled Coloration and variations, in which we provide an overall description of the known color and setae patterns for a particular species. In the material examined section abbreviations, acronyms and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of the material examined section.
Taxonomy
Traumatomutilla gemella species-group
Diagnosis. FEMALES. Females of this species group can be distinguished by a single autapomorphy: the pygidial plate with projected flange-like lateral carina restricted to apical fourth. The following combination of characters is unique to the T. gemella group females, though some characters may occur in other groups as well: body generally slender, elongate; head unarmed posterolaterally; clypeus shallowly but conspicuously bilobate on apical/ventral margin, longitudinally elevated medially; vertex and/or front frequently with medial longitudinal carina; pronotal collar with vestigial transverse rugosities anteriorly; anterior face of pronotum short, shorter than or as long as pronotal collar; lateral face of pronotum with subacute tubercle anteroventral in relation to pronotal spiracle; pronotal and propodeal spiracles virtually flat against lateral margin of mesosoma; mesosoma virtually straight laterally, at most slightly divergent anterad, not constricted anterior to propodeal spiracle; dorsal face of propodeum much longer than posterior face; scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutelar area; and apex of middle and hind femora rounded. MALES. The males can be distinguished by a unique genitalic autapomorphy: the cuspis is short, abruptly angled dorsally at midlength, with long twisted setae along the ventral surface of the apical half. The following combination of characters is unique to the T. gemella group males, though some characters may occur in other groups as well: integument black to reddish-black with contrasting black and silvery-white to silvery-golden setae patterns varying in density; head transversely subrectangular in dorsal view; parapsis and notauli vestigial, restricted to posterior margin of mesoscutum; axillae pronounced as twisted oblique and acute projections; scutellum gibbose, without posterior transverse carina, frequently with weak medial longitudinal carina anteriorly; mesopleuron simply swollen on dorsal half, without any projections or tubercles; meso and metafemora rounded apically; S2 without setae filled pit; pygidial plate strongly 90 concave apically, apical margin strongly deflected upward; hypopygium rectangular, longer than broad, lateroapical corners angulate and slightly projected. Included taxa. Traumatomutilla gemella André, 1906, Traumatomutilla andrei (Cresson, 1902), Traumatomutilla chuza Casal, 1969, Traumatomutilla diophthalma (Klug, 1821), and Traumatomutilla pucallpai Cambra, Bartholomay & Williams sp. nov. Distribution. Panama and South America (except Chile). Remarks. At first glance, females of the T. gemella species-group are remarkably similar to those of the Dasymutilla paradoxa species-group recently revised by Luz et al. (2016). They differ in the presence of flange like projections apicolaterally in the pygidial plate, absent in the D. paradoxa species-group, and the sub-petiolate T1 shape, which is globose, subcylindrical in the D. paradoxa species-group. Additionally, females of the T. gemella species-group have only one pair of integumental spots on T2 whilst those of the D. paradoxa species-group have two pairs. Males of the T. gemella species-group can be easily separated by their axilla produced posteriorly as short acute projections and the integument of T2 all black in contrast with the axillar projections connected with the lateral margins of the mesoscutellum and T2 predominantly yellowish in males of the D. paradoxa species-group. Within the other Traumatomutilla species- groups, there are no females that can be easily confused with those of the T. gemella species-group. There are males of the T. indica group that, as with the males of the T. gemella group, have the axillar projections acute and the mesopleuron unarmed. These, however, can be readily recognized by having a setae-filled pit on S2 and/or a relatively straight cuspis that lacks sinuous long setae.
Key to females of the Traumatomutilla gemella species group
1. T2 with a pair of sublateral longitudinal yellow integumental stripes arched outward at posterior third; Cerrado, Caatinga, and Restinga (Brazil) species … T. andrei (Cresson, 1902) T2 with a pair of subcircular yellowish to reddish integumental spots … 2
2. Lateral face of propodeum with micropunctures on anterodorsal third; posteroventral half of mesopleuron, metapleuron and anterodorsal third of lateral face of propodeum concealed by dense appressed silvery-golden setae; head setae completely black; Atlantic Forest (Brazil) species … T. gemella André, 1906 Lateral face of propodeum virtually unsculptured, smooth and shinning on anterodorsal third, only with sparse scattered erect silvery-white setae; mesopleuron and metapleuron clothed with silvery-white setae, dorsal fourth of metapleuron unsculptured, smooth and shinning; head setae predominantly silvery-white to silvery-golden; if head setae predominantly black, then mesosomal dorsum also with silvery-white setae restricted to narrow vestigial stripes on propodeum … 3
3. Propodeal dorsum convex throughout, not conspicuously elevated anywhere; lateral face of propodeum with sparse sculpture, predominantly unsculptured, smooth and shinning, intervals many times wider than surrounding sculpture … T. chuza Casal, 1969 Propodeal dorsum conspicuously elevated medially on posterior margin, dorsal face sharply angulate into posterior face in lateral view; if mesosoma simply convex, not angulate in lateral view, then lateral face of propodeum more or less homogenously sculptured, with intervals always narrower than surrounding sculpture … 4
4. Dorsal face of propodeum almost flat, horizontal in lateral view; head setae entirely silvery-white varying in density … T. peismatara Bartholomay & Cambra sp. nov. Dorsal face of propodeum sloping posterad, not flat, convex; at least ventral half of frons, gena and malar space with black setae … T. diophthalma (Klug, 1821)
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Key to males of the Traumatomutilla gemella species-group:
1. Pronotal dorsum predominantly concealed by dense setae, coarsely and confusedly areolate-punctate to foveolate- punctate with conspicuous interspersed micropunctures where visible; pronotum clothed exclusively with silvery-white setae throughout … T. chuza Casal, 1969 Pronotal dorsum with mostly sparse, erect setae, coarsely and confusedly areolate-punctate to foveolate-punctate; clothed predominantly with black setae, at most with silvery-white setae medially on anterior margin … 2
2. Lateral face of propodeum and metapleuron with conspicuous silvery-golden setae throughout; head clothed almost exclusively with black setae; body setae with an overall silvery-golden tone … T. gemella (André, 1906) Lateral face of propodeum and metapleuron at most with sparse scattered and erect silvery-white setae, metapleuron variable; head predominantly clothed with silvery-white setae, black setae restricted to ocellar area and/or most of vertex; body setae with an overall silvery-white tone … 3
3. Posterior margin of hypopygium projected medially and laterally with medial projection simply convex and shorter than lateral projections; S2 with interspersed micropunctures anterolaterally; propodeal dorsum at most with sparse scattered and erect silvery-white setae anterolaterally … T. peismatara Bartholomay & Cambra sp. nov. Posterior margin of hypopygium projected medially and laterally with medial projection blunt and longer than lateral projections; S2 simply foveolate-punctate throughout, without apparent interspersed micropunctures; propodeal dorsum densely clothed with appressed silvery-white setae anterolaterally … T. diophthalma (Klug, 1821)
Traumatomutilla andrei (Cresson, 1902) (Figs 1–3)
Muitlla andrei Cresson, 1902: 55, holotype (by monotypy), f#, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), type examined. Ephuta (Traumatomutilla) andrei: André, 1902: 56. Traumatomutilla andrei: André, 1904: 40.
Diagnosis. FEMALE. T2 marked with a pair of narrow longitudinal yellowish stripes abruptly curved outward posteriorly in dorsal view. MALE. Unknown. Description. FEMALE. Body length 12 mm. Head. Posterior margin virtually straight. Occipital carina conspicuously swollen and smoothly curved dorsolaterally. Vertex width 0.9 × pronotal width. Eye almost circular, its length in frontal view 1.1 × distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate, intervals aligned so as to form vestigial longitudinal carina medially starting at middle of front and extending into vertex. Mandible with conspicuous subapical tooth. Dorsal scrobal carina well-defined, separated from antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercle finely and irregularly rugose. Flagellomere 1 2.2 × pedicel length; flagellomere 2 2.0 × pedicel length. Mesosoma. Dorsal thoracic length slightly shorter than thoracic width. Mesosomal dorsum densely and coarsely areolate-punctate to foveolate-punctate, with conspicuous medial longitudinal carinae extending from anterior margin of mesonotum to posterior margin of dorsal face of propodeum; carina less defined on propodeum, sinuous, irregular; integument adjacent to longitudinal carina on mesonotum devoid of areolations, simply densely punctate. Anterior face of propodeum defined, short, as long as pronotal collar, vestigially and coarsely striated longitudinally with interspersed scattered punctures; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from well-defined raised, subangulate epaulet, anterolateral corners of pronotum subangulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with sparse interspersed micropunctures, except for blunt 92 unsculptured tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture mostly concealed by dense setation, except dorsal third asetose, smooth, unsculptured. Lateral face of propodeum with densely, coarsely and homogeneously areolate-punctate throughout. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 73:86:67:68:64. Lateral margin of mesonotum simply divergent anterior to propodeal spiracle, slightly diverging anterad. Propodeal spiracle virtually flat against lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum conspicuously elongate, dorsal face much longer than and well differentiated from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 30:73:75. Disc of T2 densely and coarsely foveolate-punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots; foveolations less defined to virtually absent posteromedially. T3–5 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with dense interspersed micropunctures where visible; T6 densely and coarsely foveolate-punctate. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, conspicuously higher anteriorly. S2 sparsely foveolate-punctate, foveolations sparser and smaller anterad and mediad; anteromedial crest-fold vestigial. S3–4 densely and coarsely foveolate-punctate with sparse micropunctures; S5–6 densely and coarsely foveolate-punctate. Pygidial plate broadly subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface mostly irregularly and confusedly rugose on apical half, interstice apparently granulose; basal half of pygidial plate simply granulose. Coloration and variations. FEMALE. Integument black, except mandibles and antennal flagellomeres reddish-brown ventrally, and T2 with a pair of longitudinal yellowish integumental stripes abruptly curved outward at posterior third. Body setae predominantly silvery-white varying in density except for the following areas with black setae varying in density: head (except ventral surface), mesosomal dorsum medially, dorsal hald of lateral face of propodeum and mesopleuron, femora apicodorsally, T1 medially, disc of T2 (except integumental stripes), fringe of T2–4 sublaterally, fringe of T5–6 medially, fringe of S4, and S5–6. MALE. Unknown. Distribution. Brazil (Bahia, Maranhão, Mato Grosso). Material examined. (3f#) Type material. Holotype, f#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. BRAZIL: Maranhão, São Luis, Floresta Sacavem, 1f#, 30.IX.1992, (CAEMA) R. Cambra & D. Quintero (MIUP); Bahia, Portello Machado [Machado Portela], 1f#, 19.VI.2015 (USNM). Remarks. The unique coloration of this species is, presently, the most reliable means of distinguishing it from other species in the group. This pattern (Fig. 1–2), with longitudinal yellowish integumental stripes on T2, though rare, it is not exclusive to T. andrei. It is also known in T. rectilineata (André, 1898) — T. trochanterata species-group — and T. ipanema (Cresson, 1902) — T. inermis species-group — which also occur in Chapada do Guimarães. Although there are no consistent structural characters to separate T. andrei from its relatives in the T. gemella species-group, no variations or intermediate color forms has been recorded so far. The rarity of this species contrasts with its apparent wide distribution in Brazil, spanning from Mato Grosso in the Midwest, to Bahia in the East and Maranhão in the Northeast. No males of the T. gemella species-group have been recorded from the same localities as the three females examined despite the fact that many of these areas are relatively well sampled in Brazil (PRB pers. obs.). This seems to be further evidence for the rarity of this species-group.
Traumatomutilla chuza Casal, 1969 (Figs. 4–15)
Traumatomutilla chuza Casal, 1969: 286, Holotype, f#, Brazil, Pará, Óbidos (AMNH), type examined.
Diagnosis. FEMALE. Propodeal dorsum evenly convex throughout, dorsal face smoothly angulate into posterior face in lateral view; lateral face of propodeum predominantly unsculptured, smooth, shinning, interval always wider than 93 surrounding sculpture. MALE. Pronotal dorsum with densely, coarsely, and confusedly areolate-punctate to foveolate- punctate with conspicuous interspersed micropunctures; sculpture of pronotum predominantly concealed by dense silvery- white setation. Description. FEMALE. Body length 10–12 mm. Head. Posterior margin virtually straight. Occipital carina conspicuously equally wide throughout. Vertex width 0.9 × pronotal width. Eye almost circular, its length in frontal view 1.5 × the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate with smooth rounded intervals. Mandible with conspicuous subapical tooth. Dorsal scrobal carina present, connected to lateral scrobal carina. Antennal tubercle finely, sparsely, and irregularly rugose to micropunctate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.3 × pedicel length. Mesosoma. Dorsal thoracic length 0.9 × width. Mesosomal dorsum densely and coarsely areolate-punctate laterad to foveolate-punctate mediad, with smooth rounded intervals. Anterior face of propodeum defined, short, shorter than pronotal collar, vestigially and coarsely striated longitudinally with interspersed scattered punctures; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, separated from well-defined raised sharp epaulet, anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Lateral face of pronotum sparsely foveolate-punctate, except for smooth blunt unsculptured tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture, micropunctate anteriorly and very sparsely and vestigially punctate along mesopleural ridge; metapleuron sculpture virtually completely concealed by dense setation, except dorsal fourth smooth, unsculptured. Lateral face of propodeum with sculpture sparsely foveolate-punctate, intervals smooth and shinning predominantly more than twice the width of surrounding sculpture. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 93:100:105:84:79. Lateral margin of mesonotum vestigially constricted anterior to propodeal spiracle, slightly diverging anterad. Propodeal spiracle vestigially pronounced from lateral margin of mesosoma; post-spiracular area present. Scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum conspicuously elongate, dorsal face much longer than and well differentiated from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 65:125:124. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, sparsely and coarsely foveolate-punctate to simply punctate with dense interspersed micropunctures where visible. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, equally high throughout. S2 densely foveolate-punctate, punctures slightly sparser posterad; anteromedial crest-fold vestigial. S3– 6 densely and coarsely foveolate-punctate with sparse micropunctures at S3–4; sculpture denser on S6. Pygidial plate broadly subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface mostly irregularly longitudinally rugose; interstice apparently granulose. MALE. Body length 10–12 mm. Head. Transversely subrectangular with posterolateral angles rounded and continuous with outline of eyes in dorsal view. Vertex width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 5.0 × DLO, IOD slightly longer than DLO. Occipital carina distinct. Head surface densely and coarsely foveolate-punctate with dense interspersed micropunctures on gena, malar space, and posterior half of vertex; Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; predominantly obscured by dense setation, coarsely and densely punctate to micropunctate where visible; apical/ventral margin with a pair of closely space short blunt inconspicuous tooth-like projections medially. Scape bicarinate. Flagellomere 1 1.6 × pedicel length; flagellomere 2 2.0 × pedicel length. Mandible obliquely tridentate apically, inner and middle teeth virtually equal, greatly reduced; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, subangulate, broadly separated from humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate to micropunctate throughout along dorsal margin, virtually unsculptured, smooth, and shinning elsewhere; virtually flat throughout. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on anterior and inner margin. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior third of mesoscutum; with medial longitudinal carina on posterior half. Scutellum convex, subglobose, with somewhat definable dorsal and posterior aces; densely and 94 coarsely areolate-punctate to foveolate-punctate; with longitudinal carina medially formed by aligned intervals on dorsal face. Axilla produced posterolaterally as acute projections, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface, except apically and along outer margin unsculptured. Metanotum virtually equally wide throughout, its surface obscured by dense setation. Propodeal dorsum convex, predominantly concealed by dense setation, densely areolate where visible; lateral face densely areolate on posterior half, vestigially areolate on anterior half; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and coarsely foveolate- punctate with interspersed dense micropunctures; mesopleuron slightly swollen on dorsal half, without any or projections; sculpture densely and coarsely foveolate-punctate with interspersed micropucntures; foveolations sparser anterad and posterad. Metapleuron predominantly micropunctured to unsculptured, smooth and shinning, except for vestigial foveolations and rugosities on dorsal and ventral fifths. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells; dark brown, slightly but conspicuously lighter on basal third. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. Ratios of width of T1, width of T2 and length of T2 45:76:58. Dorsal metasomal sculpture partially concealed by dense setation, sparsely and coarsely punctate with interspersed micropunctures where visible; pygidial plate somewhat concave, posterior margin conspicuously curved upward; surface predominantly smooth, shinning, with vestigial undefined sculpture along apical margin; weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, ending in blunt, low, concave carina. S2 sparsely and finely foveolate-punctate to punctate; foveolations conspicuously sparser posterad; anteromedial crest-fold virtually absent; sternal pit absent. S3 sparsely and finely punctate with sparse interspersed micropunctures; S4–7 sparsely foveolate-punctate. S7 longer than broad, with conspicuous medial longitudinal unsculptured area; posterior margin projected apicolaterally and medially; medial projection blunt, longer than lateral projections. Genitalia. Parapenial lobe not at all pronounced apically, simply rounded. Ratios of free length of paramere, cuspis and digitus, 56:28:13; paramere slightly sinuous in dorsal view, upcurved apically in lateral view; with dense setae ventrally at basal half; cuspis short, stout, slightly swollen medially and narrower apicad in lateral view; narrower apicad and virtually straight in dorsal view; abruptly curved dorsally in wide angle at basal third; with dense conspicuous, strongly sinuous setae on ventral surface, except at basal third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis well-developed, not sessile, slightly elongate vertically, subrounded at apical margin, densely setose along posterior margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, inconspicuously setose basodorsally; penis valve strongly concave on inner surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, with lateral pocket present on outer surface; apical distance between teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface. Coloration and variations. FEMALE. Integument black to brownish-black except for mandibles and antennal flagellomeres partially reddish-brown, and T2 with a pair of large subcircular orange integumental spots. Body setae predominantly silvery-white except for the following areas with black setae varying in density: pronotal dorsum, anterior half of mesonotum, posterior half of mesonotum medially, propodeal dorsum medially, femora apicodorsally, T1 posteromedially, disc of T2 (except over integumental spots), fringe of T2–3 medially (except inconspicuous patch of silvery-white medially), T4 sublaterally, T5–6 (except pygidial plate) laterally, fringe of S5, and S6. Some specimens may have the head completely covered with black setae (except gena and malar space), and the silvery-white setae of the mesosomal dorsum restricted to narrow inconspicuous lateral stripes on propodeum. Such specimens have the remaining silvery-white setae areas conspicuously less defined and the integumental spots of T2 have a more reddish tone rather than the usual orange. MALE. Integument black. Body setae predominantly silvery-white varying in density except for the following areas with black setae varying in density: mesoscutum, tegula, axillar projections, mesoscutellum, disc of T2 (except anterior third), fringe of T2–4 medially, T5–7, fringe of S5–6, and S7. Distribution. French Guiana, Colombia (Amazonas) and Brazil (Amapá, Amazonas, Pará, Rondônia), Ecuador, and Bolivia. 95
M aterial examined. (7f#, 21m#) Type material. Holotype, f#, BRAZIL, Pará, Óbidos (AMNH). Additional material. FRENCH GUIANA, [Cayenne]: Bélizon, 1#m, XI.2015, J. Giuglaris (MIUP); Maititi, 1#m, V.2015, J. Giuglaris (MIUP); COLOMBIA: 2#m (IAvH); Amazonas: Leticia, 1f#, 14–15.VIII.1974, M. Cooper (BMNH); La Chorrera, 1f#, 02.VIII.1976, M. Cooper (BMNH); Bolivar, Zambrano, 1#m, F. Fernandez (AMNH); BRAZIL:Amazonas: Lago Amana, 1f#, 01.XI.1980, R. Best (MIUP); Manaus: Vicinal ZF2 [Zona Franca], Km 34, trilha do igarapé em frente a base do Km34, 1f#, 22.VI.2012, J.A. Rafael (INPA); Campus Universitário, 1f#, 28.VII.1979 (INPA); EMBRAPA [Empresa Brasileira de Pesquisa Agropecuária] Amazônia Ocidental, 2#m, 2012, K. Schoeninger (INPA); Novo Airão, 02°48'58''S 60°55'18''W, AM352, Km68, 1#m, 14–28.X.2016, J.A. Rafael & F.F. Xavier-Filho (INPA); Pará, Serra Norte: Mina de Manganês, Rio Azul, 1f#, 27.VII.1983, W.L. Overal (MPEG); Estrada Manganês, 1f#, 15.V.1984, M.F. Torres (MPEG); Melgaço, ECFPn [Estação Científica Ferreira Pena] Caxiuanã, Mata da Sede, 1#m, 16.XI.1998, O. Silveia & J. Pena (MPEG); Amapá, Mazagão, Fazendinha, 1f#, 03.XII.1980, E.L. Oliveira (MPEG); Rondônia: 62 km SE [kilometers southeast of] Ariquemes: 3#m, 08.XI.1994, W.J. Hanson (EMUS); 2#m, 01.XI.1997, B.K. Dozier (EMUS); 1#m, 22.X.1997, W.J. Hanson (EMUS); 2#m, 01.XI.1997, B.K. Dozier (FSCA); ECUADOR, Sucumbios, Rio Napo, Sacha Lodge, 1#m, 14.III.1994 (EMUS); BOLIVIA, Beni: Romansos [sic!], 1km N [kilometer North of] Rio Iteneze & Rio Paragua [confluence?], 1#m, 30.VII.1964, Bouseman & Lussenhop (AMNH); Rio Itenez near Costa Marques (Brazil), 1#m, 01.IX.1964, Bouseman & Lussenhop (AMNH); Santa Cruz, Buena Vista, 1#m, 26.II.1999, F.D. Parker (EMUS); Remarks. Traumatomutilla chuza is apparently the typical representative of the T. gemella species-group in the Amazon. Two other species of this group are known in the Amazon, namely T. diophthalma in transition areas between Amazon Forest and Cerrado in the south and northeast, and T. peismatara in the western Amazon near the border between Brazil and Peru, but they are not as commonly encountered in the Amazon as T. chuza. Whilst there are records of T. diophthalma occurring in the same area as T. chuza (Pará, Brazil), there are no specimens of T. chuza known from West or South of Leticia (Amazonas, Colombia). Apart from the characters mentioned in the diagnosis, the females of T. chuza are easily recognizable by their more robust and stout body in comparison to the rather slender and elongate body of the other species within the T. gemella species-group. Casal’s original description for T. chuza was based entirely on color and setae patterns that, in light of the discovery of specimens collected in Rondônia, Brazil (DZUP), serves as a good example of how misleading these characters alone can be. The specimens from Itapuã d’Oeste (Fig. 15), though structurally identical to T. chuza, show a remarkable convergence in color and setae with T. barathra Bartholomay & Williams 2018 and T. luscoides André, 1908, which were also collected in the same area (Bartholomay et al. 2018).
Traumatomutilla diophthalma (Klug, 1821) (Figs. 16–28)
Mutilla diophthalma Klug, 1821: 318, holotype, f#, Brazil, Bahia (ZMB), type examined. Ephuta (Traumatomutilla) diophthalma: André, 1902: 55. Traumatomutilla diophthalma: André, 1904: 40. Ephuta (Traumatomutilla) angustata André, 1906: 43, Lectotype [designated here], f#, Brazil, Rio Grande do Sul (BMNH), type examined, syn. nov. Traumatomutilla angustata: Casal, 1969: 286. Traumatomutilla rastra Casal, 1969: 318, holotype f#, Brazil, Santa Catarina, Nova Teutonia (AMNH), type examined, syn. nov. Traumatomutilla diopthalma: Williams et al., 2017: 8 (misspelling).
Diagnosis. FEMALE. T2 with subcircular pair of integumental spots; lateral face of propodeum usually with sculpture sparse anterodorsad, lacking micropunctures; dorsal face of propodeum sloping posterad, conspicuously elevated posteromedially, sharply angulate into posterior face. MALE. Pronotum clothed with sparse setae, integument visible; body setae with overall silvery-white tone; dorsum of propodeum with dense areas of appressed silvery-white setae at least anterolaterally. 96
Description. FEMALE. Body length 13 mm. Head. Posterior margin virtually straight. Occipital carina evenly wide throughout. Vertex width 0.85 × pronotal width. Eye almost circular, its length in frontal view 1.6 × the distance from its ventral margin to mandibular condyle. Head ddensely coarsely and confusedly, areolate-punctate to foveolate-punctate; with conspicuous broad smooth intervals on front; sculpture denser and coarser on vertex. Genal carina present. Mandible oblique, tapering slightly apicad, conspicuously bidentate, unarmed ventrally. Dorsal scrobal carina well defined, not reaching antennal tubercles; lateral scrobal carinae present, connected to dorsal carina. Antennal tubercles irregularly rugose. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.55 × pedicel length. Mesosoma. Dorsal thoracic length virtually as long as mesosomal width. Mesosomal dorsum densely and coarsely areolate-punctate with smooth rounded intervals; sculpture overall slightly larger laterally on pronotum. Humeral carina present narrowly disconnected from slightly produced subangulate epaulet; anterolateral corners of pronotum angulate in dorsal view. Anterior face of propodeum defined, short, shorter than pronotal collar, vestigially and coarsely striated longitudinally basad, with coarse dense punctures dorsad; dorsal face roundly angulate into anterior face in lateral view. Pronotal spiracle virtually flat against lateral margin of pronotum. Lateral face of pronotum sparsely foveolate-punctate with interspersed micropunctures, except for smooth subacute unsculptured tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture, micropunctate anteriorly, sparsely and vestigially foveolate-punctate along mesopleural ridge; metapleuron sculpture virtually completely concealed by dense setation, except dorsal fourth smooth, unsculptured; with dense coarse longitudinal rugosities on dorsal margin anterior to propodeal spiracle. Lateral face of propodeum sparsely foveolate-punctate, intervals smooth and shinning predominantly as wide as surrounding sculpture. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 64:69:71:60:56. Lateral margin of mesonotum marginally constricted anterior to propodeal spiracle, slightly diverging anterad. Propodeal spiracle inconspicuously pronounced from lateral margin of mesosoma; post-spiracular area present. Scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum conspicuously elongate, dorsal face much longer than and well differentiated from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 30:68:67. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, sparsely and coarsely foveolate-punctate to simply punctate with dense interspersed micropunctures where visible; micropunctures sparser on S5, absent on S6. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal slightly concave carina. S2 sparsely foveolate-punctate, sculpture sparser posterad; anteromedial crest-fold virtually absent. S3–6 densely and coarsely foveolate-punctate with sparse micropunctures at S3–4; sculpture denser and micropunctures absent on S6. Pygidial plate subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface predominantly with irregular longitudinal rugosities; interstice apparently granulose. MALE. Body length 12–12.5 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view. Width 0.88 × pronotal width. Eye almost circular. Ocelli small; OOD 3.4 × DLO, IOD 1.5 × DLO. Occipital carina distinct. Head surface densely and coarsely punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; coarsely and densely punctate to micropunctate; apical medial margin slightly concave, with a small denticle on each side of the concavity. Scape bicarinate. Flagellomere 1 2.05 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum rounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations laterally, with a conspicuous smooth unsculptured area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on anterior third and along inner margin. Mesoscutum densely and coarsely foveolate-punctate, notaulus absent, parapsis vestigial, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely areolate-punctate to foveolate-punctate; with longitudinal irregular carina medially formed by aligned intervals. Axilla produced posterolaterally as acute projections in dorsal view, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. 97
Propodeal dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face densely and coarsely areolate, areolations less defined anterad; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum densely coarsely and confusedly punctate to micropunctate; mesopleura slightly swollen on dorsal half, without any or projections; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures anteriorly. Metapleuron foveolate-punctate ventrally, micropunctate to smooth dorsally. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated and with Rs convex, not truncate apically; three submarginal cells; dark brown, slightly but conspicuously lighter on basal third. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.48 × as wide as T2. T2 length 0.72 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly and vestigially rugose, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, slightly pronounced carina lower medially. S2 sparsely foveolate- punctate to punctate, interspersed micropunctations present anterolaterally, foveolations conspicuously sparser and larger posterad; with vestigial longitudinal anteromedial crest-fold; sternal pit absent. S3–5 sparsely and coarsely foveolate- punctate with interspersed micropunctures; S6–7 sparsely foveolate-punctate. S7 longer than broad, posterior margin projected laterally and medially, medial projection terminating in a pair of very small subacute closely spaced tooth-like on posterior margin. Genitalia. Parapenial lobe not at all pronounced posteriorly, simply rounded. Ratios of free length of paramere, cuspis and digitus, 53:31:14; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen medially and narrower posterad in lateral view; narrower posterad and virtually straight in dorsal view; abruptly curved dorsally in wide angle at anterior third; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis well-developed, not sessile, slightly elongate vertically, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, inconspicuously setose basodorsally; penis valve strongly concave on inner surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, with lateral pocket present on outer surface; apical distance between teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface. Coloration and variations. FEMALE. Integument black, except mandibles and antennal flagellomeres partially reddish- brown and T2 with a pair of orange subcircular integumental spots which vary slightly in size. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: front, genae, malar space, ventral surface of head; pronotal dorsum, mesonotum medially, scutelar area, propodeal dorsum medially; T1 medially, disc of T2 (except integumental spots), fringe of T2–5 sublaterally, T6 medially, and S6. Some specimens (T. rastra) vary slightly in the following characters: silvery-white stripes on the head reduced, not touching inner eye orbits; mesosomal silvery-white lateral stripes fully connected near the propodeal spiracles; silvery-white setae of T5 and S5 reduced. Some specimens (T. angustata) vary greatly in the following characters: predominant setae color with a silvery-golden tone; vertex entirely covered in dense silvery-golden setae (reaching dorsal area of front in some cases); silvery-golden markings of mesonotum restricted to posterior third in the form of a single transverse stripe which can be disconnected medially thus forming a pair of transverse spots; silvery-golden setae absent on fringe of T2 medially and greatly reduced in fringe of T3 medially; silvery-golden setae covering T6 (except pygidial plate) entirely. MALE. Integument black, with mandibles and flagellomeres partially reddish-brown. Head mostly with dense white setae except sparse, erect and large, black setae on postero-lateral areas of vertex and near inner eye margins; pronotum dorsum, mesoscutum, axillar projections, scutellum and tegula with black setae; dorsum of propodeum with sparse white setae, dense and decumbent antero-laterally; pronotum lateral face with white setae; mesopleura with sparse black setae near tegula other area with sparse white setae; propodeum lateral face with sparse white setae; legs with white setae except apex of meso and metafemora dorsally with black setae, T1 with white setae, dense and decumbent on dorsal face; T2 to T7 with black setae except anterior third and narrow lateral area of T2, narrow apical fringe of T2, narrow lateral areas of T3–4 with white setae; S1 to S4 with white setae; S5–S6 with white and black setae, S7 with black setae. 98
Distribution. Panama, Brazil, and Argentina. Material examined. (27#f, 6#m) Type material. Holotype of Traumatomutilla diophthalma, BRAZIL, Bahia (ZMB); Lectotype of Traumatomutilla angustata, BRAZIL, Rio Grande do Sul (HBMNH); Paralectotype (designated herer) of Traumatomutilla angustata, BRAZIL, Rio Grande do Sul (MNHN); Holotype of Traumatomutilla rastra, BRAZIL, Santa Catarina, Nova Teutônia (AMNH). Additional material. PANAMA: Panama Prov. [Province]: Chorrera, corregimiento Playa Leona, orilla río Perequete, 2#f, 19−20.III.1991, R. Cambra (MIUP); Chorrera, Llano largo, 2#f, 26−29.III.1990, A. Mena (MIUP); Parque Nacional Soberania, Camino de Cruces, 1#f, 17.II.1998, R. Cambra & A. Santos (MIUP); 1#m, reared in laboratory (MIUP); Chica, 1#m, 1−25.X.2013, (Malaise trap), Y. Cheng (MIUP); Cocle Prov. [Province]: Valle de Anton: 1#f, 6.VII.1991, R. Contreras (MIUP); 1#f, 13.VII.1991, J. Coronado (MIUP); 1#f, 9−10.I.1991, J. Coronado (MIUP); Colon Prov. [Province]: Gamboa, 1#m 16−30.IV.2016, (Malaise trap), Lezcano & Estrada (MIUP); COLOMBIA, Magdalena, P.N.N. Tayron, Neguanje, 11°20’N 74°02’W, 10m[sic!], 1#m, 28.VII.2001, R. Henriquez (IAvH); VENEZUELA: Falcon, Yaracal, 1#f, 19.VII.1986, L. Joly (MIUP); Cojedes, Hato Piñero, cr. [circa] El Baul, 1#f, 3−10.IX.1994, J. Lattke (MIUP); BRAZIL: 1#f (BMNH); Pará: 1#f (BMNH); Ponta de Pedras, 1f#, 27.X.1982, M.F. Torres (MPEG); Santarém, 1#f, IV.1919, S.M. Klages (Label: Traumatomutilla diophthalma (Cresson, nec Klug, det. Mickel 1953) (MIUP); E. [east of] Araguaia, 1#m, 19–31.I.1983, J.A. Rafael (MIUP); Serra Norte, Manganês, 1#m, 06– 09.IX.1985, Márcio Zanuto (MPEG); Mato Grosso do Sul, Cotriguaçú, -9.84’31.39’’S -58.26’26.29’’W, 245, 1f#, 20.IX.206, G. Araújo (UFMT); Goiás, Serranópolis, S-21 J0799057 TEC1 [sic!], 1#f, 10–15.I.2007 (INPA); Santa Catarina: Nova Teutônia: 1f# (MNCN); 2f#, XI.1968, F. Plaumann (DGMC, PMNH); 1f#, 05.IV.1941 (MNRJ); PARAGUAY: Caaguazu: Ypau Señorita, 1f#, 13.XII.2001, U. Dreschel (FSCA); Zudañez, 1f#, XII.1948 (UMMZ); Concepción, Parque Nacional Paso Bravo, Santa Sofia, 22° 19’ 20” S 57°10’12” W, 1#f, 28.X.2002, B. Garcete (MIUP); Paraguari, Parque Nacional Ybycui, 151 m [above sea level], 26°04’ S 56°50’ W, 1#f, 4−16.X.2004, B. Garcete (MIUP); Canindeyu, R.N.B. [Reserva Nacional Bosque] Mbaracayu, Jejui-mi, 1#f, 10−14.I.1997, B. Garcete (MIUP); ARGENTINA: Corrientes, Ituzaingo, 1f#, I.1985, M.A. Fritz (AMNH); Misiones: Puerto Esperanza, 1f#, X.1978, M.A. Fritz (AMNH); Dos de Mayo, 1#m, XII.1989, Foerster (AMNH); Entre Rios: 1#f, 5.XII.1996−15.I.1997, L. Caire (MIUP); Palmar Colón, 1#m, II.1978, M.A. Fritz (AMNH); Pronunciamiento, 1#m, Zelich (AMNH). Hosts. Hymenoptera: Apoidea: Sphecidae: Podium sp. (in laboratory); Hymenoptera: Apoidea: Crabronidae: Trypoxylon sp. (in situ). Host experiment: wooden trap nests were placed in a forested area in Panamá by RAC for two weeks, after which one of the nests was found to be occupied and the entrance closed with resin. The occupied trap was taken to the lab at MIUP and both halves of the nest were separated revealing a Podium Fabricius, 1804 larvae. The nest was once again closed so the larvae could pupate. Once the larvae reached the pupa stage, a female of T. diophthalma was placed inside the nest using forceps and the entrance was once again closed leaving the mutillid locked inside the nest for 24 hours, after which the female was removed. Approximately 30 days later a male emerged. Specimens emerged from Trypoxylon trap-nests were reared from traps placed at Amazonian forest areas in Cotriguaçú, Mato Grosso state, Brazil. Remarks. The types of Traumatomutilla rastra and Traumatomutilla diophthalma are virtually identical except for minor differences in the lateral propodeal face sculpture. This species is, to the best of our knowledge, the most widely distributed in the genus, spanning from Argentina North to Panamá. Its distribution, however, is remarkably “patchy” in between these extremes. It is not clear wheter this is due to lack of sampling or that their range is indeed disjunct, interrupted along the Amazon and resumed South of it. The latter would be a novel distribution in terms of Traumatomutilla since most species seem to be widespread throughout large areas — e.g. T. ocellaris (Klug, 1821) — or restricted to certain types of environments like T. bifurca (Klug, 1821) in Caatinga and T. guarata Casal, 1969 in Atlantic Forest). Traumatomutilla angustata specimens have a single difference in relation to other specimens of T. diophthalma which is the propodeum somewhat simply sloping posterad in lateral view as opposed to the more angulate propodeum of most T. diophthalma. The remaining structural features of T. angustata, however, are identical to the type of T. diophthalma and the males found to occur in the same areas as T. angustata have no difference from the males of T. diophthalma collected elsewhere. Therefore, we are confident that the difference in sculpture of T. angustata can be considered simply a variation and that this species is conspecific with T. diophthalma.
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Traumatomutilla gemella (André, 1906) (Figs. 29–37)
Ephuta (Traumatomutilla) gemella André, 1906: 42, lectotype [designated here], f#, Brazil, S. Paulo [sic!] (BMNH), examined. Traumatomutilla gemella: Casal, 1969, p. 285.
Diagnosis. FEMALE. Dense setae covering most of lateral face of propodeum; micropunctures present on lateral face of propodeum; body setae predominantly silvery-golden; T2 integumental spots orange. MALE. Body setae predominantly silvery-golden; dense appressed setae partially present on lateral face of propodeum and concealing most of metapleuron surface; micropunctures present anterolaterally on S2. Description. FEMALE. Body length 9–13 mm. Head. Posterior margin virtually slightly sinuous, somewhat concave sublaterally and slightly convex medially. Occipital carina conspicuously equally wide throughout. Vertex width 0.9 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate. Mandible with conspicuous subapical tooth. Dorsal scrobal carina present, irregular, separated from antennal tubercles; lateral scrobal carina virtually absent. Antennal tubercle finely and irregularly rugose. Flagellomere 1 2.1 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Dorsal thoracic length slightly smaller than mesosomal width. Mesosomal dorsum densely and coarsely areolate-punctate to foveolate-punctate, with conspicuous medial longitudinal carinae extending from anterior margin of mesonotum to posterior margin of dorsal face of propodeum; carina less defined on propodeum, sinuous, irregular. Anterior face of propodeum defined, short, shorter than pronotal collar, vestigially and coarsely striated longitudinally with interspersed scattered punctures; dorsal face roundly sub-angular with anterior face in lateral view. Humeral carina well-defined, broadly separated from well-defined low sharp epaulet, anterolateral corners of pronotum angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with sparse interspersed micropunctures, except for blunt unsculptured tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and foveolat-punctate along mesopleural ridge where visible; metapleuron sculpture virtually completely concealed by dense setation, dorsal fourth with sparser setae and medial asetose smooth area. Lateral face of propodeum with sculpture partially concealed by dense setation, foveolate-punctate where visible. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 75:82:81:65:59. Lateral margin of mesonotum simply divergent anterior to propodeal spiracle, slightly diverging anterad. Propodeal spiracle virtually flat against lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum conspicuously elongate, dorsal face much longer than and well differentiated from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 31:70:69. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots; sculpture less defined in general over integumental spots. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, slightly lower medially. S2 sparsely foveolate-punctate, punctures conspicuously smaller anteriorly; anteromedial crest-fold vestigial. S3–6 densely and coarsely foveolate- punctate with sparse micropunctures at S3–4; sculpture denser on S6. Pygidial plate broadly subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface mostly irregularly longitudinally rugose; interstice apparently granulose. MALE. Body length 10–12 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view. Width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 3.1 × DLO, IOD 1.2 × DLO. Occipital carina distinct. Head surface densely and coarsely punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; coarsely and densely punctate to micropunctate; apical/ventral margin 100 completely concealed by dense setation. Scape bicarinate. Flagellomere 1 2.4 × pedicel length; flagellomere 2 3.8 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum rounded. Anterior face of pronotum sparsely punctate with interspersed micropunctures laterally, with a conspicuous smooth unsculptured area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on anterior third and along inner margin. Mesoscutum densely and coarsely foveolate-punctate, notaulus absent, parapsis vestigial, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely areolate-punctate to foveolate-punctate; with longitudinal irregular carina medially formed by aligned intervals. Axilla produced posterolaterally as acute projections, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face densely and coarsely areolate, areolations less defined anterad; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum densely coarsely and confusedly punctate to micropunctate; mesopleura slightly swollen on dorsal half, without any or projections; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures anteriorly. Metapleuron sculpture virtually concealed by dense setation. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells; dark brown, slightly but conspicuously lighter on basal third. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly and vestigially rugose, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, slightly pronounced carina lower medially. S2 coarsely and sparsely foveolate-punctate to punctate, interspersed micropunctures present anterolaterally, foveolations conspicuously sparser and larger posterad; with vestigial longitudinal anteromedial crest-fold; sternal pit absent. S3–5 sparsely and coarsely foveolate-punctate with interspersed micropunctures; S6–7 sparsely foveolate-punctate. S7 longer than broad, posterior margin projected laterally and medially, medial projection terminating in a pair of very small subacute closely spaced tooth-like process on posterior margin. Genitalia. Parapenial lobe not at all pronounced posteriorly, simply rounded. Ratios of free length of paramere, cuspis and digitus, 63:33:19; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen sub-basally, narrower posterad, and virtually straight in dorsal view; abruptly curved in wide angle at anterior third and narrower posterad in lateral view; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with simple short setae; paracuspis well-developed, not sessile, slightly elongate vertically, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, sparsely setose dorsally at base; penis valve strongly concave on internal surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth rounded, lateral pocket present on outer margin; distance between apex of teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface. Coloration and variations. FEMALE. Head, mesosoma and metasoma integument always black, at most brownish- black, with mandibles and antennal flagellomeres partially reddish-brown and T2 with a pair of subrounded orange integumental spots varying in size; seta varying in density and predominantly silvery-golden except the following areas with black setae: head (except antennae), dorsum of pronotum, mesonotum, dorsum of propodeum medially, lateral face of pronotum, anterior half of mesopleuron, apex of meso and metafemora dorsally, most of disc of T2 (except integumental spots), most of fringe of T2–3 (except medially and laterally), narrow sublateral areas on fringe of T4, and S6. MALE. Integument black, at most brownish-black, with mandibles and flagellomeres partially reddish-brown. Body setae is predominantly silvery-golden varying in density except for the following areas with black setae varying in density: head (except apical/ventral margin of clypeus) , pronotum, mesonotum, axillar projections, scutellum, apex of meso and metafemora dorsally, posterior two thirds of T2, fringe of T2–3 (except narrow lateral areas), T4–7, most of S4– 5, and S6–7. 101
Distribution. Brazil (São Paulo, Espírito Santo, Minas Gerais, Rio de Janeiro, Paraná, and Santa Catarina). Material examined. (31f# 1m#) Type material. Lectotype, #f, BRAZIL, S. Paulo [sic!] (HBMNH); Paralectotype, #f, BRAZIL, S. Paulo [sic!] (MHNN). Additional material. Espírito Santo: Atílio Vivacqua, 1f#, 13.II.2003 (UFES); Parque Sooretama, 1f#, XI.1967, F.M. Oliveira (DZUP); Santa Teresa, 1f#, 07.XII.1964, C. Elias (DZUP); Guarapari, 1f#, 15.XI.1969, Alvarenga (DZUP); Conceição da Barra, 1f#, 15.XI.1969, C. Elias (DZUP); Corrego do Ita, 1f#, XII.1956 (MNRJ); Minas Gerais: Viçosa, 1f#, 1931 (AMNH); Nanuque, Zona Rural, 1f#, 24–29.VII.2007, F.B. Fraga (UFES); São Paulo: 1f#, Schrottky (MNHN); 4f# (MZSP); Ilha de Busios [Búzios]: 1f#, 16.X–04.XI.1963, Exp. Depto. Zool. [Expedição do Departamento de Zoologia] (MZSP); 1f#, 23–28.III.1964, Exp. Depto. Zool. [Expedição do Departamento de Zoologia] (MZSP); Teodoro Sampaio, 1#m, XII.1985, F.M. Oliviera (EMUS); Rio de Janeiro: 4f# (MZSP); Itatiaia, 700m [a.s.l.], 1f#, 04.XII.1940, J.F. Zikán (MZSP); Paraná: Antonina: Reserva Morro da Mina, 1f#, 11.XII.2006, C. Maia (DZUP); Reserva Rio Cachoeira, 1f#, 01.II.2007, A.J.C. Aguiar (DZUP); Santa Catarina: Corupá, 1f#, III.1947, A. Maller (AMNH); 1f#, XI.1946, A. Maller (AMNH); 2f#, XII.1944, A. Maller (AMNH); 1f#, II.1946, A. Maller (AMNH); 1f#, IV.1952 (MNRJ); 1f#, I.1955 (MNRJ); 1#, XII.1953 (MNRJ); Rio Vermelho, 1f#, XII.1945, A. Maller (AMNH). Remarks. The sex association was based in distribution and matching color pattern since both sexes of Traumatomutilla gemella are part of a well-defined mimicry syndrome occurring in the Atlantic Forest which has been observed and used as a basis for sex associations in multiple species such as Pappognatha patruelis (André, 1898), Hoplocrates cephalotes (Swederus, 1787), Hoplomutilla spinosa (Swederus, 1787), Atlantilla auriculata (Gerstaecker, 1874), and T. guarata Casal, 1969 (KAW pers. obs.). To the best of our knowledge, the only way to differentiate the males of T. gemella from the remaining males of the species-group are the setal characters previously mentioned in the diagnosis and identification key. Setae and color characters are also the most easily observable character to differentiate females of T. gemella especially since it appears that there are no variations whatsoever in the specimens examined. In the case of the females, however, there are reliable structural characters that can be used, namely the presence of micropunctures on the lateral propodeal face of the females.
Traumatomutilla peismatara Bartholomay & Cambra sp. nov. (Figs. 38–45)
Diagnosis. FEMALE. Lateral face of propodeum sparsely sculpture, with large smooth and shinning areas, dorsal face of propodeum virtually flat with mesonotum, sharply angled in relation to posterior face, with conspicuous elevation on posterior margin of dorsal face, head setae entirely silvery-white. MALE. Pronotal dorsum not concealed by dense appressed setation and without micropunctures, lateral face of propodeum without conspicuous areas of dense appressed setae, posterior margin of hypopygium projected medially and laterally, medial projection shorter than lateral ones. Description. FEMALE. Body length 12 mm. Head. Posterior margin virtually straight. Occipital carina evenly wide throughout and smoothly curved dorsolaterally. Vertex width 0.9 × pronotal width. Eye almost circular, its length in frontal view 1.4 × the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate- punctate to areolate-punctate, with broad longitudinal carina extending medially from posterior margin of vertex to middle of front. Mandible with conspicuous subapical tooth. Dorsal scrobal carina well-defined, irregular, separated from antennal tubercles and irregular lateral scrobal carina. Antennal tubercle finely and irregularly rugose. Flagellomere 1 1.75 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Dorsal thoracic length slightly smaller than mesosomal widht. Mesosomal dorsum densely and coarsely areolate-punctate to foveolate-punctate. Anterior face of pronotum defined, short, as long as pronotal collar, vestigially and coarsely striated longitudinally laterobasally with interspersed scattered punctures; unsculptured, smooth, shinning mediobasally; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, slightly projected dorsally, broadly separated from well-defined raised subangulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Sculpture of lateral face of pronotum virtually concealed by dense setation, except for subacute unsculptured tubercle anteroventral in relation to pronotal spiracle; mesopleuron and metapleuron sculpture concealed by dense setation, except dorsal fourth of metapleuron unsculptured, smooth, shinning. Lateral face of 102 propodeum sparsely and shallowly areolate-punctate throughout; sculpture larger and denser posterad, smaller anterad, with large and conspicuous unsculptured, smooth and shinning areas. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 82:87:86:72:65. Lateral margin of mesonotum simply divergent anterior to propodeal spiracle, slightly diverging anterad. Propodeal spiracle virtually flat against lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum conspicuously elongate, dorsal face much longer than and well differentiated from posterior face; dorsal face with conspicuous tubercle-like medial elevation at posterior margin, thus sharply differentiated from posterior margin in lateral view. Metasoma. Ratios of width of T1, width of T2 and length of T2, 33:72:66. Disc of T2 sparselyy and coarsely foveolate-punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with dense interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely foveolate-punctate. S1 sparsely punctured, surface cuneiform, ending in a rounded longitudinal carina, conspicuously higher anteriorly. S2 sparsely foveolate-punctate, foveolations sparser and smaller anterad; anteromedial crest-fold vestigial. S3–4 densely and coarsely foveolate-punctate with dense micropunctures; S5–6 densely and coarsely foveolate-punctate. Pygidial plate broadly subpyriform, defined by strong, projected, flange-like lateral carinae at apical fourth of plate; surface with predominant longitudinal, coarse and confused rugosities, interstice granulose. MALE. Body length 12 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view. Vertex width 0.9 × pronotal width. Eye almost circular. Ocelli small; OOD 3.4 × DLO, IOD 0.8 × DLO. Occipital carina distinct. Head surface densely and coarsely foveolate- punctate; interspersed micropunctures present along posterior margin, genae, and malar space. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medailly; predominantly obscured by dense setation, coarsely and densely punctate to micropunctate whre visible; apical/ventral margin with a pair of closely space short blunt tooth-like projections medially. Scape bicarinate. Flagellomere 1 1.6 × pedicel length; flagellomere 2 2.2 × pedicel length. Mandible obliquely tridentate apically, inner and middle teeth virtually equal, greatly reduced; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, subangulate, broadly separated from humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations laterally, virtually unsculptured, smooth, and shinning elsewhere; slightly longitudinally depressed medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on anterior and inner margin. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum; with medial longitudinal carina on posterior half. Scutellum convex, subglobose, with somewhat definable dorsal and posterior carina; densely and coarsely areolate-punctate to foveolate-punctate; with longitudinal carina medially formed by aligned intervals on dorsal face. Axilla produced posterolaterally as acute projections, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface, except apically and along outer margin unsculptured. Metanotum virtually equally wide throughout, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; lateral face densely areolate posterad to foveolate anterad; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and coarsely foveolate-punctate with interspersed dense micropunctures; mesopleuron slightly swollen on dorsal half, without any or projections; sculpture densely and coarsely foveolate-punctate; foveolations sparser anterad and posterad, micropunctures present anteriorly. Metapleuron predominantly micropunctured to unsculptured, smooth and shinning, except for vestigial foveolation and rugosities on dorsal and ventral fifths. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells; dark brown, slightly but conspicuously lighter on basal third. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 length 0.8 × its width. Dorsal metasomal sculpture partially concealed by dense setation, sparsely and coarsely punctate with interspersed micropunctures where visible; pygidial plate concave, posterior margin conspicuously curved upward; surface predominantly smooth, shinnig, with vestigial undefined sculpture apically; weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, ending in blunt low, slightly concave carina. S2 sparsely and finely 103 foveolate-punctate to punctate, with interspersed micropunctations on anterior third, foveolations conspicuously sparser posterad; anteromedial crest-fold virtually absent; sternal pit absent. S3–4 sparsely and finely foveolate-punctate with interspersed micropunctures; S5–7 sparsely foveolate-punctate. S7 longer than broad, with conspicuous medial longitudinal unsculptured area; posterior margin projected apicalterally, simply convex and shorter than lateral corners medially. Genitalia. Parapenial lobe not at all pronounced posteriorly, simply rounded. Ratios of free length of paramere, cuspis and digitus, 48:24:11; paramere slightly sinuous in dorsal view, upcurved posteriorly in lateral view; with dense setae ventrally at anterior half; cuspis short, stout, slightly swollen medially and narrower posterad in lateral view; narrower posterad and virtually straight in dorsal view; abruptly curved dorsally in wide angle at anterior third; with dense conspicuous, strongly sinuous setae on ventral surface, except at anterior third with simple short setae; dorsal surface with overall inconspicuous simple short setae; paracuspis well-developed, not sessile, slightly elongate longitudinally, subrounded at posterior margin, densely setose along posterodorsal margin, setae predominantly shorter than or as short as paracuspis; digitus short, slightly curved inward in dorsal view and slightly upcurved in lateral view, sparsely setose dorsally at base; penis valve strongly concave on internal surface, with closely spaced pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, lateral pocket present on outer margin; distance between apex of teeth 0.1 × length of valve; dense setae present along truncate, flat posterior margin and inconspicuous short setae present at base of subposterior tooth on outer surface. Coloration and variations. FEMALE. Integument black to brownish-black except mandible and antennal flagellomeres partially reddish-brown, and T2 with a pair of subcircular orange integumental spots. Body setae predominantly silvery- white varying in density except the following areas with black to brownish-black setae varying in density: pronotum medially, mesonotum anteromedially, scutelar area, propodeum medially, disc of T2 (except over integumental spots), and fringe of T2–3 sublaterally. The posterior transverse area of silvery-white setae on the mesonotum varies from broadly interrupted by black setae medially to complete. MALE. Integument black. Wings predominantly dark-brown infuscated, except basal third hyaline brown with blackish veins; with strong violaceous/blueish reflections. Body setae predominantly silvery-white varying in density except the following areas with black setae varying in density: vertex anteriorly, ocellar area, dorsal half of front, gena dorsally; pronotum predominantly, mesoscutum, axillar projections, mesoscutellum, mesopleuron anterodorsally, dorsal third of propodeal dorsum; dorsal and external surfaces of tibiae, femora apicodorsally; T2 disc (except anterior third), fringe of T2–5 medially, T6–7 (except pygidial plate asetose), S6 partially, and S7. Distribution. Peru (Loreto, Ucayali, Madre de Dios) and Brazil (Amazonas and Acre). Etymology. From the greek peismatára, meaning “stubborn, headstrong”, in reference to PRB’s initial stubborn denial of RAC’s conclusion that this was indeed a new species and the males from Acre state their corresponding opposite sex. Treat as an adjective in the nominative singular. Material examined. (5f#, 5m#) Type material. Holotype. 1f#, PERU, Loreto, Pucallpa, 02.X.1953, J.M. Schunke (BMNH). Paratypes. 1f#, PERU (MNHN); 1f#, Loreto, 80 km NE [kilometers northeast of] Iquitos, Yanamono River, Explorama Lodge, 7.XI.1990, R. Cambra & D. Quintero (MIUP); 1f#, Madre de Dios, Reserva Manu, Estacion Pakitza, 1−2.VII.1993, R. Cambra (MIUP); 1f#, Manu N.P. [National Park], Colapa, nr. [near] Cocha Cashu, 1992 (MIUP). Allotypes. BRAZIL, Amazonas, Rio Javari, Estirão do Equador, 4#m, M. Alvarenga (AMNH); Acre, Senador Guiomard, F.E. [Fazenda Experimental] Catuaba, 1#m, 06–26.IX.2016, J.A. Rafael & E.F. Morato (INPA). Remarks. Females of T. peismatara are more structurally similar to those of T. diophthalma but differ mainly in features of the propodeum, namely the overall shape of the dorsal propodeal face and its relation with the mesonotum in lateral view. This small difference originally raised doubts about the validity of this new species, and it was initially considered a variation of T. diophthalma. After the sex association of T. chuza, T. diophthalma, and T. gemella, a fourth male morphospecies was still left in the material examined that couldn’t be properly placed within any of the known species, especially because of the structural differences observed in the hypopygium. These males were all collected in Amazonian areas which are relatively close to the Peruvian distribution of T. peismatara and in areas where no other females of the T. gemella species-group were found. Based on this, morphological differences, and similar distribution, we hypothesize that 104 the females of T. peismatara are a distinct species and the males collected in the far west of the Brazilian Amazon are conspecific with those females.
Discussion
The T. gemella species-group is at the same time the most widely distributed in the genus and one of the rarest, with only 100 specimens found in collections and recorded from Argentina to Panama. This disparity results in paradoxical situations, such as with T. andrei that appears to be widely distributed in Brazil while at the same time being known from three specimens only. Sex associations were particularly difficult. Therefore, apart from the T. diophthalma couple reared in Panama, sex associations in this species-group relied heavily on overlapping distribution of males and females. This situation is not ideal, but the sex association hypotheses proposed here are supported by the morphological cluse provided by the reared male specimen of T. diophthalma in Panamá. Males and females of T. gemella co-occur in the Atlantic Forest and both sexes have the typical color syndrome for the region. Extensive sampling over the years in the Ducke Reserve, Manaus, Amazonas, Brazil, has produced only males and females of T. chuza. Additionally, males of T. chuza haven’t been recorded in areas outside of the Amazon so far, whilst T. diotphalma males have been recorded as far South as Argentina. Finally, through a process of elimination and considering the clear difference between males of T. chuza and T. peismatara and the known distribution of T. peismatara, the sex association for this new species became obvious. Though there are obvious differences in the cuspis, digitus, and penis valve between all males of the T. gemella species-group, it is still unclear how reliable and/or variable these characters are due to the small number of specimens available. At this early stage, however, it is evident that the penis valve of T. gemella is conspicuously more elongate with shorter posteroventral teeth unlike the other species that are stout with a more prominent narrow anterior tooth. The differences between the genitalia of T. diophthalma and T. chuza are all superficial and can only be properly determined with larger series of specimens. The biological data presented here are constitute the first records of Traumatomutilla attacking arboreal or twig- nesting hosts, albeit in trap-nests. It is important to note that females of the T. gemella species-group have the most reduced pygidial plate in the genus, with the flange-like lateral carinae restricted to the apical fifth of the plate; also, the foretarsal rake is virtually absent. Having both these structures well-developed is associated with velvet-ant species that attack ground-nesting hosts (Williams et al., 2011b; Bartholomay et al. 2018). The reduction in both the forestarsal rake and the pygidial plate in the females of the T. gemella species-group, coupled with two different instances of them being reared from two distinct non-ground nesting hosts, is strong evidence that this association is true. Additionally, it is quite remarkable that a single species, T. diophthalma, parasitizes very distinct hosts, a cockroach hunting Sphecidae (Podium) and a spider hunting Crabronidae (Trypoxylon).
Acknowledgements
We are grateful for the collection managers and curators that provided specimens for this study, including: Christine LeBeau (AMNH), Andreas Köhler (CESC), John Rawlins (CMNH), Gabriel Melo (DZUP), James Pitts (EMUS), Mercedes Paris (MNCN), Agniéle Touret-Alby (MNHN), Orlando Silveira (MPEG), Kelli Ramos (MZSP), Gavin Broad (BMNH), Brian Harris (USNM), Michael Ohl, and Viola Richter and Lukas Kirscher (ZMB). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9). 105
References
André, E. (1898) Etude sur les Mutillides du Muséum de Paris. Annales de la Société Entomologique de France, 67, 1–79. André. E. (1901) Descriptions de quelques espèces et variétés nouvelles de Mutilles d’Amerique appartenant au Museé Civique de Gênes. Zeitschrift für Hymenopterologie und Dipterologie. 1: 257–264.André, 1902 André, E. (1902) Fam. Mutillidae. In: Wytsman, P. (Ed.), Genera Insectorum, Bruxelles, Fasc. 11, pp. 1–77. André, E. (1904) Examen critique d’une nouvelle classification proposée par M. le Dr. W. H. Ashmead pour le famille des Mutillides. Revue d’Entomologie. 23(1): 27–41. André, E. (1905–1906) Nouvelles espèces de Mutillides d’Amérique. (Hym.). Zeitschrift für Systematische Hymenopterologie und Dipterologie, 5 & 6, 361–376 (1905) & 33–485 & 6, 361–376 (1905) & 33–48 + 65–80 + 161– 169 (1906).161–169 (1906). André, E. (1908) Descriptions de quelques nouveaux Mutillides du Museé National d’Hongrie. – Annales Musei Nationalis Hungarici, 6, 378–383. Bartholomay, P.R. Williams, K.A. Luz, D.R. & Oliveira, M.L. (2018) New species of Traumatomutilla André in the T. tabapua and T. integella species-groups (Hymenoptera, Mutillidae). Zootaxa 4433 (2): 361–385. Bartholomay, P.R., Williams, K.A., Lopez, V.M. & Oliveira, M.L. (2019) Revision of the Traumatomutilla americana species-group (Hymenopter: Mutillidae). Zootaxa, 4608 (1): 001–034. Brothers, D.J. (2006) Capítulo 14.4. Familia Mutillidae. In: Hanson, P.E. e Gauld, I.D. (Eds), Hymenoptera de la Región Neotropical. The American Entomological Institute, Gainesville, FL, pp. 586–593. Casal, O.H. (1969) Sobre Traumatomutilla André (Hymenoptera, Mutillidae). – Physis, 28 (77), 279–298. Cambra, R.A., Williams, K.A., Quintero, D., Windsor, D.W., Pickering, J. & Saavedra, D. (2018) Dasymutilla Ashmead (Hymenoptera, Mutillidae) in Panama: new species, sex associations and seasonal flight activity. Insecta Mundi, 0608, 1–17. Fabricius, J.C. (1804) Systema Piezatorum: secundum ordines, genera, species, adjectivis, synonymis, locis, observationibus, descriptionibus. Brunsvigae, pp. 428–439. Cresson, E.T. (1902) Descriptions of some Brazilian Mutilla. – Transactions of the American Entomological Society, 28, 1– 82. Gerstaecker, A. (1874) Mutillarum Americae meridionalis indigenarum synopsis systematica et synonymica. Archiv für Naturgeschichte, 40, 41-77, 299–328. Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in Entomology 28: 1–31. Klug, J.C.F. (1821) Entomologiae brasilianae specimen. – Nova Acta Academica Caesareae Leopoldino-Carolinae, 10 (2), 305–324. Luz D.R., Williams, K.A. & Bartholomay, P.R. (2016) The mutillid wasps of the Dasymutilla paradoxa species-group (Hymenoptera: Mutillidae). Zootaxa, 4193 (2): 361–372. Williams, K.A., Manley, D.G., Pilgrim, E.K., von Dohlen, C.D. & Pitts, J.P. (2011a) Multifaceted assessment of species validity in the Dasymutilla bioculata species group (Hymenoptera: Mutillidae). – Systematic Entomology, 36, 180–191. Williams, K.A., Brothers, D.J. & Pitts, J.P. (2011b) New species of Tobantilla Casal, 1965 and a new genus and species, Gogoltilla chichikovi gen. et sp. nov., from Argentina (Hymenoptera: Mutillidae). – Zootaxa 3064, 41–68. Williams, K.A., Manley, D.G., Deyrup, M., von Dohlen, C. & Pitts, J.P. (2012) Systematic review of the Dasymutilla monticola species-group (Hymenoptera: Mutillidae): using phylogenetics to address species-group placement and sex associations. – Zootaxa 3554, 1–29. Williams, K.A., Bartholomay, P.R. & Oliveira, M.L. (2017) Species groups of Traumatomutilla André (Hymenoptera: Mutillidae). Insecta Mundi, 0533: 1–33. Wilson, J.S., Williams, K.A., Forister, M.L., von Dohlen, C.D. & Pitts, J.P. (2012) Repeated evolution in overlapping mimicry rings among North American velvet ants. Nature Communications, 3: 1272. Wilson, J.S., Jahner, J.P., Williams, K.A. & Forister, M.L. (2013) Ecological and Evolutionary Processes Drive the Origin and Maintenance of Imperfect Mimicry. PlosONE, 8 (4), 1–7. 106
Figure legends.
Figure 1–3. Traumatomutilla Andrei (Cresson, 1902), female, holotype, lines 2mm; 1. Dorsal habitus; 2. Lateral habitus; 3. Type labels.
Figures 4–6. Traumatomutilla chuza Casal, 1969, female, holotype, lines 2mm; 4. Dorsal habitus; 5. Lateral habitus; 6. Type labels.
Figures 7–12. Traumatomutilla chuza Casal, 1969, male, line 2mm; 7. Lateral habitus; 8–12. Genitalia. 8. Dorsal view (halved); 9. Ventral view (halved); 10. Lateral/inner view (penis valve removed; 11. Cuspis, lateral/inner view (removed, not to scale); 12. Penis valve, lateral/outer view (removed, not to scale).
Figures 13–15. Traumatomutilla chuza Casal, 1969, females, color variations; 13. Bolivian Amazon; 14. Brazilian Amazon (Amazonas, state); 15. Brazilian Amazon (Rondônia).
Figures 16–17. Traumatomutilla diophthalma (Klug, 1821), female, holotype, lines 2mm; 16. Dorsal view; 17. Lateral view; 18. Type labels.
Figures 19–21. 19. Traumatomutilla angustata (André, 1906), female, paralectotype, line 2mm; 20. Paralectotype labels; 21. Traumatomutilla rastra Casal, 1969, female, holotype, line 2mm; 21. Holotype labels.
Figures 23–28. Traumatomutilla diophthalma (Klug, 1821), male, line 2mm; 23. Lateral habitus; 24–28. Genitalia. 24. Dorsal view (halved); 25. Ventral view (halved); 26. Lateral/inner view (penis valve removed; 27. Cuspis, lateral/inner view (removed, not to scale); 28. Penis valve, lateral/outer view (removed, not to scale).
Figures 29–31. Traumatomutilla gemella (André, 1906), female, paralectotype, lines 2mm; 29. Dorsal habitus; 30. Lateral habitus; 31. Paralectotype labels.
Figures 32–37. Traumatomutilla gemella (André, 1906), male, line 2mm; 32. Lateral habitus; 33–37. Genitalia. 33. Dorsal view (halved); 34. Ventral view (halved); 35. Lateral/inner view (penis valve removed; 36. Cuspis, lateral/inner view (removed, not to scale); 37. Penis valve, lateral/outer view (removed, not to scale).
Figures 38–39. Traumatomutilla peismatara Bartholomay & Cambra sp. nov., female, holotype, lines 2mm; 38. Dorsal habitus; 39. Lateral habitus.
Figures 40–45. Traumatomutilla peismatara Bartholomay & Cambra sp. nov., male, allotype, line 2mm; 40. Lateral habitus; 41–45. Genitalia. 41. Dorsal view (halved); 42. Ventral view (halved); 43. Lateral/inner view (penis valve removed; 44. Cuspis, lateral/inner view (removed, not to scale); 45. Penis valve, lateral/outer view (removed, not to scale).
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CAPÍTULO 4.9 – (formatado para submissão ao periódico ZOOTAXA)
Revision of the Traumatomutilla indica species group (Hymenoptera: Mutillidae: Sphaeropthalminae: Dasymutillini)
PEDRO R. BARTHOLOMAY1,*, KEVIN A. WILLIAMS2, ROBERTO A. CAMBRA3 & MARCIO L. OLIVEIRA1 1Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, Laboratório de Hymenoptera, Av. André Araújo, 2936, Manaus, Amazonas, Brazil. 2Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento CA 95832, USA. 3Laboratório de Biologia Comparada de Hymenoptera, Departamento de Zoologia, Universidade Federal do Paraná, Caixa Postal 19020, 81531-980 Curitiba, PR, Brazil. *Corresponding author, e-mail: [email protected]
Abstract
The Traumatomutilla indica species-group is reviewed, leaving it with 12 species known from both sexes, four known from females only and two known from males only. The following 27 junior synonyms are proposed: Traumatomutilla aemulata (Cresson, 1902) [=Mutilla caneta Cresson, 1902 syn. nov.]; Traumatomutilla centralis (Burmeister, 1875) [=Traumatomutilla fissiventris André, 1907 syn. nov.; Traumatomutilla centralis boliviana Casal, 1969 syn. nov.]; Traumatomutilla contempta André, 1908 [=Traumatomutilla alhuampa Casal, 1969 syn. nov.]; Traumatomutilla geographica (Gerstaecker, 1874) [=Traumatomutilla seabrai Casal, 1969 syn. nov.]; Traumatomutilla grossa (Gerstaecker, 1874) [=Mutilla abrupta Gerstaecker, 1874 syn. nov.; =Mutilla characterea Gerstaecker, 1874 syn. nov.]; Traumatomutilla guayaca Casal, 1969 [=Traumatomutilla tayguaya Casal, 1969 syn. nov.]; Traumatomutilla indica (Linnaeus, 1758) [=Mutilla sphegea Fabricius, 1804 syn. nov.]; Traumatomutilla parallela (Klug, 1821) [=Mutilla foveiventris Gerstaecker, 1874 syn. nov.; =Mutilla caxara Andre, 1902 syn. nov.; =Ephuta lineifera Andre, 1903 syn. nov.; =Traumatomutilla indicoides Mickel, 1952 syn. nov.; =Traumatomutilla gausapata Mickel, 1952 syn. nov.; =Traumatomutilla pillinata Casal, 1969 syn. nov.]; Traumatomutilla puella (Gerstaecker, 1874) [=Mutilla musculus Gerstaecker, 1874 syn. nov.; =Mutilla manca Cresson, 1902 syn. nov.; =Mutilla viana Cresson, 1902 syn. nov.; =Traumatomutilla peperina Casal, 1969 syn. nov.]; Traumatomutilla spectabilis (Gerstaecker, 1874) [=Mutilla funesta Gerstaecker, 1874 syn. nov.; =Mutilla melaleuca Gerstaecker, 1874; Traumatomutilla spectabilis chingona Casal, 1969 syn. nov.]; Traumatomutilla tristis (Klug, 1821) [=Mutilla almada Cresson, 1902 syn. nov.]; Traumatomutilla unimarginata (Cresson, 1902) [Mutilla cuiba Cresson, 1902 syn. nov.]; Traumatomutilla vidua (Klug, 1821) [=Mutilla graphica Gerstaecker, 1874 syn. nov.; =Mutilla scripta Gerstaecker, 1874 syn. nov.; =Traumatomutilla cachimba Casal, 1969 syn. nov.; Traumatomutilla scripta borrosa Casal, 1969 syn. nov.]. The hitherto undescribed males of Traumatomutilla guayaca Casal, 1969 and Traumatomutilla ingens (Andre, 1903) are described. The hitherto undescribed female of Traumatomutilla selligera (Gerstaecker, 1874) is described. Mutilla impetuosa Smih, 1879 is placed within Traumatomutilla. All species treated are redescribed and illustrated. Identifcation keys for males and females are also provided.
Key words: Velvet ants, Taxonomy, Neotropical, Species-groups,
Introduction
Velvet ants (Mutillidae) are a group of solitary wasps in which females are always wingless, males are usually fully winged, and whose larvae act as ectoparasitoids of encapsulated immatures of other insects, especially other solitary Hymenoptera (Brothers, 2006; Williams, 2012). With nearly 180 valid species, Traumatomutilla is one of the most 111 diverse genera of Mutillidae in the Neotropical region, with species ranging from Mexico to Argentina (Nonveiller, 1990; Williams et al. 2017). Due to the extreme sexual dimorphism generally observed in Mutillidae, Traumatomutilla has many species described based only on females (123) or males (37) and sexual associations are notoriously rare and difficult to achieve without molecular data, mating observations in situ or large series of specimens (Nonveiller, 1990; Bartholomay et al. 2019a, b). Additionally, for most of the its taxonomic history, species in Traumatomutilla have been described based solely on differences in variable color and setal patterns, with an almost complete disregard for more reliable structural characters (Gerstaecker, 1874; Cresson, 1902; Casal, 1969). Studies on the closely related Dasymutilla Ashmead, have indicated that such characters alone may not be reliable for species delimitation due to the existence of Müllerian mimicry complexes within these wasps (Williams et al. 2011, 2012; Wilson et al. 2012, 2013). As an effort to facilitate taxonomic revision of this genus, Williams et al. (2017) organized the 135 species of Traumatomutilla based on females into 14 species groups based on shared structural characters or shared combinations of structural characters. Since then, Traumatomutilla species-groups have been slowly but steadily reviewed, resulting in several synonyms, sex associations, and new species making this diverse and widespread genus better known than it has ever been (Bartholomay et al. 2018, 2019a, b). In this study we review the Traumatomutilla indica species-group, which is undoubtfully the most sampled, easily observable and readily recognizable species-group within the genus.
Materials and terminology
Approximately 2,800 specimens were examined for the currnet study; the following codens are used for institutions housing the material discussed:
AMNH - American Museum of Natural History, New York, New York, USA. ANSP - Academy of Natural Sciences, Philadelphia, Pennsylvania, USA. BMNH - British Museum of Natural History, London, England. BYU - Brigham Young University, Monte L. Bean Life Science Museum, Provo, Utah, USA. CASC - Department of Entomology, California Academy of Sciences, San Francisco, California, USA. CPDC - Centro de Pesquisas do Cacau, CEPEC, CPDC, Divisão de Zoologia Agrícola, Itabuna, Bahia, Brazil. CESC - Coleção Entomológica da Universidade de Santa Cruz do Sul, Santa Cruz do Sul, Rio Grande do Sul, Brazil. CISC - Essig Museum of Entomology, University of California, Berkeley, California, USA. CMBC - Craig M. Brabant Collection, Madison, Wisconsin, USA. CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA. CUIC - Cornell University Insect Collection, Ithaca, New York, USA. CZMA - Coleção Zoológica do Maranhão, Caxias, Maranhão, Brazil. DEI - Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany. DGMC - Donald G. Manley Collection, Florence, South Carolina, USA. DJBC - Denis J. Brothers Collection, to be deposited in Iziko South African Museum, Cape Town, South Africa (SAMC). DZUP - Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA. FMNH - Fields Museum of Natural History, Chicago, Illinois, USA. FSCA - Florida State Collection of Arthropods, Gainesville, Florida, USA. IAvH - Instituo Alexander von Humboldt, Villa de Leyva, Boyacá, Colombia. INHS - Illinois Natural History Survey, Champaign, Illinois, USA. INPA - Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil. LACM - Insect Collection, Los Angeles County Museum of Natural History, Los Angeles, California, USA. 112
LRRP - Laboratório de Sistemática e Bioecologia de Parasitóides e Predadores da Agência Paulista de Teconologia dos Agronegócios (APTA), Ribeirão Preto, São Paulo, Brazil. LSUK - Linnean Society United Kingdom, London, England. MACN - Museo Argentino de Ciencias Naturales, Buenos Aires, Buenos Aires, Argentina. MCZC - Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. MNCN - Museo Nacional de Ciencias Naturales, Madrid, Spain. MNHN - Musem Nationale d’Histoire Naturelle, Paris, France.
MNRJ - Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil. MPEG - Museu Paraense Emílio Goeldi, Belém, Pará, Brazil. Mu-Bio - Museu da Biodiversidade, Universidade Federal da Grande Dourados, Dourados, Mato Grosso, Brazil. MZSP - Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo, Brazil BMNHB - Naturhistorisches Museum Basel, Basel, Switzerland. PMAE - Royal Alberta Museum, Edmonton, Alberta, Canada. PMNH - Peabody Museum of Natural History, Yale Univeristy, New Haven, Connecticut, USA. RBINS - Royal Belgian Institute of Natural Sciences, Brussels, Belgium. RMNH - Naturalis Biodiversity Center [formerly Rijksmuseum van Natuurlijke Historie], Leiden, Netherlands. SEMC - Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA. TAMU - Texas A & M University, College Station, Texas, USA. UCDC - The Bohart Museum of Entomology, Univeristy of California, Davis, California, USA. UCRC - Entomology Research Museu, Department of Entomology, Univeristy of California, Riverside, California, USA. UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brazil. UFMG - Universidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil. UFRN - Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte, Brazil. UFRR - Universidade Federal de Roraima, Boa Vista, Roraima, Brazil. UFT - Universidade Federal do Tocantins, Palmas, Tocantins, Brazil. UMMZ - University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA. UMSP - University of Minnesota Insect Collection, St. Paul, Minnesota, USA. USNM - United States National Museum of Natural History, Simthsonian Institution, Washington D.C., USA. ZIN - Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. ZMB - Museum für Naturkunde der Humboldt-Universität, Berlin, Germany. ZMUC - Zoological Museum Univeristy of Copenhagen, Copenhagen, Denmark.
General morphological terminology, definitions, abbreviations Bartholomay et al. (2019). In the description and redescription sections we refrain from mentioning coloration and setal patterns due to the highly variable nature of these characters in Traumatomutilla. Instead, we provide a separate section titled Coloration and variations, in which we provide an overall description of the known color and setae patterns for a particular species. In the material examined section abbreviations, acronyms and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of the material examined section. The types of T. centralis (Gerstaecker, 1874), T. fissiventris André, 1907, T. graphica (Gerstaecker, 1874), T. grossa (Gerstaecker, 1874); T. scripta (Gerstaecker, 1874), T. spectabilis (Gerstaecker, 1874), and T. funesta (Gerstaecker, 1874), although selected and labeled as lectotypes by Mickel, were never published as such by the author. In this paper we maintain and officially designate those specimens selected by Mickel as the lectotypes for their respective species. Many of the species described by Cresson (1902) have more than one specimen in their type series. This is why Cresson (1916) provided a list of specimens with their respective catalog number in his collection, designating them as “types” and stating 113 that those were the specimens used in the descriptions of all his previous studies. Though these specimense were not refered to nor labeled as lectotypes by Cresson, we consider these as the lectotypes of his species and Cresson (1916) as the paper that established them as such.
Taxonomy
Traumatomutilla indica species-group
Diagnosis. FEMALES. Females of this species-group can be defined by the following combinations of characters: dorsum of mesosoma and sometimes propodeum with a medial longitudinal carina; scutelar scale usually broad and laterally connected to complete anterolateral carinae; mesonotum laterally constricted anterior do propodeal spiracles with lateral margins often medially and roundly expanded. The following characters are not exclusive of this species-groups but can also be used to define it: head unarmed posterolaterally; meso and metafemora rounded apically; T2 with conspicuous integumental spots; gena carinate; pygidial plate pyriform to subpyriform, often transversely sculptured. MALES. Males of this species group can be defined by the following combinations of characters: Mesopleuron usually tuberculate on dorsal half (sometimes simply swollen); axillar projections usually acute apically (sometimes truncate); hypopygium elongate, subrectangular, usually defined by lateral carinae. The following characters are not exclusively of this species- group but can also be used to define most of its species: S2 with conspicuous anteromedial or posteromedial pit filled with setae; scutellum with longitudinal medial carina; S1 with spiniform projection medially; cuspis slender elongate and predominantly asetose; penis valve with posterior margin projecting outward into shelf-like structure. Included taxon. T. aemulata (Cresson, 1902); T. alhuampa Casal, 1969; T. borba (Cresson, 1902); T. centralis (Burmeister, 1875); T. contempta André, 1908; T. gausapata Mickel, 1952; T. geographica (Gerstaecker, 1874); T. vidua (Klug, 1821); T. grossa (Gerstaecker, 1874); T. guayaca Casal, 1969; T. indica (Linnaeus, 1758); T. ingens André, 1903; T. mundula (Cresson, 1902); T. parallela (Klug, 1821); T. puella (Gerstaecker, 1874); T. spectabilis (Gerstaecker, 1874); T. tristis (Klug, 1821); T. unimarginata (Cresson, 1902); T. selligera (Gerstaecker, 1874); T. protuberans (Gerstaecker, 1874); T. impetuosa (Smith, 1879) comb. nov. Distribution. Widely distributed throughout South America and Trinidad (except Chile). Remarks. The Traumatomutilla indica group is one of the most diverse in the genus with various easily recognizable species, many useful structural characters; it is also perhaps the most common species-group of Traumatomutilla encountered in collections (KAW, PRB pers. obs.). Similar to what was observed with the T. gemella males, most species within the T. indica species-group are also remarkably homogeneous in genitalic characters, even when dealing with species that are obviously different in external morphology. With the exception of three species, all males have a slender, elongate and mostly asetose cuspis with minor differences in the shape of the parameres, paracuspis and/or penis valve. Body size is highly variable within this species-group that includes large-bodied species, such as T. aemulata (all known females longer than 20 mm, Fig. 1A, 1C) occurring in the same area as small species like T. puella (most females shorter than 8 mm, Fig. 22A–F). The only character that seems to be present in a consistent way for all males of the T. indica species-group is the elongate and subrectangular hypopygium that is usually defined by lateral carinae. The most variable characters are the scutellum and axillar projections. For females the most consistent character is the presence of a medial longitudinal carina on the mesonotum that can be greatly reduced, even indistinct in some species, but still observable by removing the dorsal mesosomal setae in the area. Similar to what was observed in the males, the scutellar area is highly variable with the scutelar scale and anterolateral carinae varying from virtually absent to well-defined and easily observable even under the dense mesosomal setae.
FEMALES
1. Mesonotum usually conspicuously wider than distance between pronotal spiracles (Figs. 26A, 32A, 32C, 32E, 32G), if mesonotum as wide as distance between pronotal spiracles, then scutelar scale and anterolateral carinae greatly 114 reduced, irregular, nearly indistinct; lateral face of propodeum densely sculptured throughout, at most slightly sparser posteroventrally … 2 Mesonotum usually as wide as or narrower than distance between pronotal spiracles (Figs 17A, 18A); scutelar scale variable, if mesonotum wider than distance between pronotal spiracle, then lateral face of propodeum sparsely sculptured with large areas of smooth shinning integument … 3
2. Scutelar scale and anterolateral carinae well defined; head setae black (Figs. 32A–H) … T. vidua (Klug, 1821) Scutelar scale present, highly variable, frequently reduced, anterolateral carinae usually absent, at most indistinct; head usually with a transverse band of silvery-white setae (Figs. 26A–E) … T. spectabilis (Gerstaecker, 1874)
3. Lateral face of propodeum with dense sculpture throughout (e.g. Figs. 19D, 19F, 19H); if with apparent unsculptured areas, then these are small, usually not larger than surrounding sculptured areas (e.g. Fig. 30C) … 4 Lateral face of propodeum with sparse sculpture having conspicuous unsculptured smooth and shinning areas (e.g. Fig. 24B) … 9
4. Meso- and metatibial spurs black, concolorous with legs; mesosoma with coppery-golden setal stripes (Figs. 30A, 30C) … T. unimarginata (Cresson, 1902) Meso- and metatibial spurs white, contrasting with legs; mesosoma with silvery-white setal pattern (e.g. Figs. 10A, 10D) … 5
5. Sculpture of lateral face of propodeum dense throughout, intervals indistinctly rugose, dull; propodeum elongate, sloping in lateral view, dorsal face apparently longer than posterior face (Figs. 18A, 18C, 19A –H); longitudinal medial carina on dorsal face of propodeum always conspicuous and prominent, clearly visible between dense setation … T. parallela (Klug, 1821) Sculpture of lateral face of propodeum variably dense with intervals smooth, shinning (Fig. 15B); propodeum variable, gibbose to simply convex (e.g. Fig. 15B); longitudinal medial carina on dorsal face of propodeum variable … 6
6. Dorsal face of propodeum conspicuously shorter than posterior face, which is nearly vertical in lateral view; robust species restricted to southern South America (Figs. 15A–B) … T. ingens André, 1903 Dorsal face of propodeum as long as or longer than posterior face, which is inclined and curved in lateral view (e.g. Fig. 6B); size and distribution variable … 7
7. Scutelar scale and anterolateral carinae well-defined, connected laterally; sculpture of mesonotum evenly areolate- punctate throughout, with sharp intervals; medial longitudinal carina on dorsal face of propodeum absent (Figs. 13A, 13C) … T. indica (Linnaeus, 1758) Scutelar scale and anterolateral carinae greatly reduced, indistinct, irregular, usually observable only after removing setae (e.g. Fig. 6A); sculpture of mesonotum simply punctate mediad adjacent to longitudinal carina, intervals smooth, blunt; medial longitudinal carina on dorsal face of propodeum present, frequently concealed by dense setae … 8
8. Head with transverse stripe of dense silvery-golden setae; T2 with four integumental linear yellow spots; anterior pair longitudinal and posterior pair transverse; posterior pair separated by one fourth the transverse width of one spot (known only from Chapada dos Guimarães, Mato Grosso state, Brazil) (Figs. 1A, 1C) … T. aemulata (Cresson, 1902) Head setae completely black; T2 with four subrounded to subquadrate integumental reddish to orange spots; posterior pair separated by no more than half the transverse width of one spot (known from Chaco areas in Brazil, Bolivia, Argentina and Paraguay) (Figs. 6A–E) … T. contempta (André, 1908)
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9. Longitudinal carina of mesonotum and dorsal face of propodeum well-defined, connected or nearly so at scutellar area … 10 Longitudinal carina of mesonotum and/or dorsal face of propodeum generally inconspicuous, always broadly separated near scutellar area … 12
10. Meso- and metatibial spurs black, concolorous with legs; sculpture of lateral face of propodeum with smooth shinning sculpture; anterior pair of integumental spots on T2 well-defined and posterior pair completely merged thus forming a single transverse yellow integumental stripe (Figs. 7A–E) … T. geographica (Gerstaecker, 1874) Meso- and metatibial spurs yellowish-white contrasting with legs; sculpture of lateral face of propodeum frequently with indistinct rugosistes, somewhat dull; if meso and metatibial spurs color uncertain, then posterior pair of integumental spots on T2 widely separated by more than transverse length of one spot, transversely oblique … 11
11. T2 with well-defined anterior and posterior pair of yellowish to orange integumental spots; anterior pair longitudinally sublinear; posterior pair transverse subrectangular to transversely oblique subelliptical (Figs. 22A –F) … T. puella (Gerstaecker, 1874) T2 with anterior pair of integumental spots absent, at most reduced to a slightly lighter area of integument on base of T2 with sparse silvery-white setae (visible at higher magnigication); posterior pair confluent medially, thus forming a single uninterrupted transverse yellow stripe (Figs. 17A, 17C) … T. mundula (Cresson, 1902)
12. Meso and metatibial spurs black, concolorous with legs (Figs. 4A–B, 4D–F) … T. centralis (Burmeister, 1875) Meso and metatibial spurs yellowish-white to silvery-white, contrasting with legs (e.g. Fig. 32B) … 13
13. Dorsal face of propodeum conspicuously shorter than posterior face; posterior propodeal face almost vertical, well differentiated from dorsal face; restricted to southern South America (Pampas and southern Atlantic Forest) (Figs. 8A–B, 8D–E) … T. grossa (Gerstaecker, 1874) Dorsal face of propodeum as long as or longer than posterior face (e.g. Fig. 3B); posterior propodeal face variable, poorly differentiated from dorsal face at times; widely distributed across South America … 14
14. Scutelar scale well-defined as an arched sulcus extending to propodeal spiracles laterally; mesonotal sculpture equally areolate-punctate throughout; T2 conspicuously longer than wide in dorsal view (Figs. 3A–B) … T. borba (Cresson, 1902) Scutelar scale poorly defined or if well-defined, then not reaching propodeal spiracles, at most laterally connected to anterolateral carinae; mesonotal sculpture areolate-punctate to simply punctate mediad in areas adjacent to medial longitudinal carina; T2 virtually as long as wide in dorsal view … 15
15. Lateral face of pronotum with conspicuous sharp tubercle anteroventral in relation to pronotal spiracle, clearly visible from above; pronotal collar densely, coarsely and transversely rugose throughout; anterolateral carinae on scutelar area well-defined, virtually straight (Figs. 24–B) … T. selligera (Gerstaekcer, 1874) Lateral face of pronotum at most with blunt inconspicuous swelling anteroventral in relation to pronotal spiracle, barely discernible in dorsal view; pronotal collar smooth and shining on posterior half; anterolateral carinae on scutelar scale inconspicuous to indistinct, mostly concealed by dense setation, evenly arched throughout if present … 16
16. Anterolateral corners of pronotum acutely projected in dorsal view, angulate; pronotal collar shorter than anterior face of pronotum (Figs. 10A–B, 10D–E) … T. guayaca Casal, 1969 Anterolateral corners of pronotum rounded to subangulate in dorsal view, not acute; pronotal collar longer than or as long as anterior face of pronotum (Figs. 28A–B) … T. tristis (Gerstaecker, 1874)
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MALES
1. Cuspis club like, broad, apical half laterally compressed, with outer surface concave and densely setose (e.g. Figs 16B–E) … 2 Cuspis slender, thin, not conspicuously compressed or concave at any point, with sparse setae throughout (e.g. Figs. 12C–F) … 4
2. Mesopleuron simply swollen on dorsal half (Fig. 9A) … T. grossa (Gerstaecker, 1874) Mesopleuron with distinct blunt tubercle on dorsal half (e.g. Fig. 27B) … 3
3. Scutellum gibbose, with dorsal face apparently much shorter than vertical posterior face; T2 with lateral subrounded patches of silvery-white setae basally (Fig. 16A); cuspis club-like, conspicuously broader apicad in lateral view (Fig. 16D) … T. ingens André, 1903 Scutellum roundly convex, dorsal and posterior faces less distinct, dorsal face as long as posterior face; T2 with lateral longitudinal stripes of silvery-white setae basally (Fig. 2A); cuspis slenderer, only slightly broader apicad (Fig. 2E) … T. aemulata (Cresson, 1902)
4. Meso and metatibial spurs black to brownish, concolorous with legs and/or strongly contrasting with with silvery- white setae markings of metasoma (e.g. Fig. 5A) … 5 Meso and metatibial spurs silvery-white to yellowish-white, contrasting with legs and/or having similar color to silvery-white setae markings of metasoma (e.g. Fig. 12A) … 7
5. Mesopleural tubercles well-defined, conspicuously projected, blunt; wings infuscated dark brown except at basal third, hyaline, strongly contrasting with remainder of wing; cuspis in lateral view ending in somewhat enlarged, subcapitate apex; apical/posterior margin of penis valve with well-defined shelf-like expansion towards outer surface, clearly visible in dorsal view (Figs. 5C–G) … T. centralis (Burmeister, 1875) Mesopleural tubercles reduced, dorsal half of mesopleuron with inconspicuous blunt swelling; wings hyaline- brown throughout, at most slightly darkened at apical third; cuspis evenly wide throughout, at most slightly narrower apically; apical/posterior margin of penis valve without conspicuous shelf-like expansion (e.g. Figs. 31C–G) … 6
6. S2 with well-defined subovate anteromedial pit filled with black setae; axillar projection ending in roundly truncate apex in dorsal view; free length of cuspis 4.8 × that of digitus; fringe of T2–3 entirely clothed with silvery-golden setae (known only from Chapada dos Guimarães, Mato Grosso state, Brazil) (Fig. 31A) … T. unimarginata (Cresson, 1902) S2 without pit, at most with anteromedial patch of dense silvery-white setae; axillar projections ending in sharply acute apex in dorsal view; free length of cuspis 4.2 × that of digitus; fringe of T2–3 clothed with silvery-white setae laterally and black setae medially (widespread) (Figs. 23D, 23F) … T. puella (Gerstaecker, 1874)
7. Scutellum simply convex, evenly curved throughout, dorsal face indistinguishable from posterior face; if posterior face more or less defined, then never vertical, sloping, inclined … 8 Scutellum gibbose to globose, dorsal and posterior face usually very distinct; posterior face usually almost vertical … 9
8. S2 with well-defined slightly anteromedial subovate pit filled with silvery-white setae; pronotal dorsum densely foveolate-punctate with sparse interspersed micropunctures (requires removal of pronotal setae); pronotum clothed with black setae; fringe of T2–3 mostly clothed with black setae, silvery-white setae laterally (Fig. 12A) … T. impetuosa (Smith, 1879) comb. nov. 117
S2 at most with small patch of dense silvery-white setae anteromedially; pronotal dorsum sparsely foveolate- punctate with dense micropunctures (requires removal of pronotal setae); pronotum clothed with silvery-white setae; fringe of T2–3 entirely clothed with silvery-white setae (Fig. 11A) … T. guayaca Casal, 1969
9. Axillar projections obliquely or transversely truncate apically in dorsal view … 10 Axillar projections acute apically in dorsal view … 12
10. Scutellum with posterior face inclined, sloping slightly in lateral view; axillar projection obliquely truncate apically in dorsal view, conspicuously oblique in posterior view (Figs. 27A–B, 27D–E) … T. spectabilis (Gerstaecker, 1874) Scutellum with posterior face nearly vertical; axillar projection transversely truncate apically in dorsal view, transverse in posterior view; if apparently acute in dorsal view and oblique and posterior view, then narrowing apicad in dorsal view … 11
11. Axillar projection short, virtually equally wide throughout in dorsal view; apical width of projection as wide as distance from its inner margin to mesoscutellum; penis valve with apical/posterior tooth as long as or longer than subapical/anterior tooth (Figs. 33 A–G) … T. vidua (Klug, 1821) Axillar projection elongate, narrowing apicad; apical width of projection narrower than distance from its inner margin to mesoscutellum; penis valve with apical/posterior tooth shorter than subapical/anterior tooth … (Figs. 21A–G) T. protuberans (Gerstaecker, 1874)
12. Dorsal face of mesoscutellum ending in sharp, shelf-like, transverse carina … 13 Dorsal face of mesoscutellum with longitudinal carina throughout … 14
13. Transverse carina at posterior margin of dorsal face of mesoscutellum bilobate; forewing light infuscated brown at apical third, hyaline brown at remainder of wing membrane; S2 with anteromedial pit filled with silvery-white setae; penis valve with conspicuous shelf-like outward projection at apical/posterior margin (Figs. 14A–G) … T. indica (Linnaeus, 1758) Transverse carina at posterior margin of dorsal face of mesoscutellum straight; forewing infuscated brown except at hyaline brown basal third; S2 with posteromedial pit filled with silvery-white; penis valve without shelf-like outward projection apical/posterior margin (Figs. 25A–G) … T. selligera (Gerstaecker, 1874)
14. Propodeum and pronotum usually densely clothed with silvery-white setae concealing integument, if pronotum clothed with black setae, then metasoma predominantly clothed with silvery-white setae; longitudinal medial carina of dorsal face of scutellum usually variable, wider apicad or equally wide throughout; S2 variable usually with anteromedial narrow longitudinal pit filled with setae; cuspis equally wide throughout; free length of cuspis 0.7 × free paramere length (Figs. 20A–G) … T. parallela (Klug, 1821) Pronotum always clothed with black setae; silvery-white setae of propodeum sparse, never obscuring integument; longitudinal medial carina of dorsal face of scutellum always equally wide throughout; S2 always with conspicuous anteromedial pit filled with dense silvery-white setae; cuspis conspicuously widened apicad in dorsal and lateral view; free length of cuspis 0.9 × free paramere length (Figs. 29A–K) … T. tristis (Gerstaecker, 1874)
Traumatomutilla aemulata (Cresson, 1902) (Figs. 1A–C, 2A–G)
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Mutilla aemulata Cresson, 1902: 50, lectotype [designated by (Cresson, 1916)], #f, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined. Mutilla caneta Cresson, 1902: 73, lectotype [designated by (Cresson, 1916)], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) aemulata: André, 1902: 54. Ephuta (Traumatomutilla) caneta: André, 1902: 56. Traumatomutilla aemulata: André, 1904: 40. Traumatomutilla caneta: André, 1904: 40.
Diagnosis. FEMALE. Scutelar scale and anterolateral carinae greatly reduced; dorsal face of propodeum as long as posterior face, lateral face of propodeum evenly sculptured throughout, mesonotum as wide as distance between pronotal spiracles, longitudinal setae lines of mesosoma silvery-golden anterad. MALE. Cuspis club-like broad and laterally compressed, mesopleuron distinctly tuberculate, scutellum roundly convex, T2 with anterolateral stripes of silvery-white setae. Description. FEMALE. Body length 20 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.7 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Head sculpture mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate where visible. Genal carina present, well-defined. Mandible oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, extending over antennal tubercles and connecting with well-defined lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.75 × pedicel length; flagellomere 2 1.75 × pedicel length. Mesosoma. Dorsal thoracic length 0.9 × mesosomal width. Mesosomal dorsum densely and coarsely areolate-punctate with conspicuous medial longitudinal carinae extending from anterior margin of mesonotum to scutelar area; areolations on mesonotum gradually less defined to completely absent mediad; scutelar area simply and densely punctate; sculpture of dorsal face of propodeum unaltered throughout with well- defined medial longitudinal carina. Anterior face of pronotum defined, long, as long as pronotal collar, indistinctly and coarsely striated longitudinally at base and dense coarse punctures with interspersed micropunctures elsewhere; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from low rounded epaulet, anterolateral corners of pronotum subrounded in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum, rounded. Lateral face of pronotum densely punctate with dense interspersed micropunctures; with indistinct blunt tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal third completely asetose, smooth. Lateral face of propodeum densely foveolate-punctate to areolate-punctate throughout. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 71:90:90:62:65. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae virtually absent, reduced to transverse irregular intervals delineating scutelar area which is densely and finely punctate; scabrous intervals absent on scutellar area. Propodeum gibbose, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 63:70:73. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 densely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in sub-sharp longitudinal medial carina terminating in sharp tooth posteriorly. S2 densely foveolate- punctate, with conspicuous unsculptured medial longitudinal area; anteromedial crest-fold indistinct. S3–6 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible. Pygidial plate broadly subpyriform, defined by lateral carinae at apical third of plate; surface mostly irregularly 119 longitudinally rugose; interstice apparently granulose. MALE. Body length 17 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view. Width virtually equal to pronotal width. Eye almost circular. Ocelli small; OOD 4.0 × DLO, IOD 1.35 × DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate, less densely so around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded free teeth. Scape bicarinate. Flagellomere 1 3.1 × pedicel length; flagellomere 2 4.6 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum rounded. Anterior face of pronotum sparsely punctate with interspersed micrpunctations laterally, and conspicuous smooth unsculptured area basomedially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely foveolate-punctate; with longitudinal broad carina medially. Axilla produced posterolaterally as short acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate, sculpture of lateral face gradually less defined to indistinct anterad; anterior margin mostly smooth, unsculptured; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropuncture; mesopleura with conspicuous blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures along anterior margin. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations basally. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length virtually equal to width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly, transversely and indistinctlyly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in slightly concave longitudinal carina. S2 coarsely and sparsely foveolate-punctate to punctate, with interspersed micropunctations laterally; micropunctures absent on anterior third and medially; S2 with conspicuous medial subovate pit densely filled with setae; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially, terminating in a pair of very small subacute closely spaced tooth-like structures on posterior margin. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 98:73:22; paramere marginally sinuous in dorsal view, upcurved apically in lateral view; with sparse setae ventrally at basal half; cuspis broad, long, club-like, laterally compressed at apical half, slightly wider apicad in dorsal view and conspicuously wider apicad in lateral view, outer surface somewhat concave; with strong setae on ventral, outer and dorsal margins at apical half; paracuspis well-developed, not sessile, lobe-like, virtually as wide as long with rounded apical margin and densely setose along posteroapical margin in lateral view; setae predominantly longer than or as long as paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, with externolateral pocket; apical distance between teeth 0.15 × length of valve; dense setae present along truncate posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Colorationa and variations. FEMALES. Body and appendages predominantly black to brownish-black, except most flagellomeres and mandibles partially reddish-black to reddish-brown; T2 with four linear yellow integumental spots; anterior pair longitudinal; posterior pair transverse, narrowly separated and slightly curved anterad medially. Tibial spurs brownish-yellow. Body setae predominantly silvery-white to silvery-golden varying in density, except the following areas with black to brownish-black setae varying in density: front, dorsal half of gena, posterior half of vertex, dorsum and most 120 of lateral face of pronotum, mesonotum medially, scutelar area, dorsum of propodeum medially, mesopleuron anteriorly, dorsal third of mesopleuron posteriorly, lateral face of propodeum, T1 medially, most of disc of T2, fringes of T2–4 sublaterally, fringe of T5–6 laterally, fringe of S4, S5–6, and tarsi ventrally. MALES. Integument black to brownish-black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: middle of metanotum, propodeal dorsum, hind legs partially, T1, T2 anterolaterally, fringe of T2–3 laterally, S1–3, and fringes of S2–3. Wings dark-brown, conspicuously hyaline-brown at basa third. Distribution. Brazil. Material examined. (5#f, 1m#) Type material. Lectotype of Traumatomutilla aemulata, holotype, #f, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH); Lectotype of Traumatomutilla caneta, #m, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. BRAZIL: Mato Grosso: 1#f (MZSP); Chapada [dos Guimarães: 1#, XI (ANSP); 1#, IX (ANSP); Campo, 1# (MNHN). Remarks. Traumatomutilla caneta was associated and synonymized with T. aemulata based firstly on their morphological similarity with T. grossa (#f) and T. characterea (#m) which were associated and synonymized through repeated co- occurrence in southern South America. The females (T. aemulata and T. grossa) both have a greatly reduced scutelar armature and an overall conspicuously large and stout body. The males (T. caneta and T. characterea) both are also large- bodied specimens and have a club-like laterally flattened cuspis that they also share with a third species from the dry regions of southern South America (T. ingens). Traumatomutilla caneta and Traumatomutilla aemulata are the only species in Chapada dos Guimarães with the aforementioned characters and, in fact, seem to be restricted to that area. In the midwestern Cerrado regions of Brazil, especially around Chapada dos Guimarães, certain females have dense coppery setal patterns on the mesosomal dorsum, most easily observed in T. unimarginata (Figs. 30A, 30C), but clearly present also in T. chapada (Cresson, 1902) of the T. inermis species-group. Traumatomutilla aemulata has the mesosomal stripes pale golden (Fig. 1A) which could be an intermediate state between T. unimarginata and the white stripes typical of most Traumatomutilla. There are few specimens of T. aemulata in collections and we haven’t observed any color variations; the mesosomal setal pattern is, at this point, the easiest way to recognize this species. The male also has a distinctive color character that, so far, was only observed in T. aemulata. The short and longitudinally oblique silvery-white stripes on the basal third of T2 differ from the subcircular patches of silvery-white setae that are commonly observed in other males of Traumatomutilla.
Traumatomutilla borba (Cresson, 1902) (Figs. 3A–C)
Mutilla borba Cresson, 1902: 53, lectotype [designated by Cresson (1902)], #f, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined. Ephuta (Traumatomutilla) borba: André, 1902: 54. Traumatomutilla borba: André, 1904: 40.
Diagnosis. FEMALE. Mesonotum narrower than distance between pronotal spiracles, lateral face of propodeum not evenly sculptured, scutellar scale well-defined and extending to propodeal spiracles, T2 conspicuously longer than broad and wider posteriorly than anteriorly. MALE. Unknown. Description. FEMALE. Body length 10-15 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate where visible; sculpture sparser on gena and malar space; with well defined medial longitudinal carina extending from vertex to front. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, separated from antennal tubercles and lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.1 × pedicel length; flagellomere 2 1.35 × pedicel length. 121
Mesosoma. Dorsal thoracic length slightly shorter than mesosomal width. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible with somewhat rounded intervals; medial longitudinal carina present on mesonotum and dorsal face of propodeum, interrupted near scutelar area. Anterior face of pronotum defined, virtually as long as pronotal collar; with indistinct, indistinct and coarse longitudinal striations at base and dense coarse and confused punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, broadly separated from slightly projected and sharp epaulet, anterolateral corners of pronotum subangulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded. Lateral face of pronotum densely punctate with dense interspersed micropunctures; with indistinct swelling anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely coarsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by dense setation on ventral half, completely asetose, smooth on dorsal half. Lateral face of propodeum sparsely and coarsely foveolate- punctate with smooth shinning intervals never wider than surrounding sculpture. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 60:69:69:52:53. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale present, transverse, arched, reaching propodeal spiracles; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 40:87:95. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations slightly sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around sub-sharp longitudinal medial carina equally high throughout. S2 densely foveolate-punctate, more sparsely so posteromediad; anteromedial crest-fold indistinct. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with interspersed micropunctures where visible; S5–6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Unkown. Coloration and variations: FEMALES. Integument black except flagellomeres and mandibles partially reddish-brown, and T2 with four subelliptical to subovate yellowish integumental spots. Body setae predominantly silvery-white varying in density except following areas with black setae varyin in density: front, vertex, dorsal half of gena; dorsal half of lateral face of pronotum, mesopleuron, and metapleuron; pronotal dorsum, mesoscutum medially, propodeal dorsum medially; T1 medially, disc of T2 (except integumental spots), fringe of T2–5 sublaterally, T6 (except pygidial plate) medially, S1– 5, and fringe of S2–3. Tibial spurs ywllowish-white. MALES. Unknown. Distribution. Brazil. Material examined. (2f#) Type material. Lectotype, #f, Brazil, [Mato Grosso], Chapada [dos Guimarães], Nov. [November] (CMNH); Paralectotype, #f, samel label data as holotype (CMNH). Remarks. Numerous specimens in various collections may have been erroneously identified as T. borba, since the reference specimens used thus far are remarkably different from the type. We haven’t seen any specimens that resemble T. borba outside of the type series (two specimens). There seem to be two possible explanations for this: 1. This is a rare species with restriced distribution to the Chapada dos Guimarães area, as seems to be the case with T. aemulata (Cresson, 1902); 2. This may simply be a slender version of T. spectabilis (Gerstaecker, 1874) with yellow spots, which would mirror the situation with T. grossa (Gerstaecker, 1874) and T. abrupta (Gerstaecker, 1874), discussed below. There is, however, one conspicuous structural difference between the type series of T. borba and T. spectabilis. The scutelar scale is well-defined and reaches the propodeal spiracles laterally in T. borba, which contrasts with the generally poorly defined to indistinct scutelar armature of T. spectabilis.
Traumatomutilla centralis (Burmeister, 1875) 122
(Figs. 4A–F, 5A–G)
Mutilla centralis Burmeister, 1875: 473, lectotype [designated here], #f (nec #m), Argentina, Córdova [Córdoba] (MACN), type examined. Ephuta (Traumatomutilla) centralis: André, 1902: 54. Traumatomutilla centralis: André, 1904: 40. Ephuta (Traumatomutilla) fissiventris André, 1907: 349, lectotype [designated here], #m, Argentina, Santiago Del Estero, environs d’Icano [outskirts of Icaño], E.R. Wagner (MNHN), type examined, syn. nov. Traumatomutilla fissiventris André 1908b: 213.
Diagnosis. FEMALE. Mesonotum narrower than distance between pronotal spiracles, lateral face of propodeum not evenly sculptured, medial longitudinal carinae of mesonotum and propodeum separated at scutelar area, tibial spurs black. MALE. Tibial spurs black, cuspis slender, subcapitate apically, penis valve with shelf-like outward projection on posterior margin. Description. FEMALE. Body length 13–16 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate to areolate-punctate where visible. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, reaching antennal tubercles and connected to lateral scrobal carina. Antennal tubercle irregularly and coarsely punctate-rugose. Flagellomere 1 2.4 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Dorsal thoracic length 0.9 × mesosomal width. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible; with conspicuous medial longitudinal carina extending from anterior margin of mesonotum, to scutelar area; sculpture adjacent to carina simply punctate; dorsal face of propodeum with reduced similar carina; scutelar area sculpture densely punctate Anterior face of pronotum defined, slightly longer than pronotal collar, indistinctlyly and coarsely striated longitudinally at base and with dense coarse and confused punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, narrowly separated from slightly projected rounded epaulet, anterolateral corners of pronotum subangulate in dorsal view. Pronotal spiracle strongly projected from lateral margins of pronotum, rounded. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; with indistinct blunt tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate to simply punctate along mesopleural ridge where visible; metapleuron sculpture predominantly concealed by dense setation, except narrow unsculptured and asetose smooth area on dorsal fourth. Lateral face of propodeum sparsely and shallowly foveolate-punctate to densely and coarsely areolate-punctate posterad; intervals smooth shinning medially. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 76:88:82:63:61. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad. Propodeal spiracles slughtly projected from lateral margins of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae present, irregular, connected laterally, intervals irregular, not scabrous. Propodeum convex, dorsal face virtually as long as and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 37:72:79. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser laterad and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate terminating into sharp longitudinal carina slightly higher posteriorly. S2 densely foveolate-punctate, with distinct unsculptured longitudinal medial area; anteromedial crest- fold well-defined. S3–6 sculpture mostly concealed by dense setation, densely and coarsely foveolate-punctate where visible. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with transverse coarse and confuse rugosities; interstice granulose. MALE. Body length 13-16 mm. Head. Transverse, postero-laterally swollen, 123 scarcely swollen postero-medially. Vertex width 0.95 × pronotal width. Eye almost circular. Ocelli small; OOD 4.0 × DLO, IOD 1.0 × DLO. Occipital carina distinct. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.5 × pedicel length; flagellomere 2 3.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets slightly projected from anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum sub-rounded. Anterior face of pronotum sparsely and coarsely punctate with interspersed micropunctures; sculpture sparser mediad, simply micropunctate medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Mesoscutellum convex, densely and coarsely foveolate-punctate; dorsal and posterior faces poorly distinguished; intervals aligned forming longitudinal carina throughout mesoscutellum. Axilla produced posterolaterally as oblique dentate projection, with coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setae. Propodeal dorsum convex to virtually flat basad, densely areolate; lateral face virtually unsculptured along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleuron with conspicuous sharp tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on basal fourth and indistinct rugosities on dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell slightly longer than stigma, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.8 × its width. Dorsal metasomal sculpture partially concealed by dense setae, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate basally punctate with smooth intervals, apical portion densely micropunctate, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in slightly concave and blunt longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate, without micropunctures; with conspicuous anteromedial ovate seta-filled pit; longitudinal anteromedial crest- fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a tooth-like structure on posterior margin, apex of projection bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 91:69:22; paramere marginally sinuous in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half. Cuspis slender, elongate, equally wide throughout in dorsal view and lateral view; apex subcapitate in lateral view; scattered inconspicuous long setae present apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, longer than wide with subtriangulate and sparsely setose posterodorsal margin in lateral view; setae shorther than paracuspis. Digitus short, slightly incurved in dorsal view and upcurved in lateral view; sparsely setose basodorsally, apex somewhat expanded, subcapitate in lateral view. Penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, blunt, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along truncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Integument black except mandibles and antennal flagellomeres partially reddish- brown, and T2 with four reddish integumental spots; spots highly variable in size, shape and color: large with posterior pair nearly confluent with anterior pair, to indistinct with posterior spots absent and anterior spots reduced to slightly brownish-red area on anterior margin; subrounded, subelliptical (anterior pair), subquadrate or nearly amorphous; reddish to orange. Body setae predominantly black varying in density except the following areas with silvery-white setae varying 124 in density: meso and metacoxae, ventral surface of meso and metafemora, posterior half of pronotum sublaterally, mesonotum sublaterally, propodeum sublaterally, T1, lateral areas of T2, lateral felt lines of T2, lateral margins of T2, fringe of T2–4 medially and laterally, T5–6 (except pygidial plate) medially, S1–5 (except fringe of S5); the silvery-white setae stripe of the mesosomal dorsum are variable, reaching the pronotum in some specimens and restricted to the mesonotum in others. MALE. Integument black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: hind tibiae and tarsi partially, metanotum medially, dorsum of propodeum, T1, T2 anterolaterally, fringes of T2–3 laterally, and S1–4 partly or entirely. Tibial spurs black. Wings dark- brown infuscated with basal portion partly colorless. Distribution. Bolivia, Paraguay, and Argentina. Material examined. (381#f, 57#m). Type material. Lectotype of Mutilla centralis, #f (nec #m), Argentina, Córdova [Córdoba] (MACN); Lectotype of Traumatomutilla fissiventris, #m, Argentina, Santiago Del Estero, environs d’Icano [outskirts of Icaño], E.R. Wagner (MNHN). Additional material. BOLIVIA: Santa Cruz, Cordillera, Las Juntas, Peredo,1#, II.1947 (MNCN); Cochabamba, Tarata, 1#f, 1900 (MNHN); Tarija, Villa Montes, 1#f, I.1931, Eisentraut S.G. (ZMB); Santa Cruz, Parapeti, 1#f, I.1960, Martinez (AMNH); PARAGUAY: Boquerón: Enciso, 550' [sic], 21°18'S 61°41'W, 2#f, 25–26.XI.2007, U. Dreschel (EMUS, FSCA); Mcal. [Marechal] Estigarribia, 560'[sic], 22°00'S 60°34'W, 2#f, 02–04.II.2008, U. Dreschel (EMUS, FSCA); E. [east of] La Patria, 2#f, 31.III.2006, U. Dreschel (FSCA, EMUS); Chaco: Loma Plata: 2f#, I.1956, Gerlach (AMNH, EMUS); 2#f, II.1993, Fritz (AMNH); Filadelfia: 8#f, I.1995, Arriagagda (AMNH); 2#f, X.1979, Fritz (AMNH); P.N. [Parque Nacional] Defensores del Chaco, 1#f, 05–17.XII.1982, Kochalka (DGMC); Agua Dulce, 1#f, X.1979, Fritz (AMNH); Pres. [Presidente] Hayes, Lolita, Yaragui, 3#f, 23°06'S 59°38', 05.III.2003, U. Dreschel (EMUS, FSCA); ARGENTINA: Ch. [sic] de Loria, 07.XII.1907, Jorgensen (ZMUC); Catamarca: 1#f, 9.II.1951 (AMNH); Frias, 2#f, 09.II.1951, Ross & Michelbacher (CASC); Miraflores, 1f#, II.1993, Di Lorio (AMNH); Coneta, 16 km S [kilometers South of] Catamarca, 6#m, 25.X.2003, Irwin & Parker (EMUS); Palo Labrado, 23 km S [kilometers south of] La Merced, 3#m, 24.IX.2003, Irwin & Parker (EMUS); Chaco: 2#f, IV.2003 (FSCA); Fuerte Esperanza: 8#f, XI.1978, Fritz (AMNH); 1#f, XII.1978, Fritz (AMNH); Tres Estacas, 1#f, Di Lorio (AMNH); Cordoba: 2f#, I.1974, Fritz y Martinez (MNCN); Balnearia: 3f#, II.1971,M.A. Fritz (AMNH, FSCA); 2f#, II.1970, Fritz (AMNH); Do [Departamiento] Sobremonte, 1f#, I.1974, Martinez (FSCA); Guanaco Muerto: 23#f, II.1971, Fritz (AMNH, EMUS); 15#f, II.1980, Fritz (AMNH, EMUS); 3#f, II.1980, Viana (AMNH); Cordoba, Ciudad, 1F3, XI.1970, Fritz (MNCN); Cordova, 2#f, W.M. Davis (MCZC); Cordoba, 1#f, II.1971, Fritz (AMNH); Cruz del Eje, Guanaco Muerto, 1#f, XI.1970, Martinez-Fritz (MNCN); Punilla, Valle Hermoso, 1#f, I.1943, M.J. Viana (MNCN); San Javier, La Paz, 1#f, 1–20.I.1929, C. Bruch (MNCN); Miramar, 1#f, III.1992, Fritz (AMNH); San Vicente de Cordoba, 1#f, J. Frenzel S. (ZMB); Dº [Departamiento] Calamuchita, Estancia La Granadilla, 1#f, 1974, Martinez (AMNH); Copacabana, 1#f, II.1980, Fritz (AMNH); Tanti, 1#f, II.1974, Martinez (AMNH); Las Lomitas, 1#f, 16.II.1948 (MNHN); 1#f, 07.II.1948 (MNHN); La Rioja: Patquia: (Chaco Serrana), 1#f, 06.XII.2002, L.A. Stange (FSCA); 1#f, K.J. Hayward (BMNH); XII.1932–I.1933, K.J. Hayward (BMNH); 2#f, I.1957 (LACM, EMUS); Aliumuyuna [sic], II.1993, Di Lorio (EMUS); 32#f, 1#m, I.1935, M. Gomes (SEMC, EMUS); 1#f, II.1934, M. Gomez (MNCN); Tallumuyana, 1#f, II.1993, di Lorio (AMNH); 1#f, 1#m, Mascasin, I.1959, Andrae (AMNH); Mendoza: Mendoza, 1#f, 04.I.1907, Jorgensen (ZMUC); Desaguadero, 1#f, I.1979, Williner (AMNH); Misiones: 2#f, E. Le Moult (MNCN); 2#m (RBINS); Missiones, Rio Salado, 1#m (CUIC); Salta: 1#m, 01–02.XI.1948 (BMNHB); La Viña: 11#f, 1#m, I.1984, Fritz (AMNH); 10#f, 6#m, II.1984, Fritz (AMNH); 8km N [north of] La Viña, 10#m, 26.X.2003, Irwin & Parker (EMUS); Sumalao: 5#f, III.1995, M.A. Fritz (AMNH); 1#m, 06.XI.2004, L.A. Stange (FSCA); Alemania, 7#f, 1#m, II.1983, Fritz (AMNH); Coronel Moldes: 2#f, II.1994, Fritz (AMNH); 4#m, I.1989, Fritz (AMNH, EMUS); Rosario [de] Lerma: 4#f, XII.1984, Fritz (AMNH); 1#m, XI.1984, Fritz (AMNH); Cafayate, 1#f, XII.1982, Fritz (AMNH); Cabra Corral, 1#f, II.1983, Fritz (AMNH); J.V. [Joaquin Victor] González, 2#f, XI.1984, Fritz (AMNH); San Luis: San Jeronimo, 11#f, II.1980, Fritz (AMNH); Alto Pencoso: 1#f, 22.XII.1908, Jorgensen (ZMUC); 2#m, 20.XII.1908, Jorgensen (ZMUC); Santiago del Estero: 2#f (DEI); 1#m, Konow (MNHN); Ojo de Agua, 1#f, XII.1984, Fritz (AMNH); Las Termas de Rio Hondo, 1#f, 1#m, 20.XII.1979, L.A. Stange (FSCA, DGMC); Anatuya, 9#f, III.1979, Fritz (AMNH, EMUS); Chaco de Santiago del Estero: Bañados de Rio Dulce, 60km O. D’Icano [kilometers west of Icaño], 1#f, 1909, E.R. Wagner (MNHN); Rio Dulce, 2#m (MNHN); 125
Rio Salado: 3#f, JanVIer [sic] (MNCN); Icaño, Bords du R. Salado [Banks of the Salado River], Environs D’Icano [outskirts of Icaño], 2#m, 1910, E.R. Wagner (MNHN); Tintina: 1#f, 21.XII.1898 (MNHN);1#m, 21.II.1920, Cornell Expedition (CUIC); 27k. S.W. [kilometers southwest of] Anatuya [Añatuya], 1#, 22–25.XI.1979, C. & M. Vardy (BMNH); 1k. N.E. [kilometers northeast of] Los Telares, 22–25.XI.1979, C. & M. Vardy (BMNH); Loreto, 3#f, I.1985, Fritz (AMNH); Sachayoj, 2#f, I.1993, González (AMNH); Monte Quemado, 1#m, 15.XII.1971, D.J. Brothers (DJBC); Tucuman:11 km N [kilometers north of] Cadillal, 1#f, 25.III.1990, J.G. Rozen & A. Roig (AMNH); 4 km S Capitan Caceres, 1#m, 24.X.2003, Irwin & Parker (EMUS); Las Cejas, 1#, 02.IV.1966, C.C. Porter (USNM); Lara, 4000m [meters above sea level], 1#, II.1903, G.A. Baer (MNCN); La Soledad, Cruz Alta, 1#m, 18.XII.1966, E. Bucher (AMNH); Jujuy, Ledesma, 2#f, XI.1978, Fritz (AMNH); Santa Fé, Tostado FCCN [sic] El Orden, 1#m, A.J. Giai (DJBC). An additional 136#f and 3#m from various localities in ARGENTINA were also examined (FSCA, EMUS, AMNH, DGMC, CASC, MNCN). Remarks. The sex association and synonymy of T. centralis with T. fissiventris was based on repetead co-ocurrence of both species in southern South America, particularly in Argentina where large series of Traumatomutilla were collected and in which T. centralis and T. fissiventris were the only females and males with a black meso- and metatibial spurs. The holotype of T. centralis boliviana was examined and found to be simply a variant of T. centralis with shorter mesosomal stripes and T1 completely covered by silvery-white setae. It is important to note that the male specimen labeled as the type of T. centralis in MACN is in fact Reedomutilla pubescens (Smith, 1879), which was first pointed out by Mickel (1964). Females of T. centralis are easily recognizable due to their slightly angulate and “box-like” mesosoma, similar to that of T. miniata (Gerstaecker, 1874) of the T. juvenilis species-group. The females also are a good example of the seemingly well- established color syndrome in the dry areas of Argentina, Paraguay, and Bolivia in which the body setae are predominantly black with well-defined and dense silvery-white setae markings, and the metasoma has bright red integumental spots. This pattern is subject to apparent gradients in the length of the mesosomal stripes and the size of the integumental spots of T2. Interestingly, the most conspicuous color variations found for this species appears to be restricted to the Cordoba Province in Argentina and in which the spots of T2 are reduced in sucha a way so as to appear completely absent.
Traumatomutilla contempta André, 1908a (Figs. 6A–F)
Traumatomutilla contempta André, 1908a: 198, lectotype [designated here], #f, Bolivia (MNHN), type examined. Traumatomutilla alhuampa Casal, 1969: 292, holotype, #f, Argentina, Santiago del Estero, Urutau (AMNH), type examined, syn. nov.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles, lateral face of propodeum evenly sculptured throughout, dorsal face of propodeum as long as posterior face, scutelar scale and anterolateral carinae greatly reduced, head clothed with black setae, stripes of setae on mesosomal dorsum silvery-white throughout. MALE. Unknown. Description. FEMALE. Body length 17 mm. Head. Posterior margin virtually straight. Occipital carina evenly equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view 1.15 × the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely and coarsely areolate-punctate to foveolate-punctate with somewhat scabrous intervals. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, reaching antennal tubercles and separated from lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.5 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Dorsal thoracic length 1.1 × mesosomal width. Anterior face of pronotum defined, longer than pronotal collar, with indistinct dense and coarse striations ventrad, and densely punctate dorsad; dorsal face roundly angulate into anterior face in lateral view. Humeral carina present separated from low rounded epaulet 126 virtuall flat against anterior margin of pronotum; antero-lateral corners of pronotum rounded in dorsal view. Lateral face of pronotum with dense micropunctures and few sparse punctures, and indistinct tubercle anterioventral in relation to pronotal spiracle. Pronotal spiracle slightly pronounced from lateral margin of pronotum. Mesopleuron sculpture partially concealed by dense setation; when visible, micropunctate anteriorly, and dense coarse areolate-punctate to foveolate- punctate along mesopleural ridge. Metapleuron concealed by dense setation except dorsal third asetose and unsculptured, smooth. Lateral face of propodeum densely foveolate-punctate to sparsely foveolate-puncate anterad with smooth shinning intervals, never wider than surrounding sculpture. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 72: 87: 82:63:60. Lateral margins of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutelar scale present virtually reaching propodeal spiracles laterally. Anterolateral carinae present, connected to each other thus forming a single transverse scale, connected to scutelar scale sublaterally. Propodeum convex, posterior face virtually as long as dorsal face; Metasoma. Ratios of width of T1, width of T2 and length of T2, 38:83:82. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible. S1 surface cuneiform, sparsely and coarsely punctate, with blunt longitudinal carina slightly higher posteriorly. S2 densely foveolate-punctate, sparser posteromediad; anteromedial crest-fold indistinct. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse interspersed micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Unkown. Coloration and variations. FEMALE. Integumenta black to brownish-black except mandibles and antennal flagellomeres reddish-brown ventrally, and T2 with four dark-red to orange integumental spots varying in shape and size. Body setae predominantly silvery-white varying in density, except for the following areas with black setae varying in density: head, pronotum, anterior half of mesonotum, posterior half of mesonotum medially, propodeum medially, anterior legs, meso and metafemora and tarsi, T1 medially, disc of T2 (except over integumental spots); fringe of T2–5 sublaterally, T6 laterally, fringe of S5, and S6. Certain specimens are predominantly clothed with black setae having the mesonotum, mesosomal pleurae, and lateral face of propodeum completely black; additionally, these specimens have the fringe of S2–5 completely clothed with black setae and the silvery-white setae of the metasoma reduced overall. MALE. Unknown. Distribution. Brazil, Bolivia, Paraguay, and Argentina. Material examined. (112#f) Type material. Lectotype of Traumatomutilla contempta, #f, BOLIVIA (MNHN); Holotype of Traumatomutilla alhuampa, #f, ARGENTINA, Santiago del Estero, Urutau (AMNH). Additional material. BRAZIL, Mato Grosso do Sul, 1#f (MZSP); BOLIVIA: 2#f (MNHN), Cochabamba, W. [West of] Saipina, 1#, II.1976, L. Pena (AMNH); PARAGUAY, Est. Hermoso [sic], 1#f, XI.1916, C. Olrog (MNCN); Boqueron, El Solitario, 1#f, 10.XI.2002, U. Dreschel (FSCA); 1#f, 15.XI.2002, U. Dreschel (FSCA); La Patria, 2#f, 22.VI.2003, U. Dreschel (FSCA); Nueva Asuncion, 20°42'S 61°55'W, 1#f, 17.V.2001, U. Dreschel (FSCA); Cruce Loma Plata, 2#, I.1991, Arriagada (AMNH); Chaco: Loma Plata, 2#f, I.1958, Gerlach (AMNH); Filadelfia, 1#f, I.1995, Arriagagda (AMNH); Pres. [Presidente] Hayes, Lolita, Yaragui, 11#f, 05.III.2003, U. Dreschel (FSCA); Puerto Galileo, 1#, 02–05.III.2006, U. Dreschel (FSCA); Catamarca: 1#f (MNHN); Punta [de] Balasta [Balasto], 2#f, II.1995, Arriagada (AMNH); Aconquija, Sierras áridas, 950m, 1#f, V.2005, G.E. Acuña (INPA); Santa Maria, 1#f, I.1994, Fritz (AMNH); Cordoba: Balnearia, 1#f, II.1971, M.A. Fritz (AMNH); Copacabana, 1#f, II.1980, M.A. Fritz (AMNH); Guanaco Muerto, 1#f, II.1971, M.A. Fritz (AMNH); Los Hornillos, Villa Dolores, 1#f, II.1952, M.A. Fritz (UMSP); Quilpo, 3#f, II.1980, M.A. Fritz (AMNH); Sierra de Cordoba, Capella del Monte, 2000 F [sic], 1#f, I–II.1888,J. Frenzel,ZMB; La Serranita, 1#f, Carpintero (AMNH); La Rioja: 2#f, E. Le Moult (MNCN); La Rioja, 1#, E. Giacomelli (CUIC); San Luis: Merlo, 1#f, Dureti (AMNH); San Jeronino, 1#f, M.A. Fritz (AMNH); Salta: Salta, 1#f, II.1997, Albeza (AMNH); La Viña, III.1985, Fritz (AMNH); Cafayate, 1#f, 10.II.1951, K.J. Hayward (MNCN); El Carmen, 1#f, I.1983, Fritz (AMNH); Alemania, 1#f, II.1992, Peña (AMNH); Puán, Finca del 127
Rey, 2#f, 02.XII.1952, A. Martinez (AMNH); Payogasta, 1#f, I.1993, Fritz (AMNH); Cnl. [Coronel] Moldes, Cabra Corral, 1200m [meters above sea level], 2#f, 18.III.1989, T. Osten (CESC); Chaco: 1#f, IV.2003 (FSCA); Charata, 1#f, I.1992, Di Lorio (AMNH); Tres Estacas, 1#f, Di Lorio (AMNH); Formosa, Pozo del Mortero, 1#f, 05.II.1948 (MNHN); Santiago del Estero: 1f# (DEI); Campo Gallo, 1#f, III.1945, Prosen (MNHN); Villa Unión, 1#f, III.1944, Prosen (MNHN); Loreto, 2#f, I.1985, Fritz (AMNH); Añatuya, 1#f, III.1979, Fritz (AMNH); Tucuman: 11 km N [kilometers north of] Cadillal, 2#f, 3.III.1990, J.G. Rozen & A. Roig (AMNH); La Soledad, Canete, 1#f, 06.II.1966, Bucher (AMNH); Tucuman, 1#f, Cornell Univ. [Univeristy] Expedition (CUIC); Jujuy: Pampa Blanca, 1#f, I.1983, Fritz (AMNH); Yuto, 1#f, 10.I.1966, H. & M. Townes (AEIC). An additional 35#f from various localities in PARAGUAY (AMNH, CMBC, UMSP, ZMUC) Remarks. The description of T. contempta by André (1908a) implies that there was a series of specimens in the MACN collection. We were unable to examine the MACN specimens and, therefore, we designate as lectotype the only specimen of this species labeled as “type” in André’s MNHN collection. Apart from minor color and setae characters, we found no structural differences between the types of T. contempta and T. alhuampa that would justify maintaining these as distinct species. This is the only consistently “large-bodied” species unassociated with a male in southern South America and could potentially be the female of T. protuberans. We refrain from synonymizing these species at this point for a number of reasons, which are discussed at length in the remarks of T. protuberans. The other possible explanation for this single female species in that region is that it might be a variant of T. ingens in which the body is less stout, more elongate, especially in the propodeum having the dorsal face as long as the posterior face, as opposed to the much shorter dorsal face in T. ingens. This is, however, a conspicuous and apparently consistent structural character, making this synonymy unlikely, especially considering the other minor sculpture and color differences observed between these species.
Traumatomutilla geographica (Gerstaecker, 1874) (Figs. 7A–F)
Mutilla geographica Gerstaecker, 1874: 302, holotype (by monotypy), #f, Brazil (ZMB), examined. Ephuta (Traumatomutilla) geographica: André, 1902: 55. Traumatomutilla geographica: André, 1904: 40. Traumatomutilla seabrai Casal, 1969: 283, holotype, #f, Brazil, Goiás, Aragarças (AMNH), type examined, syn. nov.
Diagnosis. FEMALE. Mesonotum narrower than distance between pronotal spiracles, lateral face of propodeum not evenly sculptured throughout, meso and metatibial spurs black, integumental spots of linear and yellowish, posterior pair confluent or nearly so. MALE.Unknown. Description. FEMALE. Body length 16 mm. Head. Posterior margin virtually straight. Occipital carina evenly equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely and coarsely foveolate punctate with indistinct scabrous intervals on front. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, reaching antennal tubercles and lateral scrobal carinae. Antennal tubercles irregularly rugose. Flagellomere 1 1.8 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Dorsal thoracic length 0.85 × mesosomal width. Mesosomal dorsum densely and coarsely areolate- punctate. Anterior face of pronotum defined slightly longer than pronotal collar, with dense indistinct and coarse longitudinal striations ventrad, coarsely and densely punctate dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, separated from projected subangulate epaulet; antero-lateral corners of pronotum subangulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum. Lateral face of pronotum densely and finely punctate with dense interspersed micropunctures, except for mostly unsculptured indistinct swelling anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and sparsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by 128 dense setation on ventral half, completely asetose, smooth on dorsal half. Lateral face of propodeum sparsely foveolate- punctate with unsculptured smooth intervals; intervals overall wider than surrounding sculpture. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 81:97:98:72:71. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post- spiracular area absent. Scutellar scale and anterolateral carinae virtually absent; scabrous intervals absent on scutellar area. Propodeum gibbose, dorsal face shorter than and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 39:83:90. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around blunt longitudinal medial carina slightly higher anteriorly. S2 densely foveolate-punctate, sparser poderomediad; anteromedial crest-fold indistinct. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse interspersed micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Unkown. Coloration and variations. FEMALE. Integument black, except antennal flagellomeres and mandibles partially reddish- brown, and T2 with four narrow linear yellow integumental spots; posterior pair variable, narrowly interrupted to broadly connected medially, thus forming single transverse line. Body setae predominantly black varying in density except the following areas with silvery-white setae varying in density: ventral third of lateral pronotal face, posteroventral half of mesopleuron, ventral half of metapleuron; posterior half of pronotum sublaterally, mesonotum sublaterally, dorsum of propodeum laterally; coxae and ventral surface of femora; T1 latearlly, lateral margins of T2, fringe of T2–4 medially and laterally, fringe of T5 laterally, T6 (except pygidial plate) medially, S1–4, and fringe of S2–3. Tibial spurs black. Distribution. Brazil. Material examined. (42 #f) Type material. Holotype of Traumatomutilla geographica, #f, BRAZIL, v. Olfers. (ZMB); Holotype of Traumatomutilla seabrai, #f, BRAZIL, Goiás, Aragarças, 06.I.1955, F.M. Oliveira (AMNH). Additional material. BRAZIL: 1#f, A.M. Parko, 13.248 [sic], 10/951 [sic] (MNRJ); Mato Grosso: Diamantino, Alto Rio Arinos, 1#f, X.1983 (MNRJ); Chavantina, Rio das Mortes, 1#f, XI.1946 (MNRJ); Rio Verde, BR-29, 1#f, XI.1960, M. Alvarenga (MPEG); Rosario Oeste, 8#f (MZSP); Utiariti,Rio Papagaios, 1#f, 1–12.XI.1966, Lenko & Pereira (MNCN); Goiás: 8#f (MZSP); Jataí, Faz. [Fazenda] Aceiro, 1#f, X.1962, Exp. Dep. Zool. [Expedição Departamento de Zoologia] (MNCN); Campinas, 1#f, XII.1936 (MNRJ); Serra da Mesa, 13°52'S 48°23'O, Campinaçu: 1#f, 18.II–02.III.1996, Silvestre, Brandão & Yamamoto (MZSP); 1#f, 01.III.1996, Silvestre, Brandão & Yamamoto (MZSP); Viannopolis, 1#f, XI.1931, R. Spitz (MNCN); Minas Gerais: 8#f (MZSP); Serra Cabral, 1#f, 1912, E. Garbo (MZSP); Aguas Vermelhas, 1#f, XII.1983 (MNRJ); Lassance, 1#f, 20–31.I.1939, Martins, Lopes & Mangabeira (MNCN); Pirapora, 1#f, 11–13.XI.1919, R.G. Harris (CUIC); São Paulo, Parque Est. [Estadual] [now, Área de Relevante Interesse Ecológico] Pé de Gigante, 1#f, 14.VII.1997, J.M. Carpenter (AMNH); Paraná, Jaguariahyva [Jaguariaíva], 1#f, I.1942, Justus (DZUP). Remarks. The types of T. geographica and T. seabrai are structurally identical, differing only in the posterior spots of T2, which are confluent in T. seabrai and narrowly separated in T. georaphica. No other color variations were observed in the specimens examined. This species is part of a color syndrome occurring in the Brazilian Cerrado and Caatinga that involves numerous species, such as T. unimarginata, T. parallela (=T. lineifera), T. mundula, T. rectilineata (André, 1898), T. ipanema (Cresson, 1902), T. solemnis (Cresson, 1902), and T. pereirai Suarez, 1960. Although T. geographica has black meso- and metatibial spurs, which facilitated other sex associations in this group, this character can also vary intraspecifically. One series of males collected in Chapada dos Veadeiros, Goiás, Brazil have the apex of the meso and metatibial spurs “blackish” and are from the same overall area that T. geographica is known to occur. Unfortunately, we haven’t seen enough distribution data to justify a sex association at this point.
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Traumatomutilla grossa (Gerstaecker, 1874) (Figs. 8A–F, 9A–G)
Mutilla grossa Gerstaecker, 1874: 73, lectotype [designated here], #f, Brazil, Cassapava [sic] (ZMB), examined. Mutilla abrupta Gerstaecker, 1874: 71, holotype (by monotypy), #f, Brazil, [Rio Grande do Sul], Alegrete (ZMB), examined, syn. nov. Mutilla characterea Gerstaecker, 1874: 319, holotype (by monotypy), #m, Brazil, [Rio Grande do Sul], Porto Alegre (ZMB), examined, syn. nov. Ephuta (Traumatomutilla) grossa: André, 1902: 55. Ephuta (Traumatomutilla) characterea: André, 1902: 54. Traumatomutilla grossa: André, 1904: 40. Traumatomutilla characterea: André, 1904: 40. Traumatomutilla abrupta: Nonveiller, 1990: 74.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles, sculpture of dorsum of mesonotum homogeneous throughout, longitudinal carina mesonotal and propodeal dorsum indistinct, dorsal face of propodeum much shorter and well differentiated from posterior face, scutelar scale and anterolateral carinae indistinct. MALE. Cuspis broad, club-like and laterally flattened, axillar projections acute, mesopleuron simply swollen on dorsal half. Description. FEMALE. Body length 17-18 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate, more sparsely and finely so on gena and malar space. Genal carina present, well-defined. Mandible oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, extending over antennal tubercles and connecting with reduced lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.7 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Length 1.2 × width. Mesosomal dorsum densely and coarsely areolate-punctate throughout; simply and densely punctate on scutelar area. Anterior face of propodeum defined, long, as long as pronotal collar, indistinctly and coarsely striated longitudinally at base with dense coarse punctures dorsally; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from low rounded epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum, rounded. Lateral face of pronotum densely punctate with dense interspersed micropunctures; indistinct blunt tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal half completely asetose, smooth. Lateral face of propodeum sparsely foveolate-punctate, with conspicuous unsculptured smooth areas. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 70:88:90:68:64. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae virtually absent, reduced to transverse irregular intervals delineating scutelar area which is densely and finely punctate; scabrous intervals absent on scutellar area. Propodeum gibbose, dorsal face much shorter than and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 38:85:84. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate- punctate to simply punctate with interspersed micropunctures where visible. S1 densely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in sub-sharp longitudinal carina equally high throughout. S2 sparsely foveolate-punctate, punctures conspicuously smaller and sparser medially; anteromedial crest-fold indistinct. S3–6 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where 130 visible. Pygidial plate broadly subpyriform, defined by lateral carinae at apical fourth of plate; surface mostly irregularly longitudinally rugose; interstice apparently granulose. MALE. Body length 16–20 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view. Width 0.7 × pronotal width. Eye almost circular. Ocelli small; OOD 4.8 × DLO, IOD 0.85 × DLO. Occipital carina distinct. Head surface densely and finely punctate, less densely so around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; sculpture concealed by dense setation; apical/ventral margin with a pair of medial subrounded free teeth. Scape bicarinate. Flagellomere 1 2.9 × pedicel length; flagellomere 2 3.6 × pedicel length. Mandible obliquely tridentate apically, medial tooth larger than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum rounded. Anterior face of pronotum sparsely punctate with interspersed micrpunctations laterally, with a conspicuous smooth unsculptured area basomedially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margin. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely foveolate-punctate; with longitudinal irregular carina medially. Axilla produced posterolaterally as short acute projections, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate, sculpture of lateral face gradually less defined to indistinct anterad; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum densely coarsely and confusedly foveolate- punctate with interspersed micropuncture; mesopleura conspicuously swollen on dorsal half, without any well-defined tubercles or projections; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations basally. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly, transversely and indistinctly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in slightly concave longitudinal carina terminating in subacute projection posteriorly. S2 coarsely and sparsely foveolate-punctate to punctate, with interspersed micropunctations anterolaterally; sculpture sparser and smalle posteromediad; S2 with conspicuous andteromedial subovate pit densely filled with setae; longitudinal anteromedial crest-fold present. S3–5 sparsely and coarsely foveolate-punctate with interspersed micropunctures; S6–7 sparsely foveolate-punctate. S7 longer than broad, well defined by lateral carinae throughout, posterior margin projected laterally and medially, medial projection longer than lateral projections and terminating in a pair of very small subacute closely spaced tooth-like structures on posterior margin. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 61:46:14; paramere virtually straight in dorsal view, except for apex slightly curved outward; upcurved apically in lateral view; with dense setae ventrally at basal half; cuspis broad, long, club-like, somewhat latearlly compressed, slightly wider apicad in dorsal and broader medially in lateral view, outer surface somewhat concave; with strong setae on ventral, outer and dorsal margins at apical half; paracuspis well-developed, not sessile, lobe-like, slightly wider than long with rounded apical margin and densely setose along posteroapical margin in lateral view predominantly, setae shorter than or as long as paracuspis; digitus short, curved inward in dorsal view and slightly upcurved in lateral view, sparsely setose dorsally at base, apex somewhat expanded in lateral view; penis valve strongly concave on internal surface, with well-defined pair of short teeth apicoventrally; apical tooth acute, subapical tooth subacute, externolateral pocket absent; apical distance between teeth 0.2 × length of valve; dense setae present along apical margin and inconspicuous setae present at base of subapical tooth on external surface. Colorationa and variations. FEMALES. Body and appendages predominantly black except most flagellomeres and mandibles partially reddish-black to reddish-brown; T2 with four sub-quadrate dark-orange to reddish spots. Tibial spurs brownish-yellow. Body setae predominantly silvery-white to silvery-golden varying in density, except the following areas 131 with black to brownish-black setae varying in density: front, dorsal half of gena, occipital area, dorsum of pronotum, mesonotum medially, scutelar area, dorsum of propodeum medially, dorsal third of lateral face of pronotum, dorsal half of mesopleuron anteriorly, lateral face of propodeum, T1 medially, most of disc of T2, fringes of T2–4 sublaterally, fringe of T5 laterally, fringe of S4–5, and tarsi ventrally. MALES. Integument predominantly black to brownish-black, except mandibles partially reddish-brown. Body setae silvery-white to silvery-golden varying in density, except following areas with black to brownish-black setae varying in density: front partially, areas around eyes, gena, malar space, ventral surface of head, lateral face of pronotum, most of mesonotum, axillar projections, anterior half of scutellum, metanotum laterally, dorsal third of mesopleuron, most of metapleuron, posterior two thirds of disc of T2, fringes of T2–5 medially, T6–7, fringe of S5, and S6–7. Wings hyaline-brownish slightly darker at apical third, costa, and around veins. Distribution. Brazil, Argentina, and Uruguay. Material examined. (86#f, 23#m) Type material. Lectotype of Mutilla grossa, #f, BRAZIL, [Rio Grande do Sul], Cassapava [Caçapava do Sul] (ZMB); holotype of Mutilla abrupta, #f, BRAZIL, [Rio Grande do Sul], Allegrette [Alegrete] (ZMB); holotype of Mutilla characterea, #m, BRAZIL, [Rio Grande do Sul], Porto Alegre (ZMB). Additional material. BRAZIL: 1#f (MNCN); Paraná: Bexiga, Cachoeira, 1#f, 22.XI.1955, Correa,S. (DZUP); Ponta Grossa, 1#f, II.1944, Justus (DZUP); Jaguariahyva [Jaguariaíva], 1#f, 28.XII.1966, Justus (DZUP); Rio Grande do Sul: 1#m (MNHN); 5#f (MZSP); Pelotas: 1#f (MNCN); 1#f, 09.II.1958, J. Lucia Mantovani & Biezanko (MNCN); 1#m, IV.1964, Mantovani, Biezanko (USNM); 1#m, 5.III.1945, Biezanko (USNM); 1#f, 05.II.1964, C.M. Biezanko (MZSP); Porto Alegre: 1#f, 1937, Pe.[Padre] Buck (CESC); 1#f, 1935, Pe. [Padre] Buck (CESC); 1#f, 15.VIII.1937, Pe. [Padre] Buck (CESC); 1#f, 01.XI.1937, Pe. [Padre] Buck (CESC); 1#f, 24.XI.1935, Pe. [Padre] Buck (CESC); 1#f, 03.X.1934, Pe. [Padre] Buck (CESC); 1#f, 03.XI.1948, Pio Buck (MNCN); São Francisco de Paula, 2#f, 24.I.1999, M.S. Barbosa (CESC); São Leopoldo, 1#f, X.1947, Pe. [Padre] Buck (MNCN); Taquara, Pituva, 1#f, 10.X.1973, F.R. Meyer (CESC); Entre Rios: Dpto. [Departamiento] Colón, Parque Nacional: 6#m (AMNH); 2#f, III.1982, Fritz (AMNH); Pronunciamiento, 1#f, XII.1957, M.R. Zelich (FSCA); Liebig, 22#f, Zelich (AMNH); Feliciano, 1#f, XII.1972, Fritz (MNCN); Io [Primero] de Mayo, 1#f, III.1957, Gontero (USNM); Cordoba, La Granja, 1#f, 12.I.1947, M.A. Fritz (UMSP); Rosario, 2#f, Dr. Stempelmann (ZMB); San Vicente de Cordoba, 1#f, J. Frenzel S. (ZMB); Patagonia, Bahia Blanca, 1#f (DEI); Catamarca, 1#f (USNM); URUGUAY: 1#f, Cornell Univ. [University] Expedition (CUIC); Menendez [sic], 2#m, I.1983, S. Guido (AMNH); Montevideo: Montevideo: 1#f, 1#m (MNHN); 1#f, IX.1820 (MNHN); Treinta y Tres, Rio Olimar Chico, 25 km WSW [kilometers West by southwest of] Treinte-y-tres [Treinta y Tres], 1#m, 19.IV.1963, J.K. Bouseman (AMNH); Quebrada de los Cuervos, 1#f, XII.1952, Carbonell (AMNH); Isla Patrulla, 1#f, V.1965, Gambardella (AMNH); Artigas: 1#f, Carbonell (AMNH); Aº [arroyo] Cuaró, 1#f, IV.1960, L. do Ximenez (AMNH); Colonia, Colonia, 1#f, 05.I.1943, P.A. Berry (CISC); Rio Negro, 1#f, Rincón de Porrua, IV.1952, Acosta y Lara (AMNH); Cerro Largo, Paso de Aguiar, 1#f, XII.1952, Carbonell (AMNH); Tacuarembó, Puntas,Aº [arroyo] Laureles, 1#ff, XII.1965, Carbonell (AMNH). An additional 16#f and 8#m from various localities in ARGENTINA were also examined (AMNH, LACM, USNM, MNCN, DGMC). Remarks. The association between T. grossa and T. characterea was based on repeated co-occurrence of both species in the same areas of southern South America, especially in Uruguay, where they were the only species of Traumatomutilla seen in certain areas. We have seen specimens that vary considerable in the overall aspect of the body and there appears to be more slender form of T. grossa that was formerly named T. abrupta. The types of both species are identical in structural and color characters. Even though the type of T. grossa is remarkably more robust and stouter than the slender T. abrupta, the proportion of the mesosoma and metasoma are consistent. Additionally, this species seems to be restricted to the Pampas and adjacent areas, being found only in Argentina, Uruguay and Brazil, with its northernmost record in São Francisco de Paula in Rio Grande do Sul state, Brazil.
Traumatomutilla guayaca Casal, 1969 (Figs. 10A–F, 11A–F)
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Traumatomutilla guayaca Casal, 1969: 284, holotype, #f, Ecuador, Esmeraldas, Parr. San Mateo (AMNH), examined. Traumatomutilla tayguaya Casal, 1969: 285, holotype, #f, Colombia, Valle Let. Ang. [sic] (AMNH), examined, syn. nov.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles, anterolateral corners of pronotum sharply angulate, scutelar scale and anterolateral carinae indistinct, dorsal face of propodeum as long as posterior face, lateral face of propodeum mostly unsculptured smooth and shinning. MALE. Mesopleuron simply swollen on dorsal half, axillar projections acute, scutellum simply convex without distinguishable dorsal and posterior faces, S2 without setae- filled pit, pronotum with dense micropunctures. Description. FEMALE. Body length 10–14 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 1.1 × the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate to areolate-punctate where visible; sculpture conspicuously sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, broadly separated from antennal tubercles and indistinct lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible; simply micropunctate mediad; with poorly defined medial longitudinal carina; dorsal face of propodeum with medial longitudinal carina; scutelar area simply punctate. Anterior face of pronotum defined, slightly longer than pronotal collar, indistinctly and coarsely striated longitudinally basad and with dense coarse and confused punctures dorsad; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from slightly projected rounded epaulet, anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; with indistinct blunt tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and sparsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal half completely asetose, smooth. Lateral face of propodeum sparsely foveolate-punctate, with large smooth, shinning intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 62:69:68:50:47. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post- spiracular area absent. Scutellar scale and anterolateral carinae indistinct, reduced to irregular transverse outlines on scutelar area; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face longer than and slightly angulate rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 35:85:91. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around blunt longitudinal medial carina slightly higher medially. S2 densely foveolate-punctate, more sparsely and finely so posteromediad; anteromedial crest-fold indistinct. S3–6 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate where visible. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Body length 13 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 4.8 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface virtually concealed by dense setation, sparsely and finely punctate where visible. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; surface densely concealed by setation; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Flagellomere 1 1.8 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. 133
Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctures, marginally concave basomedially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, densely and finely foveolate- punctate; with longitudinal unsculptured area anteromedially; dorsal and posterior faces poorly distinguished. Axilla produced posterolaterally as short acute projection, with slightly concave and coarse dense foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, partially concealed by dense setation, densely areolate where visible; sculpture of lateral face indistinct to absent anterad; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum densely micropunctate; mesopleuron with indistinct blunt swelling on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; simply micropunctate anterad; sparsely foveolate punctate with smooth intervals posterad. Metapleuron smooth throughout, except for indistinct areolations on basal fourth, micropunctate on dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. Ratios of width of T1, width of T2 and length of T2, 50:98:78. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible; Pygidial plate indistinctly granulose apicad, unsculptured and smooth basad, slightly broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in longitudinal slightly concave carina with short spine-like projection posteriorly. S2 sparsely foveolate-punctate to punctate; sculpture sparser posteromediad; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 64:52:13; paramere virtually straight in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half; cuspis thin, slender, elongate, equally wide throughout in dorsal view, slightly narrower apicad in lateral view; with scattered inconspicuous long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, longer than wide, with subrounded and sparsely setose posterodorsal margin in lateral view; setae longer than or as long as paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, apparently asetose basodorsally, apex somewhat expanded, subcapitate in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, with externolateral pocket; apical distance between teeth 0.1 × length of valve; sparse setae present along subtruncate posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Integument black except mandibles and antennal flagellomeres partially reddish- brown, and T2 with four yellowish integumental spots; Spots varying in shape and color: linear to subrectangular or elliptical; yellowish to orange. Body setae predominantly silvery-white varying in density except the following areas with black setae varying in density: frons, pronotal dorsum, dorsal half of lateral face of pronotum, mesonotum laterally, dorsal half of mesopleuron, propodeal dorsum laterally, lateral face of propodeum, T1 medially, disc of T2 (except over integumental spots), fringe of T2–5 sublaterally, T6 (except pygidial plate) laterally, and S1–5: some specimens have the silvery-white areas better defined and somewhat denser than others, the head completely covered with silvery-white setae, and the mesosomal lines of silvery-white setae broader and longer. MALES. Integument black to brownish-black except mandibles partially reddish-brown. Body setae predominantly silvery-white varying in density, except following areas with black setae varying in density: propodeum posterolaterally; mesoscutum, axillar projections, scutellum basally, posterior half of T2, fringe of T2 partially, T5–7 (except pygidial plate) medially. Tibial spurs yellowish-white. Wings mostly light brown infuscated, basal third hyaline, apical third conspicuously darker and with faint purplish reflections. Distribution. Colombia, Venezuela, French Guiana, and Ecuador. 134
Material examined. (16#f, 25#m) Type material. Holotype of Traumatomutilla guayaca, #f, ECUADOR, Esmeraldas, Parr. San Mateo (AMNH); Holotype of Traumatomutilla tayguaya, #f, Colombia, Valle Let. Ang. [sic] (AMNH); Additional material. VENEZUELA: 1#f (MNHN); Bolivar, 50 km ESE [kilometers east by southeast of] Ciudad Bolivar, Santa Rita Ranch, 1#f, 19.VI–08.VII.1999, T. Alten (UCRC); Ribero, Cariaco, 1#f, Santschi (MNHN); Guárico, Hato Masaguaral, 44km S Calabozo, 1#f, 20–28.V.1985, Menke & Carpenter (EMUS); Aragua: 2 km N [kilometers North of] Ocumare de la Costa, 1#m, 31.III.1981, Hollenberg & Menke (USNM); Puerto de Cata, 3#m,10.VI.1976, Vincent & Menke (USNM); Zulia, El Tucuco, 45 km SW [kilometers southwest of] Mathiques, 1#m, 5.VI.1976, Vincent & Menke (USNM); COLOMBIA: 6#m (IAvH); Magdalena: 6#m (IAvH); 26 km E [kilometers east of] Santa Maria, 2#m, 15.II.1979, R.C. Wilkerson (FSCA); Rio Frio, 1#m, 26.VII.1927, G. Salt (BMNH); Meta, PNN La Macarena: Mesetes, 3°22'N 74°02'W, 580m [meters above sea level], 1#m, 23.XII.1993, F. Fernandez (IAvH); La Curia, 3°20'N 73°53'W, 550m [meters above sea level], 1#m, 25.XII.1995, F. Fernandez (IAvH); Res. [Reserva] La Naca Rena [La Macarena] Mpio [sic], 1#f, 17.VII.1988, W. Cubillos (AMNH); Res. [Reserva] Nat. [Natural] El Caduceo, H.C. [sic], Savanna, 03°40N 73°39'W 1400'[sic], 03–08.I.2012 (EMUS); Cord. [Cordillera] Macarena, 01–15.III.1976, M. Cooper (BMNH); 1#f, 15–26.II.1976, M. Cooper (BMNH); Bolívar, Sambrano [Zambrano], 2#f, VIII.1992, M. Andrea (AMNH); M. forestral [sic], 9°37'N 74°54'W, 10m [above the groun], 1#m, 23.IV.1993, F. Fernandez (IAvH); Guajira, Dibulla, 1#f, Forel (MNHN); Vichada, PNN [Parque Nacional Natural El] Tuparro, Cumarida, 315m [meters above sea level], 1#m, 30.XII.1998, W. Villalba (IAvH); Madre de Diós, PNN [Parque Nacional Natural] Tayrona Neguanje, 11°20'N 74°02'W, 10m [above the ground], 21.III–05.IV.2001, R. Henriquez (CSCA); Arauca, Tame., 2#f, 20–27.VII.1976, M. Cooper (BMNH); FRENCH GUIANA, [Cayenne], Cayenne, 1#f (MNHN). Remarks. There are four widespread species of the T. indica species-group in the northwestern Amazon, T. indica, T. selligera, T. parallela, and T. guayaca. After the sex associations of T. indica, T. selligera, and T. parrallela were established (detailed in their respective remarks sections), the only male and female of this group to be left unassociated in the northwestern Amazon were T. guayaca and its previously undescribed male. Therefore, the sex association of T. guayaca was achieved through a process of elimination as well as repeated co-ocurrence in multiple Colombian and Brazilian areas. Additionally, T. guayaca is the only species of Traumatomutilla to be recorded west of the Andes in Ecuador, apart from T. vitelligera (Gerstaecker, 1874). The holotypes of T. guayaca and T. tayguaya where examined, compared, and found to be structurally identical differing only the mesosomal stripes of silvery-white setae which are broader and longer in T. tayguaya, and the head setae which silvery-white only on the vertex in T. guayaca. Though this species also superficially resembles and can occur in the same areas as T. selligera, it is distinct when comparing the scutellar armature and sculpture of the lateral propodeal face in the females and the scutellum of the males.
Traumatomutilla impetuosa (Smith, 1879) n. comb. (Figs. 12A–G)
Mutilla impetuosa Smith, 1879: 220, holotype [by monotypy], #m, Brazil, Para [sic] (BMNH), examined. Mutilla impetuosa: André, 1902: 73 (incertae sedis).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half, axillar projections acute, scutellum simply convex, pronotal sculpture with sparse micropunctures. Description. FEMALE. Unknown. MALE. Body length 11 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.85 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 3.6 × DLO, IOD 1.4 × DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate; conspicuously denser and coarse on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Antennae lost. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or 135 ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations laterally, simply micropunctate sublaterally, and virtually unsculptured smooth and shinning medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, without dorsal and posterior faces, densely and coarsely foveolate-punctate; with conspicuous longitudinal flat unsculptured area anteromedially. Axilla produced posterolaterally as short acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, partially concealed by dense setation, densely areolate where visible; lateral face virtually unsculptured, smooth and shinning on anterior half; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleuron with short blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on ventral margin and indistinct rugosities on dorsal margin. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length 0.7 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible. Pygidial plate missing. S1 longitudinally elevated medially, terminating in low longitudinal slightly concave carina; carina higher posteriorly. S2 coarsely and sparsely foveolate-punctate to punctate, with micropunctures laterally and conspicuous elongate setae filled pit medially; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate- punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 58:48:15; paramere virtually straight in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half; cuspis thin, slender, elongate, equally wide throughout in dorsal view, slightly narrower apicad in lateral view; with scattered inconspicuous long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, somewhat petiolate, virtually as long as wide, subtriangulate and with densely setose posterodorsal margin in lateral view; setae longer than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, subcapitate in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short acute teeth posteroventrally; externolateral pocket present, greatly reduced; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Unknown. MALES. Integument black to brownish-black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: clypeus, ventral surface of tibiae and basitarsi, ventral surface of meso and metafemora, coxae, propodeal dorsum, T1, basal third of T2, lateral margins and felt line of T2, fringe of T2–3 laterally, S1–4, and fringe of S2–3. Tibial spurs yellowish-white. Wings dark brown infuscated throughout, with faint purplish reflections at apical third of forewings. Distribution. Brazil. Material examined. (1m#) Type material. Holotype (by monotypy), #m, Brazil, Para [sic] (BMNH). Remarks. This male is quite unremarkable and though its differences in relation to other males of the T. indica species- group can be considered slight variations, there are no obvious putative females that can be safely associated with it. Any association is also especially difficult because the only distribution information about T. impetuosa is “Brazil, Para” which means that the type locality could be any area in the current states of Pará, Amapá, and possibly Amazonas and Roraima which, between 1850 and 1879 (when the specimen was likely collected) were all part of a single state. Comparing male 136 morphology allows us to hypothesize that this is either an extreme variant of the male of T. guayaca, for which we have not yet found an intermediate form, or the male of an undescribed female.
Traumatomutilla indica (Linnaeus, 1758) (Figs. 13A–C, 14A–G)
Mutilla indica Linnaeus, 1758: 583, holotype [by monotypy], #f, Columbia [sic] (LSUK), examined (by photograph). Mutilla diadema Fabricius, 1787: 311, holotype [by monotypy], #f, Suriname [synonymized by Mickel, (1952)] (ZIN) Mutilla sphegea Fabricius, 1804: 435, holotype [by monotypy], #m, [Guiana], Essequibo [sic], ZMUC 00241654, (ZMUC), examined (by photograph), syn. nov. Ephuta (Traumatomutilla) sphegea: André, 1902: 56. Ephuta (Traumatomutilla) indica: André, 1902: 55. Traumatomutilla sphegea: André, 1904: 40. Traumatomutilla indica: André, 1904: 40.
Diagnosis. FEMALE. Mesonotum narrower than distance between pronotal spiracles, lateral face of propodeum evenly sculptured, scutelar scale and anterolateral carinae well-defined, longitudinal carina on mesonotal and propodeal dorsum well-defined, dorsal face of propodeum longer than posterior face. MALE. Mesopleuron tuberculate on dorsal half, axillar projection acute, scutellum angulate with well-defined dorsal and posterior faces, dorsal face of scutellum terminating in transverse bilobate carina. Description. FEMALE. Body length 12–18 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate where visible; sculpture sparser on gena and malar space; with micropunctures on malar space and ventral half of gena. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, reaching antennal tubercles and separated from poorly defined lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.3 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible; with conspicuous medial longitudinal carina extending from anterior margin of mesonotum to scutelar area; sculpture of mesonotum slightly less defined around medial longitudinal carina. Anterior face of pronotum defined, virtually as long as pronotal collar, indistinctly and coarsely striated longitudinally at base and with dense coarse and confused punctures dorsad; dorsal face roundend into anterior face in lateral view. Humeral carina well-defined, broadly separated from low subangulate epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle strongly projected from lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with sparse indistinct micropunctures and with indistinct blunt tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely finely punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal half completely asetose, smooth, unsculptured. Lateral face of propodeum denseley foveolate-punctate; sculpture sparser posteroventrad. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 69:92:85:64:61. Lateral margin of mesonotum strongly constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma. Scutellar scale well-defined, connected laterally to anterolateral carinae; anterolateral carinae connected, thus forming single transverse carina reaching propodeal spiracles laterally; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 40:84:78. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent 137 laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around blunt longitudinal medial carina equally high throughout. S2 densely foveolate-punctate, sculpture sparser posterad; anteromedial crest-fold greatly reduced. S3–6 sculpture mostly concealed by dense setation, densely and finely foveolate- punctate where visible. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with coarse and irregular rugosities; interstice granulose. MALE. Body length 12–18 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 4.0 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate; conspicuously denser and coarser on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; surface completely concealed by dense setation. Scape bicarinate. Flagellomere 1 1.9 × pedicel length; flagellomere 2 2.5 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations laterally, simply micropunctate sublaterally, and virtually unsculptured smooth and shinning medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum gibbose, densely and coarsely foveolate-punctate; with well-defined dorsal and posterior faces; dorsal face flat, ending in transverse bilobate carina at posterior margin. Axilla produced posterolaterally as short acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, completely concealed by dense setation; lateral face densely areolate; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and shallowly punctate with dense interspersed micropunctures; mesopleuron with short conspicuous tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron predominantly unsculptured, smooth, shinning, except dorsal fifth indistinctly rugose and basal fifth coarsely foveolate- punctate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length 0.8 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate irregularly, transversely and indistinctly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in longitudinal carina with conspicuous subacute spine-like projection posteriorly. S2 coarsely and densely foveolate-punctate to punctate, with conspicuous elongate setae filled pit anteromedially; longitudinal anteromedial crest-fold present. S3–6 sparsely foveolate-punctate; S7 densely foveolate- punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projection bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 106:28:75; paramere mostly straight and curved outward apically in dorsal view; upcurved apically in lateral view; with sparse setae ventrally at basal half; cuspis slender, virtually straight, equally wide in dorsal and lateral view; with short, ventrally directed setae at apical fourth; paracuspis well-developed, not sessile, lobe-like, virtually as wide as long with rounded and densely setose posteroapical margin in lateral view, setae shorter than or as long as paracuspis; digitus short, curved inward in dorsal view and upcurved in lateral view, setose dorsally at basad, apex somewhat expanded in lateral view; penis valve strongly concave on internal surface, with well-defined pair of short acute teeth apicoventrally; externolateral pocket present; posteropical margin truncate, conspicuously “shelf-like” in dorsal view; apical distance between teeth 0.1 × length of valve; dense setae present along posteroapical margin and inconspicuous setae present at base of subapical tooth on outer surface. 138
Coloration and variations: FEMALES. Integument black except mandibles and antennal flagellomeres partially reddish- brown, and T2 with four linear to subrectangular yellowish spots. Body setae predominantly silvery-white varying in density except the following areas with black setae varying in density: frons, dorsal half of gena, pronotum (except ventral third of lateral face), anterior half of mesonotum, posterior half of mesonotum medially, dorsal third of mesopleuron, propodeum medially, lateral face of propodeum, T1 medially, disc of T2 (except over integumental spots), fringe of T2–4 sublaterally, T5–6 (except pygidial plate) laterally, fringe of S5, and S6. Certain specimens have the lateral areas of silvery-white setae on the mesonotum extending and virtually covering the lateral expansion of the mesonotal margin or slightly advancing over scutelar area between the propodeal spiracles. MALES. Integument black. Body setae predominantly silvery-white varying in density except the following areas with black setae varying in density: mesosocutum, tegulae, axillar projections, dorsal face of scutellum, posterior half of T2, T5–7 (interspersed with silvery- white setae on T5–6), fringe of S6, and S7. Distribution. Belize, Colombia, Trinidad, Venezuela, Guiana, Suriname, French Guiana, Brazil, and Peru. Material examined. (48#m, 459#f) Type material. Holotype of Mutilla indica, #f, Columbia [sic] (LSUK); Holotype of Mutilla sphegea, #m, [Guiana], Essequibo [sic], Smidt, ZMUC 00241654 (ZMUC). Additional material. BELIZE, Benque Viejo, 1#f, Father Stanton (MCZC); COLOMBIA: 2#f, 302-40 [sic] (MNHN); 2#m (MNHN); Meta: PNN [Parque Nacional Natural] Sierra de La Macarena, Caño Curía, Sendero Cachicamos, 3º21'N 73º56'W, 460m [meters above sea level], 2#f, 21.XII.2002, M. Duarte (IAvH); 1#f, 24.II.2003, W. Villalba (IAvH); PNN [Parque Nacional Natural] Tinigua, Caño Nevera, 2º11'N 73º48'W, 390m [meters above sea level], 5#f, 7.II.2002, C. Sánchez (IAvH); Cundinamarca, PNN [Parque Nacional Natural] Sumapaz, Porvenir Marayal, 3º48'N 73º49'W, 535m [meters above sea level], 2#f, 30.IV.2002, H. Vargas (IAvH); TRINIDAD, St. [Saint] George, St. [Saint] Augustine, 1#m, 15.VI.1976, J.S. Noyes (BMNH); VENEZUELA, Monagas, 42km SE [kilometers southeast of] Maturín: 3#f, 03.VII.1958, Arnold Menke (EMUS); 1#f, 09.VII.1958, Arnold Menke (EMUS); 1#m, 8.VII.1958, A.S. Menke (LACM); 1#m, 08.VII.1958, Arnold Menke (EMUS); Bolivar, Canaima, 1#f, 28.IX.1974, C. Murtaugh (UMSP); 115 km N [kilometers north of] Santa Elena de Uairen, 1#f, 07.III.1987, Miller & Stange (FSCA); 2 km E [kilometers east of] Kavanayen, 1#f, 06.III.1987, R.B. Miller & L.A.Stange (FSCA); Valle Ariba, El Pajui, 1#f, 25.XI.1970, D. Fairchild (FSCA); Caracas, Canaima, 28–30.XII.1968, A. Michelbacher (CISC); Monagas, 42 km SE [kilometers southeast of] Maturin, 2#f, 18.VII.1958, A.S. Menke (FSCA); GUIANA: 1#m, A.W. Bartlett (BMNH); jct. Cuyani Mazaruni [junction of Cuyani-Mazaruni rivers], Essequibo Rivers, 2#f, 1927, R.E. Fuglestad (CISC); Georgetown, 2#f, III.1955 (MNRJ); Demerara, Rio Demerara, 1#f, 1823, Janson & Sons (MNCN); Dubulay Ranch, 05°40.95'S 57°51.52'W, 1#f, 1#m, 15–18.I.1999, M. Sharkey & B. Brown (EMUS); Blairmont, 1#m, 1923, H.E. Box (USNM); Camaria, 1#m, 30.VII.1924, W. Myers (AMNH); Mahaica, 1#m, 20.VI.1927 (CUIC); Rockstone, 1#m, 9.VII.1911 (AMNH); SURINAME: 5#f (MZSP); 2#f (MCZC, MNHN); Paramaribo, 1#f, 24.X.1997, P.H. v. Doesburg (MNCN); Sandry Field, 1#f, 05–08.IX.1943, D.G. Hall (USNM); Tibitl, Savanne, 1#f, 05.I.1949, Surinam exp. [expedition] (MNCN); Marowinje, Christian Kondre, 3#f, 1963, B. Malkin (CASC); Coebiti, 1#f, 04–14.VIII.1980, B.V. Ridout (BMNH); Kwakoegron Saramaccan, 1#f, 08.VI.1927 (USNM); FRENCH GUIANA: 1#f (MNHN); Cayenne, 4#f, 2#m (MNHN, ZMUC, DEI); Maroni, 4#f, 1917, E. Le Moult (MNCN); Kourou, 1#f, 1914, R. Benoist (MNHN); 1#m, 4.VI.1977, D. Roubik (SEMC); Roches de Kourou, 2#f (MNHN); Kourou, Degrad Saramanka, Km5, 4#m, IX.2005, D. Faure (EMUS); Riv. [Riviére] De Kourou, 4#f, E. Le Moult (MNCN); Nouveau Chantier, 1#f (MNCN); Agarouany [sic], 1#f, 15.I (MNHN); Gourdonville: 1#f, Juillet [July] (MNHN); 1#f, X.1914, R. Benoist (MNHN); Paranaiba, 3#f, I.1880 (MCZC); Pariacabo, Riv. [Riviere] De Kourou, 1#f, VI, Le Moult (MNCN); Montsinery, 04°55.995'N 52°29.689'W, 1#f, 23.I.2003, V. Soon (EMUS); Chavrein, Baixo Maroni, 1#m, VIII (MNHN); cote Atlantique, camp Warana, 2#m, 13.VII.1996, BRAET (DGMC); Wismar, 1#m, 4.VI.1934, S. Dinkus (AMNH); BRAZIL: 3#f (ZMUC, ANSP); Amapá, R. [Rio] Oiapoque, 2#f, V.1959 (MNRJ); Roraima, PARNA [Parque Nacional] Viruá, Vicinal na Estrada Perdida, 01°28'N 60°58'O, 1#m, 24.XII.2015–07.I.2016, REDE BIA [Biodiversidade na Amazônia], Rafael et al. (INPA); Amazonas, Manaus, Reserva Ducke, AM010, Km26, 02°55'46.3''S 59°58'29.3''W: 1#f, 03.VI.2016, M.A.S. Nascimento (INPA); 1#m, 29.IX–10.X.2005,Aguiar, A.,INPA; Sítio Zé Nildo, AM010, Km 35, Ramal Cinco, 02°50'54''S 59°56'02.42''W, 1#f, 16.VI.2016, F.J. Telias (INPA); Pará: Mangabeira, Mocajuba, 1#f, VIII.1953, O. Rego (FSCA); Barcarena,Caripi, 01°29'40''S 48°42'35''W, Mata, 1#m, 13–22.xi.2001, I.S. Gorayeb, A. Tavares, N. Bittencourt, 139
J.O. Dias (MPEG ); Cach. [Cachoeira] Paciencia, A. [sic] Cuminá, 1#f, 27.X.1956 (MNRJ); Rio Xingu Camp. [sic], 60 km S [kilometers South of] Altamira, 1#m, 8.X.1986, Spangler & Flint (USNM); Belém, 3#f, VI.1924, F.X. Williams (UMSP); Ilha de Marajó, Igarapé Taperebá, 1#f, VII.1958, Vanzolini (MNCN); Ananindeua, 1#f, 1976 (MNRJ); Maranhão: Caxias, Reserva Ecológica do Inhamum, Povoado Coités, 04°54'41''S 43°25'29''W, 1#f, 27–29.V.2011, J.A. Silva & R.M. Servian (CZMA); Barreirinha, Parque Nacional Lençóis Maranhenses, 02°39'80''S 42°49'88''O, 1#f, 07.V.2016, J.A. Rafel e F.L. Oliveira (INPA); Centro Novo do MA [Maranhão], Reserva Biológica do Gurupi, 03°14'05''S 46°41'83''W, 2#f, 01–06.III.2011, F. Limeira-de-Oliveira & D.W.A. Marques (CZMA); São Pedro da Água Branca, Fazenda Santa Rosa, 05°07'07''S 48°15'19''W, 1#f, 06.XII.2001, J.A. Rafael, F.L. Oliveira & J. Vidal (INPA); Rio Grande do Norte: Mossoró, Faz. [Fazenda] Sta. [Santa] Júlia, 05°01'10''S 37°22'56''W, 12#m, 03.III.2007, D.R.R. Fernandes e equipe, (LRRP); João Câmara, Cauçu, 1#f, 05.V.2012, Lopes, M.C.A. (UFRN); Macaíba, Escola de Agricola Jundiaí, 1#f, 02.XI.2010, Mesquita, A.D.S. (UFRN); Natal, Praia de Búzios, 1#f, 18.VI.2009, James, H. (UFRN); Nísia Floresta, Estrada para FLONA [Floresta Nacional], 1#f, 22.IV.2005, Azevedo, D.L.O. (UFRN); Parnamirim, EMPARN [Empresa de Pesquisas Agropecuárias do Rio Grande do Norte], 1#f, 11.III.2011, Rufener, M.C. (UFRN); Piauí, Parque Nacional de Sete Cidades, 04.09909°S 41.70952°W, Sede 193 m.a. [sic], 19–25.IV.2012, Oliveira & Somavilla (INPA); Ceará: 5#f (MZSP); Carquejo, 3#f, III.1963, Dirings (MZSP); Russas, Perímetro Irrigado, 04°59'39.14''S 37°58'46''W, 1#f, 16.IX.2013, D. Nogueira (INPA); Paraíba, Santa Luzia, Brandão Junco, 1#f, VIII.1956 (MNRJ); Mato Grosso, Barra do Tapirapé, 2#f, 01–15.I.1966, B. Malkin (AMNH); Kingu [Xingú] River, 1#f, 1962, J. Goddard (BYU); Bahia: Tremembe, 1#f, 24.VI.1997, J.R.M. Santos (CPDC); Ilha Boipera, Cairu, 1#f, 31.XII.2000, J. Fabricio (CPDC); Jequié, 1#f, 12.II.2002, J.R.M. Santos (CPDC); Sergipe, Villa Nova [Neópolis], 1#f, XII.1934 (MNRJ); Espírito Santo, Praia das Neves, 1#f, 27.IV.1998, Raw, A. (DZUP); Rio de Janeiro: Ponta Monica, 1#, 1.II.1977 (MNRJ); Cabo Frio, 1#f, 5.I.1984 (MNRJ); Niteroi, Itaipu-Assu, 1#f, 11.XI.1945 (MNRJ); Santa Catarina, 5#f (MZSP); PERU: Lima, Valle Chanchamayo, 1#f, 1939, P. Vaquero (UMSP). An additional 333#f and 7#m from various localities VENEZUELA, SURINAME, BRAZIL, and PERU have also been examined (FSCA, CASC, EMUS, DEI, ZMB, INPA, MNRJ, FSCA, MNCN, LACM, FMNH, BMNH, MZSP, CZMA, ZMUC, UFRN) Remarks. This is perhaps the most easily recognizable and one of the least variable species of Traumatomutilla. Though T. indica occurs from Belize to Santa Catarina state in Brazil, no variations were observed for the males and only slight differences in the setal pattern of the mesosomal dorsum of the females were recorded. The association and synonymy of T. sphegea and T. indica was based firstly on couples collected in Trinidad along with couples of T. selligera which are somewhat restricted to the northern areas of the Amazon whilst T. indica and T. sphegea are found throughout the Amazon and, in smaller numbers, in other South American environments. Day (1979) did not provide any clues regarding the type of M. indica even though he provided valuable info on T. americana (Linnaeus, 1758), which was also described by Linnaeus and would likely be housed in the same collection. Even though the specimen at LSUK is labelled as having “no type status”, it is placed in the list of Linnean types of that institution and it matches the morphology of all reference material provided by André and Mickel, who likely studied the type to identifiy their own specimens. Given the aforementioned data and that Linnaues (1758) did not provide any locality information other than “habitat in Indiis”, and the specimen housed at LSUK is almost certainly the holotype of T. indica.
Traumatomutilla ingens (André, 1903) (Figs. 15A–C, 16A–F)
Ephuta (Traumatomutilla) ingens André, 1903: 452, holotype, #f, Argentina, Santiago del Estero (MNHN), examined. Mutilla tristis Burmeister, 1875 (nec Klug, 1821): 471, #f [nec #m], [synonymizd by André (1908c)] (MACN). Traumatomutilla ingens: André, 1904: 40.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles, sculpture of lateral face of propodeum sparser anteroventrad, tibial spurs white, dorsal face of propodeum much shorter than virtually vertical posterior face. 140
MALE. Cuspis club-like, laterally flattened, and conspicuously broader apicad in lateral view, scutellum gibbose with dorsal face shorter than posterior face, mesopleuron tuberculate on dorsal half, axillar projections acute. Description. FEMALE. Body length 20 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.9 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Head sculpture mostly partially concealed by dense setation, densely and coarsely foveolate-punctate to punctate where visible; sculpture less dense and coarse on gena and mala space. Genal carina present, well-defined. Mandible oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, extending over antennal tubercles and narrowly separated from poorly defined and irregular lateral carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.55 × pedicel length; flagellomere 2 1.65 × pedicel length. Mesosoma. Length 0.8 × width. Mesosomal dorsum densely and coarsely areolate-punctate with smooth intervals; medial longitudinal carina on mesonotum present, well-defined, not reaching scutelar area; medial longitudinal carina present on dorsal face of propodeum; areolations on mesonotum slightly less defined adjacent to longitudinal carina; scutelar area simply and densely punctate. Anterior face of pronotum defined, long, as long as pronotal collar, indistinctly and coarsely striated longitudinally ventrad, densely and coarsely punctured dorsad; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from slightly projected and rounded epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle narrowly projected from lateral margin of pronotum, rounded. Lateral face of pronotum densely punctate with dense interspersed micropunctures, slightly swollen anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely coarsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by dense setation, except dorsal third completely asetose, smooth. Lateral face of propodeum densely foveolate-punctate to throughout with smooth shinning intervals, never wider than surrounding foveolations. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 64:85:83:63:58. Lateral margin of mesonotum constricted anterior to propodeal spiracle, strongly diverging anterad, mesonotum with expanded lateral margin. Propodeal spiracle slightly projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae indistinct, irregular, reduced to transverse irregular intervals narrowly delineating scutelar area which is densely and finely punctate; scabrous intervals absent on scutellar area. Propodeum gibbose, dorsal face virtually apparently shorter than posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 21:42:41. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures throughout; foveolations slighlty sparser and larger laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, completely concealed by dense setation. S1 densely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in sub-sharp longitudinal medial carina slightly higher posteriorly. S2 densely and finely foveolate-punctate, with conspicuous unsculptured medial longitudinal area; anteromedial crest-fold indistinct. S3–6 sculpture completely concealed by dense setation. Pygidial plate broadly subpyriform, defined by lateral carinae at apical third of plate; surface mostly irregularly longitudinally rugose; interstice apparently granulose. MALE. Body length 20 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margin subparallel posterior to eyes, not contiguous with eyes outline, 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 3.9 × DLO, IOD 1.1 × DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate, less densely so around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, slightly convex dorsomedially, virtually flat ventromedially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded free teeth. Scape bicarinate. Flagellomere 1 2.3 × pedicel length; flagellomere 2 2.9 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum rounded. Anterior face of pronotum densely punctate with interspersed micrpunctations throughout. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and finely foveolate- 141 punctate, parapsis present, notaulus vesitigial, both reduced to posterior half of mesoscutum. Scutellum subglobose, with well-defined dorsal and posterior faces, densely and coarsely foveolate-punctate; with broad longitudinal unsculptured “carina-like” area medially. Axilla produced posterolaterally acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate, sculpture of lateral face gradually less defined to indistinct anterad; dorsal and posterior faces indistinguishable. Lateral face of pronotum densely and coarsely punctate with dense interspersed micropunctures; mesopleura with conspicuous rounded blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate- punctate to foveolate-punctate with interspersed micropunctures along anterior margin. Metapleuron sparsely micropunctate to smooth throughout, except for basal fourth densely and coarasely areolate-punctate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.4 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; pygidial plate irregularly, transversely and indistinctly rugose apicad, granulose basad, broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in slightly concave blunt longitudinal carina, slightly higher posteriorly. S2 coarsely and sparsely foveolate- punctate to punctate, with interspersed micropunctations laterally, conspicuous medial subovate pit densely filled with setae; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially, terminating in a pair of very small subacute closely spaced tooth-like structures on posterior margin. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 60:37:11; paramere marginally sinuous in dorsal view, upcurved apically in lateral view; with sparse setae ventrally on basal half; cuspis broad, ellongate club-like, laterally compressed at apical half, slightly wider apicad in dorsal view and conspicuously wider apicad in lateral view, outer surface somewhat concave; with strong setae apically on ventral, outer and dorsal margins; paracuspis well- developed, not sessile, lobe-like, subrounded, virtually as wide as long with rounded posterior margin and sparsely setose along posterior margin in lateral view predominantly, setae overall shorter than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded in lateral view, knob-like; penis valve with inner surface strongly concave, with well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, externolateral pocket present, reduced; apical distance between teeth 0.1 × length of valve; dense setae present along truncate posterior margin and inconspicuous setae present at base of subposterior tooth on external surface. Colorationa and variations. FEMALES. Body and appendages predominantly black to brownish-black, except most flagellomeres and mandibles partially reddish-black to reddish-brown; T2 with four longitudinally subovate orange to reddish integumental spots. Body setae predominantly black varying in density, except for the following areas with silvery-white varying in density: posterior half of mesoscutum laterally, mesopleuron posteroventrally, metapleuron predominantly, propodeal dorsum laterally; T1 laterall, T2 disc (except areas in between integumental spots), fringe of T2–3 medially and laterally, fringe of T4–6 medially, S1–3, and fringe of S2–3. Tibial spurs yellowish-white. MALES. Integument black, except antennal flagellomeres and mandibles partially reddish-brown. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: metanotum medially, propodeal dorsum, anterolateral corners of T2, fringe of T2–4 laterally, S1–4, and fringe of S2–3. Tibial spurs yellowish-white. Wings light brown infuscated, except basal third hyaline, with faint purplish reflections. Distribution. Paraguay and Argentina Material examined. (21#f, 8#m) Type material. Lectotype, #f, Argentina, Santiago del Estero (MNHN). Additional material. BRAZIL: 1#f, Kriechbaum (MNHN); Mariposa [sic], 1#f, 1915, M. Pic (MNHN); Minas Gerais, Sertão de Diamantina, Fazenda das Melancias, 1#f, 10.XI.1902, E. Gounelle (MNCN); PARAGUAY, Chaco, Teniente Enciso, 1#f, 02.XI.2001, G. Arriagada (FSCA); ARGENTINA: Santiago del Estero: 1#f, Konow (MNHN); Banks of the Salado river, outskirst of Icaño: 2#f, 1910, E.R. Wagner (MNHN); 1#, XII.1910, E.R. Wagner (MNHN); 25 km north of Icaño, Marshes 142 of the Salado river, Paso de Don Jose, 2#f, decembre, janvier, 1910, E.R. Wagner (MNHN); Rio Salado, 6#f, Décembre (MNCN); Salta, La Viña: II.1984, Fritz (AMNH); 1#f, III.1984, Fritz (AMNH); 1#f, 22–25.XII.1983, M. Wasbauer (AMNH); 3#m, II.1984, Fritz (AMNH); 1#m, II.1993, Fritz (EMUS); 1#m, II.1988, Fritz (AMNH); 1#m, II.1987, Fritz (AMNH); 1#m, I.1985, Fritz (AMNH); 1#m, I.1984, Fritz (AMNH). Remarks. Traumatomutilla ingens is the third species in a small apparent sub-group, which includes T. aemulata and T. grossa, and is characterized by large and stout bodied species in both sexes and males with a highly modified club-like cuspis. The sex association was made possible due to distribution and “morphological clues” provided by T. aemulata and T. grossa. A large bodied male morphospecies was found in relatively large numbers in Salta, Argentina, having the typical club-like cuspis of T. characterea, from which it differs mainly in the tuberculate mesopleuron (simply swollen in T. characterea). Coincidentally, the only large-bodied female to occur in the same area was T. ingens, which shares the overall body shape, indistinct scutelar armature, and mesosomal sculpture of T. grossa. Burmeister (1875) identified a few females in MACN as T. tristis (Klug, 1821) and described the male of that species based on three males, all from Argentina. André (1908c) disassociated the sexes, stating that the females belonged to his recently described species T. ingens but the males labeled as T. tristis by Burmeister could not be confidently associated with the females because two of them belonged to T. foveiventris and the third belonged to T. characterea.
Traumatomutilla mundula (Cresson, 1902) (Figs. 17A–C)
Mutilla mundula Cresson, 1902: 54, lectotype [designated by Cresson (1916)], #f, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH) Ephuta (Traumatomutilla) mundula: André, 1902: 55. Traumatomutilla mundula: André, 1904: 40.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles; lateral face of propodeum sparsely sculptured, with wide conspicuous smooth intervals; longitudinal carinae of mesonotum/dorsal face of propodeum indistinct; integumental spots of T2 yellowish, linear, anterior pair greatly reduced to absent, posterior pair confluent, froming single transverse line. MALE. Unknown. Description. FEMALE. Body length 8-12 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view 1.15 × the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate where visible; sculpture sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, extending over antennal tubercles and narrowly separated from lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.3 × pedicel length; flagellomere 2 2.0 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible with somewhat rounded intervals; medial longitudinal carina present on mesonotum, extending from posterior margin of pronotum, narrowly interrupted in scutelar area, and nearly connecting with similar carina on dorsal face of propodeum; sculpture of mesonotum slightly less defined around medial longitudinal carina; scutelar area sculpture densely and coarsely punctate. Anterior face of pronotum defined, slightly longer than pronotal collar; with indistinct, indistinct and coarse longitudinal striations at base and dense coarse and confused punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, broadly separated from slightly projected rounded epaulet, anterolateral corners of pronotum subrounded in dorsal view. Pronotal spiracle projected from lateral margin of p ronotum, rounded. Lateral face of pronotum densely punctate with dense interspersed micropunctures; with indistinct swelling anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and sparsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by dense setation on ventral 143 half, completely asetose, smooth on dorsal half. Lateral face of propodeum sparsely and coarsely foveolate-punctate along anterior and posterior margin, with conspicuous unsculptured area medially bearing longitudinally oblique indistinct rugosities. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 72:91:87:69:64. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae virtually absent; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 41:89:98. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around sub-sharp longitudinal medial carina equally high throughout. S2 densely foveolate-punctate, more sparsely and finely so anteromediad; anteromedial crest-fold indistinct. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with interspersed micropunctures where visible; S5–6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Unkown. Coloration and variations: FEMALES. Integument black to brownish-black, except mandibles and antennal flagellomeres partially reddish-brown, and T2 with single linear transverse yellowish integumental stripe on posterior fourth. Some specimens have a pair of variably developedand longitudinally linear similar stripe on anterior third. Tibial spurs yellowish-white. Body setae predominantly silvery-white varying in density, except the following areas with balck setae varying in density: MALES. Unknown. Front, vertex, and dorsal half of gena; anterior half of pronotum, posterior half of pronotum medially and laterally, and lateral face of pronotum predominantly; mesonotum medially and laterally; dorsal half of mesopleuron and metapleuron; propodeal dorsum medially; T1 medially, disc of T2 (except over integumental spots), fringe of T2–5 sublaterally, T6 (except pygidial plate) laterally; fringe of S5, and S6. Distribution. Brazil. Material examined. (6#f) Type material. Lectotype: #f, BRAZIl, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Paralectotype: f#, same label data as lectotype (CMNH). Additional material. BRAZIL, Mato Grosso, Chapada dos Guimarães, 1#f, 26.III.2001, Sousa, W.O. (DZUP); Goiás, Campinas, 2#f, XII.1936 (MNRJ); Minas Gerais, 1#f (MZSP). Remarks. Females of Traumatomutilla mundula are similar in structure to T. puella (=T. peperina and T. manca), differing only in the integumental spots of T2 for which the anterior pair is absent or greatly reduced, and the posterior pair form a single transverse band (Fig. zzz). No definitive intermediate color forms between T. mundula and T. puella have been recorded (e.g. specimens with posterior spots separated, reduced, and posterior spots more elongated). Additionally, T. mundula is known mainly from the same areas and type locality as T. manca for which there are several specimens, including the type series, and from which no variation approximates the color pattern of T. mundula. Therefore, we maintain T. mundula as a discrete species, but will not be surprised if discovery of the male or intermediate forms lead to their eventual synonymy with T. puella.
Traumatomutilla parallela (Klug, 1821) (Figs. 18A–C, 19A–J, 20A–G)
Mutilla parallela Klug, 1821: 315, holotype [by monotypy], #f, Brazil, Para [sic] (ZMB), examined. Mutilla optata Cresson, 1902: 53, lectotype [designated by Cresson (1916)], #f, Brazil (CMNH), examined. Mutilla almada Cresson, 1902: 74, holotype [by monotypy], m#, Brazil, [Pará], Santarém (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) almada: André, 1902, 54. Ephuta (Traumatomutilla) optata: André, 1902: 55. 144
Ephuta (Traumatomutilla) parallela: André, 1902: 55. Ephuta (Traumatomutilla) lineifera André, 1903: 451, lectotype [designated here], Brazil (MNHN), examined, syn. nov. Traumatomutilla almada: André, 1904, 40. Traumatomutilla lineifera: André, 1904, 40. Traumatomutilla optata: André, 1904, 40. Traumatomutilla parallela: André, 1904, 40. Traumatomutilla indicoides Mickel, 1945: 42, holotype, #f, Bolivia, Santa Helena [sic] (USNM), examined, syn. nov. Traumatomutilla gausapata Mickel, 1952: 129, holotype #f, Guyana, Rupununi savannah [sic] (UMSP), examined, syn. nov. Traumatomutilla pillinata Casal, 1969: 285, holotype, #female, Brazil, Goiás, Aragarças (AMNH), examined, syn. nov.
Diagnosis. FEMALE. Distance between pronotal spiracles conspicuously wide than mesonotal width; propodeal spiracles strongly projected from lateral margin of mesosoma; lateral face of propodeum densely sculptured, intervals oblqieuly and indistinctly striate; MALE. Cuspis elongate, slender; mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; axillar projections acute; scutellum gibbose with well-defined dorsal and posterior faces; pronotum and propodeum with dense silvery-white setae obscuring integument. Description. FEMALE. Body length 17 mm. Head. Posterior margin slightly convex. Occipital carina evenly equally wide throughout. Vertex width 0.7 × pronotal width. Eye almost circular, its length in frontal view 1.2 × the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely, coarsely and confusedly areolate-punctate to foveolate-punctate with irregular intervals; sculpture overall sparser on gena and malar space; front with irregular longitudinal carinadorsomedially. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, reaching antennal tubercles, connected to irregular lateral scrobal carinae. Antennal tubercle coarsely punctate to irregularly rugose. Flagellomere 1 1.8 × pedicel length; flagellomere 2 1.4 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum densely and coarsely areolate-punctate, with irregular intervals and narrow longitudinal carina extending virtually uninterrupted from anterior margin of mesonotum to posterior margin of dorsal propodeal face; sculpture adjacent to medial carina densely punctate. Anterior face of pronotum defined slightly longer than pronotal collar, with dense indistinct and coarse longitudinal striations ventrad, coarsely and densely punctate dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina prominent, narrowly from produced subangulate epaulet; antero-lateral corners of pronotum subangulate in dorsal view. Pronotal spiracles strongly produced from lateral margins of pronotum. Lateral face of pronotum densely punctured with dense interspersed micropunctures, except low micropunctate swelling antero-ventral in relation to pronotal spiracle. Mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate along mesopleural ridge where visible. Metapleuro concealed by dense setation on ventral half, unsculptured and smooth to indistinctly rugose on dorsal half. Lateral face of propodeum densely and coarsely foveolate- punctate, with indistinctly rugose intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 92:128:120:87:92. Lateral margin of mesosoma strongly constriced anterior to propodeal spiracle, strongly divergent anterad thus expanded laterally. Propodeal spiracle strongly projected from lateral margin of mesosoma. Post-spiracular area absent. Scutellar scale present as wide as and connected laterally to anterolateral carinae which are connected thus forming a single straight transverse carina. Cariniform reticulations on scutelar area absent. Propodeum convex, dorsal face shorter than and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 51:112:113. Disc of T2 partially concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures where visible; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; micropuncutres nearly absen on T6. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around short longitudinal medial carina equally high throughout. S2 densely foveolate- punctate; foveolations denser and larger laterad; anteromedial crest-fold well-defined. S3–4 sculpture mostly concealed by 145 dense setation, densely and coarsely foveolate-punctate with sparse interspersed micropunctures where visible; S5–6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Body length 13–16 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.85 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 5.8 × DLO, IOD 1.5 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate with dense interspersed micropunctures; sculpture conspicuously sparser around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, virtually flat elsewhere; densely and confusedly punctate; apical/ventral margin with a pair of short subrounded free teeth medially. Scape bicarinate. Flagellomere 1 1.7 × pedicel length; flagellomere 2 1.9 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum subangulate. Anterior face of pronotum sparsely punctate with interspersed micropunctures throughout. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum slightly gibbose, with well-defined dorsal and posterior faces; densely and coarsely foveolate-punctate with broad medial longitudinal, slightly elevated, “carina-like” area on dorsal face; unsculptured area conspicuously broadened apicad. Axilla in dorsal view produced posterolaterally as short acute projection, coarsely and densely foveolate-punctate dorsally. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, partially concealed by dense setation, densely areolate where visible; slightly depressed sublaterally at base; sculpture of lateral face less defined to indistinct along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and indistinctly punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt tooth-like projection on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; simply micropunctate anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on ventral fourth and indistinct rugosities in dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate coarsely and indistinctly rugose with granulose interstices, broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in longitudinal carina with short spine- like projection posteriorly. S2 coarsely and sparsely foveolate-punctate to punctate, with indistinct micropunctures laterally; sculpture slightly sparser posteromediad; without setae filled pit, at most with small anteromedial tuft of silvery- white setae; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 sparsely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projection bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 76:53:22; paramere marginally sinuous in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half; cuspis slender, elongate, equally wide throughout in dorsal view and lateral view; with scattered inconspicuous long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, longer than wide with subtriangulate and sparsely setose posterodorsal margin in lateral view; setae shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, blunt, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. 146
Coloration and variations. FEMALE. Integument black to brownish-black, except mandibles and antennal flagellomeres apartially reddish-brown, and T2 with four yellowish integumental spots; T2 spots shape and size highly variably: posterior pairs from transversely rectangulate to linear, narrow, completely confluent medially, thus forming single transverse line on T2; anterior pair from longitudinally subelliptical to linear, elongate, separated from posterior pair by a fourth of its length. Spots color varying from bright yellow to opaque orange. Body setae highly variable, usually predominantly silvery-white varying in density, except the following areas with black setae varying in density: head, pronotal dorsum, lateral face of pronotum (except ventral third), mesonotal dorsum medially, propodeal dorsum medially, T1 medially, disc of T2 (except over integumental spots), fringe of T2–5 sublaterally (sometimes extending to lateral margins of T5); T6 medially (except pygidial plate); fringe of S5, and S6. Some specimens have a transveser band of silvery-white setae covering the vertex; the silvery-white longitudinal lateral stripes of silvery-white setae vary in width and length, from reaching the midlle of pronotal dorsum to ending before pronotal spiracles. MALE. Integument black to brownish-black, except flagellomeres and mandibles partially reddish-brown. Body setae predominantly silvery-white varying in density, except following areas with black setae varying in density: mesoscutum, axillar projections, scutellum, posterior half to posterior third of T2, fringe of T5 medially, T6–7 (except pygidial plate), fringe of S6, and S7. Tibial spurs yellowish-white. Wings light brown infuscated on apical third, hyaline with brown veins elsewhere. Distribution. Colombia, Venezuela, Brazil, Peru, Bolivia, Paraguay, and Argentina. Material examined. (307#f, 54#m) Type material. Holotype of Mutilla parallela, #f, BRAZIL, Para [sic] (ZMB); holotype of Mutilla almada, m#, Brazil, [Pará], Santarém (CMNH); lectotype of Ephuta (Traumatomutilla) lineifera, f#, BRAZIL (MNHN); holotype of Traumatomutilla indicoides, #f, BOLIVIA, Santa Helena [sic] (USNM); holotype of Traumatomutilla gausapata, #f, GUYANA, Rupununi savannah [sic] (UMSP); holotype of Traumatomutilla pillinata, #female, BRAZIL, Goiás, Aragarças (AMNH). Additional material. COLOMBIA, Vichada, Gaviotas Sabana, Mpio. [sic] del Viento 170m [above sea level], 1#m, II.1995, Cortez & Mora (IAvH); VENEZUELA: Monagas: 42 km SE [kilometers southeast of] Maturin, 1#f, 03.VII.1958, A.S. Menke (LACM); 1#f, 09.VII.1958, A.S. Menke (FSCA); Zamora: Libertad, 1#f, 1923, Mayeul Grisol (MNHN); Suapure, Caura River, 1#f, 25.VI.1900 (EMUS); 1#f, 09.IX.1899 (EMUS); 1#f, 03.XII.1900 (EMUS); BRAZIL: Minas Gerais: Berizal, Faz. [Fazenda] Veredão, em flores, 850m (above sea level), 15°39’54’’S 41°39’56’’W, 1m#, 11.XII.2012, J.A. Rafael (INPA); Araxá, 1f#, 6.XI.1965, Elias,C. & Elias, T. (DZUP); Paraopeba N.o proc.16/38 [sic], 1f# (MNRJ); Belo Horizonte, 3f#, 06–11.V.1993, S.D. Gaimari (INHS); Uberlândia, Cachoeira do Mirandão, 1f#, 7.IX.2005, Rocha-Filho, L.C. (DZUP); Janaúba, Poção de Santa Cruz, 1f#, 26.I.2008, Melo, G.A.R. & Costa (DZUP); Berizal, Faz. [Fazenda] Veredão, 1f#, 14.XII.2007, Grossi, P., Rafael, Parizotto, D. (DZUP); Bom Despacho, Bom Despacho, 1f#, VII–IX.2000, Ramos, L.S., Marinho, C.G.S. (CPDC); Campos de Diamantina, Fazenda do Riacho Fundo, 1f#, XII.1902, E. Gounelle (MNCN); Sertão de Diamantina, Fazenda das Melancias, 1f#, 10.XI.1902 (MNCN); Lagoa Santa, 1f#, Reinhardt (ZMUC); Lassance, 1f#, 05.VI.1911 (UMSP); Santa Rita do Cedro, 1f#, 1904, E. Gounelle (MNCN); 15 km SE [southeast] de Riacho dos Machados, 1f#, 12.IV.1998, Melo, G.A.R. (DZUP); Ponto dos Volantes, 14 km ao S [South] de Ponto dos Volantes, 1f#, 12.II.2010, Melo, G.A.R., Parizotto, D., Grossi, P. (DZUP); Mato Grosso: Rosário do Oeste, 1f#, 1970, Hummelgen (DZUP); Serra Roncador, 1f#, 11.VII.1968, Laroca & Azevedo (DZUP); Gleba Uirapuru, 1f#, 12.VII.1972, Mielke & Brown (DZUP); Barra do Tapirape, 1f# (MZSP); Utiariti, 1f# (MZSP); Estação Ecológica Iquê, 1f#, 16.V.1984, P.T. Nascimento (MPEG); Nova Lacerda, 14°28'38''S 59°33'30''W, 1f#, 27.IV.2006, J.A. Rafael & F.F. Xavier Filho (INPA); Nova Mutum, Fazenda Buriti, 1f#, 9.VII.1997, Ribeiro, G.C., Mendes, H.F. (DZUP); marg. esq. [margem esquerda] rio Sucuriu, Faz. [Fazenda] Canaã, Três Lagoas, 1f#, IV.1967, F. Lane (MZSP); Andradinha, 2f#, VIII.1971, Oliveira, F.M. (DZUP); P. N. [Parque Nacional] do Xingu, Jacaré, 1f#, XI.1961, Alvarenga & Werner (DZUP); Confluencia Xingu Kuluene, 2f#, VI.1947 (MNRJ); Santo Antônio da Barra [sic], 1f# (MNHN); Vacaria [sic], 1f#, XII.1922 (DEI); Sinop, 12°31'S 55°37'W, 1m#, X.1974, M. Alvarenga (EMUS); Cáceres, 1m#, 27.III.1985,Elias, C. (DZUP); Chapada dos Parecis, 1m#, 15.XII.2001 (EMUS); Mato Grosso do Sul: Chapada dos Guimarães, Chapada dos Guimarães, 1f#, 17.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); Porto Murtinho, 1f#, XI.1929 (DEI); Selviria, UNESP [Universidade Estadual de São Paulo] Farm, 1f#, 16.I.1990, C.A.H. Flechtmann (EMUS); Tres lagoas, Horto Rio Verde, Champion, 1f#, 27.X.1994, C.A.H. Flechtmann (EMUS); Corumbá, Imbirissú, 1f#, 01.XII.1960 (MNCN); Maranhão: Mirador, Parque Estadual do Mirador: Base da Geraldina, 06°37'25''S 45°52'08''W, 147
1f#, 30.VII–06.VIII.2006, F. Limeira-de-Oliveira, T.T.A. Silva & A.A. Santos (CZMA); Base dos Cágados, 06°48’29’’S 45°06’34’’W, 1m#, 27.IX–02.X.2011, F. Limeira-de-Oliveira & D.W.A. Marques (CZMA); Cândido Mendes, Fazenda Sete Irmãos, 01°51’37’’S 49°46’10’’W, 1m#, 03–06.X.2008, F. Limeira-de-Oliveira, J.A. Rafael & P.A.M. Moraes (CZMA); Bom Jardim, Reserva Biológica do Gurupi, 1f#, 17–27.I.2010, F. Limeira-de-Oliveira, J.T. Câmara & M.B. Aguiar Neto (CZMA); Caxias, Reserva Ecológica do Inhamum, 1f#, 10.VIII.2005, F. Limeira-de-Oliveira et al. (CZMA); Carolina, Balneario Urupuxete, 2f#, 08–09.I.2008, F. Limeira-de-Oliveira (CZMA); Imperatriz, 1f#, 1.VII.1974, Exp. Dept. Zoo. [Expedição Departamento Zoologia] - UFPR [Universidade Federal do Paraná] (DZUP); Roraima: Boa Vista, Cbio [sic] UFRR [Universidade Federal de Roraima], 1f#, 11.IX.2014, P.C.S. Barroso (UFRR); Serra do Tepequém, Estrada do Igarapé do Barata, 2f#, 2m#, 14.XII.2015, Rafael et al. (INPA); ESEC [Estação Ecológica] Maracá, L4 - 2500 - P.50 – 250 [grid coordinates], 1f#, 24.II.2007 (INPA); Alto Alegre, ESEC Maracá, 03°21'59''N 61°26'04''W, 8m#, 01– 15.IV.2016, R. Boldrini e J.A. Rafael (INPA); Serra Grande, 2f#, 11–20.X.1992, D.W. Davis (EMUS); Amajari, Estância Ecológica do SESC, Serra do Tepequém, 03°44'46''N 61°43'40''W, 1m#, 15–31.III.2016, R. Boldrini e J.A. Rafael (INPA); Caracaraí, P.N. [Parque Nacional] Viruá, 01°29'23.3''N 61°00'08''W, 4m#, 19.IV.2015, J.A. Rafael, R.A. Heleodoro, D.M.M. Mendes, D.W.A. Marques e C. Maldaner (INPA); Pacaraima, 1m#, 05–08.III.1988, Eq. [Equipe] J.A. Rafael (INPA); Rondônia: Rancho Grande, 62 km S [South of] Ariquemas, 1f#, 08–20.IV.1997, A. Rehn & C. Alexander (UCDC); Vilhena, 1f#,17.XII.1986, Elias,C. (DZUP); 62 km SW [southwest of] Ariquemas, nr [near] Fzda [Fazenda] Rancho Grande: 1f#, 03–15.XII.1996, J.E. Eger (FSCA); 5m#, 08–20.XI.1994, W.J. Hanson (EMUS); Pará: Mangabeira, Mocajuba, 25f# (MNRJ); 1f#, III.1955 (MNRJ); Gorotire, Xingú, 2f#, 18.IV.1983, W. Overal (MPEG); [Santarém], Altér do Chão, 1f#, 16.X.1987, Izeldinha Miranda (INPA); [Ilha de] Marajó, Soure, 1m#, O. Bertram S.V. (ZMB); Goiás: 24 mi E [miles east of] Formosa, 1f#, 16.V.1956, F.X. Truxal (LACM); Leopoldo Bulhões, 1f#, XII.1933, Spitz (MNCN); Serra da Mesa, 13°52'S 48°23'O, 1f#, Campinaçu, 02–15.XII.1995, Silvestre, Diaz & Campaner (MZSP); Trindade, 3f#, IX.1892 (MNHN); Jatai, 1f#, XI.1971, Oliveira, F.M. (DZUP); Niquelândia,14°01'S 48°18'O, 1f#, 24.IX–06.X.1995 (MZSP); Rio Grande do Norte: Extremoz, 1f#, 07.IX.2007, Barca, R.R.B. (UFRN); Macaíba, E.A.J. [Escola Agrícola Jundiaí] UFRN [Universidade Federal do Rio Grande do Norte], 1f#, 07.V.2010, Castro, E.S. (UFRN); Parnamirim, Mata do Jiqui, 1f#, 28.V.2008, Lima, D.C. (UFRN); Tocantins: Ilha do Bananal, 1f#, 1931 (MNHN); Miracema, P12 e CM [sic], UHE [Usina Hidrelétrica] Lajeado, 1f#, 30.X.2001, Equipe de Resgate (UFT); Palmas, 1f#, 05.VI.2011, Fabiana Maione (UFT); Porto Nacional, 1f#, 12.VII.2005, Parente S.G.G. (UFT); São Paulo: Ilha Solteira, urban area, 1f#, 26.IX.1992,Vilamaior, P.S.L. (EMUS); Estação Raiz da Serra, 1f#, XII.1907, H. Luederwaldt (MZSP); Cajuru, 1f#, 30.XII.1985, Moure, J.S. (DZUP); Distrito de Sousas, Jundiaí, 1f#, 03.VIII.1987 (UFMG); Estrada Itú-Pau D'Alho, Km 4, Cerrado, 1f#, 30.X.1960 (MNCN); Jundiahy [Jundiaí], 1f#, 1899 (CUIC); Ribeirão Preto, 1f#, X.1953 (MNCN); Horto Florestal Navarro de Andrade, Rio Claro: 2f#, 14.VII.1987 (UFMG); 2f#, 24.VI.1987 (UFMG); T. [Teodoro] Sampaio, Pq.Est. [Parque Estadual] Morro do Diabo, 1f#, 5.XII.2010, Melo, G.A.R. & Luz, D. (DZUP); Presidente Epitácio, 1f#, 10.X.1954 (MZSP); Faz. [Fazenda] Itaquerê, Nova Europa, 1f#, 03.X.1964 (EMUS); Distrito Federal: Brasília: FLONA [Floresta Nacional] de Brasília, Gleba 1 [sic], 1f#,14.XI.2012, Luz, D. (DZUP); Bahia: Riachão das Neves, BR135, Faz. [Fazenda] Jacuba, 12°00'08"S 45°57'53"W, 1f#, 22.I.2009, Figueiredo, Almeida, Cezar (MZSP); Vitoria da Conquista, Poço Escuro, 14°50’28’’S 04°05’16’’W, 1f#, 12.I.2001, Maia, J.R. (CPDC); Amazonas, Manaus, AM010, Km29, EMBRAPA [Empresa Brasileira de Pesquisas Agropecuárias] Amazônia Ocidental, 1f#, 13.V.2016 (INPA); Paraná: Piraquara, Rede Entomológica, 1f#, 24.IV.2003, Maria, V.A. (DZUP); Diamante do Norte, Estação Ecológica Caiuá, 1f#, 28.I.2012, Dias, F. (DZUP); Ceará, Viçosa do Ceará, estrada para Padre Vieira, Km 3 'N [sic], 1f#, 28.V.2013, Dolibana & Pessoa (DZUP); Barbalha:1m#, V.1969, M. Alvarenga (EMUS); 2m#, V.1969, M. Alvarenga (FSCA); Espírito Santo, Linhares, Sooretama, 1f#, I.1949 (MNRJ); PERU: El Campamiento, Col. Perone, 1f#, 04–07.VI.1920, Cornell Univ. [University] Expedition (CUIC); Chanchamayo, 2f#, 22.VII.1948 (UMMZ); Ucayali, Pucallpa, 200m [above sea level], 2f#, 11–20.XI.1964, J. Schunke (EMUS); BOLIVIA: Beni: Guayaramerin, 2f#, XII.1956, M.A. Fritz (AMNH); Versailles, 1f#, 19.VII.1964, J.K. Bouseman & J. Lussenhop (AMNH); La Paz, Mupiri, Arroyo Tuhiri, 508m [above sea level], 15°17'26''S 68°15'46''W, 1f#, 12.IV.2004, S.D. Gaimari (CSCA); San Miguel, 2f#, XI.1906, Dr. T. Staecker S.V. (ZMB); Yungas, Guanay, Uyapi, 2f#, X.1995, Gerlach (AMNH); Santa Cruz: 3.7 mi SSE [miles south-southeast of] Buena Vista, Hotel Flora y Fauna, 2f#, 02–13.II.2000, B.K. Dozier (FSCA); 40 km NW [northwest of] Santa Cruz, Potrerillo de 148
Guenda, 1f#, 22.XI–12.XII.2005, B.K. Dozier (FSCA); Buena Vista, 09.I.1991, Carpenter & Wenzel (AMNH); Santa Cruz Botanical Gardens, 1f#, 15.XII.2004, Gino Nearns (CMBC); PARAGUAY: San Pedro, Rio Ypane, Cororo, 8f#, XII.1983, M.A. Fritz (AMNH); Tupi Loma, 23°30'S 56°45'W, 1f#, 11–20.XII.2006, U. Dreschel (EMUS); Concepcion, Cororo, 1f#, II.1993, Arriagada (AMNH). An additional 143f# and 67m# from various localities in BRAZIL, PERU, BOLIVIA, and PARAGUAY have also been examined (AMNH, ANSP, INHS, UCRC, DZUP, ZMUC, UFRR, MZSP, DGMC, MNHN, MNRJ, EMUS, MNCN, CZMA, INPA, MPEG, FSCA, LACM, ANSP, RBINS, UFRN, UFT, UFMG, DEI, UMMZ, CUIC, BMNH, CMBC, MCZC).
Remarks. After the sex association of T. indica and T. selligera, the repeated coocurrence of T. almada and the four female species synonymous with T. parallela were the main basis for associationg and synonymizing these sepcies. Females of T. parallela aredistinct in the shape of their mesosoma, with pronounced pronotal and propodeal spiracles, as well as the densely sculptured and obscurely striate lateral propodeal face. The type specimens of T. parallela, T. lineifera, T. indicoides, T. gausapata, and T. pillinata are structurally identical, differing only in setal and color patterns.
Traumatomutilla protuberans (Gerstaecker, 1874) (Figs. 21A–G)
Mutilla protuberans Gerstaecker, 1874: 318, holotype [by monotypy], #m, Argentina, Catamarca (MLUH), examined. Ephuta (Traumatomutilla) protuberans: André, 1902, 55. Traumatomutilla protuberans: André, 1904, 40.
Diagnosis. FEMALE. Unknown. MALE. Cuspis elongate, slender; mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; scutellum gibbose with nearlt vertical posterior face; axillar projections transversely truncate, conspicuously narrower apicad in dorsal view; penis valvewith posterior tooth conspicuously shorter than anterior tooth. Description. FEMALE. Unknown. MALE. Body length 20 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.75 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 5.8 × DLO, IOD 2.0 × DLO. Occipital carina distinct. Head surface densely and coarsely punctate with sparse interspersed micropunctate; conspicuously denser and coarse on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 2.6 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations except medially mostly unsculptured smooth and shinning. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus present and parapsis indistinct, reduced to posterior half of mesoscutum. Scutellum gibbose, with well-defined dorsal and posterior faces densely and coarsely foveolate-punctate; with irregular somewhat elevated “carina-like” longitudinal medial are on dorsal face. Axilla produced posterolaterally as short and narrow truncate projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; lateral face with sculpture indistinct on anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and indistinctly rugose-punctate with dense interspersed micropunctures; mesopleuron with large blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on basal third 149 and indistinct rugosities on apical fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.55 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible. Pygidial plate irregularly, transversely and indistinctly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in longitudinal carina with conspicuous subacute spine-like projection posteriorly. S2 coarsely and sparsely foveolate-punctate to punctate, with reduced anteromedial pit filled with setae; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate- punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, rounded. Ratios of free length of paramere, cuspis and digitus, 67:58:15; paramere marginally sinuous in dorsal view, curved outward apically in dorsal view and upcurved apically in lateral view; with dense setae ventrally at basal half; cuspis thin, slender, elongate, slightly wider apically, equally wide throughout in lateral view except apex conspicuously wider, subcapitate; with dense long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, virtually as long as wide, subtriangulate and with densely setose posterodorsal margin in lateral view; setae as long as or shorter than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, subcapitate in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute shorter than subacute subposterior tooth; externolateral pocket well-developed; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations. FEMALES. Unknown. MALES. Integument black to brownish-black. Body setae predominantly blackvarying in density, except following areas with black setae varying in density: posterior face of propodeum, T1, basal third of T2, lateral margins of T2, fringe of T2–3 laterally, and S1–4. Distribution. Argentina. Material examined. (1m#) Type material. Holotype, #m, ARGENTINA, Catamarca (MLUH). Remarks. The peculiar axillar projection and strongly produced anterior tooth of the penis valve in T. protuberans set this specimen apart from every other male of the T. indica species-group examined in this study. We have not seen any other specimen with this character combination except this holotype. Two possible explanations for this single male include: 1. Based on distribution and size, this is a putative male for T. contempta in Argentina; 2. Based on morphological similarities, this could be an unusual variant of T. spectabilis or T. vidua. Either way, both hypotheses can only be confimerd with more specimens with the same characters of T. protuberans or specimens that bear intermediate characters between this species and others.
Traumatomutilla puella (Gerstaecker, 1874) (Figs. 22A–I, 23A–K)
Mutilla puella Gerstaecker, 1874: 72, holotype [by monotypy], #f, Brazil, Brandt S. (ZMB), examined. Mutilla musculus Gerstaecker, 1874: 320, holotype [by monotypy], #m, Brazil, Rio de Janeiro [sic] (MLUH), examined, syn. nov. Mutilla manca Cresson, 1902: 52: lectotype [designated by Cresson (1916)], #f, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Mutilla viana Cresson, 1902: 77: lectotype [designated by Cresson (1916)], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) puella: André, 1902: 55. 150
Ephuta (Traumatomutilla) musculus: André, 1902: 55. Ephuta (Traumatomutilla) manca: André, 1902: 55. Ephuta (Traumatomutilla) viana: André, 1902: 56. Traumatomutilla puella: André, 1904: 40. Traumatomutilla musculus: André, 1904: 40. Traumatomutilla manca: André, 1904: 40. Traumatomutilla viana: André, 1904: 40. Traumatomutilla peperina Casal, 1969: 298: holotype, #f, Argentina, Cordoba, San Janvier (AMNH), examined, syn. nov.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles; lateral face of propodeum sparsely sculptured, with wide conspicuous smooth intervals; longitudinal carinae of mesonotum/dorsal face of propodeum indistinct; integumental spots of T2 yellowish, well separated, anterior and posterior pair well-developed. MALE. Cuspis slender, elongate; mesopleuron simply swollen to indistinctly tuberculate on dorsal half; meso and metatibial spurs black to brownish, concolorous with legs; wings hyaline brown throughout; axillar projections acute, S2 without medial pit filled with setae. Description. FEMALE. Body length 8-14 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.85 × pronotal width. Eye almost circular, its length in frontal view 1.25 × the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate to areolate-punctate where visible; sculpture sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, reaching antennal tubercles and separated from poorly defined lateral scrobal carina. Antennal tubercle irregularly rugose. Scape densely punctured. Flagellomere 1 2.2 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate- punctate where visible; with conspicuous medial longitudinal carina extending from anterior margin of mesonotum, through scutelar area and connecting with similar carina on dorsal face of propodeum; sculpture of mesonotum slightly less defined around medial longitudinal carina; scutelar area sculpture virtually equal to that of mesonotum. Anterior face of pronotum defined, slightly shorter than pronotal collar, indistinctly and coarsely striated longitudinally at base and with dense coarse and confused punctures with interspersed micropunctures elsewhere; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, broadly separated from low rounded epaulet, anterolateral corners of pronotum subrounded in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; with indistinct blunt tubercle on ventral margin anterior to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate to areolate-punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal third completely asetose, smooth. Lateral face of propodeum sparsely and shallowly foveolate-punctate along anterior margin, more densely and deeply so along posterior margin, medial area with indistinct traces of longitudinally oblique rugosities. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 74:81:80:64:60. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale and anterolateral carinae virtually absent, reduced to transverse irregular intervals delineating slightly larger areolations; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 35:76:79. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around sub-sharp longitudinal medial carina equally high throughout. S2 densely foveolate-punctate, more sparsely and 151 finely so anteromediad; anteromedial crest-fold indistinct. S3–6 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate where visible. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Body length 11-13 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.85 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 4.5 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate; conspicuously denser and coarse on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Flagellomere 1 1.6 × pedicel length; flagellomere 2 4. × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations, except medially simply micropunctate along dorsal margin and with conspicuous smooth unsculptured area basomedially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely foveolate-punctate; with longitudinal carina medially. Axilla produced posterolaterally as short acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; sculpture denser and coarser anterolaterad; lateral face virtually unsculptured on anterior half, posterior half with coarse to indistinct sculpture; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleura with simply swollen on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations basally. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible; lateral margins of T2 asetose and unsculptured, smooth. Pygidial plate irregularly, transversely and indistinctly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in slightly blunt longitudinal slightly concave carina. S2 coarsely and sparsely foveolate-punctate to punctate, without micropunctures or setae filled pit; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 63:44:13; paramere slightly sinuous and conspicuously curved outward apically in dorsal view, upcurved apically in lateral view; with sparse setae ventrally at basal half; cuspis slender, elongate, equally wide throughout in dorsal view and lateral view; with scattered long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, subtriangulate in lateral view, virtually as wide as long with subtrinagulate and sparsely setose posterior margin in lateral view; setae as long as or shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Body and appendages predominantly black to brownish-black, except most flagellomeres ventrally and mandibles partially reddish brown; T2 always with four integumental spots varying in size and 152 ranging from orange-red to yellow; anterior spots longitudinal, linear, varying in extension; posterior spots ranging from obliquely subovate to transversely sublinear/subrectangular. Head setae ranging from being predominantly black with malar space and ventral surface of head with silvery setae to having a dense subrounded spot of silvery-white setae on the middle of vertex which can extend slightly into front. Setae on mesosomal dorsum always predominantly black with two lateral stripes of dense silvery-white setae extending, uninterrupted, from posterior margin of pronotum to apex of propodeum. Setae of lateral face of pronotum, mesosomal pleurae and lateral face of propodeum silvery-white ventrally and black dorsally varying in density and extension. Metasomal setae predominantly black, except the following areas with silvery-white setae varying in density: T1 laterally; lateral areas, lateral margins, lateral felt lines and integumental spots of T2; fringes of T2–3 medially and laterally; fringes of T4–6 medially; S1–5; fringes of S2–3. Certain specimens have silvery-white setae, though usually reduced, on the fringes of T4-5 laterally. Seta on the legs varies from being predominantly silvery-white to predominantly black. Tibial spurs yellowish-white to brownish. MALES. Integument black to brownish-black except mandibles partially reddish-brown. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: propodeum posterolaterally; T1 and basal third of T2; fringes of T2–3 laterally; T4 laterally; S1–4; and tarsi. Tibial spurs yellowish-white to brownish. Wings dark-brown throughout, veins darker than wing membrane. Distribution. Brazil, Bolívia, Paraguay, and Argentina. Material examined. (96f#, 9m#) Type material. Holotype of Mutilla puella, #f, BRAZIL, Brandt S. (ZMB); holotype of Mutilla musculus, #m, BRAZIL, Rio de Janeiro [sic] (MLUH); lectotype of Mutilla manca, #f, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH); lectotype of Mutilla viana, #m, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH); holotype of Traumatomutilla peperina, #f, ARGENTINA, Cordoba, San Janvier (AMNH). Additional material. BRAZIL: Rondônia, 5f# (MZSP); Mato Grosso: Rosário Oeste, 6f# (MZSP); Chapada dos Guimarães: 1f#, 27.X.1961, Oliveira, F.M. (DZUP); 1f#, 30.I.1965, Laroca (DZUP); 3f#, III [sic] (ANSP); 1f#, IV [sic] (ANSP); 1f#, X [sic] (ANSP); Campo, 1f#, sept. [september] (MNHN); Cáceres, 1f#, 27.III.1985, Elias,C. (DZUP); Barra do Tapirape, 5f# (MZSP); Utiariti, 5f# (MZSP); Mato Grosso do Sul: Três Lagoas, Faz. Dr. José Mendes, 1f#, 21.X.1964, Exp. Dep. Zool. [Expedição Departamento Zoologia] (MZSP); Três Lagôas, 8f# (MZSP); Goiás: Jataí, 31f# (MZSP); Mineiro, 1f# (MNHN); Minas Gerais, Araguari, 1f#, X.1931 (MNCN); São Paulo, 5f# (MZSP); BOLIVIA, Santa Cruz de La Sierra, Chiquitos, 1f#, 1834, D'Orbigny (MNHN); PARAGUAY: Concepcion, Cororo, 1f#, II.1993, Arriagada (AMNH); San Pedro, Rio Ypane, Cororo: 7f#, XII.1983, M.A. Fritz (AMNH); 1f#, XI.1983, M.A. Fritz (AMNH); 3f#, X.1979, Viana (AMNH); 1f#, XI.1979, Fritz (FSCA); Kanindeyu, Tava Yopoi, 26.X–04.XI.2007, U. Dreschel (FSCA); ARGENTINA, Cordoba, Ycho Cruz, 1f#, II.1975, Williner (AMNH). Remarks. The sculpture of the female mesonotal dorsum varies greatly in this species. Cresson’s type series of T. manca (aprox. 60 specimens), includes some specimens with the sculpture virtually homogeneous throughout, becoming slightly confused around the medial longitudinal carina as in the typical specimens of T. puella and T. peperina. In several other cases from the same series, however, most of the medial sculpture of the mesonotum is obliterated, being simply and densely punctate which is a character that appears to be associated with larger specimens (PRB pers. obs.). Similarly, the sculpture of the lateral face of the propodeum varies from being mostly unsculptured and smooth to being comparatively densely foveolate-punctate in each series examined. There are not, to our knowledge, any consistent characters to separate these females and the males repeatedly collected with them in large series are consistent with the characters of T. viana, with no apparent variations in setae or color. Slight structural variations apparently associated with larger specimens have also been recorded for males, which can have the dorsal half of the mesopleuron simply swollen in smaller specimens or bearing a conspicuous, albeit small, blunt tubercle in larger specimens. Every specimen of T. viana examined is identical in genitalia, structure, setae, and color to the holotype of T. musculus. Additionally, we have observed that the tibial spurs color varies from brownish to yellowish-white within the type series of T. viana and T. manca which further supports the association based on distribution.
Traumatomutilla selligera (Gerstaecker, 1874) 153
(Figs. 24A–B, Figs. 25A–G)
Mutilla selligera Gerstaecker, 1874: 319, holotype [by monotypy], #m, Columbia [sic] (ZMB), examined. Ephuta (Traumatomutilla) selligera: André, 1902: 56. Traumatomutilla selligera: André, 1904: 40.
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles; lateral face of propodeum predominantly unsculptured, smooth and shinning; anterolateral corners of pronotum sharply angulate in dorsal view; lateral face of propodeum with conspicuous sharp tubercle; anterolateral carinae of scutelar area virtually straight; MALE. Cuspis elongate, slender; mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; axillar projections acute; scutellum gibbose with well-defined dorsal and posterior faces, dorsal face with straight transverse carina on posterior margin; penis valve without shelf-like outward projection on posterior margin. Description. FEMALE. Body length 12-16 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and finely foveolate-punctate where visible; sculpture conspicuously sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, reaching antennal tubercles and narrowly separated from lateral scrobal carina. Antennal tubercle coarsely punctate to irregularly rugose. Flagellomere 1 2.4 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate- punctate laterad; densely punctate mediad, with conspicuous medial longitudinal carina extending from anterior margin of mesonotum to scutelar area; dorsal face of propodeum with vestiges of similar carina posteriorly. Anterior face of pronotum defined, slightly longer than pronotal collar, indistinctly and coarsely striated longitudinally basad and with dense coarse and confused punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well- defined, broadly separated from slightly projected and rounded epaulet, anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle projected from lateral margin of pronotum, subacute. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures and conspicuous subacute tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate to areolate-punctate along mesopleural ridge where visible; metapleuron sculpture partially concealed by dense setation, dorsal third completely asetose, smooth, unsculptured. Lateral face of propodeum sparsely foveolate- punctate, with broad unsculptured intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 79:89:87:65:63. Lateral margin of mesonotum strongly constricted anterior to propodeal spiracle, strongly diverging anterad. Propodeal spiracle projected from lateral margin of mesosoma; Scutellar scale present, well-defined, connected to anterolateral carinae; anterolateral carinae connected, thus forming single transverse carina extending to propodeal spiracles; scabrous intervals absent on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 33:75:71. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around blunt longitudinal medial carina equally high throughout. S2 densely foveolate-punctate; sculpture sparser posteromediad; anteromedial crest-fold present. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate where visible with interspersed micropunctures; S6 densely and coarsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with coarse and irregular rugosities; interstice granulose. MALE. Body length 18 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 4.8 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface 154 densely and finely punctate with sparse interspersed micropunctate; conspicuously denser and coarse on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Flagellomere 1 1.8 × pedicel length; flagellomere 2 2.2 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctations laterally, simply micropunctate sublaterally, and virtually unsculptured smooth and shinning medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum gibbose, densely and coarsely foveolate-punctate; with well-defined dorsal and posterior faces; dorsal face longitudinally elevated medially, not flat, ending in transverse straight carina at posterior margin. Axilla produced posterolaterally as short acute projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; lateral face virtually unsculptured on anterior margin; dorsal face rounded into and poorly distinguished from posterior face; slightly depressed laterobasally. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleuron with short blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on basal fourth and indistinct rugosities on apical fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0. 5 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and coarsely punctate with interspersed micropunctures where visible. Pygidial plate irregularly, transversely and indistinctly rugose, broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in longitudinal carina with conspicuous subacute spine-like projection posteriorly. S2 coarsely and sparsely foveolate- punctate to punctate, witht micropunctures laterally and conspicuous elongate setae filled pit posteromedially; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 64:48:14; paramere virtually straight in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half; cuspis thin, slender, elongate, equally wide throughout in dorsal view, slightly wdier apically in lateral view; with scattered inconspicuous long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, virtually as long as wide, subrounded and with sparsely setose posterodorsal margin in lateral view; setae shorter than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, subcapitate in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subacute; externolateral pocket greatly reduced; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Integument black, except basal half of mandibles and most antennal flagellomeres ventrally reddish-brown; T2 with four yellowish integumental spots. Body setae predominantly black: except the following areas with silvery-white setae varying in density: transverse arcuate band on vertex; ventral third of lateral pronotal face; ventral half o mesopleuron and metapleuron; postero-lateral corners of mesonotum; dorsal face of pronotum laterally; legs; integumental spots, lateral areas, lateral felt lines, and lateral margins of T2; fringe of T2–4 medially and laterally; fringe of T5–6 medially; S1–4, and fringes of S2–4. MALES. Integument black to brownish-black. 155
Body setae predominantly silvery-white varying in density, except following areas with black setae varying in density: head, pronotum, mesoscutum, axillar projections, scutellum, metanotum laterally, mesopleuron dorsally, dorsal surface of femora, posterior two thirds of T2, fringe of T5–6 medially, T6 (except pygidial plate), fringe of S5–6, and S7. Tibial spurs yellowish-white. Wings light brown infuscated, except basal third hyaline, with faint purplish reflections on apical third. Distribution. Colombia, Trinidad, Venezuela, and French Guyana. Material examined. (86f#, 56m#) Type material. Holotype, #m, Columbia [sic] (ZMB). Additional material. COLOMBIA: 3m# (IAvH); Magdalena: 3m# (IAvH); PNN [Parque Nacional Natural] Tayrona Zaino, 11°20'N 74°02'W, 50m [above sea level], Malaise [trapa], M789 [sic], 1f#, 17.X–10.XI.2000, R. Henriquez (EMUS); 10-15km E [east of] Santa Marta, 1f#, 04–05.XII.1976, M. Cooper (BMNH); PNN SN [Parque Nacional Natural Sierra Nevada], Santa Maria, Filo Cartagena, 1m#, 1994, R. Olmos (IAvH); Meta: 2m# (IAvH); PNN [Parque Nacional Natural] Macarena, Borde Rio Guejar, 03°20'N 73°56'W, 460m [above sea level], M2609 [sic], Malaise [trap], 24–28.XII.2001, D. Campos (EMUS); Meta: PNN [Parque Nacional Natural] La Macarena: Carretera, 580m [above sea level], 1m#, 18.X.1991, W.Cubillos (IAvH); Rio Guejar, 3°20'N 73°53'W, 580m [above sea level], 1m#, 12.XI.1991, W.Cubillos (IAvH); San Juan de Arama: 3°20'N 73°53'W, 750m [above sea level],1m#, F. Fernandez (IAvH); 2f#, 13.VII.1988, W. Cubillos (AMNH); 580m [above sea level], 1m#, 04.X.1991, Carret, W.C. (EMUS); Rio Samsa, 3°20'N 73°53'W, 580m [above sea level], 1m#, 18.X.1991, W.Cubillos (IAvH); Puerto Lleras, Loma Linda, 03°18'N 73°22'W, 300m [above sea level], 2f#, R. Rondoh (EMUS); Res. Nat. [Reserva Natural] El Caduceo: 03°40'N 73°39'W, 1400', HC, savanna, 1f#, 03–08.I.2012 (EMUS); 03°40'N 73°39'W, 1400', MT, forest, 1f#, 03–08.I.2012 (EMUS); 1m#, 3.I.2012, K.A. Williams (EMUS); Carimagua, 1m#, 8.III.1979, J. de Porre (FSCA); Cord. [Cordillera] Macarena, 1f#, 15–28.II.1976, M. Cooper (BMNH); Arauca: Tame: 1f#, 17–18.VII.1976, M. Cooper (BMNH); 1f#, 01–07.VII.1976, M. Cooper (BMNH); 1f#, 21–29.VI.1976, M. Cooper (BMNH); 3f#, 08–17.VII.1976, M. Cooper (BMNH); TRINIDAD: 13 km S [South of] Arima: 13m#, 22.VI.1993, S & J Peck (EMUS); Farm Pecks, 1m#, 12–22.vi.1993 (FSCA); 2km N [north of] Talparo,Quesnell Farm, rainforest FIT, 50m, 18f#, 22;VI–08.VII.1993, S.J. Peck (EMUS); 14 km S [South of] Arima, 20m#, 12.VI.1993, S & J Peck (EMUS ); 11km SE [southeast of] Arima: 3m#, 28.VI.1993, S & J Peck (EMUS); Arena For. Res., 80m, 3f##, 28.VI–08.VII.1993, S & J Peck (EMUS); 2f#,13.VI–08.VII.1993, Pecks (EMUS); 16f#, 12–22.VI.1993, Pecks (EMUS); Arima Valley, Arena Forest Reserve: 1f#, 09.IV.1951 (AMNH); 1f#, 19.VI.1991, H.L. Dozier (FSCA); 2 km E [east of] San Rafael, Arena Forest Reserve, 1f#, 01.VII.1999, G.B. Edwards (FSCA); St. [Saint] George County, Arena Forest Preserve, 1f#, 05.VI.2000, M.E. Berres (CMBC); Paria Matelot Trail, 1f#, 14.I.1999, Ann Berres (CMBC); U.S. N.A.S. [sic], 1f#, 03.III.1943, S.G. Pederson (UMSP); Couva-Tabaquite, Talparo: 1f#,11.VI.1991, Byrd Dozier (FSCA); 1f#, 10.VII.1988, H.L. Dozier (FSCA); 1f#, 07.VII.1988, H.L. Dozier (FSCA); 1f#, 13.VI.1991, H.L. Dozier (FSCA); VENEZUELA: Barinas:40km SE [southeast of] Socopo, 150m [above sea level], 1f#, 07.II.1970, S.L. Wood (EMUS); 40 km E [east of] Canton, 1f#, 08.III.1970, S.L. Wood (BYU); Socopo, 1f#, 07.II.1970, S.L. Wood (BYU); Bolívar: 20 km W Guasipati, 1m#, 24.VI.1987, S & J Peck (PMAE); Tumeremo, 20 km S [south of], 1m#, 24.VI.1987, S & J Peck (PMAE); Suapure: 1f#, II–III.1999 (EMUS); Caura River: 1f#, 14.VIII.1899, E.A. Klages (EMUS); 1f#, 21.IX.1899, E.A. Klages (EMUS); 1f#, 20.IX.1899, E.A. Klages (EMUS); Monagas: Caripito: 2f#, 30.VIII.1942 (AMNH); 2f#, 06.IV.1942 (AMNH); Tachira: Santa Barbara, 1f#, VII.1981, Williner (AMNH); Rubio, 1f#, X.1988, D. Havrenek (FSCA); Zulia, Los Angeles del Tucuco, 1m#, 15.VI.1981, Hollenberg & Menke (USNM); Caracas, Caracas, 1f#, Styrech (ZMUC); Zamora, Libertad, 2f#, 1923, Mayeul Grisol (MNHN); Rancho Grande, nr. [near] Maracay, 1f#, 22.VI.1946 (AMNH); FRENCH GUYANA: Lunier River, 2f#, 1899, F. Geay (MNHN). Remarks. The sex association of T. selligera was based on couples collected in Trinidad, as well as large numbers of both sexes co-ocurring in various areas in the northern Amazon. There are several female specimens of T. selligera in various collections identified as T. aequinidialis by Mickel, PRB, and KAW. This was a name given by Mickel to the females of T. selligera, considered as a putative new species by the author at the time. The anterolateral angles of the pronotum and the sculpture of the lateral propodeal face in the females seem to be unique within the species-group and are only approximated by the females of T. guayaca which differ in the structure of the lateral pronotal face and scutelar armature. 156
The males of T. selligera, on the other hand, are similar in structure to those of T. tristis, differing mainly in structure of the mesoscutellum.
Traumatomutilla spectabilis (Gerstaecker, 1874) (Figs. 26A–F, 27A–K)
Mutilla spectabilis Gerstaecker, 1874: 73, lectotype [designated here], f#, Brazil, S. Paulo [São Paulo (sic)] (ZMB), examined. Mutilla funesta Gerstaecker, 1874: 316, lectotype [designated here], m#, Brazil, [Minas Gerais], Lagoa Santa (MLUH) syn. nov. Mutilla melaleuca Gerstaecker, 1874: 316, holotype [by monotypy], m#, Brazil (ZMB), examined, syn. nov. Ephuta (Traumatomutilla) spectabilis: André, 1902, 56. Ephuta (Traumatomutilla) funesta: André, 1902, 55. Ephuta (Traumatomutilla) melaleuca: André, 1902, 55. Traumatomutilla spectabilis: André, 1904, 40. Traumatomutilla funesta: André, 1904, 40. Traumatomutilla melaleuca: André, 1904, 40. Traumatomutilla spectabilis chingona Casal, 1969. 294, holotype, f#, Argentina, Cordoba, Villa del Dique (AMNH), examined, syn. nov.
Diagnosis. FEMALE. Mesonotum usually wide than distance between pronotal spiracles; scutelar scale and anterolateral carinae greatly reduced, nearly indistinct; head usually with transvers band of appressed silvery-white setae. MALE. Cuspis elongate, slender; mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; scutellum gibbose; axillar projections obliquely truncate; posterior face of scutellum sloping in lateral view. Description. FEMALE. Body length 18 mm. Head. Posterior margin virtually straight. Occipital carina evenly wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 1.1 × the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely and coarsely foveolate punctate with micropunctures. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, reaching antennal tubercles and lateral scrobal carinae. Antennal tubercles irregularly rugose. Flagellomere 1 2.3 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Mesosoma slightly wider than longer. Mesosomal dorsum densely and coarsely areolate-punctate with irregular intervals; with longitudinal medial carina extending from anterior margin of mesonotum. Anterior face of pronotum defined virtually as long as pronotal collar, with dense indistinct and coarse longitudinal striations ventrad, coarsely and densely punctate dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, narrowly separated from epaulet; epaulet virtually flat against anterior margin of pronotum; antero-lateral corners of pronotum subrounded in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum. Lateral face of pronotum densely and finely punctate with dense interspersed micropunctures, except for mostly unsculptured indistinct swelling anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and sparsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by dense setation on ventral half, completely asetose, smooth on dorsal half. Lateral face of propodeum densely and coarsely foveolate- punctate with smooth intervals, at most as wide as surrounding foveolations; foveolations conspicuously smaller anteriorad. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 65:75:80:59:58. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, mesonotum with expanded lateral margins. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale present, irregular, connected to indistinct anterolateral carinae; anterolateral carinae marginally connected. Propodeum convex, dorsal face shorter than and poorly 157 distinguished from posterior face; dorsal face with irregular longitudinal carina medially. Metasoma. Ratios of width of T1, width of T2 and length of T2, 53:113:116. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, sparsely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely foveolate-punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around blunt longitudinal medial carina slightly higher posteriorly. S2 sparsely foveolate-punctate, sculpture sparser poderomediad; anteromedial crest-fold well-defined. S3–5 sculpture mostly concealed by dense setation, densely and coarsely foveolate-punctate with sparse interspersed micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with coarse and irregular rugosities; interstice granulose. MALE. Body length 16-20 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 5.0 × DLO, IOD 1.45 × virtually equal to DLO. Occipital carina distinct. Head surface densely and finely punctate with sparse interspersed micropunctate; conspicuously denser and coarse on front. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely punctate; apical/ventral margin with a pair of medial subrounded subsessile teeth. Scape bicarinate. Flagellomere 1 2.2 × pedicel length; flagellomere 2 3.2 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from short humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely and coarsely punctate with interspersed micropunctations; sculpture sparser mediad, simply micropunctate medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum convex, densely and coarsely foveolate-punctate; dorsal and posterior faces poorly distinguished; basal intervals of dorsal face indistinctly aligned forming rudimental longitudinal carina at base of scutellum. Axilla produced posterolaterally as obliquely truncate projection, with coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex to virtually flat basad, densely areolate; lateral face virtually unsculptured on along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleura with conspicuous blunt tubercle on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; areolations/foveolations gradually sparser anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on basal fourth and indistinct rugosities on dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.4 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate irregularly, transversely and indistinctly rugose, slightly broader than long, weakly defined by parallel carinae apicolaterally, interstice apparently granulose. S1 longitudinally elevated medially, terminating in slightly concave and blunt longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate, without micropunctures; with conspicuous anteromedial setae filled pit; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 densely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 75:56:21; paramere slightly sinuous and conspicuously curved outward apically in dorsal view, upcurved apically in lateral view; with sparse setae ventrally at basal half; cuspis slender, elongate, equally wide throughout in dorsal view and lateral view; with scattered long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, subtriangulate in lateral view, slightly longer than wide with subtriangulate and densely setose posterodorsal margin in lateral view; setae 158 as long as or shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, blunt, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations. FEMALE. Integument black to brownish-black except mandibles and antennal flagellomeres partially reddish-brown, and T2 with four reddish integumental spots. Posterior spots of T2 variable in size, separated by more than spot width to half spot width. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: front, posterior margin of vertex, dorsal half of gena, pronotal dorsum, lateral face of pronotum (except ventral third); mesonotum medially, dorsal propodeal face medially, dorsal half of meso and metapleuron, T1 medially, disc of T2 (except over integumental spots), fringe of T2–4 sublaterally, fringe of T5 laterally (sparse silvery-white setae sometimes present laterally), T6 (except pygidial plate) laterally, and S5–6. Some specimens have the vertex virtually all black with indistinct spots of silvery-white setae laterally and the lateral silrvery-white setae stripes of the mesosomal dorsum uninterrupted. MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: anterior/ventral margin of clypeus, hind tibiae and tarsi partially, metanotum medially, dorsum of propodeum, T1, T2 anterolaterally, fringes of T2–3 laterally, S1 partially, and fringe of S2 partially. Tibial spurs yellowish-white. Wings dark-brown infuscated with conspicuous blueish/purplish reflections. Distribution. Brazil, Bolívia, Paraguay, and Argentina. Material examined. (175f#, 56m#) Type material. Lectotype of Mutilla spectabilis, f#, BRAZIL, S. Paulo [São Paulo (sic)] (ZMB); lectotype of Mutilla funesta, m#, BRAZIL, [Minas Gerais], Lagoa Santa (MLUH); holotype of Mutilla melaleuca, m#, BRAZIL (ZMB); holotype of Traumatomutilla spectabilis chingona, f#, ARGENTINA, Cordoba, Villa del Dique (AMNH). Additional material. BRAZIL, 1m# (ZMUC); São Paulo: 3f# (DEI); 1f#, X.1950 (MNRJ); 1f#, II.1935 (MNRJ); 40f# (MZSP); 1f#, Dr. O. Staudinger (MNCN); 1f# (MNRJ); Baruer: 1f#, 5.III.1955 (MNRJ); 1f#, IV.1957, K. Lenko (USNM); 1m#, 28.II.1956 (AMNH); 1m#, V.1957, K. Lenko (USNM); São Paulo: 1f#, 24.III.1926 (DEI); 1f#, 07.II.1922 (DEI); Ipiranga: 1f#, I.1914 (MNRJ); 1f#, I.1907, Salva M. Torres (MZSP); Jundiaí: 1f#, 24.II.1899, Bezon (MNCN); 1f#, 23.II.1899, Schrottky (MNHN); 1f#, 1899, Schrottky (MNHN); 1f#, 16.II.1899, Schrottky (MNHN); Alto da Serra, 1m# (MZSP); 1m#, II.1929, R. Spitz (MZSP); Cotia: 2f# (DEI); 1f# (ZMB); Horto Florestal Navarro de Andrade, Rio Claro, 1f#, 17.VI.1987, R.P. Martins (UFMG); 1f#, 22.III.1924, G.R. Fischer (MZSP); Piracicaba, 1f#, IV.1898, Bikego [sic] (MZSP); Rio Claro, 1f#, B. Dias (AMNH); Santa Branca, 1f#, I.1962, R.F.O. Azevedo (MNCN); São Carlos, 1f#, I.1962, L. Stowbunenko (MNCN); Campinas, 1m#, 1903, Hempel (MNHN); Rio de Janeiro: 1m# (ZMUC); 10f#, Reinhardt (MNCN); Butantan, 1f#, 26.III.1922 (DEI); Paraná: 5f# (MZSP); [Palmira] RPPN [Reserva Particular do Patrimônio Natural] Papagaio Velho, 1m#, 09.I.1966, Moure J.S. (DZUP); Ponta Grossa: Parque Estadual da Vila Velha: 1f#, 5.IV.2013, Williams, K.A. (DZUP); 1f#, 9.IV.2003, Melo, G.A.R, Gonçalves,R. (DZUP); Mato Grosso: 8f# (MZSP); Chapada dos Guimarães: 8f#, 12.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 2f#, 18.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 5f#, 13.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 1f#, 14.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 1f#, 19.I.1961, J & B Bechyne (MPEG); 1m#, Oct. [October] (MNHN); Mato Grosso do Sul: Maracaju: 1f#, 09.III.1937, S. Lana (MPEG); 1f#, III.1937, J. Lane (MNCN); 1f#, III.1937, Shannon Lane (MZSP); Corumbá, 1f# (MNCN); Minas Gerais: 1f#, Reinhardt (ZMUC); Uberaba: 1f# (DEI); 1f# (RBINS); 1f#, E. Le Moult (MNCN); Passos: 1m#, X.1961, Elias, C. (DZUP); 1m#, 16–21.XII.1963, Elias, C. (DZUP); 1m#, X.1961, Elias, C. (DZUP); 1m#, 5–10.XI.1961, Elias, C. (DZUP); São Roque de Minas, 1f#, 12–13.I.1992, Moure, J.S., Camargo, Sergei, Silvis (DZUP); Araguari, 1f#, X.1931, R. Spitz (MNCN); Barro Alto, 1f#, XI.1931, J. Blaser (USNM); Campos de Diamantina, Fazenda do Riacho Fundo, 1f#, XII.1902, E. Gounelle (MNCN); Araxá, 1m#, 10.XII.1965,Elias, C. & Elias, T. (DZUP); Ibiraci, 1m#, X.1961, Elias, C. (DZUP); Barbacena, 1m#, 26.XII.1905, Ducke (MNHN); Santa Catarina: 5f# (MZSP); Nova Teutônia: 8f# (MNRJ); 1f#, VII.1968, F. Plauman (PMNH); 3f#, XII.1968, F. Plaumann (PMNH); 1f#, I.1969, F. Plaumann (PMNH); 5f#, III.1969, F. Plaumann (PMNH); 1f#, IV.1969, F. Plaumann (PMNH); 1f#, 21.XII.1952, F. Plaumann (USNM); 1m#, XI.1969, F. Plaumann 159
(PMNH); Goiás: 1m# (MNHN); Jataí, 1m# (MNHN); Campinas, 2f#, XII.1936 (MNRJ); Distrito Federal: Brasília: FLONA [Floresta Nacional] de Brasília:, Gleba 1 [sic]:, 1f#, 14.XI.2012, Luz, D. (DZUP); 1f#, 13.XI.2012, Luz, D. (DZUP); Pará: 1m#, V.1903, Ducke (MNHN); Óbidos, 1m#, 1929 (USNM); Prainha, 1m#, 20.V.1903, Ducke (MNHN); Espírito Santo, Praia das Neves, 1f#, 27.IV.1998, Raw, A. (DZUP); BOLIVIA, Chuquisaca, Zudanez, 1f#, III.1976, Pena (AMNH); PARAGUAY: Alto Paraguay, Cuaacupe, 1f#, 15.XII.1945, P.A. Berry (CISC); Paraguari: E [east] of Ybycui, 1f#, 02.IV.2006, U. Dreschel (FSCA); La Rosada, 1f#, 27.XII.2008, U. Dreschel (FSCA); [Paraguari], Cerro Pelado, 1m#, 04.XII.1935 (DEI); [Cordillera]: San Bernardino, 1f#, K. Fiebrig (USNM); 1m#, K. Fiebrig S.V. (ZMB); San Pedro: Rio Ypane: Cororo: 1m#, II.1979, Fritz (AMNH); 2m#, XI.1979, Fritz (AMNH); 10m#, XII.1983, Fritz (AMNH); [Guairá]: Villarica, 1f#, XI, Sternitzcky (USNM); 3m#, I.1938, Slg. H. Köller (DEI); Caaguazu, 1m#, XII.1977, M.A. Fritz (FSCA); Santa Barbara [sic], 1m#, 02.III.1948, Schade (MNHN); ARGENTINA: 2º Par. [sic], 2f#, 08.II.1925, Weyrauch (UMSP); Geraldina, 1f#, 10.II.1927, Weyrauch (UMSP); Corrientes: Ituzaingo: 1f#, XI.1985, M.A. Fritz (AMNH); 1f#, XI.1979, Viana (AMNH); 2m#, III.1982, Fritz (AMNH); 1m#, I.1985, Fritz (AMNH); Córdoba: [La] Granja, 3f#, 07.II.1925 (UMSP); 2f#, 10.I.1922 (UMSP); Dolores,1000m [above sea level], 1f#, 01–17.XII.1997, C. Porter (FSCA); Alta Gracia, 1f#, 13.I.1921, C. Bruch (UMSP); San Antonio, 1f#, II.1980, Williner (AMNH); Santa Fé: Rio San Javier, Estancia La Noria, 3m#, 22.XII.1911, G.E. Bryant (BMNH); Tostado FCCN [sic], El Orden, 1m#, A.J. Giai (SEMC); Las Garzas, Margens do Rio Las Garzas, 25km a oeste de Ocampo, Chaco, 1m#, 1903, E.R. Wagner (MNHN); Misiones: Punto Esperanza, 1f#, XII.1976, Fritz (AMNH); Rio Paraná, Tiju-Cuare, Prés. [Presidente] San Ignacio, 3f#, III–IV.1911, E.R. Wagner (MNHN); Entre Rios, Federacion, 1m#, XII.1981, Fritz (AMNH); Santiago del Estero, Icaño, Margens do Rio Salado, arredores de Icaño, 1m#, XI.1909, E.R. Wagner (MNHN). Remarks. As with most species of the Traumatomutilla indica species-group, the sex association of T. spectabilis with T. funesta relied heavily on repeated co-ocurrence of both sexes in multiple locations. This particular association, however, could only be achieved after almost every other species from the edges of the Brazilian and Bolivian Amazons to the Argentinian Chacho was associated with its respecteive opposite sex. Females of T. spectabilis can be frequently confused with those of T. vidua since both species have the mesonotum wider than the distance between pronotal spiracles. This character, however, tends to vary in smaller specimens of T. spectabilis, which frequently have the mesonotum as wide as the distance between the pronotal spiracels. Nonetheless, these species can be distinguished based on scutelar armature, which is usually reduced in T. spectabilis, with only the scutelar scale being well-defined, as opposed to the conspicuous and well-defined anterior transverse carinae of T. vidua females. Additionally, males of T. spectabilis can be confused with those of T. vidua and T. protuberans. Males of T. vidua are the easiest to distinguish from T. spectabilis based on their broad transversely truncate axillar projections, as opposed to the obliquely truncate and somewhat twisted axillar projections of the latter. Males of T. protuberans on the other hand have the cuspis and penis valve remarkably different from those of T. spectabilis (see description, plates and remarks of T. protuberans).
Traumatomutilla tristis (Klug, 1821) (Figs. 28A–C, 29A–K)
Mutilla tristis Klug, 1821: 318, holotype [by monotypy], f#, Brazil, Para [sic] (ZMB), examined. Mutilla foveiventris Gerstaecker, 1874: 316, holotype [by monotypy], #m, Brazil, Sello (ZMB), examined, syn. nov. Mutilla caxara Cresson, 1902: 73, holotype [by monotypy], Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) foveiventris: André, 1902: 55. Ephuta (Traumatomutilla) caxara: André, 1902: 56. Ephuta (Traumatomutilla) tristis: André, 1902, 56. Traumatomutilla foveiventris: André, 1904, 40. Traumatomutilla caxara: André, 1904, 40. Traumatomutilla tristis: André, 1904, 40.
160
Diagnosis. FEMALE. Mesonotum as wide as distance between pronotal spiracles; lateral face of propodeum sparsely sculptured, with conspicuous smooth and shinning intervals, often wider than surrounding sculpture; anterolateral corners of pronotum subrounded in dorsal view, not angulate; lateral pronotal face at most with blunt swelling; anterolateral carinae of scutelar scale vesitigial, frequently concealed by dense setation; MALE. Cuspis elongate, thin, widened apicad; mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; axillar projections acute; scutellum gibbose, dorsal and posterior faces well defined, with equally wide longitudinal medial carina on dorsal face. Description. FEMALE. Body length 12–14 mm. Head. Posterior margin virtually straight. Occipital carina evenly equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal virtually equal to the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely and coarsely foveolate-punctate; denser and coarser on vertex; with interspersed sparse, with with sparse micropunctures on gena and malar space. Head with a longitudinal carina extending from middle of vertex to dorsal half of frons. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, narrowly separated from antennal tubercle and lateral scrobal carinae. Antennal tubercle coarsely punctate to irregularly rugose. Flagellomere 1 2.5 × pedicel length; flagellomere 2 2.1 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Pronotal and mesonotal dorsum densely and coarsely areolate-punctate with irregular intervals laterad, and simply densely punctate mediad; mesonotum with narrow longitudinal carina extending medially from anterior margin of mesonotum to scutelar area; dorsal face of propodeum with similar carina throughout; Anterior face of pronotum defined slightly longer than pronotal collar, with dense indistinct and coarse longitudinal striations ventrad, coarsely and densely punctate dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina prominent, narrowly from epaulets; epaulets virtually flat against anterior margin of pronotum; antero-lateral corners of pronotum subrounded in dorsal view. Pronotal spiracles slightly produced from lateral margin of pronotum. Lateral face of pronotum densely punctured with dense interspersed micropunctures, except low micropunctate swelling antero-ventral in relation to pronotal spiracle. Mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly, sparsely and shallowly foveolate- punctate along mesopleural ridge where visible. Metapleuron concealed by dense setation on ventral half, unsculptured and smooth on dorsal half. Lateral face of propodeum sparsely and finely foveolate-punctate anteriad to densely and coarsely foveolate-punctate posterad; Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 61:79:80:61:60. Lateral margin of mesosoma strongly constriced anterior to propodeal spiracle, strongly divergent anterad thus expanded laterally. Propodeal spiracle strongly projected from lateral margin of mesosoma. Scutellar scale greatly reduced, nearly absent, present only as an slightly elevated arcuate outline on the integument extending to propodeal spiracles on both sides. Anterolateral carinae absent; scabrous intervals absent on scutelar scale. Propodeum convex in lateral view, dorsal face poorly distinguished from and virtually as long as posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 34:76:78. Disc of T2 densely and coarsely foveolate-punctate with dense interspersed micropunctures; foveolations sparser and larger laterally and over integumental spots; micropunctures absent over integumental spots. T3–6 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; micropuncutres nearly absent on T6. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate around short longitudinal medial carina equally high throughout. S2 densely foveolate- punctate; foveolations denser and larger posteromediad; anteromedial crest-fold well-defined. S3–5 sculpture mostly concealed by dense setation, densely and coarsely foveolate-punctate with sparse interspersed micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface with coarse and irregular rugosities; interstice granulose. MALE. Body length 16 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 5.3 × DLO, IOD 1.45 × DLO. Occipital carina distinct. Head surface densely and finely punctate with interspersed micropunctures; sculpture conspicuously sparser around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, virtually flat elsewhere; densely and confusedly punctate; apical/ventral margin with a pair of short subrounded free teeth medially. Scape bicarinate. Flagellomere 1 2.1 × pedicel length; flagellomere 3 × pedicel length. Mandible 161 obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum subangulate. Anterior face of pronotum coarsely and densely punctate with interspersed micropunctures throughout; sculpture conspicuously sparser mediad; surface virtually flat throughout. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum subglobose, densely and coarsely foveolate-punctate; with conspicuous dorsal and posterior faces; posterior face almost vertical; dorsal face with medial longitudinal, slightly elevated, “carina-like” area extending into posterior face. Axilla in dorsal view produced posterolaterally as short acute projection, with coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; sculpture of lateral face less defined to indistinct along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and indistinctly punctate with sparse interspersed micropunctures; mesopleura with conspicuous short subacute spine-like projection on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; simply micropunctate anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on ventral fourth and indistinct rugosities in dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate coarsely micropunctate apically, broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in longitudinal carina with short spine-like projection posterioly. S2 conspicuously flat medially, coarsely and sparsely foveolate-punctate to punctate, with micropunctures laterally; sculpture sparser posteromediad; with conspicuous subovate antero-medial setae filled pit; longitudinal anteromedial crest-fold present. S3– 6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 sparsely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projection bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 78:71:25; paramere virtually straight and slightly curved outward apically in dorsal view, upcurved apically in lateral view; with sparse setae ventrally at anterior half; cuspis slender, elongate, posterior half conspicuously widening apicad in dorsal view, slightly narrower subapically in lateral view; with scattered long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, subtriangulate in lateral view, longer than wide with subtriangulate and sparsely setose posterodorsal margin in lateral view; setae as long as or shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, blunt, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations. FEMALE. Integument black to brownish-black, except mandibles and antennal flagellomeres partially reddish-brown, and T2 with four reddish integumental spots. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: ventral half of gena, malar space, ventral third of lateral face of pronotum, ventral half of meso- and metapleuron, lateral face of propodeum, mesoscutum, laterally, propodeal dorsum laterally, femora ventrally, T1 laterally, lateral areas of T2, lateral felt lines of T2, lateral margins of T2, fringe of T2–3 medially and laterally, fringe of T4–6 medially, and S1–4 ( except fringe of S4). MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: metanotum medially, metapleuron basally, propodeal dorsum predominantly, legs partially, T1, T2 anterolatearlly, fringe of T2–4 laterally, S1– 4, and fringe of S2–3. Tibial spurs yellowish-white. Wings dark-brown infuscate throughout, with faint blueish/purplish reflections. 162
Distribution. Brazil, Bolivia, Paraguay, and Argentina. Material examined. (97f#, 14m#) Type material. Holotype of Mutilla tristis, f#, Brazil, Para [sic] (ZMB); holotype of Mutilla foveiventris, #m, BRAZIL, Sello (ZMB); holotype of Mutilla caxara, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. BRAZIL: 3f# (ZMUC); Pará: 5f# (MZSP); Santarém, 1f# (RBINS); Alto Parú D'Oeste, [Aldeia] Tiriós: 3f#, I–II.1963, Machado e Pereira (MNCN); 2f#, II.1963, Machado & Pereira (CASC); 2f#, I– II.1968, Machado & Pereira (MZSP); 1f#, 01.II.1963, Machado & Pereira (MPEG); Óbidos: Le long de la plage, 1m#,13.I.1936, W. Adam (RBINS); Rondônia: 1f# (RBINS); Mato Grosso: 2f#, 1886, P. Germain (MNHN); Barra do Tapirape: 5f# (MZSP); 2f#, 30.XII.1962, B. Malkin (MPEG); 1f#, 1–2.II.1964, R. Malkin (MZSP); 1f#, II.1964, R. Malkin (MZSP); Rosário Oeste: 1f#, XI.1971, Dirings (MZSP); 1f#, Dirings (MZSP); 25f# (MZSP); 1f#, Cuiabá, 17– 21.I.1965, Laroca (DZUP); Cáceres, 1f#, 11.II.1985, Elias,C. (DZUP); Descalvados, Rio Paraguay, 1f#, 28.VIII.1931, J.A.G. Rehn (ANSP); Diamantina, Facienda São João, 1f#, 05.II.1981, C. Young (MPEG); São Paulo: Horto Florestal Navarro de Andrade: Rio Claro: 1f#, 03.VIII.1987, R.P. Martins (UFMG); 1f#, 30.VI.1987, R.P. Martins (UFMG); 2m#, 26.X.1987, R.P. Martins (UFMG); 1m#, 12.X.1987, R.P. Martins (UFMG); Jundiaí, Serra do Japi, 1f#, 22.IV.1988, R.P. Martins (UFMG); Campinas, 1m# (CUIC); Tocantins, Porto Nacional: 1f#, 09.X.2002, M. Bragança (UFT); 1f#, 1955 (MNRJ); Espírito Santo, Praia das Neves, 1f#, 27.IV.1998, Raw, A. (DZUP); BOLIVIA: Rosario [sic], Lake Rogagua, 1f#, XI, M.R. Lope (USNM); Santa Cruz: 40 km NW [northwest of] Santa Cruz, Potrerillo de Guenda: 22.XI– 12.XII.2005, B.K. Dozier (FSCA); 1f#, 09–28.XI.2006, B.K. Dozier (FSCA); 1f#, 16–22.X.2006, Wappes, Nearns, Eya (FSCA); Santa Cruz [de la Sierra]: 500m [above sea level], 1f#, 01.XII.1955, Zischko (MNCN); 2f#, V.1955, Zischke (AMNH); 1f#, J. Steinbach (MCZC); 1f#, 13.III.1954 (MNRJ); 6 km N [north of] Pedro Lorenzo, 1f#, 02.III.1999, L.A. Stange (FSCA); Saavedra, 1f#, 12–16.III.1986, C.J. Pruett (FSCA); Buena Vista, 2f#, 1928, J. Steinbach (CUIC); Beni, Reyes, 1f#, X.1921, W.M. Mann (USNM); PARAGUAY: 1m# (MNHN); Pto. [Puerto] Pablo [sic], 4m#, XI–XII.1936 (RBINS); Distrito Capital: Asunción: 1f#, Schrottky (MNHN); 1m#, 1891 (MNHN); Chaco, Teniente Enciso, 1f#, 02.XI.2001, G. Arriagada (EMUS); Alto Paraguay, Bahia Negra Peichiuta, 1f#, 18–24.VI.1988, V.D. & B. Roth (CASC); [Guairá], San Bernardino, 1f#, K. Fiebrig S.V. (ZMB); Guairá,Villarica, 1f#, III.1923 (ZMB); Caaguazu, Caaguazu, 1m#, XI.1977, Fritz (AMNH). ARGENTINA: Geraldina [sic], 2f#, 10.II.1937 (UMSP); Cordoba, Dolores, ca. [circa] La Cumbre, 1f#, 14.III–07.IV.1996, C. Porter (FSCA); Alta Gracia, 1f#, III.1921 (AMNH); Freyre, 2f#, IV.1992, Williner (AMNH); Chaco, Las Brenas, 1f#, 16.IV.1944, P.A. Berry (CISC); Formosa, Gran Guardia, 1f#, J. Foerster (LACM); Mendoza, Mendoza, 1f#, 04.I.1908, Jorgensen (ZMUC); Salta, La Viña, 1f#, II.1993, M.A. Fritz (AMNH); Santa Fé, Josefina, 1f#, IV.1993, Williner (AMNH). Remarks. Strangely, no conspicuous color variations were observed for either sex of T. tristis, even though this widespread species overlaps with differently colored communities of mutillids in the various parts of its distribution. As with the sex association of T. spectabilis mentioned above, females of T. tristis and males of both T. foveiventris and T. caxara could only be associated through a process of elimination after males and females co-occurring with these species were paired off. As mentioned previously, males of T. tristis can be confused with those of T. vidua and T. spectabilis, from which they are separated based predominantly in the shape of the axillar projection and genitalia characters.
Traumatomutilla unimarginata (Cresson, 1902) (Figs. 30A–C, 30A –G)
Mutilla unimarginata Cresson, 1902: 54, lectotype [desginated by Cresson, (1916)], f#, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined. Mutilla cuiba Cression, 1902. 71, holotype [by monotypy], m#, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) unimarginata: André, 1902: 56. Ephuta (Traumatomutilla) cuiba: André, 1902: 56. Traumatomutilla unimarginata: André, 1904: 40. 163
Traumatomutilla cuiba: André, 1902: 40.
Diagnosis. FEMALE. Mesonotum narrower than distance between pronotal spiracles; meso and metatibial spurs black; mesonotum with two lateral longitudinal stripes of dense appressed coppery setae; lateral face of propodeum densely sculputured; MALE. Cuspis slender, elongate; mesopleuron tuberculate on dorsal half; axillar projections roundly truncate; S2 with anteromedial pit filled with setae. Description. FEMALE. Body length 12–14 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view 1.1 × the distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate where visible; sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, broadly separated from antennal tubercles and lateral scrobal carina. Antennal tubercle coarse punctate to irregularly rugose. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Mesosoma 0.85 × as long as wide. Mesosomal dorsum sculpture partially concealed by dense setation, densely and coarsely areolate-punctate where visible; with conspicuous medial longitudinal carina extending uninterrupted from anterior margin of mesonotum posterior margin of dorsal propodeal face. Anterior face of pronotum defined, slightly longer than pronotal collar, indistinctly and coarsely striated longitudinally at base and with dense coarse and confused punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, narrowly separated from rounded epaulet, anterolateral corners of pronotum rounded in dorsal view. Pronotal spiracle projected from lateral margins of pronotum, rounded. Lateral face of pronotum densely and coarsely punctate with interspersed micropunctures, and small subacute tubercle anteroventral in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely foveolate-punctate to simply punctate along mesopleural ridge where visible; metapleuron sculpture partialy concealed by dense setation, except dorsal half unsculptured and asetose. Lateral face of propodeum sparsely and irregularly foveolate-punctate to punctate; intervals smooth, dull thoruhgout; sculpture denser posterad. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 52:75:74:58:59. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad. Propodeal spiracles slightly projected from lateral margins of mesosoma. Scutellar scale and anterolateral carinae absent. Propodeum convex, dorsal face slightly longer than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 39:90:96. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser laterad and over integumental spots; micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate- punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate-punctate terminating into sharp longitudinal carina equally high throughout. S2 densely foveolate-punctate, with distinct unsculptured longitudinal medial area; anteromedial crest-fold well-defined; sculpture overall sparser mediad. S3–6 sculpture mostly concealed by dense setation, densely and coarsely foveolate-punctate where visible. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with transverse coarse and confused rugosities; interstice granulose. MALE. Body length 13 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.8 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 4.5 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface sparsely and finely punctate with dense interspersed micropunctures; sculpture conspicuously sparser around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, virtually flat elsewhere; densely and confusedly punctate; apical/ventral margin with a pair of short subrounded free teeth medially. Scape bicarinate. Flagellomere 1 1.5 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, virtually flat against anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum sparsely punctate with interspersed micropunctures throughout; virtually unsculptured, smooth and shinning 164 medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis indistinct, reduced to posterior half of mesoscutum. Scutellum convex, dorsal and posterior faces indistinct; densely and coarsely foveolate-punctate. Axilla in dorsal view produced posterolaterally as short acute projection, coarsely and densely foveolate-punctate dorsally. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate, slightly depressed sublaterally at base; sculpture of lateral face less defined to indistinct along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and indistinctly punctate with sparse interspersed micropunctures; mesopleuron with short blunt swelling/projection on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; simply micropunctate anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on ventral fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.8 × its width. Dorsal metasomal sculpture mostly concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate coarsely and indistinctly granulose apically, broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in longitudinal carina with short blunt projection posteriorly. S2 finely and densely foveolate- punctate to punctate; sculpture slightly sparser posteromediad; with well-defined subovate anteromedial setae filled pit; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate-punctate with sparse interspersed micropunctures; S7 sparsely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projection bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 62:48:11; paramere sinuous in dorsal view, apex upcurved in lateral view and outcurved in dorsal view; with sparse setae ventrally at anterior half; cuspis thin, slender, elongate, equally wide throughout in dorsal view and lateral view; with scattered inconspicuous long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, virtually as wide as long with rubrounded and sparsely setose posterodorsal margin in lateral view; setae shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, subcapitate in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations: FEMALES. Integument black, except mandibles and antennal flagellomeres partially reddish-brown, and T2 with four yellowsih, narrow, linear integumental spots; anterior pair longitudinal; posterior pair narrowly confluent thus forming apparent single travserve line. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density (coppery on mesosomal dorsum): malar space, ventral third of lateral pronotal face, mesosomal dorsum laterallly, ventral half of mesopleuron and metapleuron, lateral propodeal face, legs (except tarsi); T1 laterally, T2 (except disc); fringe of T2–4 medially and laterallly; fringe of T5 medially, T6 (except pygidial plate) medially, and S1–5 (except fringe of S5). MALES. Integument black. Body setae predominantly black varying in density, except following areas with silvery-white to silvery-golden setae varying in density: posterior half of T1, basal margin of T2, lateral margins of T2, fringe of T2–3, and fringe of S2–3 laterally. Distribution. Brazil and Paraguay. Material examined. (22f#, 1m#) Type material. Lectotype of Mutilla unimarginata, f#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH); holotype of Mutilla cuiba, m#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. BRAZIL: Mato Grosso: Chapada dos Guimarães: 1f#, 12.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 6f#, 18.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 3f#, 16.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 1f#, 14.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 2f#, 16–25.VII.2001, Alexandre, R., Ribeiro, K. (DZUP); APM-Mauso [sic], Areia Branca, 2f#, 26.III.2001, Sousa, W.O. 165
(DZUP); Fazenda Buriti, 1f#, 18.XI.1982, M. Zanuto & W. Overal (MPEG); 2f#, X [sic] (ANSP); Campo, 1f#, Nov. [November] (MNHN); Paraná: Curitiba, S. [São] José dos Pinhais, 1f#, 27.VIII.2003, Maria, V.A. (DZUP); PARAGUAY, 1f# (CUIC). Remarks. The sex association between T. unimarginata and T. cuiba is evident since these are the only species of the T. indica species-group with distinctly black spurs (males and females of T. puella have variably yellow to dark yellow- brown spurs) in the Chapada dos Guimarães area in Mato Grosso state, Brazil. As previously mentioned, the females of T. unimarginata possess a conspicuous color syndrome typical of the Chapda dos Guimarães area, in which the mesosomal stripes are coppery. This pattern is observed in T. aemulata, T. unimarginata and T. chapada and also some members of undescribed species in Ephuta Say, 1836 and Pertyella Mickel, 1952 (KAW & PRB pers. obs.). Of these similarly colored species, T. unimarginata has the darkest, thickest, and most red-tinged mesosomal stripes that, at first glance from above, give the appearance of the whole mesosoma being reddish, a common pattern for the genus Timulla Ashmead, 1899 and others (Wilson et al. 2015). Our knowledge of the male of T. unimarginata is unfortunately still limited to the type specimens of T. cuiba, even though females have been collected recently and even found outside of Chapada dos Guimarães. As with T. puella, the female mesosomal sculpture varies; certain specimens have the dorsal sculpture uniform areolate-punctate throughout, while others have the areolations obliterated adjacent to the medial longitudinal carina.
Traumatomutilla vidua (Klug, 1821) (Figs. 32A–L, 33A–G)
Mutilla vidua Klug, 1821. 313, holotype [by monotypy], m#, Brazil, Para [sic] (ZMB), examined. Mutilla graphica Gerstaecker, 1874: 74, lectotype [designated here], f#, Brazil, Rio de Janeiro [sic1] (ZMB), examined, syn. nov. Mutilla scripta Gerstaecker, 1874: 74, lectotype [designated here], f#, Brazil, [Rio Grande do Sul], Allegrette [Alegrete] (ZMB), examined, syn. nov. Ephuta (Traumatomutilla) vidua: André, 1902, 56. Ephuta (Traumatomutilla) graphica: André, 1902, 55. Ephuta (Traumatomutilla) scripta: André, 1902, 55. Traumatomutilla vidua: André, 1904, 40. Traumatomutilla graphica: André, 1904, 40. Traumatomutilla scripta: André, 1904, 40. Traumatomutilla cachimba Casal, 1969: 285, holotype, f#, Brazil, Pará, Cachimbo (AMNH), examined, syn. nov. Traumatomutilla scripta borrosa Casal, 1969: 290, holotype, f#, Argentina, Tucumán, San Pedro de Colalao (AMNH), examined, syn. nov.
Diagnosis. FEMALE. Mesonotum wider than distance between pronotal spiracles; scutelar scale and anterolateral carinae conspicuous, well-defined, clearly visible amognst dense setation; head setae always black; MALE. Cuspis slender, elongate, mesopleuron tuberculate on dorsal half; meso and metatibial spurs yellowish-white; scutellum with well-defined dorsal and posterior faces; axillar projections short, broadly and transversely trunctate; posterior tooth of penis valve longer than anterior tooth. Description. FEMALE. Body length 15–18 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Sculpture partially concealed by dense setae, densely and coarsely foveolate-punctate where visible, with subscabrous intervals at front; sculpture sparser on gena and malar space, with sparse interspersed micropunctures; with well defined medial longitudinal carina dorsally on front. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth. Dorsal scrobal carina well defined, connected to lateral scrobal carina. Antennal tubercle irregularly rugose. Flagellomere 1 2.1 × pedicel length; flagellomere 2 1.3 × pedicel length. Mesosoma. Mesosoma slightly broader than long. Mesosomal dorsum sculpture partially concealed 166 by dense setation, densely and coarsely areolate-punctate where visible with subscabous intervals; medial longitudinal carina present on mesonotum and dorsal face of propodeum, interrupted near scutelar area. Anterior face of pronotum defined, virtually as long as pronotal collar; with indistinct, indistinct and coarse longitudinal striations basad and dense coarse, and confused punctures dorsad; dorsal face anguletaly rounded into anterior face in lateral view. Humeral carina well-defined, separated from epaulet; epaulet virtually flat against anterior margin of pronotum; anterolateral corners of pronotum subangulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; with small blunt tubercle anteroventrally in relation to pronotal spiracle; mesopleuron sculpture mostly concealed by dense setation, micropunctate anteriorly and densely coarsely foveolate-punctate along mesopleural ridge where visible; metapleuron sculpture concealed by dense setation on ventral half, completely asetose, smooth on dorsal half. Lateral face of propodeum sparsely and coarsely foveolate-punctate anterad, densely and coarsely foveolate-punctate posterad; intervals smoother and wider anterad. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 61:76:77:56:55. Lateral margin of mesonotum constricted anterior to propodeal spiracle, diverging anterad, expanded. Propodeal spiracle projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale present, transverse, arched, reaching propodeal spiracles connected laterally with anterolateral carinae; anterolateral carinae well-defined, wider than scutelar scale, reaching propodeal spiracles laterally; scabrous intervals present on scutellar area. Propodeum convex, dorsal face virtually as long as and rounded into posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 39:95:91. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations slightly sparser and micropunctures almost absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely and coarsely punctate. S1 surface cuneiform, densely, coarsely and confusedly foveolate- punctate around sharp longitudinal medial carina slightly projected posteriorly. S2 sparsely foveolate-punctate, more sparsely so posteromediad; anteromedial crest-fold well-defined. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with interspersed micropunctures where visible; S5–6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface with transverse coarse and irregular rugosities; interstice granulose. MALE. Body length 14-20 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view, 0.75 × as wide as pronotal width. Eye almost circular. Ocelli small; OOD 3.8 × DLO, IOD slightly longer than DLO. Occipital carina distinct. Head surface densely and finely punctate with interspersed micropunctures; sculpture conspicuously sparser around ocelli. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and confusedly punctate; apical/ventral margin with a pair of medial subrounded free teeth. Scape bicarinate. Flagellomere 1 2.2 × pedicel length; flagellomere 2 3.0 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, broadly separated from humeral carina, anterolateral angles of pronotum subangulate. Anterior face of pronotum coarsely and densely punctate laterally, micropunctate sublaterally and virtually unsculptured medially; slightly longitudinally concave medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-punctate, notaulus and parapsis present, reduced to posterior half of mesoscutum. Scutellum subglobose, densely and coarsely foveolate-punctate; with conspicuous dorsal and posterior faces; posterior face almost vertical; dorsal face with medial longitudinal “carina-like” area indistinctly extending into posterior face. Axilla in dorsal view produced posterolaterally as short truncate projection, with conspicuous flat coarsely and densely foveolate-punctate dorsal surface. Metanotum slightly wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, densely areolate; sculpture of lateral face less defined to indistinct along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely and indistinctly punctate-rugose with dense interspersed micropunctures; mesopleura with conspicuous blunt spine-like projection on dorsal half; mesopleural sculpture densely and coarsely areolate-punctate to foveolate-punctate with interspersed micropunctures; simply 167 micropunctate anterad. Metapleuron sparsely micropunctate to smooth throughout, except for indistinct areolations on ventral fifth and indistinct rugosities in dorsal fifth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.85 × its width. Dorsal metasomal sculpture partially concealed by dense setation, densely and finely punctate with interspersed micropunctures where visible. Pygidial plate irregularly, and indistinctly rugose laterally, granulose elsewhere, broader than long, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in large blunt spine-like projection. S2 finely and sparsely foveolate-punctate to punctate, with micropunctures laterally and postero-medially; with conspicuous subovate antero-medial setae filled pit; longitudinal anteromedial crest-fold present. S3–6 sparsely and coarsely foveolate- punctate with sparse interspersed micropunctures; S7 sparsely foveolate-punctate with well-defined medial longitudinal unsculptured area; longer than broad, well defined by lateral carinae throughout, posterior margin projected medially into a single tooth-like structure on posterior margin; apex of projections bilobate. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 85:68:21; paramere virtually straight and slightly curved outward apically in dorsal view, upcurved apically in lateral view; with sparse setae ventrally at basal half; cuspis slender, elongate, equally wide throughout in dorsal view and lateral view; with scattered long setae apically and sparse inconspicuous short setae elsewhere; paracuspis well-developed, not sessile, subtriangulate in lateral view, virtually as long as wide with subtriangulate and densely setose posterodorsal margin in lateral view; setae shorther than paracuspis; digitus short, slightly curved inward in dorsal view and upcurved in lateral view, sparsely setose basodorsally, apex somewhat expanded, knob-like in lateral view; penis valve with inner surface strongly concave, and well-defined pair of short teeth posteroventrally; posterior tooth acute, subposterior tooth subrounded, blunt, with externolateral pocket; apical distance between teeth 0.1 × length of valve; dense setae present along subtruncate, shelf-like posterior margin; inconspicuous setae present at base of subposterior tooth on external surface. Coloration and variations. FEMALES. Integument black to brownish-black, except for mandibles and antennal flagellomeres partially reddish-brown and T2 with four narrowly sublinear to broadly subrectangular integumental spots varying from yellowish to reddish. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: malar space (also black occasionally); ventral third of lateral pronotal face; ventral half of meso and metapleuron; mesosmal dorsum usually with a pair of lateral stripes on mesonotum and propodeum broadly interrupted on scutelar area, or a pair of longitudinal uninterrupted stripes extending from posterior margin of propodeum to halfway into pronotal dorsum, rarely with stripes restricted only to propodeal dorsum; legs predominantly (completely black occasionally); T1 laterally; T2 (except disc); fringe of T2–4 medially and laterally; fringe of T5 medially; T6 (except pygidial plate) medially; and S1–4 (except fringe of S4 occasinally). MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except following areas with silvery-white setae varying in density: apical/ventral margin of clypeus, metanotum medially, dorsum of propodeum, ventral surface of femora, inner surface of tibiae, T1, T2 anterolaterally, lateral margins of T2 posteriorly, fringe of T2–4 laterally, S1, and S2–4 indistinctly. Tibial spurs yellowish-white. Wings dark-brown infuscated throughout with faint blueish/purplish reflections. Distribution. Colombia, Brazil, Ecuador, Bolivia, Paraguay, and Argentina. Material examined. (432f#, 101m#) Type material. Holotype of Mutilla vidua, m#, BRAZIL, Para [sic] (ZMB); lectotype of Mutilla graphica, f#, BRAZIL, Rio de Janeiro [sic1] (ZMB); lectotype of Mutilla scripta, f#, BRAZIL, [Rio Grande do Sul], Allegrette [Alegrete] (ZMB); holotype of Traumatomutilla cachimba, f#, BRAZIL, Pará, Cachimbo (AMNH); holotype of Traumatomutilla scripta borrosa, f#, ARGENTINA, Tucumán, San Pedro de Colalao (AMNH). Additional material. COLOMBIA: Antiochia, Medellín, 1f#, Gerard (ZMB); Amazonas, Monkey Island near Leticia, 1m#, 21.IX.1979, M.A. Tidwell (DGMC); BRAZIL: 1f# (ZMUC); South Brazil [sic], 1f#, X.1999, K.G. Ross (EMUS); 1f#, n.o. proc. 10/49 [sic], 1f#, X.1949 (MNRJ); Rio Machados [sic], 1m# (DEI); 1m# (RMNH); Pará: 5f# (MZSP); Óbidos, Colônia São Tomé, 1°50'46"S 55°32'24"W, 1f#, 21.IX.2010, Almeida, J. (MZSP); Óbidos, 3#m, VII.1959, F.M. Oliviera (AMNH); Santarém, 1f# (MNCN); Belterra, Alter do Chão, 2°29'38"S 54°57'26"W, 1f#, 16.IX.2010, Expedição Mocorango (MZSP); Alto Parú D'Oeste, Tiriós, 2f#, I–II.1963, Machado e Pereira (MNCN); Almeirim [Arrayolos], 1m#, 168
24.IV.1903, Ducke (MNHN); Juruti, Jaratuba, Rio Mamuru, Área 2, 03°11'32,6''S 56°34'51,4''O, 2m#, 29.IX.2009, O.T. Silveira, S.S. Silva & J. Pena (MPEG); Monte Alegre: Serra do Elerê, Gruta Pedra Pintada, 1f#, 15.III.1994, O. Silveira & A. Henriques (MPEG); Cerrado, 1m#, 16–20.I.1992, A.L. Henriques (MPEG); Gorotire, 51W8S, 1f#, 09.VI.1978, D.A. Posey (MPEG); Tumukumake, Aldeia Araibá, 1m#, 05.II.1981, E. Oliveira (MPEG); Amazonas: Parintins, 1f#, Knudsen, S.G. (ZMB); São Paulo de Olivença, 1m#, X–XI.1878, M. de Mathan (MNHN); Borba (Lago Acara), 2f#, X.1943 (MNRJ); Humaitá, Rio Ipixuna, 07°31'18''S 63°20'48.8''W, 1f#, 02.X.2014, J.A. Rafael, F.F. Xavier F°, R.M. Vieira, R.H. Aquino (INPA); Amapá: Calçoene, 2m#, 30.X.1978, W. França (MPEG); Mazagão, Fazendinha, 1m#, 03.XII.1980, E.L. Oliveira (MPEG); Rondônia: Rio Guapore at Rio Baures, 1m#, 26.IX.1964, Bouseman & Lussenhop (AMNH); Vilhena, 1m#, 23.II.1961, J. & B. Bechyné (MPEG); 62 km SW [southwest of] Ariquemas, nr [near] Fzda [fazenda] Rancho Grande, 1m#, 06–15.XII.1990, D.A. Rider & J.E. Eger (FSCA); Itapua do Oeste, FLONA [Floresta Nacional] do Jamari: 2f#, 12.VI.2013, Luz, D., Rosa, B., Williams, K.A. (DZUP); 3m#,11.VI.2013,Luz, D.; Rosa, B.; Williams, K.A. (DZUP); Mato Grosso: Chapada dos Guimarães: 1f#, 15.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 1m#, 18.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); Rosario Oeste, 2f#, I.1976 (MNRJ); Cáceres, 1f#, 12.II.1985, Elias, C. (DZUP); Salôbra, 1f#, 19–30.I.1940, F. Lane (MZSP); marg. [margem] esq. [esquerda] rio Sucuriu, Faz. [Fazenda] Canaã, Três Lagoas, 1f#, X.1966, F. Lane (MZSP); São Paulo: Rio Claro, 1m#, B. Dias (AMNH); Teodoro Sampaio, 1m#, 15.II.1999, Melo, G.A.R. (DZUP); Reserva de Jatai, Luis Antônio, 1f#, 16.X.1999, Melo, G.A.R. (DZUP); Tambau, 1f#, I.1971, Silva, L.C. (DZUP); Distrito de Sousas, Campinas, 1f#, 26.VIII.1988, R.P. Martins (UFMG); Faz. [Fazenda] Itaquerê, Nova Europa, 1f#, 14.XII.1964, K. Lenko (MZSP); Guzolandia, Tres Irmaos ranch, 1f#, 31.III.1999, A.F. Santos (EMUS); Porto Cabral,Rio Paraná, 1f#, 01–25.IV.1944, Trav. Fo. & Carrera & E. Dente (MNCN); Rio Grande do Sul: Pelotas, 1f#, 04.II.1966, C. Biezanko (MZSP); Caçapava do Sul, Pedra do Segredo, 1f#, 07.III.2010, P. Bartholomay (CESC); Ijui, 1f#, 20.II.2007, Muller, G.A. (DZUP); Porto Alegre, Morro do Coco, 1f#, 25.I.1962 (CESC); Candelária, Ponte do Império, 1m#, 28.XII.2004, A. Köhler & M. Hermes (CESC); Camaquã, Ilha de Três Bocas, 1f#, 01.II.1976 (BMNH); Espírito Santo: Cariacica, 1f#, 21.II.2001, Coser, C. (UFES); Vitória, 1f#, 22.X.1981 (UFES); Baixo Guandu: 1f#, 15.VIII.1970, Elias,C. & Elias, C.T. (DZUP); 1m#, 23–31.III.1970, Elias, C. & Elias, T. (DZUP); Santa Teresa, 1f#, 24–31.IV.1967, Elias,C. & Elias,C.T. (DZUP); Conceição da Barra, 1f#, 1968, Elias,C. & Elias, C.T. (DZUP); Pedro Canário, 1f#, 12.X.1971, Enoqui (DZUP); Colatina, 3#f, I.1970 (MNRJ); Campus da UFES [Universidade Fedral do Espírito Santo], Vitória, 1f#, 26.XI.1988, E.M. Freitas (UFES); Corrego D'Água, Aracruz, 1f#, 17.VI.2001, Bertazzo, K. (UFES); Duas Bocas, Cariacica, 1f#, 03.III.2001, Gomes, I.T. (UFES); Barra do Jucu, Vila Velha, 1f#, 19.VIII.1987, F. Vieira (UFES); Linhares, Sooretama, 3f#, I.1949 (MNRJ); Morro Mestre Álvaro [Alvares], Serra, 1f#, 16.IV.1987, A.P. Aguiar (UFES); PEPCV, Setiba, 1f#, 22.XI.2000, Kawada (UFES); São Francisco Batatal, Alfredo Chaves, 1f#, 23.II.2004, Fraga, F.B. (UFES); Bahia: 1f#, 28.III.2001, J.R. Maia (CPDC); Vitoria da Conquista, 1f#, I.1963 (MNRJ); Barrolandia, 1f#, III.1999, J.R.M. Santos (CPDC); Belmonte, 1f#, 15.V.2004, J.H.C. Delabie (CPDC); Una, 1f#, 21.VII.1997, J.R. Maia (CPDC); Zona Rural,T[eixeira]. de Freitas, 1f#, 30.VII.2003, Fraga, F.B. (UFES); Mato Grosso do Sul: Porto Murtinho, 2f#, XI.1929 (DEI); Três Lagôas, 5f# (MZSP); Anastácio, Fazenda Boa Esperança, 1f#, 16– 26.II.2008, Bossi, R. & Bervian, C. (MZSP); Corguinho, Taboco, RPPN [Reserva Particular do Patrimônio Natural] Quintas do Sol, 284m [above sea level], 39°46'26''S 55°14'38''W, Cerrado, 1f#, 30.X.2011, Silva, R.J. (Mu-Bio); Jardim, Sta. [Santa] Maria, 250m [above sea level], 21°32'46''S 56°55'29''W, 1f#, II.2008, Carbonari, V. (Mu-Bio); Aquidauana, malaise 09, 20°26'07"S 55°39'33"W, 2m#, 11–26.X.2011, Lamas & Nihei (MZSP); Pedra Azul, 1m#, XI.1972, Alvarenga & Seabra (AEIC); Minas Gerais: nr. [near] Timoteo, 1m#, 30.IX.1997, E.R. DePaula (EMUS); 11 km S de Divino das Laranjeiras, 1f#, 26.XII.2000, Melo, G.A.R. (DZUP); Cássia, 1f#, 13.II.1963, Elias,C. (DZUP); Uberlândia, Faz. [Fazenda] Três Irmãos, 1f#, 19.XI.2004, Probio (DZUP); Zona Rural,Nanuque, 1f#, 24–29.VII.2007, Fraga, F.B. (UFES); 12km a N [norte] de Aguas Vermelhas, Faz. [Fazenda] Faceiro, 1m#, 14.XII.2012, Melo, G.A.R. (DZUP); Goiás: Rio Araguaya, A22 [sic], 1f# (MNRJ); Faz. [Fazenda] Aceiro, Jataí, 1f#, X.1962, Exp. Dep. Zool. [Expedição Departamento de Zoologia] (MZSP); Faz. [Fazenda] Cachoeirinha, Jataí, 1f#, X.1963, Exp. Dep. Zool. [Expedição Departamento de Zoologia] (MZSP); Serranópolis, Pousada das Aranas, 1f#, 3.VII.2008, Melo, G.A.R. (DZUP); Paraná: P. [Parque] Estadual de Vila Velha: 3f#, 7.III.2003, Melo, G.A.R & Gonçalves, R. (DZUP); 1m#, 7.XII.2002, Melo, G.A.R & Gonçalves,R. (DZUP); Maranhão, Imperatriz, 1f#, 15.VII.1974, Exp. Dept. Zoo. [Expedição Departamento Zoologia] 169
UFPR [Universidade Federal do Paraná] (DZUP); Tocantins, Arraias, P1 RV MD [sic], 1f#, 24.II.2002, Filho A.R.S. e H.S. Morais (UFT); Acre, 64f# (MZSP); ECUADOR, Pichinche, 1f#, Expedición al Pacífico (MNCN); PERU: 1f# (BMNH); [Pasco], Paucartambo, 2f#, J. Clermont (MNCN); [San Martin], Tarapoto, 1f# (MNCN); BOLIVIA: Santa Cruz: El Carmen, 1f#, 25–27.II.1954, C. Gans & F. Pereira (MZSP); 40 km NW [northwest] Santa Cruz, Potrerillo de Guenda, 1f#, 19.XII.2004, Gino Nearns (CMBC); 4–6 km SSE [south southeast] Buena Vista, Hotel Flora y Fauna, 1f#, 02–12.II.2000, J.E. Wappes (TAMU); Santa Cruz: E [East] of Warnes,1m#, XII.1983, R.C. Wilkerson (FSCA); 5 km ESE [east southeast] Warnes, Hotel Rio Selva, 1f#, 13.X.2000, B.K. Dozier (FSCA); Cordillera, Las Juntas, 1f#, II.1947, Peredo (MNCN); Beni: Rio Itenez, 4 km above Costa Marques (Brazil), 1f#,12–18.IX.1964, J.K. Bouseman & J. Lussenhop (AMNH); near Rio Mapiri, 1m#, IX.1921, W.M. Mann (USNM); La Paz: San Buenaventura, Ixiamas rd. [road] km 23, Hacienda Chiquitos, 1f#, 13.VII.2003, Robertson & Blahnik (UMSP); Santa Ana de Alto Beni, 300m [above sea level], 2f#, 15–16.IV.1979, M. Cooper (BMNH); Guanay, Uyapi, 1m#, X.1995, Gerlach (AMNH); PARAGUAY: Pto. [Puerto] Pablo [sic]: 2m#, XI–XII.1936 (RBINS); 3f#, XI–XII.1936 (RBINS); Paraguari: La Rosada, 560', 26°06'S 56°50'W, 4f#, 27–30.XII.2008, U. Dreschel (EMUS); Ybycui National Park, 1f#, 12–24.IV.1980, P.J. Spangler (USNM); 1m#, 22.I.1981, H. Ferreira (USNM); Presidente Hayes, Puerto Galileo, 230', 25°04'S 57°52'W, 1f#, 05–08.III.2008, U. Dreschel (EMUS); San Pedro: Rio Ypane: Cororo: 1f#, XI.1983, M.A. Fritz (AMNH); 1m#, II.1994, Arriagada (AMNH); Villarica, 1f#, 15.X.1924 (DEI); Chaco, San Bernardo, 1f#, di Iorio (AMNH); Caaguazu, Zudanez, 1f#, II.1949 (UMMZ); Distrito Capital, Assomption [Asunción], 1f# (RBINS); ARGENTINA: Chaco: Tres Estatas, 1f#, di Iorio (AMNH); San Bernardo: 1f#, Di Lorio (EMUS); 2m#, DiLorio (AMNH); Chaco Austral, Ingênio Las Palmas, 1f# (DEI); 100 km NW [northwest] Resistencia, NP [National Park] Chaco, 1m#, 12.XII.1990, S & J Peck (PMAE); Tres Estacas, 1m#, II.1991, DiLorio (AMNH); Cordoba: Dolores, ca. [circa] La Cumbre, 1f#, 08.III.1995, C. Porter (FSCA); Cordoba, 1f#, 1919, E. Giacomelli, (MNCN); Villa Cabrera, 1f#, 1919, E. Giacomelli (MNCN); Capillo del Monte, 1f#, 1928, E. Giacomelli (MNCN); Corrientes: Perrugorria, 1f#, 13.IV.1950 (MNHN); Ytuzaingo, 1f#, III.1982, Fritz (FSCA); 7m#, III.1982, Fritz (AMNH); Manantiales, 1f#, I.1960, I. Apostol (MNCN); Paso de la Patria, 1m#, 20.I.1972, D.J. Brothers (DJBC); Entre Rios: [Pueblo] Liebig: 1f#, II.1979, Zelich (AMNH); 1m#, III.1995 (EMUS); Colon, Ita Cora, 1f#, II.1997, L. Caire (EMUS); Palmar Colon, 5m#, Fritz (AMNH); Formosa: Estancia Guaycolec, 25 km N [north of] Formosa, 2f#, 26.II–10.III.1999, Heydon & Ledford (UCDC ); Est. [Estancia] Guaycolsec, 25 km N Formosa, 2m#, 10.III.1999, Heydon & Ledford (UCDC); Jujuy: San Pedro de Jujuy, 1f#, 02.V.1968, C.C. Porter (USNM); Yuto, 2m#, 10.I.1966, H & M Townes (AEIC); Mendoza, Mendoza, 1f# (MNHN); La Rioja, 3f#, 1919, E. Giacomelli (MNCN); Misiones: 1f#, E. Le Moult (MNCN); Rio Paraná, Tiju-Cuare, Missão de San Ignácio, 2f#, 1911, E.R. Wagner (MNHN); Santa Fé: 1m#, 11.IX.1942 (CUIC); Huerta Gr. [sic1], 1m#, I.1910 (UMSP); Las Garzas, 1f#, E. Le Moult (MNCN); Santiago del Estero, Rio Salado, 12f#, 1916, E. Le Moult (MNCN); Tucuman: Tucuman: 1f#, 1.III.1902, J. Steinbach S.V. (ZMB); 2m#, 1.III.1902, J. Steinbach S.V. (ZMB); Catamarca, Punta [de] Balasto: 1m#, I.1992, Fritz (AMNH); 7m#, XII.1976, Fritz (AMNH); URUGUAY: 1f#, J. Wyman (MCZC); 1f#, Cornell Univ. [University] Expedition (CUIC); Montevideo, Isern, 1f#, 1862–1865, Expedición al Pacífico (MNCN); Rio Negro, R. [Rio] Grande, km 287, ruta 3, 2m#, I.1965, Carbonell (AMNH). An additional 212f# and 26m# from various localities in BRAZIL, BOLIVIA, PARAGUAY, and ARGENTINA have also been examined (LACM, MNRJ, MNCN, MZSP, FSCA, DZUP, ZMB, AMNH, USNM, CMNH, MPEG, MNHN, CESC, ZMUC, AEIC, ZMB, BMNH, Mu-Bio, UFES, CASC, CMBC). Remarks. The synonymy and consequent sex association of T. graphica and T. vidua was made evident by the fact that typical forms of both species occur from Colombia to Argentina with scarcely any variation. Three fomerly recognized species, T. cachimba, T. scripta and T. scripta borrosa, were subsequently recognized to share identical structural features with the most commonly seen form, T. graphica. Males of T. vidua are virtually unaltered in color or structure throughout their range, with specimens from the Amazon being identical to those collected in Argentina. The similarities between T. vidua, T. spectabilis, and T. protuberans are intriguing and indicate that further synonymy may be eventually needed or that these are closely related. While both sexes of T. spectabilis can approximate these structures, they always appear to be somewhat deformed or poorly developed versions of the well-developed broad, flat, truncate axillar projections of T. vidua males or the discrete transverse carinae anterolateral to the scutellar scale of T. vidua females. Males of T. 170 protuberans similarly have their axillar projections appearing to be a poorly developed version of those seen in T. vidua males.
Discussion
This is the largest Traumatomutilla species-group in number of species and number of specimens examined. Issues such as the relation between T. spectabilis, T. protuberans and T. vidua, however, demand even more specimens to validate if the provided morphological diagnostic features are valid. Though an argument could be presented that theses three species belong to one highly varible species distributed throughout South America, the current hypotheses are consistent with the available material. Additionally, a hypothesis that these species are all conspecific would begin to invalidate the use of characters that are consistent and unvariable in other species in the T. indica species-group and that have been efficiently used to separate species within Traumatomutilla as a whole. Likewise, additional females of T. contempta, T. geographica, and T. mundula are still needed to determine if these are distinct species for which the male is unknown, or they are conspecific variants of other species. The only stable structural characters that are found throughout the T. indica group are the longidutinal carina on the mesonotum of females and the elongate rectangular hypopygium defined by lateral carinae of males. The longitudinal carina seems to be correlated with changes in the mesonotal sculpture; species that have this carina well-developed often have the sculpture simply and densely punctate in the areas adjacent to it, while species with a reduced carina have the mesonotum uniformly areolate-punctate throughout. Although this carina is found in every species of the T. indica species-group, albeit reduced or indistinct in some cases, it is not exclusive to this species-group and can be found in a reduced form in some individuals from the T. gemella, T. quadrinotata and T. juvenilis species-groups. Similarly, the elongate hypopygium of the males, though conistent within the T. indica species-group, is approximated by males of the T. quadrinotata species-group, which differ by lacking the lateral carinae defining the hypopygium. In addition to these characters, females of the T. indica, T. juvenilis, T. gemella, and T. quadrinotata species-groups are the only groups with a subpyriform pygidial plate where the lateral carinae are dorsally obliterated or obscured by setae. Likewise, males of the T. indica, T. juvenilis and T. quadrinotata species-groups almost always have the dorsal half of the mesopleuron projected into a blunt or acute tubercle, except for two species in the T. indica species-group;this feature can also be found in males of the T. bifurca and T. vitelligera species-groups (Bartholomay et al. 2019b). Additionally, the acute axillar projections found in all but four species of the T. indica species-group can only be found in the males of the T. gemella, T. bifurca and T. virginalis species-groups (Bartholomay et al. 2019b, KAW & PRB pers. obs.). These similarities, found in both sexes of the T. indica species-group, indicate that it is likely more closely related to the T. juvenilis, T. gemella, and especially the T. quadrinotata species-group, than to any other species-group in the genus. As observed in the T. juvenilis species-group (Bartholomay et al. in press), the sex associations achieved here were made possible first and foremost by discovery of one or more irrefutable associations based on distribution. This then provided vital morphological clues to decipher the remaining associations in the group. For example, once both sexes of T. grossa were discovered, the unique cuspis in the genitalia of the male allowed us to discover two additional male forms with the same character. Specific females that occurred in the same area as these male morphopecies were then compared with females of T. grossa and found to be similar in size, sculpture, and structure to this species, making them obvious candidates for association with the aforementioned males. Certain couples within the T. indica species-group demonstrate that morphological parallels between sexes can be used as additional clues to associate males with females. The males of T. vidua are unique within the T. indica species- group for having broad transversely truncate axillar projections that are mirrored by the well-defined transverse anterolateral carinae of the females, also unique when compared with other species in the group. Additionally, both sexes of T. centralis and T. unimarginata have the meso- and metatibial spurs black, while the same structure varies from brownish to yellowish-white in series of both sexes of T. puella collected in the same area. Situations like these indicate 171 that, even though these wasps are extremely sexually dimorphic, there are indeed morphological parallels between sexes that provide vital supporting evidence for associating the sexes. This being the most ubiquitous, commonly observed, and commonly collected species-group of Traumatomutilla, we are confident that the plates, keys and descriptions provided herein will allow others to identify these wasps and provide the much-needed additional specimens required to further elucidate this still poorly understood genus.
Acknowledgements
We are grateful for the collection managers and curators that provided specimens for this study, including: Christine LeBeau (AMNH), Robert Zuparko (CASC), Andreas Köhler (CESC), John Rawlins (CMNH), Stephan Blank (DEI), Gabriel Melo (DZUP), James Pitts (EMUS), Mercedes Paris (MNCN), Agniéle Touret-Alby (MNHN), Felipe Vivallo (MNRJ), Orlando Silveira (MPEG), Kelli Ramos (MZSP), Fernando Silvestre (MuBio-UFGD), Gavin Broad (BMNH), Lynn Kimsey (UCDC), Fernando Silveira (UFMG), Robin Thomson (UMSP), Brian Harris (USNM), Michael Ohl, Viola Richter and Lukas Kirscher (ZMB), Lars Vilhelmsen (ZMUC), Denis Brothers (DJBC), Donald Manley (DGMC), Wouter Dekoninck (RBINS) and Karla Schneider (MLUH). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9).
References
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Figure 1A–C. Traumatomutilla aemulata (Cresson, 1902), lectotype, female, line 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 2A–G. Traumatomutilla caneta (Cresson, 1902), holotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 3A–C. Traumatomutilla borba (Cresson, 1902), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels.
Figures 4A–F. A–C. Traumatomutilla centralis (Burmeister, 1875), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–E. Traumatomutilla centralis boliviana Suárez, 1960, female, line 2mm; D. Dorsal habitus; E. Lateral habitus; F. Traumatomutilla centralis, females, color variant.
Figures 5A–F. Traumatomutilla fissiventris André, 1907, lectotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 6A–F. A–C. Traumatomutilla contempta André, 1908, holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla alhuampa Casal, 1969, holotype, female, line 2mm; D. Dorsal habitus; C. Lateral habitus; F. Type labels.
Figures 7A–F. A–C. Traumatomutilla geographica (Gerstaecker, 1874), holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla seabrai Casal, 1969, holotype, female, line 2mm; D. Dorsal habitus; C. Lateral habitus; F. Type labels.
Figures 8A–F. A–C. Traumatomutilla grossa (Gerstaecker, 1874), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla abrupta (Gerstaecker, 1874), holotype, female, line 2mm; D. Dorsal habitus; C. Lateral habitus; F. Type labels.
Figures 9A–G. Traumatomutilla characterea (Gerstaecker, 1874), lectotype, dorsal habitus, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 10A–F. A–C. Traumatomutilla guayaca Casal, 1969, holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla tayguaya Casal, 1969, holotype, female, line 2mm; D. Dorsal habitus; C. Lateral habitus; F. Type labels.
Figures 11A–F. Traumatomutilla guayaca Casal, 1969, male, line 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 12A–G. Traumatomutilla impetuosa (Smith, 1879) comb. n., holotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve 174 removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 13A–C. Traumatomutilla indica (Linnaeus, 1758), holotype, female, line 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 14A–G. Traumatomutilla sphegea (Fabricius, 1804), holotype, male, 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 15A–C. Traumatomutilla ingens (André, 1903), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels.
Figures 16A–F. Traumatomutilla ingens (André, 1903), male, line 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 17A–C. Traumatomutilla mundula (Cresson, 1902), lectotype, female, line 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 18A–C. Traumatomutilla parallela (Klug, 1821), holotype, female, line 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 19A–J. A–B. Traumatomutilla lineifera (André, 1903), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C–D. Traumatomutilla gausapata Mickel, 1952, female, line 2 mm; C. Dorsal habitus; D. Lateral habitus; E–F. Traumatomutilla indicoides Mickel, 1952, female, line 2mm; E. Dorsal habitus; F. Lateral habitus; G–H. Traumatomutilla pillinata Casal, 1969, holotype, female, line 2mm; G. Dorsal habitus; H. Lateral habitus; I. Traumatomutilla lineifera (André, 1903), type labels. J. Traumatomutilla pillinata Casal, 1969, type labels.
Figures 20A–G. Traumatomutilla almada (Cresson, 1902), holotype, male, 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 21A–G. Traumatomutilla protuberans (Gerstaecker, 1874), holotype, male, 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 22A–I. A–B. Traumatomutilla puella (Gerstaecker, 1874), holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C–D. Traumatomutilla manca (Cresson, 1902), lectotype, female, line 2 mm; C. Dorsal habitus; D. Lateral habitus; E–F. Traumatomutilla peperina Casal, 1969, holotype, female, line 2mm; E. Dorsal habitus; F. Lateral habitus; G. Traumatomutilla puella (Gerstaecker, 1874), type labels; H. Traumatomutilla manca (Cresson, 1902), type labels; I. Traumatomutilla peperina Casal, 1969, type labels.
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Figures 23A–K. A–C. Traumatomutilla viana (Cresson, 1902), lectotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Dorsal habitus; D–F. Traumatomutilla musculus (Gerstaecker, 1874), holotype, male, line 2 mm; D. Lateral habitus; E. Type labels; F. Dorsal habitus; G–K. Traumatomutilla viana (Cresson, 1902), lectotype, male; G. Genitalia (halved), dorsal view; H. Genitalia (halved), ventral view; I. Genitalia (halved, penis valve removed), lateral/inner view; J. Cuspis (removed, not to scale), lateral/inner view; K. Penis valve (removed, not to scale), lateral/outer view.
Figures 24A–B. Traumatomutilla selligera (Gerstaecker, 1874), female, line 2 mm; A. Dorsal habitus; B. Lateral habitus.
Figures 25A–G. Traumatomutilla selligera (Gerstaecker, 1874), holotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 26A–F. Traumatomutilla spectabilis (Gerstaecker, 1874), lectotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla spectabilis chingona Casal, 1969, holotype, female, line 2mm; D. Dorsal habitus; C. Lateral habitus; F. Type labels.
Figures 27A–K. A–C. Traumatomutilla melaleuca (Gerstaecker, 1874), holotype, male, line 2 mm; A. Lateral habitus; B. Dorsal habitus; C. Type labels; D–F. Traumatomutilla tapera (Cresson, 1902), holotype, line 2 mm; D. Lateral habitus; E. Dorsal habitus; F. Type labels; G–K. Traumatomutilla melaleuca (Gerstaecker, 1874), holotype, male; G. Genitalia (halved), dorsal view; H. Genitalia (halved), ventral view; I. Genitalia (halved, penis valve removed), lateral/inner view; J. Cuspis (removed, not to scale), lateral/inner view; K. Penis valve (removed, not to scale), lateral/outer view.
Figures 28A–C. Traumatomutilla tristis (Klug, 1821), holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels.
Figures 29A–K. A–C. Traumatomutilla floccosa (Gerstaecker, 1874), holotype, male, line 2 mm; A. Lateral habitus; B. Dorsal habitus; C. Type labels; D–F. Traumatomutilla caxara (Cresson, 1902), holotype, male, line 2 mm; D. Lateral habitus; E. Dorsal habitus; F. Type labels; G–K. Traumatomutilla floccosa (Gerstaecker, 1874), holotype, male; G. Genitalia (halved), dorsal view; H. Genitalia (halved), ventral view; I. Genitalia (halved, penis valve removed), lateral/inner view; J. Cuspis (removed, not to scale), lateral/inner view; K. Penis valve (removed, not to scale), lateral/outer view.
Figures 30A–C. Traumatomutilla unimarginata (Cresson, 1902), lectotype, female, line 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 31A–G. Traumatomutilla cuiba (Cresson, 1902), holotype, male, line 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 32A– L. A–B. Traumatomutilla cachimba (Casal, 1969), holotype, female, line 2 mm; A. Dorsal habitus; B. Lateral habitus; C–D. Traumatomutilla scripta (Gerstaecker, 1874), lectotype, female, line 2 mm; C. Dorsal habitus; D. Lateral habitus; E–F. Traumatomutilla scripta borrosa Casal, 1969, holotype, female, line 2mm; E. Dorsal habitus; F. Lateral habitus; G–H. Traumatomutilla graphica (Gerstaecker, 1874), lectotype, female, line 2mm; G. Dorsal habitus; H. Lateral habitus; I. Traumatomutilla cachimba (Casal, 1969), holotype labels; J. Traumatomutilla scripta (Gerstaecker, 176
1874), lectotype labels; K. Traumatomutilla scripta borrosa Casal, 1969, holotype labels; L. Traumatomutilla graphica (Gerstaecker, 1874), lectotype labels.
Figures 33A–G. Traumatomutilla vidua (Klug, 1821), holotype, male, 2 mm; A. Lateral habitus; B. Type labels; C. Genitalia (halved), dorsal view; D. Genitalia (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view. 177
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CAPíTULO 4.10 – (formatado para submissão ao periódico ZOOTAXA)
The Traumatomutilla quadrinotata species group (Hymenoptera, Mutillidae)
PEDRO R. BARTHOLOMAY1,*, KEVIN A. WILLIAMS2, ROBERTO A. CAMBRA3 & MARCIO L. OLIVEIRA1 1Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, Laboratório de Hymenoptera, Av. André Araújo, 2936, Manaus, Amazonas, Brazil. 2Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento CA 95832, USA. 3Museo de Invertebrados G. B. Fairchild, Estafeta Universitaria Apartado 00017, Universidad de Panamá, Panamá 0824, República de Panamá *Corresponding author, e-mail: [email protected]
Abstract
The Traumatomutilla quadrinotata species-group is reviewed, leaving it with six species known from both sexes, four known from females only, and two known from males only. The following junior synonyms are proposed: Traumatomutilla austera (Gerstaecker, 1874) [=Mutilla sigillata Gerstaecker, 1874 syn. nov.]; Traumatomutilla chrysozona (Gerstaecker, 1874) [=Mutilla lugubrina Dalla Torre 1897 syn. nov.; =Ephuta dives André, 1906 syn. nov.]; Traumatomutilla quadripustulata (Klug, 1821) [=Mutilla pruinosa Smith, 1855 syn. nov.; =Mutilla maraca Cresson, 1902 syn. nov.]; Traumatomutilla sancta (Gerstaecker, 1874) [=Mutilla solemnis Cresson, 1902 syn. nov.]; Traumatomutilla incerta (Spinola, 1841) [=Mutilla dentata Smith, 1879 syn. nov.; =Mutilla sodalicia Kohl, 1882 syn. nov.; =Traumatomutilla tabatinga Casal, 1969 syn. nov.; =Traumatomutilla dignitosa Mickel, 1952 syn. nov.]; Traumatomutilla pompiliformis (Gerstaecker, 1874) [=Mutilla serra Cresson, 1902 syn. nov.]; Traumatomutilla infernalis [=Mutilla floccosa Gerstaecker, 1874]. The hitherto undescribed males of T. ameliae Casal, 1969 and T. quadrinotata (Klug, 1821), are fully described and illustrated. A new species, T. tetratrauma Bartholomay & Williams sp. nov., is described based on couples from the Atlantic Forest of Brazil. All previously described species are redescribed and illustrated. Identification keys for males and females are also provided.
Key words: Neotropical, Taxonomy, Velvet Ants, Sphaeropthalminae, Dasymutillini
Introduction
The recent work of Williams et al. (2017) subdivided Traumatomutilla André, 1901 into 14 species-groups based on species known from females that shared unique characters or combinations of characters. Up to that point Traumatomutilla had 183 valid species and subspecies ranging from Mexico to Argentina and, due to the extreme sexual dimorphism typical of velvet-ants, 133 of those species were known only from females, 48 only from males, and 2 were known from both sexes (Nonveiller, 1990; Williams et al. 2017). The species-group construct allowed for the study of this diverse genus to be compartimentalized and resulted in a series of recent revisions that redescribed, synonymized, and/or associated the sexes of more than half of the previously known species (e.g. Bartholomay et al. 2018, 2019a, b). With only three groups left to revise (T. quadrinotata, T. inermis, and T. trochanterata species- groups) and after the revision of the diverse Traumatomutilla indica species-group (in press), the genus is left with only one species-group that has consistently larged-bodied females (Traumatomutilla quadrinotata species-group) and a handful of males with similar size (PRB & KAW pers. obs.). In the present study we review the Traumatomuitlla quadrinotata species-group and associate these large-bodied males with said group.
Materials and terminology
Approximatelly 640 specimens were studied in the current work and the following codens are used for institutions housing the material discussed:
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AMNH - American Museum of Natural History, New York, New York, USA ANSP - Academy of Natural Sciences, Philadelphia, Pennsylvania, USA BMNH - British Museum of Natural History, London, England CASC - Department of Entomology, California Academy of Sciences, San Francisco, California, USA CPDC - Comissão Executiva do Plano da Lavoura Cacaueira, Ilhéus, Bahia, Brazil CISC - Essig Museum of Entomology, Department of Entomological Sciences, University of California, Berkeley, California, USA CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA CUIC - Cornell University Insect Collection, Department of Entomology, Ithaca, New York, USA CZMA - Coleção Zoológica do Maranhão, Caxias, Maranhão, Brazil DZUP - Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil DEI - Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany DGMC - Donald G. Manely Collections, Florence, South Carolina, USA EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA FMNH - Fields Museum of Natural History, Chicago, Illinois, USA FSCA - Florida State Collection of Arthropods, Gainesville, Florida, USA HBMNH - Hungarian Natural History Museum, Budapest, Hungary IAvH - Insituto Alexander Von Humboldt, Villa de Leyva, Colombia IEPA - Instituto de Pesquisas Científicas e Tecnológicas do Estado do Amapá, Macapá, Amapá, Brazil INPA - Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil LACM - Insect Collection, Los Angeles County Museum of Natural History, Los Angeles, California, USA MCZC - Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA MNCN - Museo Nacional de Ciencias Naturales, Madrid, Spain MNHN - Muséum Nationale d’Histoire Naturelle, Paris, France MNRJ - Museu Nacional do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil MPEG - Museu Paraens Emílio Goeldi, Belém, Pará, Brazil MRSN - Museo Regionale de Scienze Naturali, Torino, Italia MZSP - Museu de Zoologia da Universidade de São Paulo, São Paulo, São Paulo, Brazil BMNHW - Naturhistoriska Riksmuseet, Stockholm, Sweden RBINS - Royal Belgian Institute of Natural Sciences, Brussels, Belgium TAMU - Texas A & M University, College Station, Texas, USA UAIC - Univeristy of Arizona, Tucson, Arizona, USA UCDC - The Bohart Museum of Entomology, University of Californis, Davis, California, USA UEFS - Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brazil UMSP - University of Minnesota Insect Collection, St. Paul, Minnesota, USA UNEMAT - Universidade do Estado de Mato Grosso, Tangará da Serra, Mato Grosso, Brazil USNM - United States National Museum of Natural History, Simthsonian Institution, Washington D.C., USA. ZMB - Museum für Natrukunde Berlin, Berlin, Germany ZMUC - Zoological Museum University of Copenhagen, Denmark
General morphological terminology, definitions, abbreviations, and measurements followed that of Bartholomay et al. (2019). In the description and redescription sections we refrain from mentioning coloration and setal patterns due to the highly variable nature of these characters in Traumatomutilla. Instead, we provide a separate section titled Coloration and variations, in which we provide an overall description of the known color and setae patterns for a particular species. In the material examined section abbreviations, acronyms and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of the material examined section.
Taxonomy 188
Key to females of the T. quadrinotata species-group.
1. Lateral face of metapleuron and propodeum almost entirely concealed by dense golden appressed setation (Figs. 12B, 18B) sculpture sparsely foveolate punctate with densely micropunctate intervals where visible; vertex always clothed with black setae (Figs. 12A, 18A) … 2 Lateral face of metapleuron and propodeum generally smoother, at most with sparse erect silvery-white setae (e.g. Fig. 7C); sculpture entirely exposed, sparsely foveolate-punctate with smooth shinning unsculptured intervals (e.g. Figs. 1C, 8B, 14B); vertex sometimes clothed with silvery-white setae (e.g Fig. 8D) … 3
2. Frons with golden patch of golden setae (Fig. 18B); anterolateral carinae present in scutellar area; dorsal mesosomal setae more or less uniform in length (Fig. 18B) … T. tetratrauma Bartholomay & Williams sp. nov. Head setae entirely black (Figs. 12A–B); anterolateral carinae absent in scutellar area; pronotum and anterior portion of mesonotum with numerous long erect setae (Fig. 12B) … T. quadrinotata (Klug, 1821)
3. T2 marked with two round orange to reddish integumental spots (Figs. 8A, 8D, 8F, 8H); mesonotum with transverse silvery band, sometimes largely obliterated (Figs. 8A, 8D, 8F, 8H); occipital carina slightly but conspicuously swollen dorsolaterally (northern Amazon) … T. incerta (Spinola, 1841) T2 generally with four yellow to red integumental spots (e.g. Figs. 1A, 7A, 14A), anterior spots sometimes reduced or obliterated, posterior spots sometimes enlarged and confluent; mesonotum setae generally either entirely black (e.g. Fig 14A) or with lateral longitudinal stripes of silver to golden setae (e.g. Figs. 17A, 17D); occipital carina equally wide throughout …4
4. Head and mesosomal dorsum with setae entirely black, silvery-white setae greatly reduced on entire body, often completely absent (Figs. 4A–B; 14A–B) … 5 Silvery-white setae always conspicusouly present at least on propodeal dorsum and/or vertex (e.g. Fig. 3E) … 6
5. Post-mesonotal tubercle present, sometimes reduced but still conspicuous; lateral face of propodeum usually with large unsculptured smooth area posteriorly; silvery-white setae present, though greatly reduced, on different areas of T2–4, S1–4, and often on metapleuron (Fig. 14B) (Amazon) … T. quadripustulata (Klug, 1821) Post-mesonotal tubercle absent; lateral face of propodeum usually uniformly and densely areaolate-punctate to foveolate-punctate; body setae virtually all black (Figs. 4A–B); if different color setae present, then of a coppery- golden tone and reduced to inconspicuous patches on fringes of T2–4 medially and/or laterally (Fig. 5B) (Cerrado) … T. chrysozona (Gerstaecker, 1874)
6. T2 virtually devoid of sculpture anterolaterally, smooth, shinning and asetose (Figs. 20A, 20C); in addition to dense short black and silvery-white setae, entire body (except mandibles, antennae, and tarsi) clothed with dense conspicuously long brownish setae (Figs. 20A, 20C) … T. ursina (Gerstaecker, 1874) T2 at most with sparser sculpture and setation anterolaterally; body setae usually of equal length throughout, either black or silvery-white, never brownish (e.g. Figs. 1A–C) … 7
7. Post-mesonotal tubercle absent, at most with slight swelling of lateral margin posterior to mesonotal projection; T2 with orange integumental spots (sometimes dulled by time) (Figs. 7A–C) … T. funebris (Gerstaecker, 1874) Post-mesonotal tubercle present, often reduced but still conspicuous; T2 integumental spots yellowish or reddish (e.g. Figs. 3A, 17A) … 8
8. Integumental spots of T2 with scatterd foveolations usually restricted to the edges (Fig. 17A); spots yellowish in color, often linear in shape (Fig. 17D) … T. sancta (Gerstaecker, 1874) 189
Integumental spots of T2 with sparse foveolations throughout; spots reddish, always subquadrate or subrectangular (e.g. Figs. 1A, 3E) … 9
9. Lateral face of propodeum densely areolate-punctate with smooth flat intervals; intervals conspicuously broader along posterior margin of lateral propodeal face; vertex and mesonotum conspicuously clothed with silvery- white to silvery-golden appressed setae (Figs. 3A, 3E) … T. austera (Gerstaecker, 1874) Lateral face of propodeum densely areolate-punctate with sharp homogeneous intervals (Fig. 1C); vertex and mesonotum clothed with black setae only (Fig. 1A) … T. ameliae Casal, 1969
Key to males of the T. quadrinotata species-group.
1. Apex of cuspis with short setae (e.g. Figs. 19B–E) … 2 Apex of cuspis with long setae (e.g. Figs. 13B–E) … 3
2. Pronotum sparsely and finely foveolate-punctate with micropunctate intervals, concealed by dense silvery- white setation (Fig. 9A) (predominantly found in the Amazon) … T. incerta (Spinola 1841) Pronotum densely and coarsely foveolate-punctate without micropunctured intervals, sculpture mostly exposed, setae black (Fig. 19A) (predominantly found in Atlantic Forest areas) … T. tetratrauma Bartholomay & Williams sp. nov.
3. Lateral face of propodeum and metapleuron clothed with dense appressed silvery-golden setae (Fig. 13A) … T. quadrinotata (Klug, 1821) Lateral face of propodeum at most with sparse erect silvery-white setae (e.g. Fig. 2A); metapleuron at most with sparse appressed black or silvery-white setae (e.g. Fig. 11A) … 4
4. Body setae predominantly black with coppery-golden areas (Figs. 5A, 6A) … T. chrysozona (Gerstaecker, 1874) Body setae almost entirely black (e.g. Fig. 10J) or predominantly silvery-white with black areas (e.g. Figs. 15A, 15C) … 5
5. Wings brown, dark-brown on apical third, hyaline-brown elsewhere; pronotum and T2 clothed with extensive silvery-white setae (Figs. 15A, 15C) … T. quadripustulata (Klug, 1821) Wings entirely dark-brown with strong violaceous and blueish reflections; pronotum and T2 mostly clothed with black setae (e.g. Figs. 10B, 10J) … 6
6. Silvery-white setal markings of propodeum and T2 dense, completely obscuring integument beneath (Fig. 2A) … T. ameliae Casal, 1969 Silvery-white setal markings of propodeum and T2 sparse (e.g. Fig. 11J), if apparently dense, then never completely obscuring integument beneath (e.g. Fig. 11B) … 7
7. Cuspis length approximately 0.65 × paramere length (Figs. 11C–E, 11K–M); paracuspis relatively well- developed, almost node-like (Figs. 11F, 11N); posterior margin of penis valve subtruncate (Figs. 11G, 11O); silvery- white setae present throughout propodeal dorsum (Fig. 11B) … T. pompiliformis (Gerstaecker, 1874) Cuspis length approximately 0.85 × paramere length (Figs. 10C–E, 10K–M); paracuspis poorly-developed, lobe-like (Figs. 10F, 10N); posterior margin of penis valve more or less evenly convex (Figs. 10G, 10O); silvery- white setae restricted to posterior half of propodeal dorsum (Fig. 10J) … T. infernalis (Gerstaecker, 1874)
Traumatomutilla quadrinotata species-group
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Diagnosis. FEMALES. lateral margins of the mesonotum projected laterally into blun tubercles; head unarmed on posterior margin of vertex; scutelar scale narrow, usually lacking anterolateral carinae; pygidial plate narrow, side strongly convergent basad, subpyriform. MALES. Mesopleuron always tuberculate on dorsal half; axillar projections always truncate; S2 always lacking setae filled pit; hypopoygium elongate, subrectangular, never defined by lateral carinae; cuspis slender, elongate, mostly asetose; paracuspis poorly developed, lobe-like. Included taxa. Traumatomutilla ameliae Casal, 1969, T. austera (Gerstaecker, 1874), T. chrysozona (Gerstaecker, 1874), T. funebris (Gerstaecker, 1874), T. incerta (Spinola, 1841), T. infernalis (Gerstaecker, 1874), T. pompiliformis (Gerstaecker, 1874), T. quadrinotata (Klug, 1821), T. quadripustulata (Klug, 1821), T. sancta (Gerstaecker, 1874), T. tetratrauma Bartholomay & Williams sp. nov., and T. ursina (Gerstaecker, 1874). Distribution. Widely distributed in South America from Colombia to Argentina. Remarks. The T. quadrinotata species-group is a more or less consistently large-bodied (>10 mm) group in both sexes. The anterolateral carinae in the scutelar area — albeit partially concealed by dense setation — and the dorsolaterally expanded occipital carina observed in T. tetratrauma were not included in the original diagnosis of the T. quadrinotata species-group by Williams et al. (2017). Apart from these two new characters, the remaining characters for the species-group are relatively consistent and non-variable for both sexes, most notably so for males, which apart from the length of certain genitalia setae and sculpture of the pronotal dorsum, can only be distinguished based on color and setae characters. The most conspicuous and reliable character for identifying females is the lateral expansion of the mesonotum which varies slightly from broad and blunt (e.g. T. funebris and T. quadrinotata) to narrow and tuberculiform (e.g. T. ameliae and T. sancta). The post-mesonotal tubercle is a character that was first observed in T. poranga Bartholomay & Williams, 2018 and, though it can be found in four species of the T. quadrinotata species-group, it is most conspicuous in T. quadripustulata.
T. ameliae Casal, 1969 (Fig. 1A–C, 2A–F)
Traumatomutilla ameliae Casal, 1969: 291, holotype, #f, Argentina, Tucumán, S. [San] Pedro de Colalao, I.1949, Arnau (AMNH), examined.
Diagnosis. FEMALE. Occipital carinae equally wide throughout; anterolateral carinae absent on scutelar scale; lateral face of propodeum evenly and densely sculptured with sharp intervals; T2 with two pairs of red integumental spots; body setae silvery-white in color; setae markings of mesosomal dorsum restricted to propodeum. MALE. Apex of cuspis with long setae; body setae silvery-white in color; wings dark brown infuscated throughout with strong violaceous and blueish reflections; setae markings of propodeum and T2 dense, concealing integument beneath. Description. FEMALE. Body length 13-21 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view slightly longer than distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate, less densely so on malar space. Genal carina well-defined. Mandible oblique, tapering slightly towards apex with small subapical tooth. Dorsal scrobal carina present, well-defined, narrowly disconnected from antennal tubercles, connected to lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 1.9 × pedicel length; flagellomere 2 1.4 × pedicel length. Mesosoma. Length 0.85 × width. Mesosomal dorsum mostly concealed by dense setation, densely and coarsely areolate-punctate with apparent sharp to scabrous intervals where visible. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, coarsely striated longitudinally throughout with dense coarse punctures dorsomedially; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, projected dorsally, broadly separated from conspicuously projected angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle strongly projected from lateral margin of pronotum, rounded, bulging. Lateral face of pronotum sparsely punctate with dense micropunctures except at smooth conspicuous subacute tubercle anterior to pronotal spiracle; mesopleuron densely micropunctate anteriorly, densely and coarsely foveolate-punctate to areolate-punctate ventrad on mesopleural ridge; metapleuron unsculptured smooth shinning on dorsal third, densely, coarsely and confusedly areolate on basal third, and concealed by dense setation elsewhere. Lateral face of propodeum densely and coarsely areolate-punctate 191 throughout with sharp intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 70:83:87:60:60. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into sharp process; with very small conspicuous blunt post-mesonotal tubercle. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area vestigial. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; scabrous intervals absent on scutellar area. Propodeum somewhat gibbose, dorsal face shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 28:64:67. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures where visible; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely foveolate-punctate. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 densely and coarsely foveolate- punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold vestigial. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S4 densely foveolate-punctate; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface irregularly rugose; interstice apparently granulose. MALE. Body length 13–18 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes, contiguous with eye outline in dorsal view. Width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 4.4 × DLO, IOD 1.4 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate, with sparse interspersed micropunctures along posterior margin of vertex; sculpture sparser and finer posterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; sculpture concealed by dense setation; apical/ventral margin with a pair of medial short subacute free teeth. Scape bicarinate. Flagellomere 1 1.9 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, subangulately projected from anterior margin of pronotum, separated from well-defined pronounced humeral carina, anterolateral corners of pronotum angulate. Anterior face of pronotum, with sparse fine to dense coarse punctures laterad, mostly unsculptured mediad; with medial longitudinal slightly concave smooth area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate to pucntate. Mesoscutum densely and finely foveolate-punctate, parapsis and notaulus reduced to posterior half of mesoscutum; with medial longitudinal carina on posterior half. Mesoscutellum somewhat gibbose, densely and coarsely areolate-punctate to foveolate-punctate, with well-defined dorsal and posterior faces; dorsal face as broad as long, with intervals aligned basally so as to form irregular longitudinal carina restricted to dorsal face. Axilla produced posterolaterally as obliquely truncate projection, with inner margin straigth; projection coarsely and densely foveolate-punctate except at unsculptured apical third. Metanotum wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, mostly concealed by dense setation, densely areolate where visible; sculpture of lateral face absent along anterior margin; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely and vestigially punctate with sparse interspersed micropunctures; mesopleura with conspicuous acute spine-like projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron micropunctate throughout, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 0.9 × as long as wide. Dorsal metasomal sculpture, except pyigidium, partially concealed by dense setation, densely and finely punctate with dense interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by somewhat arched carinae laterally; surface irregularly granulose to rugose with interspersed coarse punctures apicad. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate; sculpture sparser mediad; anteromedial crest-fold present, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced posteriorly, subacute. Ratios of 192 free length of paramere, cuspis and digitus, 83:58:22. Paramere virtually straight in dorsal view, upcurved posteriorly in lateral view; virtually asetose except for sparse scattered inconspicuous setae throughout. Cuspis narrow, elongate, virtually straight and slightly sinuous throughout in dorsal view, upcurved and slightly wider posterad in lateral view, with conspicuous tuft of long setae at apex, inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, slightly longer than wide, with subacute projection posteroventrally on posterior margin, projection with single setae, otherwise asetose. Digitus slightly incurved in dorsal view and upcurved in lateral view, apex subcapitate in lateral view, with short inconspicuous setae on dorsal surface. Penis valve strongly concave on inner surface, with well-defined pair of acute teeth posteroventrally, without defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along convex posterior margin and inconspicuous setae present at base of anterior tooth on outer margin, setae longer ventrad, posterior margin sloping dorsad. Coloration and variations. FEMALE. Integument black, except mandible and antennal flagellomeres partially reddish-brown, and T2 with four large reddish integumental spots. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: coxae, ventral surface of meso and metafemora, mesopleuron posteroventrally, metapleuron, propodeal dorsum laterally, and lateral face of propodeum; T1 entirely, integumental spots, lateral areas, lateral felt line and lateral margins of T2, fringe of T2– 4 medially and laterally, fringe of T5, and T6, except pygidial plate medially; S1–4, and fringes of S2–3. MALE. Integument black, except antennal flagellomeres partially reddish-brown. Body setae predominantly black varying in density except the following areas with silvery-white setae varying in density: propodeum, legs predominantly; T1, anterior half to anterior third of T2, lateral felt lines and lateral margins of T2, fringe of T2–5 medially and laterally (vestigial medially on T2–5 and laterally on T5); S1–5, except fringe of T5. Distribution. Bolivia, Paraguay, and Argentina. Material examined. (15#f, 5#m) Type material. Holotype, #f, ARGENTINA, Tucuman, S. [San] Pedro de Colalao, I.1949, Arnau (AMNH). Additional material. BOLIVIA, Santa Cruz, 12km N [kilometers north of] Camiri, 19°56.67'S 03°31.34'W, 1#f, 27.II–07.III.1999, M.E. Irwin & F. Parker (EMUS); PARAGUAY: Boqueron, E. [East] of La Patria, 1#f, s30.III.2006, U. Dreschel (FSCA); Chaco: Filadelfia: 2#f, I.1995, Arriagada (AMNH); Filadelfia, 1#f, I.1995, Arriagada (EMUS); Loma Plata, 2#f, I.1956, Gerlach (AMNH); Pres [Presidente]. Hayes, Lolita, Yaragui, 390', 23°06'S 59°38'W, 1#f, 23–27.XI.2007, U. Dreschel (FSCA); ARGENTINA: Salta: La Viña, 1#f, XII.1992, M.A. Fritz (AMNH); 1#f, I.1984, M.A. Fritz (AMNH); 1#f, I.1988, M.A. Fritz (AMNH); 2#m, XII.1992, Fritz (AMNH); Rosario Lerme [Rosario de Lerma]: 1#f, II.1993, M.A. Fritz (AMNH); 2#m, 21–23.XII.1983, M. Wasbauer (UCDC); El Carmen, 1#m, I.1983, Fritz (AMNH); Sumalao, 1#f, III.1991, Fritz (EMUS); Alemania, 1#f, II.1983, M.A. Fritz (AMNH); Santiago del Estero, Chaco de Santiago del Estero, Rio Salado, 1#f (MNCN); Catamarca, Santa Maria, 1#m, I.1994, Fritz (AMNH). Remarks. The sex association of T. ameliae was based on distribution since both sexes were the southernmost records of the T. quadrinotata species-group and have been collected in the same areas across Argentina. Both sexes have consistent and apparently unvariable color and setae characters, typical of northern Argentinean mutillids. Based on structure, especially the lateral propodeal face sculpture and lateral mesonotal expansions, females of T. ameliae are similar to T. sancta.
Traumatomutilla austera (Gerstaecker, 1874) (Figs. 3A–G)
Mutilla austera Gerstaecker, 1874: 70. Holotype female, Brazil, [São Paulo], Salto Grande (ZMB), examined. Mutilla sigillata Gerstaecker, 1874: 70. Holotype, #f, Brazil, [Rio Grande do Sul], Cassapava [Caçapava do Sul] (ZMB), type examined, syn. nov. Ephuta (Traumatomutilla) austera: André, 1902, p. 54. Ephuta (Traumatomutilla) sigillata: André, 1902, p. 56. Traumatomutilla austera: André, 1904, p. 40. Traumatomutilla sigillata: André, 1904, p. 40.
193
Diagnosis. FEMALE. Occipital carina equally wide throughout; anterolateral carinae absent on scutelar scale; body setae silvery-white in color; lateral face of propodeum densely but unevenly sculptured, with conspicuous unsculptured areas; T2 with two pairs of reddish integumental spots; mesonotum with a pair of posterolateral oblique dense setae patches. MALE. Unknown. Description. FEMALE. Body length 14–16mm. Head. Posterior margin virtually straight. Occipital carina evenly wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 1.1 × the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate; sculpture sparser on gena and malar space. Genal carina present. Mandible oblique, tapering slightly towards apex, with small subapical tooth, unarmed ventrally. Dorsal scrobal carina well defined, reaching antennal tubercles and narrowly separated from lateral scrobal carinae. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.2 × pedicel length; flagellomere 2 1.75 × pedicel length. Mesosoma. Mesosoma 0.9 × as long as wide. Mesosomal dorsum densely and coarsely areolate-punctate with irregular intervals. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, coarsely striated longitudinally basad with dense coarse punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, projected dorsally, broadly separated from slightly projected rounded epaulet; anterolateral corners of pronotum angulate in dorsal view. Pronotal spiracle slightly projected from lateral margins of pronotum, rounded. Lateral face of pronotum densely foveolate-punctate with micropunctures, except low blunt tubercle anteroventral in relation to pronotal spiracle; mesopleuron mostly concealed by dense setation, densely micropunctate anteriorly, and densely and coarsely foveolate-punctate to areolate-punctate on mesopleural ridge where visible; metapleuron completely concealed by dense setation, except dorsal fourth unsculptured smooth and shinning. Lateral face of propodeum densely and coarsely areolate-punctate throughout with dull intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 68:76:84:57:60. Lateral margin of mesonotum strongly constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt process; post-mesonotal tubercle present, inconspicuous. Propodeal spiracle strongly projected from side of mesosoma. Post-spiracular area absent. Scutellar scale present, greatly reduced, as narrow as surrounding sculpture; anterolateral carinae absent. Propodeum gibbose in lateral view, dorsal face shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 35:79:78. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed coarse micropunctures where visible; sculpture sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, virtually concealed by dense setation, densely foveolate-punctate where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, equally high throughout. S2 densely and coarsely foveolate-punctate, sculpture conspicuously sparser mediad; anteromedial crest-fold present. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose. Coloration and variations. FEMALE. Integument black, except mandibles partially and antennal flagellomeres ventrally reddish-brown, and T2 with four reddish to orange integumental spots. Body setae predominantly silvery- golden, except the following areas with black setae varying in density: malar space, gena, vertex posterolaterally, ventral surface of head; pronotum, mesonotum anteromedially, mesopleuron anteriorly, scutelar area, propodeal dorsum medially; T1 medially, disc of T2 (except integumental spots), fringe of T2–4 sublaterally, fringe of T5 medially, T6 medially, fringe of S4, and S5–6. Some specimens have the silvery-golden marking of the mesonotum larger, nearly confluent medially; silvery-golden setae greatly reduced on propodeal dorsum, nearly absent; venter of metasoma completely clothed with black to brownish-black setae, T6 completely clothed with black to brownish-black setae; medial marking of silvery-golden setae on T3 greatly reduced, nearly absent. MALE. Unknown. Distribution. Brazil. Material examined. (36#f) Type material. Lectotype of Mutilla austera, #f, BRAZIL, [São Paulo], Salto Grande, Sello S. (ZMB); Holotype of Mutilla sigillata, #f, BRAZIL, [Rio Grande do Sul], Casapava [Caçapava do Sul]. Sello S. (ZMB); Additional material. BRAZIL: 1#f (ZMUC); Goiás: 4#f (MZSP); 1#f, Goyasz, n.o proc. [número de procedência] 18/242 (MNRJ); 1#f, [unintelligible label] (MNHN); Leopoldo Bulhões, 1#f, XII.1939, Spitz (MNCN); 194
Trindade, 1#f, 1952 (MNHN); Goiânia, 1#f, XII.1942, Gonçalvez (AMNH); Distrito Federal: 1#f (MZSP); Brasília, 1#f, Bráulio Dias (AMNH); Minas Gerais: 1#f (MZSP); 2#f, Reinhardt (ZMUC); Campos Gerais, 1#f (MNHN); Sertão de Diamantina, Fazenda das Melancias, 1#f, 10.XI.1902, E. Gounelle (MNCN); Sete Lagoas, 1#f, Reinhardt (ZMUC); Uberaba: 2#f (DEI); 1#f, E. Le Moult (MNCN); Ibiá, 1#f, 1965, Elias,C. (DZUP); Passos, 1#f, 16– 22.I.1963, Elias,C. (DZUP); São Paulo: 2#f (MZSP); 2#f, [unintelligible label], 1914 (MNHN); Jundiaí, 2#f, 15.XII.1960 (MNRJ); Iperó [sic!], George Oeterer, 1#f, 22.XI.1961, F. Grossmann (MNCN); Ipiranga, São Paulo, 1#f, II.1910, H. Luederwaldt (MZSP); Corumbai, 1#f, IX.1963 (DZUP); Paraná: 1#f (MZSP); Carambehy [Carambeí], 1#f, 27.VI.1937, Westerman (MZSP); Ponta Grossa: 1#f, III.1946, Justus (DZUP); Parque Estadual da Vila Velha, 1#f, 3.III.2012, Amaral-Neto, L.P. (DZUP). Remarks. Traumatomutilla austera has a distinct color pattern (Figs. 3A–B, 3E–F) that varies slightly in the mesosomal dorsum but has no intermediate forms with other known patterns. Certain specimens have patches of somewhat dense appressed silvery-white setae on the lateral face of the proprodeum that are also found in T. funebris and, more notably, in the Atlantic Forest species (T. quadrinotata and T. tetratrauma). The structure of T. austera however, is different from these species, being finer and homogeneous as opposed to the overall coarser sculpture of most species within the T. quadrinotata species-group. Based on distribution, T. austera is one of four possible candidates for association with T. infernalis or T. pompiliformis.
Traumatomutilla chrysozona (Gerstaecker, 1874) (Figs. 4A–B, 5A–H, 6A–G)
Mutilla lugubris Burmeister, 1854. (nec Fabricius; #f nec #m): holotype, #f, Brazil, [Minas Gerais], Queluz [Conselheiro Lafayate] or Ouro Preto [preoccupied name, renamed by Dalla Torre (1897)] (location unknown). Mutilla chrysozona Gerstaecker, 1874: 315. Holotype, #m, Brazil, São Paulo, Sello. S. (ZMB), examined. Mutilla burmeisteri Gerstaecker, 1874: 316, holotype, #m, Brazil, [Minas Gerais], Queluz [Conselheiro Lafayate] or Ouro Preto (location unknown). syn. nov. Mutilla lugubrina Dalla Torre, 1897: 55 (new name for Mutilla lugubris) syn. nov. Ephuta (Traumatomutilla) burmeisteri: André, 1902, 54. Ephuta (Traumatomutilla) chrysogona: André, 1902, 54 (misspelling). Ephuta (Traumatomutilla) lugubris: André, 1902, 55 (misspelling). Traumatomutilla burmeisteri: André, 1904, 40. Traumatomutilla chrysozona: André, 1904, 40. Traumatomutilla lugubrina: André, 1904, 40. Ephuta (Traumatomutilla) dives André, 1906: 66, lectotype [designated here], #m, Brazil, Piauhy [Piauí] (BMNH), type examined, syn. nov. Traumatomutilla dives: Nonveiller, 1990: 77.
Diagnosis. FEMALE. Occipital carina equally wide throughout; anterolateral carinae absent on scutelar scale; body setae almost entirely black, at most with vestigial coppery setae in some metasomal fringes; T2 with two pairs of dark red integumental spots. MALE. Apex of cuspis with long setae; body setae black with conspicuous coppery areas. Description. FEMALE. Body length 15 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and slightly swollen laterally. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate- punctate to areolate-punctate, less densely so on malar space. Genal carina well-defined. Mandible oblique, tapering slightly towards apex with small subapical tooth. Dorsal scrobal carina present, well-defined, narrowly disconnected from antennal tubercles. Lateral scrobal carina virtually absent. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.6 × pedicel length; flagellomere 2 1.9 × pedicel length. Mesosoma. Length 0.8 × width. Mesosomal dorsum mostly concealed by dense setation, densely and coarsely areolate-punctate with apparent sharp to scabrous intervals where visible. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, coarsely striated longitudinally throughout with dense coarse punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, slightly projected dorsally, broadly separated from conspicuously projected 195 angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded, bulging. Lateral face of pronotum sparsely punctate with dense micropunctures except at smooth conspicuous subacute tubercle anterior to pronotal spiracle, distance between tubercles wider than distance between pronotal spiracles; mesopleuron mostly concealed by dense setation, densely micropunctate anteriorly, and densely and coarsely foveolate-punctate to areolate-punctate on mesopleural ridge where visible; metapleuron completely concealed by dense setation, except small posterior area on dorsal fourth unsculptured smooth and shinning. Lateral face of propodeum mostly concealed by dense setation, densely and coarsely areolate-punctate throughout with interspersed micropunctures where visible; intervals dull and blunt where visible. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 70:83:87:60:60. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt process; post-mesonotal tubercle absent. Propodeal spiracle strongly projected from lateral margin of mesosoma; post- spiracular area indistinguishable. Scutellar scale present, as wide or wider than surrounding sculputre; anterolateral carinae present, approximately twice as wide as scutelar scale; scabrous intervals vestigial on scutellar area. Propodeum simply convex, dorsal face indistinguishable from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 28:64:67. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate- punctate to punctate with dense interspersed coarse micropunctures where visible; sculpture sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, virtually concealed by dense setation, densely foveolate-punctate where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 densely and coarsely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold present. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate- punctate with sparse micropunctures where visible; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose. MALE. Body length 14–16 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes, but not contiguous with eye outline in dorsal view. Width 0.85 × pronotal width. Eye almost circular. Ocelli small; OOD 4.3 × DLO, IOD 1.0 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate; sculpture sparser and finer posterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely foveolate-punctate medially and along apical/ventral margin laterally; apical/ventral margin with a pair of medial short subacute free teeth medially. Scape bicarinate. Flagellomere 1 2.1 × pedicel length; flagellomere 2 2.9 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, sharply projected from anterior margin of pronotum, separated from well-defined humeral carina, anterolateral corners of pronotum not angulate. Anterior face of pronotum, with sparse fine punctures laterad with interspersed micropunctures, mostly unsculptured mediad; with medial longitudinal slightly concave smooth area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate to areolate-punctate with somewhat sharp intervals. Mesoscutum densely and finely foveolate-punctate, parapsis reduced to posterior half of mesoscutum, notaulus absent. Scutellum sloping throughout, somewhat depressed medially, without defined dorsal and posterior faces, densely and coarsely areolate-punctate to foveolate-punctate; anterior intervals somewhat aligned so as to form vestigial irregular longitudinal carina medially. Axilla produced posterolaterally as obliquely truncate projection, with inner margin slightly curved inward apicad in dorsal view; projection coarsely foveolate-punctate basad, unsculptured, smooth, shinning apicad. Metanotum slightly wider laterad, its surface obscured by dense setation. Propodeal dorsum convex, mostly concealed, densely areolate; sculpture of lateral face absent along most of anterior margin; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely and vestigially punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron partially concealed by dense setation, micropunctate where visible, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. 196
Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 0.9 × as long as wide. Dorsal metasomal sculpture, except pygidial plate, partially concealed by dense setation, densely and finely punctate with sparse interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by parallel carinae laterally; surface densely micropunctate throughout. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate; sculpture sparser mediad; anteromedial crest-fold present, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 96:83:22. Paramere virtually straight with slightly outcurved apex in dorsal view, upcurved posteriorly in lateral view; virtually asetose except for sparse scattered inconspicuous setae throughout, setae more evident in lateral view. Cuspis narrow, elongate, virtually straight throughout and slightly wider posterad in dorsal view, virtually stright and slightly wider posterad in lateral view, with conspicuous tuft of long setae at apex, inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, broader than long, posterior margin simply convex, with inconspicuous sparse setae. Digitus slightly incurved in dorsal view and upcurved in lateral view, soncpisuously narrower posterad in lateral view, apex subcapitate, with short inconspicuous setae on dorsal surface. Penis valve strongly concave on inner surface, with well-defined pair of teeth posteroventrally, posteriormost tooth acute, anterior tooth subacute to blunt, with poorly- defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along truncate posterior margin and inconspicuous setae present at base of anterior tooth on outer margin, posterior margin setae longer ventrad. Coloration and variations. FEMALE. Integument black, except mandibles and antennal flagerllomeres partially reddish-brown, and T2 with four large dark-orange to dark-red integumental spots. Body setae virtually entirely black varying in density, except the following areas with vestigial traces of coppery-golden to silvery-golden setae: lateral margin of T2, fringe of T2 medially and laterally, fringe of T3 laterally, fringe of S3 laterally, fringe of T4 medially and laterally. MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except the following areas with coppery-golden setae: posterior half of T1, anterior margin of T2, fringe of T2–4, fringe of S2–3, and fringe of S4 laterally. Certain specimens may have coppery-golden setae on posterior half of scutellum and covering dorsal face of propodeum, T1, and anterior third of T2 entirely. Distribution. Brazil Material examined. (36#f, 19#m) Type material. Lectotype of Mutilla chrysozona, #m, BRAZIL, S. Paul [São Paulo], Sello S. (ZMB); Lectotype of Mutilla dives, #m, BRAZIL, Piauhy [Piauí] (HBMNH). Additional material. BRAZIL: 2#f (DEI); Alto da Serra [sic!], 1#f, II.1929, F. Spitz (MZSP); Piauí, 1#f, Mocszary (MNHN); Minas Gerais: 915m [above sea level], Passa Quatro, 2#m, 27.II.1922, J. Zikán (MZSP); Poços de Caldas: 1#m, XI.1961, Elias, C. (DZUP); 1#f, 29.XII.1969 (MNRJ); Campo do Saco, 1#f, 12.XII.1967 (MNRJ); Seminario, 1#f, 10.XII.1967 (MNRJ); Mo S [Serra de Santo] Domingos, 1#f, 18.XII.1967 (MNRJ); Serra do Caraça, 2#m (MZSP); 1880m [above sea leve], 1#m (MZSP); 2#m, 27.XI–05.XII.1972, Exp. Mus. Zool. [Expedição Museu de Zoologia] (MZSP); 1880m, 1#m, XI.1961, Kloss, Lenko, Martins & Silva (MZSP); 1#m, I.1970, F.M. Oliviera (AEIC); Barbacena,1#f, 15.XII.1905, Ducke (MNHN); 6 km NE [kilometeres northeast] de Careacu, 1#m, 10.XII.2012, Melo, G.A.R; Grossi, P. (DZUP); São Paulo: 18#f (MZSP); 1# (DEI); 1#f, 01.XI.1920 (DEI); San Pablo [sic!], 1#f (AMNH); Ipiranga: 1#m (MZSP); 1#m, XI.1919 (MZSP); 1#m, 30.III.1936 (CASC); 1#f, 31.V.1950, R. Spitz (MNCN); 1#f, I.1907, S.M. Torres (MZSP); Santo Amaro: 1#m I.1949, J. Lane (MZSP); 1#f, VI.1932, Lane (MNCN); 1#m (MZSP); Jundiaí: 1#m, 12.XII.1899, Schrottky (MNHN); 1#m, 15.XII.1899, Schrottky (MNHN); Barueri, 1#f, 28.XII.1965, K. Lenko (MZSP); Cotia, 1#f (DEI); Capital [São Paulo], 1#f, 20.V.1956, L. Tzarasios & P. Nogueira (CASC). Remarks. The association and synonymy of T. chrysozona with T. lugubrina was initially confusing, as the females (T. lugubrina) usually have exclusively black setae, while the males are extensively covered with brilliant coppery setae. Many females in São Paulo, however, have traces of coppery setae that are the tint as seen in males from São Paulo, and differ from the setal markings of other members of the T. quadrinotata species-group. One additional male, T. dives from Piaui and Minas Gerais, is identical to the type of T. chrysozona in every aspect except it has the propodeal dorsum clothed with dense coppery setae. Though we haven’t seen any intermediate forms between T. chrysozona and T. dives, we consider T. dives to be conspecific with T. chrysozona, as their external morphology and genitalia are indistinguishable. Further, although females of T. lugubrina are known from Minas 197
Gerais, males of the typical form are apparently restricted to São Paulo. We therefore hypothesize that T. dives is a northern color variant of T. chrysozona. Burmeister (1854) described both sexes of Mutilla lugubris based on at least two specimens from Queluz (present day Conselheiro Lafayte) and Ouro Preto in Minas Gerais. Gerstaecker (1874) stated that he was unable to locate the male specimen of M. lugubris, considered both sexes as distinct species, and described the male as a new species, M. burmeisteri, without actually seeing the specimen. The original description of the male of M. lugubris in Burmeister (1854) mentioned that the male has a narrow reddish band on the abdomen (meaning metasoma), and Gerstaecker (1874) placed M. burmeisteri in a subcategory that included only males with black spurs and reddish markings on the metasoma, probably based on Burmeister’s original description of the reddish band. The black spurs are evident in T. chrysozona and it’s very likely that the narrow reddish band mentioned by Burmeister (1854) is in fact his interpretation on the coppery setae clothing the fringe of T2–3, especially since he used the same word “rubra” to define the coppery metasomal marking of Hoplocrates cephalotes (Swederus, 1787). Based on this, we place T. burmeisteri as a junior synonym of T. chrysozona.
Traumatomutilla funebris Gerstaecker, 1874 (Figs. 7A–C)
Mutilla funebris Gerstaecker, 1874: p. 75, holotype, #f, Brazil, Minas Gerais, v. Langsdorf S. (ZMB), examined. Ephuta (Traumatomutilla) funebris: André, 1902: p. 55. Traumatomutilla funebris: André, 1904: p. 40.
Diagnosis. FEMALE. Occipital carinae equally wide throughout; anterolateral carinae lacking on scutelar area; post mesonotal tubercle absent; lateral face of propodeum densely but not evenly sculptured, with conspicuously wide intervals; body setae patterns silvery-white in color, restricted to propodeum on mesosomal dorsum. MALE. Unknown. Description. FEMALE. Body length 15–17 mm. Head. Posterior margin straight, with occipital equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view virtually equal to the distance from its ventral margin to mandibular condyle. Front, vertex, and gena densely and coarsely foveolate- punctate with irregular intervals; sculpture sparser on malar space. Genal carina present. Mandible oblique, tapering slightly towards apex with small subapical tooth, unarmed ventrally. Dorsal scrobal carinae well defined, narrowly separated from antennal tubercles, connected to well-defined lateral scrobal carina. Antennal tubercle coarsely punctate to irregularly rugose. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Mesosomal length 1.5 × width; pronotum 0.85 × as wide as mesothorax. Mesosomal dorsum densely and coarsely areolate-reticulate with sharp irregular intervals. Humeral carina well developed, slightly projected apically, separated from projected subangulate epaulet; anterolateral corners of pronotum angulate in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Anterior face of pronotum coarsely punctate dorsad to vestigially, coarsely and longitudinally striate ventrad. Lateral face of pronotum sparsely and coarsely punctured, with dense micropunctures and conspicuous blunt tubercle anteroventral in relation to pronotal spiracle. Mesopleuron partially concealed by dense setation, micropunctate anteriorly and densely coarsely foveolate-punctate along mesopleural margin where visible. Metapleuron partially concealed by dense setation, except at dorsal fourth unsculptured smooth. Lateral face of propodeum partially concealed by dense setation, densely and coarsely areolate-punctate, with vestigially rugose intervals. Post-spiracular area absent. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 66:76:86:58:56. Lateral margin of mesonotum strongly constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt tooth-like process; with small inconspicuous tubercle posterior to lateral process. Propodeal spiracle strongly projected from lateral margin of propodeum; post-spiracular area absent. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; intervals irregular on scutelar area, not scabrous. Propodeum gibbose, dorsal face much shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 33:75:78. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 predominantly concealed by dense setation, densely and coarsely foveolate-punctate to 198 simply punctate with interspersed micropunctures where visible; T6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate. S1 sparsely, coarsely and confusedly foveolate- punctate, surface cuneiform, ending in short blunt longitudinal carina, equally high throughout. S2 densely and coarsely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold present, reduced. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose. Coloration and variations. FEMALE. Integument black except for mandibles and antennal flagellomeres partially reddish-brow, and T2 with two pairs of subovate (anterior pair) and subquadrate (posterior pair) orange integumental spots; older or poorly conserved specimens may have the spots with darker more reddish tones. MALE. Unknown. Distribution. Brazil. Material examined. (4#f) Type material. Holotype, #f, BRAZIL, Minas Gerais, v. Langsdorf S. (ZMB). Additional material. BRAZIL: 2#f (MNHN); Minas Gerais: Serra do Caraça, 1380m [above sea leve], 1#f, XI.1961, Kloss, Lenko, Martins & Silva (MNCN); Serra do Cipó, Monte Ribeiro, 1#f, 06.II.1939, Lopes & Tupinambá (MNCN). Remarks. All specimens examined are from higher altitude areas (1300–1700 m a.s.l.) transitioning between Cerrado and Atlantic Forest in Minas Gerais state, Brazil. Although the holotype has no indication of the specific location in which it was collected, it is also from Minas Gerais. This high-altitude distribution could be purely coincidental, but it is worth pointing out that Minas Gerais is one of the most relatively well sampled areas in Brazil for velvet ants and no specimens of T. funebris were recorded in lower altitude areas so far. The large orange spots on T2 (darkened by age on the holotype) and indistinct appressed silvery-golden setae on the lateral propodeal face of T. funebris are reminiscent of the pattern observed in T. quadrinotata. Both species are nonetheless distinct structurally, especially on the sculpture of the lateral propodeal face and shape of the lateral mesonotal projections. Further collecting in higher altitude areas of Minas Gerais may reveal the male of this rare species.
Traumatomutilla incerta (Spinola, 1841) (Figs. 8A–H, 9A–G)
Mutilla incerta Spinola, 1841: p. 93, lectotype [designated by Mickel (1937), p. 196], #f, [French Guiana], Cayenne, D. Lebas (MRSN). Mutilla dentata Smith, 1879: 219, holotype [by monotypy], #m, Brazil, [Amazonas], St. Paulo [São Paulo de Olivença] (BMNH), examined, syn. nov. Mutilla sodalicia Kohl, 1882: p. 490, holotype [by monotypy], #f, Brazil, Amaz. [sic] (BMNHW), type examined, syn. nov. Ephuta (Traumatomutilla) sodalicia: André, 1902, p. 56. Traumatomutilla sodalicia: André, 1902: 40. Mutilla incerta: André, 1902: 73 (incertae sedis). Traumatomutilla incerta: Mickel, 1937, p. 196. Traumatomutilla weyrauchi Mickel, 1945: p. 30, holotype, #f, Peru, Oxapampa, 1800m [above sea level] (UMSP), [synonymized by Quintero & Cambra (1996)]. Traumatomutilla dignitosa: Mickel, 1952, p. 131, holotype, m#, Guiana, Bartica, Bartica District, Wm. Beebe (UMSP), type examined, syn. nov. Traumatomutilla tabatinga Casal, 1969: p. 287, holotype, #f, Brazil, Amazonas, Tabatinga, XI.1958, F.M. Oliveira (AMNH), type examined, syn. nov.
Diagnosis. FEMALE. Occipital carina slightly but conspicuously swollen dorsolaterally; anterolateral carinae present on scutelar scale, partially obscured by dense setation; lateral face of propodeum predominantly smooth and shinning, sculpture sparse; mesonotum with transverse band of silvery-white setae on posterior half; T2 with a pair of reddish to orange integumental spots. MALE. Apex of cuspis with short setae; pronotum densely clothed with appressed silvery- white; pronotal sculpture with dense micropunctures. 199
Description. FEMALE. Body length 13–20 mm. Head. Posterior margin virtually straight. Occipital slightly but conspicuously swollen dorsolaterally. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Head sculpture partially concealed by dense setation, densely and finely foveolate-punctate with sharp intervals where visible. Genal carina well-defined. Mandible oblique, tapering slightly towards apex with small subapical tooth. Dorsal scrobal carina present, well- defined, narrowly separated from antennal tubercles and lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.6 × pedicel length; flagellomere 2 1.9 × pedicel length. Mesosoma. Dorsal thoracic length 0.75 × maximum dorsal thoracic width. Mesosomal dorsum partially concealed by dense setation, densely and finely foveolate-punctate with sharp intervals where visible. Anterior face of propodeum defined, short, virtually as long as pronotal collar, finely striated longitudinally throughout with sparse interspersed fine punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, strongly projected dorsally, narrowly connected to conspicuously projected angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum. Lateral face of pronotum with sparse scattered punctures and dense micropunctures except at smooth conspicuous acute tubercle anteroventral in relation to pronotal spiracle; mesopleuron partially concealed by dense setation, densely micropunctate anteriorly, densely and coarsely foveolate-punctate on mesopleural ridge where visible; metapleuron completely concealed by dense setation, except small dorsal fourth unsculptured smooth and shinning. Lateral face of propodeum predominantly unsculptured smooth and shinning, with sparse scattered punctures. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 57:84:74:57:58. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt process; post-mesonotal tubercle absent. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area indistinguishable. Scutellar scale present, as wide or wider than surrounding sculputre; anterolateral carinae present, vestigial, approximately twice as wide as scutelar scale; scutellar area with irregular intervasl, not scabrous. Propodeum convex in lateral view, dorsal face indistinguishable from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 34:73:66. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed coarse micropunctures where visible; sculpture sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; micropucntures sparser on T4 and absent on T5; T6, except pygidial plate, virtually concealed by dense setation, densely foveolate-punctate where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 densely and coarsely foveolate-punctate, sculpture conspicuously sparser and smaller anteromediad; anteromedial crest-fold vestigial. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate- punctate with sparse micropunctures where visible; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface densely and irregularly rugose; interstice apparently granulose. MALE. Body length 14–18 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes, but not contiguous with eye outline in dorsal view. Vertex width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 3.6 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Head surface sparsely and finely punctate; sculpture dense and coarser on frons. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely foveolate-punctate medially and along apical/ventral margin laterally; apical/ventral margin with a pair of medial short subacute teeth medially. Scape bicarinate. Flagellomere 1 1.7 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than medial tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, slightly projected from anterior margin of pronotum, separated from well-defined humeral carina, anterolateral corners of pronotum subrounded. Anterior face of pronotum, sparsely and finely punctate with sparse micropucntures, except medioventrally smooth and shinning; evenly flat troughout. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum predominantly concealed by dense setation, densely and coarsely foveolate-punctate with interspersed micropuncutres where visible. Mesoscutum densely and finely foveolate-punctate, parapsis and notaulus present, reduced to posterior half of mesoscutum. Scutellum subquadrate to subglobose, with well-defined dorsal and posterior 200 faces, elevated medially into longitudinal crest extending from anterior margin of dorsal fface to posterior face, densely and coarsely areolate-punctate to foveolate-punctate. Axilla strongly produced transversely truncate projection, with inner margin slightly curved inward apicad in dorsal view; projection coarsely foveolate-punctate basad, unsculptured, smooth, shinning apicad. Metanotum slightly wider laterad, its surface obscured by dense setation. Propodeal dorsum convex, sculpture of dorsal face concealed by dense setation, posterior face densely areolate; lateral face densely areolate along posterior margin and dorsal third, virtually unsculptured smooth and shinning elsewhere. Lateral face of pronotum sparsely and vestigially punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron partially mostly unsculptured smooth and shinning, except for vestigial rugosities on dorsal fifth and coarse areolateion on ventral fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 0.8 × as long as wide. Dorsal metasomal sculpture, except pygidial plate, partially concealed by dense setation, densely and finely punctate with sparse interspersed micropunctures where visible; sculpture sparser and less defined in apical segments, denser posteromediad onT2; pygidial plate slightly longer than broad, weakly defined by parallel carinae apicolaterally; surface vestigially rugose, appearing smooth and sinning, except apical margin densely punctate and setose. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate; anteromedial crest-fold and sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 63:51:19. Paramere slightly sinuous and posteriorly outcurved in dorsal view, upcurved apically in lateral view, with inconspicuous sparse setae throughout, setae more evident on ventral surface. Cuspis elongate, slender, virtually straight and conspicuously tapered posterad in dorsal view, slightly upcurved apically and equally wide throughout in lateral view, with short sparse inconspicuous setae apically, virtually astose elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with flat dorsally sloping posterior margin, with sparse setae on posterior margin, setae as long as or longer than paracuspis length. Digitus slightly incurved in dorsal view and upcurved in lateral view, conspicuously narrower posterad in lateral view, with short inconspicuous setae on dorsal surface. Penis valve strongly concave on inner surface, with well-defined pair of acute teeth posteroventrally, with poorly-defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along truncate posterior margin and inconspicuous setae present at base of anterior tooth on outer margin, setae on posterior margin longer ventrad, posterior margin sloping dorsad. Coloration and variations. FEMALE. Integument black, except mandibles and antennal flagellomeres partially reddish-brown, and T2 with two orange to reddish integumental spots. Spots vary in size and slightly in shape, from small (separated by half spot width) to large (virtually confluent medially), and from almost perfectly circular to subcircular. Body setae predominantly silvery-white varying in density except the following areas usually with black setae varyin in density: ventral half of frons, gena, malar space, pronotum, anterior half of mesonotum, propodeum, T1 medially, disc of T2 (except over integumental spots), fringe of T2–3 medially, fringe of T4–5 sublaterally, T6 laterally, fringe of S5, and S6. Head setae varying from vestigial medial spot of silvery-white setae to entire head clothed with silvery-white setae. Some specimens may have the fringe of T3 entirely clothed with silvery-white setae and/or vestigial and scattered silvery-white setae on lateral margins of propodeal dorsum. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density except for the following areas with black setae varying in density: ventral hald of frons (interspersed with silvery-white), mesoscutum, axillar projection, dorsal face of scutellum, disc of T2 (except anterior third), fringe of T2–3 medially, T4–7 (except pygidial plate and vestigial silvery-white setae laterally on T4–6), fringe of S5, and S6–7. Some specimens have the silvery-white setae areas conspicuously denser and usually the fringe of T2–3 entirely clothed with dense silvery-white setae. Wings dark brown with strong vioalceous and blueish reflections. Tibial spurs yellowish-white. Distribution. Colombia, Venezuela, Surinam, Guiana, French Guiana, Brazil, Ecuador, Peru, and Bolivia. Material examined. (63#f 35#m) Type material. Lectotype of Mutilla incerta, #f, [FRENCH GUYANA], Cayenne, D. Lebas (MRSN); holotype of Mutilla dentate, #m, BRAZIL, [Amazonas], St. Paul [São Paulo de Olivença] (BMNH); holotype of Mutilla sodalicia, #f, Brazil, Amaz. [sic] (BMNHW); holotype of Traumatomutilla dignitosa, #m, GUIANA, Bartica, Bartica District, Wm. Beebe (UMSP); holotype of Traumatomutilla tabatinga, #f, BRAZIL, 201
Amazonas, Tabatinga, XI.1958, F.M. Oliveira (AMNH). Additional material. COLOMBIA: 2#m (IAvH); Rio Negro [sic!], Ost Columb. [sic!], 1#f (MNCN); Putomayo: Mocoa, 1#m, 04.VII.1978, M. Cooper (BMNH); Santa Rosa, Kofan Indian village, 1#m, 10.X.1970, B. Malkin (FMNH); PNN [Parque Nacional Natural] La Paya, Resguardo Cecílio Cocha, 00°11'S 74°55'W, 200m [above sea level, 1#m, 20–24.I.2003, C. Sarmiento (IaVH); Caquetá: PNN [Parque Nacional Natural] Chiribiquete: Rio Cuñare, 0°30'N 72°37'W, 300m [above sea leve],1#m, 01–05.xi.2000, E. González y M. Ospina (FSCA); Rio Cuñare-Amu, 0°13'N 72°25'W, 300m [above sea level], 1#m, 28.II–03.III.2001, M. Ospina & E. González (EMUS); Meta, PNN [Parque Nacional Natural] Tinigua, Vda. [sic!] Bajo Raudal, 02°16'N 73°48'W, 460m [above sea level], 1#m, 01–15.III.2003, C. Sánchez (IaVH); VENEZUELA, Amazonas, Alto Rio Sialba, 510m [above sea level], 01°43'10''N 64°31'50''W, 1#f, 24–25.II.1989, T. Lattke (FSCA); GUYANA: Kartabo, 1#f, VII–VIII.1920, W.M. Wheeler (MCZ); jct. [juncture] Cuyani/Mazaruni, Essequibo Rivers, 1#f, 1927, R.E. Fuglestad (CISC); Bartica, 1#m, 1.VI.1924 (AMNH); FRENCH GUIANA: Cayenne, 1#f (ZMUC); Chavrein, 1#f, 1914, R. Benoist (MNHN); Gourdonville, 1#f, 1914, R. Benoist (MNHN); Maroni, St. [Saint] Laurent du Maroni, 1#f, E. Le Moult (MNCN); SURINAME, Brokopondo, Brownsberg Nature Park, Irene Val Trail, 04°95'N 55°18'W, 400m [above sea level], 1#m, 25.VIII–02.IX.2007, G. Kung, A. Kreuter (EMUS); BRAZIL: Pará: 3#f (MZSP); Óbidos: 1#f, XII.1965, Maller,A (DZUP); 1#f, III.1958 (USNM); 1#f (USNM); 1#f, VII.1959 (MNRJ); Santarém: 2#f (RBINS); 1#m (MNHN); 1#f, 1923 (DEI); Altér do Chão, 1#m, 12–13.IX.1991, J. Vidal (INPA); Oriximina, 1#f, 02.IX.1919, H.S. Parish (FMNH); Jacareacanga, 2#m, XII.1968, M. Alvarenga (AEIC); Tucurui, 8#m, I.1979, M. Alvarenga (AEIC); Amapá: Oiapoque, (Retiro) Ambiente de mata, 1#f, 23.X.2001, J. Madson (IEPA); Serra Lombard, Limão, 1#f, 26.VIII.1961, J. & B. Bechyné (MPEG); Amazonas: Manaus: 4#f (RBINS); 1#f, VII.1962, Oliveira, F.M. (DZUP); Bairro Flores, 1#f, 13–20.IV.2009, D. Storck-Tonon (INPA); INPA 1 [Campus 1], Bosque da Ciência, 03°05'50''S 59°59'16''W, 1#f, 10.VI.2017, M.A. Bento (INPA); Reserva Ducke: Trilha principal, 1#f, 03.X.2016, G. Amora (INPA); 2.93308°S 5897116°W, 1#f, 19.V.2012, Medeiros (INPA); 02°55'48''S 59°58'31''W, 1#f, 08.VI.2013, Saraiva, J.F. (INPA); Teffe [Tefé]: 1#f, 05.II.1920, H.S. Parish (FMNH); 1#m, 09.ix.1909,Ducke (MNHN); Tonantins, Amazon River, 2#f, VII.1923, S.M. Klages (CMNH); 2#f, VIII.1923, S.M. Klages (CMNH); Manacapuru, Amazon River, 1#f, IX.1923, S.M. Klages (CMNH); Rio Javari, Estinao [Estirão do] Equador, M. Alvarenga (AMNH); Rondônia: 62km SE [kilometers southeast of] Ariquemes: 1#f, 17–24.III.1989, W.J. Hanson (EMUS); 1#f, 05–16.XI.1996, W.J. Hanson (EMUS); Ouro Preto d'Oeste, 1#m, 20.IX.1987, Elias, C. (DZUP); Mato Grosso, Sinop, 12°31'S 55°37'W, 1#m, X.1975, F.M. Oliviera (EMUS); ECUADOR: 2#f (MNCN); Napo, Tiputini Biodiversity Station, 1#f, 20.VII.1999, M.J. Miller (UAIC); Archidona, 1#f, R. Haensch (MNCN); Misahualli, nr. [near] Tena: 1#f, 10.I.1989, B.J. Nichols (UMRM); 1#f, 13–20.VI.1998, C. & K. Messenger (UNSM); 1#f, 06–19.X.2001, C. Brammer (EMUS); Rio Aguarico, 1#f, III.1992, E.S. Ross (CASC); 21–25km E [east of] Atahualpa, 1#f, 27–31.IX.1997, F.T. Hovore (UCDC); 25km E [east of] Atahualpa, 01°00'S 78°00'W, 01–06.X.1997, F.T. Hovore (EMUS); Sucumbios River, Sacha Lodge, 1#m, 3–23.VI.1994 (EMUS); Sucumbios: Rio Napo: Sacha Lodge: 220–230m [above sea level], 00°30'S 76°30'W, 1#m, 27.VIII–10.IX.1994, P. Hibbs (EMUS); 1#m, 13.VII.1994, P. Hibbs (EMUS); 270m [above sea leve], 0°30'S, 75°30'W, 1#m, 13–25.II.1994, P. Hibbs (EMUS); PERU: Santo Domingo [sic!] 1# (BMNH); [Madre de Diós], Pakitza, 1#f, 23–28.II.1992, B.V. Brown (USNM); Loreto, Amazon Camp, Rio Momon, nr [near] Iquitos, 1#f, 01–10.XII.1982, E.S. Ross (CASC); Junin, Satipo, 1#f, 18.VII.1939, W.F. Walsh, Jr. (MCZ); Lima: Valle Chanchamayo: 1#f, 1939, P. Vaquero (UMSP); 2#f, 1914 (MNHN); 2#f, Thamm S. (ZMB); 1#m, 21.VII.1949, J.M. Schunke (BMNH); Huánuco, Tingo Maria, 1#f, 03.I.1970, J.C. Schuster (FSCA); 2#m, 23.X.1946, Pallister et al. (AMNH); vic. [vicinity] Tingo Maria, 1#f, 01–05.VI.1999, W.Hanson S.Keller (EMUS); Monson Valley, 2#m, 16.XI.1954 (EMUS); BOLIVIA: Region Chapare, 400m [above sea level], 1#f, 20.III.1907, Zischka (MNCN); Tumupasu, 1#f, 1921, W.M. Mann (USNM); Yungas de Coroico, 2#f, 1914 (MNHN). Remarks. The sex association between T. incerta and T. dentata is evidenced by the distribution of both species which, to the best of our knowledge, are the sole representatives of the T. quadrinotata species-group in the northwestern Amazon Forest. Even though we were unable to access to lectotype of T. incerta, numerous specimens examined were identified as this species and compared with the type by Mickel which designated the lectotype (Mickel, 1937). The lectotype designation was later confirmed by Pagliano (2005), who also officially designated the paralectotypes for T. incerta and provided a photograph of said type. Close examination of T. dentata and T. dignitosa revealed that the difference between the axillar projections (Mickel 1952) is not consistent; this, coupled with their virtually identical genitalia characters, and only marginally different setal characters, is sufficient basis for synonymizing these males. Casal (1969) stated that T. tabatinga differed from T. weyrauchi (then a valid species) by 202 having transverse irregular rugosities on the pygidial plate, which was the main difference between T. incerta and T. weyrauchi mentioned by Mickel (1945). Cambra & Quintero (1996) studied a paratype of T. weyrauchi and concluded that it could not be distinguished from T. incerta. Therefore, the pygidial sculpture was shown to be inadequate for differentiating species. After examining the types of T. tabatinga and T. sodalicia, we conclude that these species are also synonymous with T. incerta and T. weyrauchi. They are all basically identical in structure, differing only in the size of the T2 integumental spots and the presence or absence of silvery-white setae on the head, with numerous intermediate specimens found between each form.
Traumatomutilla infernalis (Gerstaecker, 1874) (Figs. 10A–P)
Mutilla infernalis Gerstaecker, 1874: 318, lectotype [designated here], #m, “S. Brasilien” [Southern Brazil], Sello S. (ZMB), type examined. Mutilla floccosa Gerstaecker, 1874: 314, holotype (by monotypy), #male, Brazil, Sello S. (ZMB), type examined, syn. nov. Ephuta (Traumatomutilla) infernalis: André, 1902, 55. Ephuta (Traumatomutilla) floccosa: André, 1902, 55. Traumatomutilla infernalis: André, 1904, p. 40. Traumatomutilla floccosa: André, 1904, p. 40.
Diagnosis. FEMALE. Unknown. MALE. Cuspis with long setae apically; cuspis length 0.85 × length of paramere; posterior margin of penis valve evenly convex; lateral face of propodeum with sparse erect silvery-white seta; metapleuron setae black; wings strongly dark brown infuscated throughout, with strong violaceous and blueish reflections; propodeum and metasoma silvery-white setae sparse, never obscuring integument completely. Description. FEMALE. Unknown. MALE. Body length 12 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes. Vertex width 0.86 × pronotal width. Eye almost circular. Ocelli small; OOD 4.3 × DLO, IOD 1.0 × DLO. Occipital carina distinct. Head surface punctate; sculpture sparser and finer posterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely foveolate-punctate, coarser mediad; apical/ventral margin with a pair of medial short subacute free teeth medially. Scape bicarinate. Flagellomere 1 2.1 × pedicel length; flagellomere 2 2.6 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets moderate, connected towell-defined humeral carina by feeble interrupted carina, anterolateral corners of pronotum subangulate. Anterior face of pronotum, with coarse dense punctures laterally, micropuncutures sublaterally, and medial longitudinal, slightly concave, smooth unsculptured area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate mediad to areolate-punctate laterad with somewhat sharp intervals. Mesoscutum densely and finely foveolate-punctate, parapsis and notaulus reduced to posterior half of mesoscutum. Mesoscutellum “box-like”, with conspicuously horizontal dorsal face roundly angulate into nearly vertical posterior face; dorsal face shorter than posterior face; sculpture densely and coarsely areolate- punctate to foveolate-punctate. Axilla produced posterolaterally as obliquely truncate projection in dorsal view; projection coarsely foveolate-punctate basad, unsculptured, smooth, shinning apicad. Metanotum slightly wider laterad, its surface obscured by dense setation. Propodeal dorsum convex, slightly depressed sublaterally, mostly concealed by dense setation, densely areolate where visible; sculpture of lateral face absent along most of anterior margin; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely obscurely punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron micropunctate, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 0.85 × as long as wide. Dorsal 203 metasomal sculpture, except pygidial plate, partially concealed by dense setation, sparsely and finely punctate with sparse interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by parallel carinae laterally; surface densely micropunctate throughout. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina, carina slightly higher posteriorly. S2 sparsely and finely foveolate-punctate to punctate with few scattered micropunctures latearlly; sculpture sparser mediad; anteromedial crest-fold present, sternal pit absent. S3–7 sparsely and finely foveolate- punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 118:101:33. Paramere virtually straight in dorsal view, apex slightly curved outward, slighlty narrower posterad, upcurved posteriorly in lateral view, virtually asetose except for sparse scattered inconspicuous setae troughout, setae more evident on ventral surface. Cuspis slender, elongate, slightly curved outward and tapered posterad in dorsal view, upcurved posterad and slightly wider apically in lateral view, with tuft of conspicuous long setae at apex and sparser scattered inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with slightly flattened sparsely setose posterior margin, setae longer than paracuspis. Digitus short, slightly curved inward in dorsal view, strongly tapered posterad and with apex slightly upcurved in lateral view, setose anterodorsally. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, lateral pocket on outer margin greatly reduced, apical distance between teeth 0.1 × length of valve, dense setae present along convex posterior margin and inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. Integument black. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: posterior half of propodeal dorsum, T1, fringe of T2 laterally, and anterior third of S2. Distribution. Brazil, Bolivia, and Paraguay. Material examined. (8#m) Type material. Lectotype of Mutilla infernalis, #m, “S. Brasilien” [Southern Brazil], Sello S. (ZMB); holotype of Mutilla floccosa, #m, Brazil, Sello S. (ZMB). Additional material. BRAZIL: Mato Grosso do Sul, Tres Lagoas, Horto Rio Verde, 1#m, 19.VII.1994, Fletchmann (FEIS); BOLIVIA: Pando, Guayaramerin, 3#m, XII.1956, Fritz (AMNH); PARAGUAY: San Pedro: Rio Ypane: Cororo: 1#m, 05–09.XII.1983, M. Wasbauer (UCDC); 1#m, XI.1979, Fritz (AMNH). Remarks. Traumatomutilla infernalis is virtually indistinguishable from T. pompiliformis apart from the length of the cuspis in relation to the length of the paramere and minor characters of the penis valve and setae pattern. Based on distribution this could be a putative male for T. sancata.
Traumatomutilla pompiliformis (Gerstaecker, 1874) (Figs. 11A–P)
Mutilla pompiliformis Gerstaecker, 1874: 314, holotype (by monotypy), #m, Brazil, Sello S. (ZMB), type examined. Mutilla serra Cresson, 1902: 77, holotype (by monotypy), #m, Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), type examined, syn. nov. Ephuta (Traumatomutilla) pompiliformis: André, 1902, 55. Ephuta (Traumatomutilla) serra: André, 1902, 56. Traumatomutilla pompiliformis: André, 1904, p. 40. Traumatomutilla serra: André, 1904, p. 40.
Diagnosis. FEMALE. Unknown. MALE. Apex of cuspis with long setae, approximately 0.65 × length of paramere; posterior margin of penis valve truncate; lateral face of prpodoeum with sparse erect silvery-white seta; metapleuron setae black; silvery-white setae present throughout propodeal dorsum; wings strongly dark brown infuscated throughout, with strong violaceous and blueish reflections. Description. FEMALE. Unknown. MALE. Body length 19 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes. Vertex width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 3.4 × DLO, IOD 0.8 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate; sculpture sparser and finer posterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally 204 immediately below antennal insertion, conspicuously convex medially; densely and coarsely foveolate-punctate, coarser mediad; apical/ventral margin with a pair of medial short subacute free teeth medially. Scape bicarinate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 2.6 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, sharply projected from anterior margin of pronotum, separated from well-defined humeral carina, anterolateral corners of pronotum subangulate. Anterior face of pronotum, with coarse dense punctures laterally, micropuncutures sublaterally, and medial longitudinal, slightly concave, smooth unsculptured area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate mediad to areolate-punctate laterad with somewhat sharp intervals. Mesoscutum densely and finely foveolate-punctate, parapsis and notaulus reduced to posterior half of mesoscutum. Scutellum “box-like”, with conspicuously horizontal dorsal face roundly angulate into nearly vertical posterior face; dorsal face shorter than posterior face; sculpture densely and coarsely areolate-punctate to foveolate-punctate. Axilla produced posterolaterally as obliquely truncate projection in dorsal view; projection coarsely foveolate-punctate basad, unsculptured, smooth, shinning apicad. Metanotum slightly wider laterad, its surface obscured by dense setation. Propodeal dorsum convex, slightly depressed sublaterally, mostly concealed by dense setation, densely areolate where visible; sculpture of lateral face absent along most of anterior margin; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely and vestigially punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron micropunctate, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 0.9 × as long as wide. Dorsal metasomal sculpture, except pygidial plate, partially concealed by dense setation, sparsely and finely punctate with sparse interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by parallel carinae laterally; surface densely micropunctate throughout. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina, carina slightly higher posteriorly. S2 sparsely and finely foveolate-punctate to punctate with few scattered micropunctures latearlly; sculpture sparser mediad; anteromedial crest-fold present, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 77:52:18. Paramere virtually straight in dorsal view, apex slightly curved outward, slughlty narrower posterad, upcurved posteirorly in lateral view, virtually asetose except for sparse scattered inconcpsiucous setae troughout, setae more evident on vetral surface. Cuspis slender, elongate, slightly curved inward and tapered posterad in dorsal view, upcurved posterad and virtually straight throughout in lateral view, with tuft of conspicuous long setae at apex and sparser scattered inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with rounded and setose posterior margin, setae shorter than or longer than paracuspis. Digitus short, slightly curved inward in dorsal view, strongly tapered posterad and with apex abruptly curved dorsally in lateral view, setose anterodorsally. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, with well-defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along truncate posterior margin and inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. Integument black. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: anterior/ventral margin of clypeus; metanotum, prodeal dorsum; legs predominantly (occasionally all black); T1, anterior third, lateral areas, lateral felt lines, and lateral margins of T2; fringe of T2 laterally, fringe of T3 medially and laterally, fringe of T4–5 laterally, and S1–4. Wings dark-brown throughout with strong violaceous and blueish reflections. Distribution. Brazil and Bolivia. Material examined. (25#m) Type material. Holotype of Mutilla pompiliformis, #m, BRAZIL, Sello S. (ZMB); holotype of Mutilla serra, #m, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. BRAZIL: Minas Gerais, Berizal, Faz. [Fazenda] Veredão, 15°39’54’’S 41°39’56’’W, 850m [above sea level], 2m#, 12.XII.2012, Arm. [Armadilha] Malaise, J.A. Rafael & E.J. Grossi (INPA); BRAZIL: Rondônia: Vilhena: 1#m, 19.XI.1986, Elias, C. (DZUP); 1#m, 4.XI.1986, Elias, C. (DZUP); Bahia, Cerrado near Lençóis, 2#m, 15.XII.2002 205
(UEFS); Mato Grosso: Chapada dos Parecis, 2#m, 1.XII.2001, A Foucart (EMUS); Sinop, 12°31'S 55°37'W, 1#m, X.1974, M. Alvarenga (AEIC); Andradinha, 1#m, VIII.1971, Oliveira, F.M. (DZUP); Cáceres, 1#m, 10.III.1985, Elias, C. (DZUP); Chapada dos Guimarães, 1#m, 12–18.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 30km N [north of] Uirapuru, Usine Alcomat, 14°15'50.80'S 59°14'02.05''W, 1#m, 01–15.XII.2002, A. Foucart (EMUS); Minas Gerais: Pedra Azul, 800m [above sea level], 1#m, XI.1972, Alvarenga & Seabra (AEIC); 12km a N de [north of] Aguas Vermelhas, Faz. [Fazenda] Faceiro, 1#m, 14.XII.2012, Melo, G.A.R. (DZUP); BOLIVIA: Santa Cruz: Buena Vista, 17°27'58''S 63°39'63''W, 1#m, 20.II.1999, L.A. Stange (FSCA); Gral. [General] Saavedra Est. [Estación] Experimental, 2#m, XII.1973, Porter & Stange (FSCA); Buena Vista, 1#m, 1928, Steinbach (CUIC); 3 km N [north of] Brazilio, 1750' [sic!] 18°06.82'S 63°10.51'W, 1#m, 27.II–8.III.1999, Irwin & Parker (EMUS); Beni, Romanos, 1 km N [north of junction of] Rio Itenez & Rio Paragua, 1#m, 30.VII.1964, Bouseman & Lussenhop (AMNH). Remarks. This species can only be separated from T. infernalis on certain genitalia and setae pattern characters. Likewise, it can be a candidate for a sex association with T. sancta based on its known distribution.
Traumatomutilla quadrinotata (Klug, 1821) (Figs. 12A–C, 13A–F)
Mutilla quadrinotata Klug, 1821: 316, holotype, #f, Brazil, Bahia (ZMB), type examined. Mutilla micans Lepeletier, 1845: 622, lectotype [designated by Mickel (1937)], #f, Brazil (MRSN) [synonymized by Mickel (1937)]. Ephuta (Traumatomutilla) quadrinotata: André, 1902: 55. Traumatomutilla quadrinotata: André, 1904: 40.
Diagnosis. FEMALE. Occipital carina equally wide throughout, anterolateral carinae present on scutelar scale; lateral face of propodeum and metapleuron completely cover with dense appressed golden setae; pronotal setae usually conspicuously denser and longer than remainder of mesosoma. MALE. Apex of cuspis with longe setae; body setae black and silvery-golden; lateral face of propodeum and metapleuron with conspicuous patches of sparse appressed golden setae. Description. FEMALE. Body length 16-20 mm. Head. Posterior margin slightly and angularly concave. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Head densely, coarsely and confusedly foveolate-punctate to areolate-punctate with sharp irregular intervals. Genal carina present, well-defined. Mandible oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, reaching antennal tubercles and narrowly disconnected from lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 2.0 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma. Length 0.8 × width. Mesosomal dorsum densely, coarsely and confusedly areolate-punctate to foveolate punctate throughout, with sharp and irregular intervals; sculpture smaller, denser and more confused on pronotum; mesonotum with medial intervals aligne so as to form vestigial and irregular longitudinal carina. Anterior face of propodeum defined, virtually as long as pronotal collar, with vestigial coarse longitudinal striae basally and dense coarse punctures dorsally; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, projected dorsally, narrowly separated from conspicuously projected subangulate epaulet, anterolateral corners of pronotum angulate in dorsal view. Pronotal spiracle virtually flat against lateral margin of pronotum. Lateral face of pronotum sparsely punctate to foveolate- punctate with dense interspersed micropunctures except at conspicuous subacute tubercle anteroventral in relation to pronotal spiracle; mesopleuron, metapleuron and lateral face of propodeum completely concealed by dense setation. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 60:68:83:54:53. Lateral margin of mesonotum strongly constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt tooth-like process; with small inconspicuous tubercle posterior to lateral process. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area vestigial. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; intervals irregular on scutellar area, not scabrous. Propodeum gibbose/convex, dorsal face slightly shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and 206 length of T2, 38:94:87. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate where visible. S1 sparsely, coarsely and confusedly foveolate- punctate, surface cuneiform, ending in short blunt longitudinal carina, equally high throughout. S2 sparsely foveolate- punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold well-defined. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical third of plate; surface irregularly rugose; interstice apparently granulose. MALE. Body length 13–17 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; convergent immediately behind eyes. Width 0.85 × pronotal width. Eye almost circular. Ocelli small; OOD 4.9× DLO, IOD 1.2 × DLO. Occipital carina distinct. Head surface sparsely and coarsely punctate, with sparse interspersed micropunctures along posterior margin of vertex; sculpture sparser and finer posterad, denser and coarser anterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; sculpture concealed by dense setation; apical/ventral margin with a pair of medial short subacute free teeth. Scape bicarinate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, subangulately projected from anterior margin of pronotum, separated from well-defined pronounced humeral carina, anterolateral angles of pronotum subrounded. Anterior face of pronotum coarsely punctate laterally, micropunctate sublaterally, and mostly unsculptured with slightly concave longitudinal area medially. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate laterad, sculpture sparser and finer mediad. Mesoscutum densely and finely foveolate-punctate, parapsis vestigial, notaulus absent; with vestigial medial longitudinal carina. Scutellum globose to gibbose, densely and coarsely areolate-punctate to foveolate-punctate, with well-defined dorsal and posterior faces; both faces slightly convex; dorsal face, with intervals aligned so as to form irregular longitudinal carina extending medially from anterio margin of dorsal face into middle of posterior face. Axilla produced posterolaterally as obliquely truncate projection coarsely and densely foveolate- punctate except at apical third unsculptured. Metanotum wider laterally, its surface obscured by dense setation. Propodeal dorsum evenly convex, surface mostly concealed by dense setation, densely areolate-puncticulate where visible; sculpture of lateral face mostly concealed by dense setation and gradually less defined to vestigial anteroventrad; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely finely punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron mostly concealed by dense setation sparsely micropunctate to smooth throughout where visible, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 virtually as long as wide. Dorsal metasomal sculpture, except pyigidium, partially concealed by dense setation, sparsely and finely punctate with dense interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by somewhat arched carinae laterally; surface irregularly micropunctate, somewhat granulose, sculpture coarser apicad. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina terminating. S2 sparsely and finely foveolate-punctate to punctate; sculpture slightly sparser mediad; S2 with reduced anteromedial crest-fold, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 70:53:17. Paramere virtually straight in dorsal view, apex slightly curved outward, upcurved posteriorly in lateral view, virtually asetose except for sparse scattered inconspicuous setae troughout, setae more evident on vetral surface. Cuspis slender, elongate, slightly curved inward and virtually straight throughout in dorsal view, upcurved posterad and slightly broader posterad in lateral view, with tuft of conspicuous long setae at apex and sparse scattered inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with irregular 207 almos serrate and setose posterior margin, setae longer than paracuspis. Digitus short, strongly curved inward in dorsal view, somewhat dorsoventrally flattened, evenly upcurved in lateral view, apex more abruptly curved and subcapitate, setose anterodorsally. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without well-defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along convex posterior margin and inconspicuous setae present at base of anterior tooth on outer surface, setae on posterior margin longer ventrad. Colorations and variations. FEMALES. Integument black, except mandibles and antennal flagellomeres reddish- brown ventrally, and T2 with four large orange integumental spots. Body setae predominantly golden, except following areas with black to brownish-black setae varying in density and length: head; procoxae; pronotum; mesonotum anteriorly; propodeal dorsum medially; T1 medially; disc of T2 (except integumental spots); fringe of T2 medially; fringe of T3–5 sublaterally; S6. MALES. Head and mesosomal integument black to brownish-black, except antennal flagellomeres and mandibles partially reddish-brown. Body setae predominantly golden varying in density, except following areas with black to brownish-black setae varying in density: head, dorsum of foretibae and femora, pronotum, mesopleuron anteriorly, mesoscutum, axillar projections, dorsal face of scutellum, posterior two thirds of T2, fringe of T2–3 medially, T4–7, fringe of S2–6, and S5–7. Wings dark-brown except at basal third hyaline-brow. Tibial spurs yellowish-white. Distribution. Brazil. Material examined. (151#f 37#m) Type material. Holotype of Mutilla quadrinotata, #f, Brazil, Bahia (ZMB). Other material. BRAZIL: 8#f (ANSP, ZMUC, DEI, MNCN); 1#m (MNRJ); Pernambuco: San Sabrador [sic!], 1#f (ANSP); Caruaru, 4#m, V.1972, Lima, J.M. (DZUP); Bahia: 1#f (CPDC); Barra do Choca, 1#f, 17.I.2001, J.R. Maia (CPDC); Barrolandia, 1#f, III.2005, J.R.M. Santos (CPDC); Boa Nova, 1#f, 28.I.2004, E. Mariano (CPDC); Itamarajó, 2#f, 11.XI.2004, Valmir Paulino (CPDC); Encruzilhada, 980m [above sea level], 1#m, XI.1974, M. Alvarenga (TAMU); Santa Terezinha, Serra da Jibóia, 1#f, 08.IV.2001, M.F. Soares (CPDC); Serra das Lonas, Serra das Lonas, 2#f, III.2009, Camacho, A. (MZSP); 2#f, Cachimbo,1890 (DEI); Espírito Santo: 6#f (MZSP, MNCN); Santa Teresa, 1#f, 03.IV.2004, E. Mariano (CPDC); 1#f, 7.XII.1964, Elias,C. (DZUP); 1#m, 13.I.1970, Elias, C., Elias, CT. (DZUP); Linhares: 1#f, 6.I.1971, Elias,C. (DZUP); 1#m, X.1972, M. Alvarenga (CMNH); 1#f, V.1973 (MZSP); 1#m, XII.1975, Elias, C. (DZUP); Parque Sooretama: 7#f (MNRJ); 1#f, XI.1967, Oliveira, F.M. (DZUP); Conceição da Barra, 1#f, 4.IX.1969, Elias,C., Elias,C.T. (DZUP); Colatina, 1#f, 13.XII.1967, Elias,C.T. (DZUP); 1#f, XII.1954 (MNRJ); Faz. [Fazenda] Usina Paineiras, P1 [sic!], 20°56'29''S, 41°03'06''O, Itapemirim, 3#m, 19–26.XI.2010, M.T. Tavares e eq. [equipe] (UFES); Res. [Reserva] Biol. [Biológica] Córrego do Veado, Água Limpa, 18°21'S 40°09'O, Pinheiros, 2#m, 27.XI–06.XII.2011, M.T. Tavares e eq. [equipe] (UFES); Joatuba, Faz. [Fazenda] Betzel, 19°50'25''S 40°49'40''O, 280-430 [sic!], Laranja da Terra, 14#m, 05–12.x.2012, M.T. Tavares e eq. [equipe] (UFES); Conceição da Barra: 1#f, VII.1969 (MZSP); FLONA [Floresta Nacional] Rio Preto, 1#f, 21–25.III.2005 (UFES); D. [Domingos] Martins: 1#f, I.1962, Elias,C. (DZUP); Zona Rural, 1#f, XII.1999 (UFES); Corrego do Ita, 1#f, XI.1957 (MNRJ); Chaves, 1#f, 11.IX.1963, J. Baskin (USNM); Aracruz, 1#f, 15.IX.2000 (UFES); Sítio Benincá, 18°56'S 40°27'O, Vila Valério, 1#m, 14–28.IX.2011, C.O. Azevedo e eq. [equipe] (UFES); Reserva Biol. [Biológica] Duas Bocas, Vitória, 1#f, 21.VII.2001 (UFES); Minas Gerais: 1#f (MNCN); 1#f, Reinhardt (ZMUC); Faz. [Fazenda] Montes Claros, Ipanema, 1#f, 30.IV.1998 (DZUP); Paineiras, n.o proc 10/963 [sic!], 1#f, X.1963 (MNRJ); Viçosa: 1#f, 1931, E.C. VanDyke (CASC); 1#f, 1931, Y. Mexia (CASC); Mar de Espanha: 1#f, 04.X.1905 (MNCN); 2#f, 20.XI.1892 (DEI); Mato Grosso do Sul, Dourados, 1#f, I.1976, Lorenzoni (DZUP); Rio de Janeiro: 30#f (MZSP); 1#f, 15.XI.1926 (MNCN); Rio de Janeiro: Rio de Janeiro: 1#f (ZMUC); 1#, 6.III.1932 (MNRJ); 2#, 30.II.1940 (MNRJ); 1#f, 25.XII, A.F. Porter (FMNH); Dist. Federal [sic!], n.o. proc. 10/954, 1#f, X.1954 (MNRJ); 1#f, III.1937, R.C. Shannon (USNM); Floresta do Tijuca: 2#f (MNRJ); 1#f, III.1963, C.A. Campos Seabra (AMNH); 1#f, III.1951, C.A. Campos Seabra (USNM), 1#f, VI.1951, C.A. Campos Seabra (USNM); Est. Guanabara [Rio de Janeiro], Corcovado, 1#f, 27.I.1906, H. Luederwaldt (MZSP); 1#m, II.1962, Alvarenga, Seabra (DZUP); 1#f, III.1959, Alvarenga, Seabra (DZUP); 1#f, III.1934 (MNRJ); Repressa Rio Grande, 1#f, X.1960 (MNRJ); 1#f, 5.II.1967, Oliveira, F.M. (DZUP); 2#f, 30.IV.1967, Oliveira, F.M. (DZUP); 1#f, 5.X.1960, Oliveira, F.M. (DZUP); M. [Morro] Sta. [Santa] Barbara, 1#f, 16.III.1952 (MNRJ); Horto do J. [Jardim] Bot. [Botânico], 1#f, 12.II.1984 (MNRJ); Itatiaia: 1#f, 17.I.1924, J.F. Zikan (CASC); 1#f, 20.VII.1944, J.F. Zikan (CASC); 1#f, 21.II.1926, J.F. Zikan (CASC); 1#f, 07.III.1948, J.F. Zikán (MZSP); Petropolis: 1#f, IV.1952, C. Novais (USNM); 1#f, 1938 (MNRJ); 1#f, III.1952 (MNRJ); Grotão de Independencia, 1#f, 1935 (MNRJ); Ilha Grande, 1#f, 1944 (MNRJ); 1#f, 17.II.2003, D.J. Souza (CPDC); Angra dos 208
Reis: 1#f, 11.XI.1972 (MNRJ); Jabuhyba [Jabuíba], 1#f, X.1936 (MNRJ); Jussaral, 2#f, IV.1934 (MNRJ); 1#f, X.1934 (MNRJ); Mendes, 1#f, 1917 (MNCN); [Teresópolis], Serra dos Orgaos, 1#f, XI.1940 (MNRJ); Estr. [Estrada] Sumare, 1#f, 7.II.1955 (MNRJ); Floresta do Macaco, 1#f, XI.1960 (MNRJ); "Sitio las Luas" cua formiguero de "Saríva" Parada de Mendes [sic!], 1#f, 24.I.1936 (MNRJ); São Paulo: 5#f (MZSP); T. [Teodoro] Sampaio, Pq. [Parque] Est. [Estadual] Morro do Diablo, 1#f, 6.XII.2010, Melo, G.A.R., Luz, D. (DZUP); Caraguatatuba, Res. [Reserva] Flor. [Florestal], 40m, 1#f, 2.IV.1962 (MNCN); Guarujá, Ilha de Santo Amaro, 1#f, 20.VI.1961 (MNCN); Ubatuba, PESM-Nucl. [Parque Estadual Serra do Mar – Núcleo] Picinguaba, ponto 2, 23d20'01"S 44d49'57"W: 1#m, 18.IV.2010, Perioto, N.W. (MZSP); 1#m, 18.II.2010, Perioto, N.W. (MZSP); 1#m, 18.XII.2009, Perioto, N.W. (MZSP); Piraquara, Mananciais da Serra, 1#m, 10.II.2006, L.C. Rocha-Filho (DZUP); 1#f, 10.I.2003, Melo, G.A.R, Costa, M. (DZUP); 1#f, 2.II.2003, Garcia,E.Q. (DZUP); Antonina, Reserva Cachoeira, 1#f, 1.XII.2006, Maia, C. (DZUP); Campo Largo, 1#f, 10.IV.2005, Melo, G.A.R., Aguiar, A. (DZUP); Santa Catarina: 3#f (MZSP, MNCN); S. [São] Bento, 1#f, II.1952, A. Maller (USNM); Mafra, 2#f, IV.1942, A. Maller (AMNH); Corupa: 1#f, V.1940, A. Maller (AMNH); 1#f, IV.1941, A. Maller (AMNH); 1#f, XI.1942, A. Maller (AMNH); 1#f, III.1941, A. Maller (AMNH); Rio Vermelho, 2#f, (MZSP). Remarks. Traumatomutilla quadrinotata is perhaps one of the most common and easily identifiable species in the Brazilian Atlantic Forest. The sex association was based on the reliable color pattern of males and females of the Atlantic forest species, which has been used as a basis for sex association in various species in the region (Bartholomay et al. in prep.). The males of T. quadrinotata and T. tetratrauma are remarkably similar in coloration and external morphology, differing only in the length of the setae at the apex of the cuspis. Therefore, associating sexes for T. quadrinotata was only possible after the discovery of T. tetratrauma and its similarities with T. incerta (detailed in the remarks of T. tetratrauma). Although it is common for some specimens to completely or partially lose some of the body setae, the conspicuously long erect pronotal setae on the females of T. quadrinotata seems to be a consistent and exclusive character for this species (PRB & KAW pers. obs.).
Traumatomutilla quadripustulata (Klug, 1821) (Figs. 14A–C, 15A–D, 16A–G)
Mutilla quadripustulata Klug, 1821: 316, lectotype [designated by Mickel (1964)], f#, Brazil, Para [sic] (ZMB), examined. Mutilla pruinosa Smith, 1855: 43, holotype [by monotypy], #m, Brazil, Para [sic] (BMNH), examined, syn. nov. Mutilla maraca Cresson, 1902: 78, lectotype [designated by Cresson (1916)], #m, Brazil, [Pará], Santarém (CMNH), examined, syn. nov. Ephuta (Traumatomutilla) quadripustulata: André, 1902: 55. Ephuta (Traumatomutilla) maraca: André, 1902: 55. Mutilla pruinosa: André, 1902: 74 (incertae sedis). Traumatomutilla quadripustula: André, 1904: 40. Traumatomutilla maraca: André, 1904: 40.
Diagnosis. FEMALE. Occiptial carina equally wide throughout; anterolateral carinae absent on scutelar scale; post- mesonotal tubercle conspicuously; body setae predominantly black with inconspicuous silvery-white areas on metasoma; T2 usually with two pairs of small yellowish integumental spots. MALE. Apex of cuspis with longe setae; pronotal sculpture not at all concealed by dense setation; T2 entirely covered with silvery-white setae, less densely so on posterior half. Description. FEMALE. Body length 16-18 mm. Head. Posterior margin slightly and angularly concave. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 1.3 × the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate- punctate to areolate-punctate. Genal carina present, well-defined. Mandible oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, reaching antennal tubercles and disconnected from lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 1.9 × pedicel length; flagellomere 2 1.6 × pedicel length. Mesosoma. Length 0.85 × width. Mesosomal dorsum densely and coarsely areolate-punctate throughout, intervals sharper posteromediad and blunt laterad; with vestigial irregular longitudinal carina 209 anteromedially. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, vestigial coarse striae longitudinally basomedially, and sparse coarse punctures dorsomedially; micropunctate anterior to epaulets; dorsal face roundly angulate into anterior face in lateral view. Humeral carina well-defined, slightly projected dorsally, broadly separated from conspicuously projected subangulate epaulet, anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded, bulging. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures except at smooth conspicuous subacute tubercle anterior to pronotal spiracle; mesopleuron densely micropunctate anteriorly, sparsely punctate to foveolate-punctate to areolate-punctate ventrad on mesopleural ridge; metapleuron unsculptured smooth shinning on dorsal fourth, densely and coarsely areolate on basal fourth, and with dense micropunctures elsewhere. Lateral face of propodeum densely and coarsely foveolate-punctate anterad; sparsely foveolate-punctate posterad; conspicuous smooth shinning, somewhat granulose areas along posterior margin. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 69:79:83:60:61. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt tooth-like process; with very small conspicuous tubercle posterior to lateral process. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area vestigial. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; scabrous intervals vestigial on scutellar area. Propodeum gibbose, dorsal face much shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 42:99:98. Disc of T2 densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 sparsely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold vestigial. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S4 densely foveolate-punctate; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose; rugae vestigial to absent apicad. MALE. Body length 9–15 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; convergent immediately behind eyes. Width 0.9 × pronotal width. Eye almost circular. Ocelli small; OOD 3.6 × DLO, IOD 1.25 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate, with sparse interspersed micropunctures along posterior margin of vertex; sculpture sparser and finer posterad, denser and coarser anterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; sculpture concealed by dense setation; apical/ventral margin with a pair of medial short subacute free teeth. Scape bicarinate. Flagellomere 1 1.7 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, subangulately projected from anterior margin of pronotum, separated from well-defined pronounced humeral carina, anterolateral angles of pronotum angulate. Anterior face of pronotum mostly unsculptured, with sparse punctures and interspersed micropunctures laterally. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate. Mesoscutum densely and finely foveolate-punctate, parapsis virtually absent, notaulus reduced to posterior half of scutum; with medial longitudinal carina on posterior half. Scutellum globose to gibbose, densely and coarsely areolate-punctate to foveolate-punctate, with well-defined dorsal and posterior faces; dorsal face broader than long, with intervals aligned so as to form irregular longitudinal carina extending medially from anterio margin of dorsal face into middle of posterior face. Axilla produced posterolaterally as obliquely truncate projection, with inner margin conspicuously curve inward apically; projection coarsely and densely foveolate-punctate except at apical third unsculptured. Metanotum wider laterally, its surface obscured by dense setation. Propodeal dorsum convex, slightly depressed dorsolaterally, surface mostly concealed by dense setation, densely areolate where visible; sculpture of lateral face gradually less defined to vestigial anteroventrad; dorsal face indistinguishable from posterior face. Lateral face of pronotum sparsely finely punctate with sparse interspersed micropunctures; mesopleura with conspicuous subacute spine-like projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron sparsely micropunctate to smooth throughout, except 210 basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 virtually as long as wide. Dorsal metasomal sculpture, except pyigidium, partially concealed by dense setation, densely and finely punctate with dense interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by somewhat arched carinae laterally; surface irregularly micropunctate, somewhat granulose, sculpture coarser apicad. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina terminating. S2 sparsely and finely foveolate-punctate to punctate; sculpture slightly sparser mediad; S2 with reduced anteromedial crest-fold, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe not at all pronounced apically, subacute. Ratios of free length of paramere, cuspis and digitus, 69:48:20. Paramere virtually straight in dorsal view, upcurved posteriorly in lateral view, virtually asetose except for sparse scattered inconspicuous setae troughout, setae more evident on vetral surface. Cuspis slender, elongate, slightly curved inward and tapered posterad in dorsal view, upcurved posterad and slightly broader posterad in lateral view, with tuft of conspicuous long setae at apex and sparse scattered inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with subangulately concave and vestigially setose posterior margin, setae shorter than paracuspis. Digitus short, strongly curved inward in dorsal view, somewhat dorsoventrally flattened, evenly upcurved in lateral view, apex more abruptly curved and subcapitate, setose anterodorsally. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without well-defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along convex posterior margin and inconspicuous setae present at base of anterior tooth on outer surface, setae on posterior margin longer ventrad. Colorations and variations. FEMALES. Integument black except for mandibles and antennal flagellomeres partially reddish-brown, and T2 with four usually small and yellowish integumental spots. Spots of T2 highly variable in size, shape and color: separated by more than spots width to separated by hald a spot width; subcircular to subquadrate; yellowish to orange or reddish. Body setae almost exclusively black, except for the following areas vestigially and sparsely clothed with silvery-white setae often with interspersed black setae: mesopleuron posteromedially, coxae, ventral surface of femora and tibiae, inner surface of basitari, lateral areas of T2, lateral felt lines of T2, lateral margins of T2, fringe of T2–5 medially and laterally (usually absent medially on fringe of T3), T6 medially (except pygidial plate), and S1–5. Tibial spurs reddish-brown. MALES. Head and mesosomal integument black; legs and metasoma brownish-black; antennae and mandibles partially reddish-brown. Body setae predominantly silvery-white varying in density, except following areas predominantly or with interspersed black to brownish-black setae: vertex and front partially, dorsum of pronotum, mesonotum, axillar projections, scutellum, fringe of T3 medially, T4–7, fringe of S5–6, and S7. Wings hyaline-brown, slightly infuscated on apical third and small costal patch basal in relation to pterostigma. Certain specimens might have the pronotal dorsum completely black or with interspersed black and silvery-white setae and/or the posterior half of T2 with sparse black setae. Wings infuscated-brown on apical third, hyaline-brown elsewhere, without any noticeable reflections. Tibial spurs white. Distribution. Brazil. Material examined. (45#f 47#m) Type material. Lectotype of Mutilla quadripustulata, f#, BRAZIL, Para [sic] (ZMB); holotype of Mutilla pruinosa, #m, BRAZIL, Para [sic] (BMNH); lectotype of Mutilla maraca, #m, BRAZIL, [Pará], Santarém (CMNH). Additional material. Pará: 4#f (MZSP); 1#f, 24.X.1901 (MPEG); 1#f, 13.I.1902 (MPEG); 1#f, 16.VIII.1900 (MPEG); BR 014, Km 92, 1#f, XII.1960 (MNCN); Santarém, 4#f (ANSP); 1#f (FSCA); 3#f (CMNH); 4#f, VI.1919, S.M. Klages (CMNH); Amazonen-stroomgebied, 1#f (CSCA); Fazenda Taperinha, 1#f, 01– 11.II.1968 (MPEG); 2#f, Mararu (CMNH); Mangabaira, Mocajuba: 2#f, VIII.1953, O. Rego (AMNH); 1#f, III.1953, O. Rego (LACM); 3#f, X.1953, O. Rego (USNM); 1#f, VIII.1953 (MNCN); 1#f, XI.1952, O. Rego (FSCA); 5#f (MNRJ); Benevides: PA-408 Km 06, 1#f, 26.VII.1982 (MPEG); Benevides: Fazenda Morelândia: 1#m, 02.VII.1988, F.F. Ramos (MPEG); 1#m, 11.XII.1983, Jarbas (MPEG); 1#m, 20.VI–02.VII.1988, F.F. Ramos (MPEG); 1#m, 04– 06.VIII.1988, F.F. Ramos (MPEG); 1#m, 30.VI–02.VII.1988, F.F. Ramos (MPEG); Benevides, CPDC, 1#m, 11.VIII.1994, J.A. Pena (MPEG); Melgaço, ECFPn [Estação Científica Ferreira Pena] Caxiuanã: 1#f, 24.IV.1995 (MPEG); SMC1 [sic!], Ig. [Igarapé] Curuazinho, 1#f, 28.III.1998 (MPEG); Mata da Sede, 1#m, 16.XI.1998, O. Silveira & J. Pena (MPEG); Percurso, 1#m, 26.IV.1999, O. Silveira & J. Dias (MPEG); FLONA [Floresta Nacional] 211
Tapajós, 1#m, 11–14.IX.1997, M. Henriques (MPEG); 1#m, 02–05.IX.1997, M. Henriques (MPEG); Juruti: Alcoa: Área beneficiamento bauxite: 02°29'32.10''S 56°09'20.04''W: 1#m, 21–24.VI.2008, E. Monteiro-Santos, R.L. Trindade, Domingos Guimarães, L.A. Quaresma (MPEG); 1#m, 09–13.XII.2007, E. Monteiro-Santos, R.L. Trindade, Domingos Guimarães, L.A. Quaresma (MPEG); Jaratuba, Rio Mamuru, Área 2, 03°11'32,6''S 56°34'51,4''O, 1#m, 29.IX.2009, O.T. Silveira, S.S. Silva & J. Pena (MPEG); Mosqueiro, 1#m, 12.XII.1988, L.B. Albuquerque (MPEG); 1#m, 17.XII.1983, L.B. Albuquerque (MPEG); Serra Norte, Est. [Estrada] Manganês, 1#m, 10.VI.1983, F.F. Ramos (MPEG); 1#m, 08.VI.1983, F.F. Ramos (MPEG); 1#m, 09–12.IX.1985, F.F. Ramos (MPEG); 1#m, 06–09.IX.1985, W. França (MPEG); N1 Mata [sic!], 1#m, 03–05.XI.1985, J. Dias (MPEG); Carajás, Est. [Estrada] Manganês, 1#f, 24.IV.1983 (MPEG); Pedras, Rio Cuminá, 1#f, 01.XI.1969 (MPEG); Marabá, 1#m, X.1974, J.F. Reinert (FSCA); Belém, Parque Ambiental, Sede, 1#m, 14–18.VI.2004, A.L. Nunes e equipe (MPEG); Bujaru, 1#m, 27.XII.1985 (MPEG); Paragominas, Fazenda Cachoeira do Rio Vermlho, 1#m, 15–18.I.1991, R.B. Neto (MPEG); Vitória do Xingú, Margem direita Rio Xingú, 1#m, 22–24.XI.2000, R. Santos & J. Dias (MPEG); Pacoval, 1#m, 07–17.XII.2008 (EMUS); Jacareacanga, 1#m, XI.1968, Moacir Alvarenga (EMUS); Mutum, 1#m, 01–06.XII.2008 (EMUS); Tucuruí, 1#m, I.1979, M. Alvarenga (EMUS); Maranhão, Bom Jardim, Reserva Biológica do Gurupi, 1#f, 17–27.I.2010 (CZMA); Rondônia: 1#f (RBINS); Ouro Preto D'Oeste, Linha 62, Km 16.11, 1#m, 13.XI.1984, F.F. Ramos (MPEG); 62 km SE [southeast of] Ariquemes, 1#m, 8.XI.1994, W.J. Hanson (EMUS); Vilhena: 1#m, 22.XII.1986, Elias, C. (DZUP); 1#m, 27.XII.1986, Elias, C. (DZUP); 2#m, 4.XI.1986, Elias, C. (DZUP); 1#m, 27.XI.1986, Elias, C. (DZUP); 1#m, 17.XII.1986, Elias, C. (DZUP); 1#m, 22.XII.1986, Elias, C. (DZUP); Mato Grosso, Sinop, 5#m, II.1976, M. Alvarenga (AEIC); 12°31'S 55°37'W, 1#m, X.1974,M. Alvarenga (EMUS); Tangará da Serra, Fazenda Promissão, 14°30'01''S 57°17'30''W, 346m asl [above sea level], Floresta semidecidual, 2#m, 01.X.2017, Ferreira, J.V.A. (UNEMAT); BOLIVIA, Chuquisaca, 5 mi S [miles south of] Camarga, 1#f, 19.II.1951 (AMNH). Remarks. After the sex association and synonymies of T. incerta were recognized, the only species found in the Amazon, particularly the Southeastern Amazonian areas were T. quadripustulata, Mutilla pruinosa and T. maraca. The types of M. pruinosa and T. maraca are identical in external and genitalic morphology. They have been repeatedly been collected in the same areas as females of T. quadripustulata. As the two largest Traumatomutilla females with almost entirely black setal coloration, T. chrysozona (formerly T. lugubrina) and T. quadripustulata were easily mistaken for one another. Previous researchers separated T. chrysozona from T. quadripustulata mainly using T2 spot color: yellow in T. quadripustulata and reddish in T. chrysozona. The T2 integumental spots in T. quadripustulata, however, vary greatly in size and color. Various easily overlooked features are now recognized to differentiate these forms. First, T. quadripustulata females have a post-mesonotal tubercle that is usually associated with a slenderer and more elongate mesosoma than females of T. chrysozona. The most consistent setal features are the always present, though often inconspicuous, silvery-white setae patches on the metasoma of T. quadripustulata, compared against the rarely present and usually reduced coppery setae patches on the fringes of T2–3 in T. chrysozona. Finally, true T. quadripustulata specimens have been found only in Southeastern Amazonian areas, while T. chrysozona has a more southern distribution predominantly in eastern Cerrado areas of Brazil.
T. sancta (Gerstaecker, 1874) (Figs. 17A–F)
Mutilla parallela Burmeister (nec Klug) 1854: 24, holotype (by monotypy), #f, Brazil, [Minas Gerais], Lagoa Santa (ZMB), examined (preoccupied name). Mutilla sancta Gerstaecker, 1874: 303 [new name for Mutilla parallela (Burmeister, 1854)]. Mutilla solemnis Cresson, 1902: 49, holotype [by monotypy], #f, Brazil, Minas [Minas Gerais] Car. [sic!] (CMNH), type examined, syn. nov. Mutilla sodalis Cresson, 1902: 52, lectotype [designated by Cresson (1902)], Brazil, [Mato Grosso], Chapada [dos Guimarães] (CMNH), examined, [synonymized by Mickel (1964)]. Ephuta (Traumatomutilla) sancta: André, 1902: 55. Ephuta (Traumatomutilla) solemnis: André, 1902: 56. Traumatomutilla sancta: André, 1904: 40. Traumatomutilla solemnis: André, 1904: 40. 212
Diagnosis. FEMALE. Occipital carina equally wide throughout; anterolateral carinae absent on scutelar scale; lateral face of propodeum densely but unevenly sculptured, with conspicuous unsculptured areas; T2 with four pairs of linear to subrectangular narrow yellowish integumental spots. MALE. Unknown. Description. FEMALE. Body length 21–23 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 1.1 × the distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate. Genal carina well-defined. Mandible oblique, tapering slightly towards apex with small subapical tooth. Dorsal scrobal carina present, well-defined, separated from antennal tubercles, narrowly connected to lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 1.9 × pedicel length; flagellomere 2 1.4 × pedicel length. Mesosoma. Dorsal thoracic length 0.85 × its width. Mesosomal dorsum mostly concealed by dense setation, densely and coarsely areolate-punctate with apparent sharp to scabrous intervals where visible. Anterior face of propodeum defined, as lon as than pronotal collar, coarsely striated longitudinally throughout with dense coarse punctures dorsomedially; dorsal face rounded into anterior face in lateral view. Humeral carina well- defined, projected dorsally, broadly separated from conspicuously projected angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle projected from lateral margin of pronotum, bulging. Lateral face of pronotum sparsely punctate with dense micropunctures except at smooth conspicuous subacute tubercle anteroventral in relation to pronotal spiracle; mesopleuron partially concealed by dense setation, densely micropunctate anteriorly, and dense vestigial foveolate-punctate on mesopleural ridge; metapleuron unsculptured smooth shinning on dorsal fourth, densely, coarsely and confusedly areolate-punctate on basal fourth, and concealed by dense setation elsewhere. Lateral face of propodeum densely and coarsely areolate-punctate throughout with blunt vestigially rugose intervals. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 52:59:63:46:45. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt process; with small conspicuous blunt post-mesonotal tubercle. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area absent. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; scabrous intervals present on scutellar area. Propodeum convex, dorsal face shorter than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 34:75:76. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures where visible; foveolations sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate- punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, densely foveolate-punctate. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, with equally high irregular edges; S2 densely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold present. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S5–6 densely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose. Coloration and variations. FEMALE. Integument black, except mandible and antennal flagellomeres partially reddish-brown, and T2 with four large yellow integumental spots; anterior pair always longitudinal linear, posterior pair transverse subrectangular and broadly separated to transverse linear narrowly interrupted medially. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: head, except vertex frequently with transverse line of silvery-white setae which can be absent or reduced to few scattered setae; pronotum, mesopleuron anteriorly, anterior half of mesoscutum, posterior half of mesoscutum medially, lateral face of propodeum partially, propodeal dorsum medially; legs, except dorsal surface of tibiae and apicodorsal surface of femora; T1 and disc of T2 medially (except integumental spots), fringe of T2–5 sublaterally, T6 laterally (except pygidial plate), Fringe of S5, and S6. Distribution. Venezuela, Brazil, Bolivia, and Paraguay Material examined. (102#f) Type material. Holotype of Mutilla sancta, #f, BRAZIL, [Minas Gerais], Lagoa Santa (ZMB); holotype of Mutilla solemnis, #f, BRAZIL, Minas [Minas Gerais] Car. [sic!] (CMNH); lectotype of Mutilla sodalis, #f, BRAZIL, [Mato Grosso], Chapada [dos Guimarães] (CMNH). Additional material. VENEZUELA, Baixo 213
Sarare [sic!], 33 96 [sic!], 1#f, F. Gray (MNHN); BRAZIL: 1#f (ANSP); Maranhão, Mirador, Parque Estadual do Mirador, Base da Geraldina, 1#f, 27.X–01.XI.2008, M.B. Aguiar-Neto & A.L. Costa (CZMA); Mirador, E14 [sic!], 1#f (INPA); Codó, E13 [sic!], 1#f (INPA); Rondônia, 5#f (MZSP); Bahia, R. [Rio] de Contos [Contas], 1#f, 01.V.1993, Argolo, A.J.S. (CPDC); Encruzilhada, 2#f, 12.XII.2007, Grossi, Rafael & Parizotto (DZUP); Lençois, 1#f, 28.II.2001, Maia, J.R. (CPDC); Mato Grosso: 1#f, 1886, P. Germain (MNHN); Diamantina, 1#f, XI–XII.1973 (MZSP); Chapada dos Guimarães: 1#f, 8.II.1986, Exp. Dept. Zoo. [Expedição Departamento de Zoologia UFPR [Univerisdade Federal do Paraná] (DZUP); 1#f, 26.III.2001, Sousa, W.O. (DZUP); 1#f, 18.XI.2013, Melo, G.A.R., Luz, D.R., Williams, K.A. (DZUP); 1#f, X (ANSP); 1#f, XI (ANSP); 1#f, I (ANSP); P. N. [Parque Nacional] do Xingu, Jacaré: 4#f, 30.II.1965, Alvarenga & Werner (DZUP); 2#f, XI.1961, Alvarenga & Werner (MPEG, MNCN); Cáceres, 1#f, 27.III.1985, Elias,C. (DZUP); Utiariti, 5#f (MZSP); Rio Papagaios, Utiariti, 1#f, 1–12.XI.1966, Lenko & Pereira (MZSP); Rosario Oeste, 3#f (MZSP); Andradinha, 1#f, VIII.1971, Oliveira, F.M. (DZUP); 30km N [north of] Uirapuru, Usine Alcomat, 14°15'50.80''S 59°14'02.05''W, Chapada dos Parecis, 1#f, 01–15.XII.2002, A. Foucart (EMUS); Goiás: 2#f, 1914 (MNHN); 3#f (MZSP); Jataí, 1#f, 1914 (MNHN); Serra da Mesa, 13°52'S 8°23'O, Campinaçu, 3#f, 18.II–02.III.1996, Silvestre, Brandão & Yamamoto (MZSP); Minas Gerais: 2#f (MZSP); 1#f (ZMUC); Pirapora, 1#f, XI.1975 (MNRJ); Lagoa Santa, 1#f, Reinhardt (ZMUC); 15 km SE [sudeste] de Riacho dos Machados, 3#f, 12.IV.1998, Melo, G.A.R (DZUP); Rio Pardo de Minas, 1#f, 10.I.1952 (MNRJ); Pedra Azul, 700m [above sea level], 2#f, XI.1972 (MNRJ); Barro Alto, 1#f, XI.1931 (DGMC); Lassance, 1#f, III.1935, D.M. Cochran (USNM); Ponto dos Volantes,14 km ao S [sul] de Ponto dos Volantes, 1#f, 12.II.2010, Melo, G.A.R., Parizotto, D., Grossi, P. (DZUP); Sertão de Diamantina,Fazenda das Melancias, 1#f, X–XI.1902, E. Gounelle (MNCN); Três Lagôas, 3#f (MZSP); São Paulo: 13#f (MZSP); Ainores, 1#f, X.1947, J. Bras (AMNH); Rio Claro, 1#f, II.1971, B. Dias (AMNH); Rio de Janeiro, 1#f, Reinhardt (ZMUC); BOLIVIA: Santa Cruz, Buena Vista, 4#f, 1928, J. Steinbach (CUIC); La Junta, 2#f, II.1947, Peredo (USNM); Beni, Cavinas, 1#f, I.1922, W.M. Mann (USNM); Guayaramerin, 1#f, XII.1956, M.A. Fritz (AMNH); PARAGUAY: Boqueron, El Solitario, 22°50'S 61°59'W, 1#f, 10.XI.2002, U. Dreschel (FSCA); El Solitario, 22°50'S 61°59'W, 1#f, 10.XI.2002, U. Dreschel (FSCA); Pres. [Presidente] Hayes, Lolita, Yaragui: 1#f, 10.I.2002, U. Dreschel (FSCA); 1#f, 25.I.2005, U. Dreschel (FSCA); San Pedro, Rio Ypane, Cororo, 1#f, II.1991, Arriagada (EMUS); 1#f, 26.I.1965, Williner (AMNH); 1#f, XI.1983, M.A. Fritz (AMNH); 1#f, II.1991, Arriagada (AMNH); Concepción, Cororo, 2#f, II.1993, L.K. Arriagada (AMNH); San Bernardino, 1#f, V.1918, K. Fiebrig S.V. (ZMB). Remarks. Based on distribution T. sancta is another species that could potentially be associated with T. pompiliformis or T. infernalis, though we refrain from doing so at this point due to the lack of any other putative males for T. austera, T. funebris, or T. ursina. These also partly overlap in distribution with T. pompiliformis. The color pattern and distribution of T. sancta is typical of the dry savannah like areas of Cerrado and Caatinga in Midwestern and Northeastern Brazil, which is why the record from Venezuela is likely to be mislabeled. Aside from the integumental spots of T2, there are no differences between T. sancta and T. solemnis. There are, however, no obvious intermediate forms between the relatively short and subrectanuglar spots of T. sancta and the elongate linear spots of T. solemnis. We have, however, seen examples of similar patterns forming a continuum between various Traumatomutilla species, such as T. parallela (Klug, 1821) from the T. indica species-group (PRB & KAW pers. obs.).
T. tetratrauma Bartholomay & Williams sp. nov. (Figs. 18A–B, 19A–F)
Diagnosis. FEMALE. Occipital carinae slightly swollen dorsolaterally, anterolateral carinae present on scutelar area, lateral face of propodeum with dense micropucntures in between sparse foveolations, frons usually clothed with golden setae. MALE. Apex of cuspis with short setae; lateral face of propodeum and metapleuron with conspicuous patches of appressed golden setae. Description. FEMALE. Body length 15 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and slightly swollen laterally. Vertex width 0.75 × pronotal width. Eye almost circular, its length in frontal view virtually equal to distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate- punctate to areolate-punctate, less densely so on malar space. Genal carina well-defined. Mandible oblique, tapering slightly towards apex with small subapical tooth. Dorsal scrobal carina present, well-defined, narrowly disconnected from antennal tubercles. Lateral scrobal carina virtually absent. Antennal tubercle coarsely and irregularly rugose. 214
Flagellomere 1 2.6 × pedicel length; flagellomere 2 1.9 × pedicel length. Mesosoma. Length 0.8 × width. Mesosomal dorsum mostly concealed by dense setation, densely and coarsely areolate-punctate with apparent sharp to scabrous intervals where visible. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, coarsely striated longitudinally throughout with dense coarse punctures dorsad; dorsal face rounded into anterior face in lateral view. Humeral carina well-defined, slightly projected dorsally, broadly separated from conspicuously projected angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded, bulging. Lateral face of pronotum sparsely punctate with dense micropunctures except at smooth conspicuous subacute tubercle anterior to pronotal spiracle, distance between tubercles wider than distance between pronotal spiracles; mesopleuron mostly concealed by dense setation, densely micropunctate anteriorly, and densely and coarsely foveolate-punctate to areolate-punctate on mesopleural ridge where visible; metapleuron completely concealed by dense setation, except small posterior area on dorsal fourth unsculptured smooth and shinning. Lateral face of propodeum mostly concealed by dense setation, densely and coarsely areolate-punctate throughout with interspersed micropunctures where visible; intervals dull and blunt where visible. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 70:83:87:60:60. Lateral margin of mesonotum conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into blunt process; post-mesonotal tubercle absent. Propodeal spiracle strongly projected from lateral margin of mesosoma; post- spiracular area indistinguishable. Scutellar scale present, as wide or wider than surrounding sculputre; anterolateral carinae present, approximately twice as wide as scutelar scale; scabrous intervals vestigial on scutellar area. Propodeum simply convex, dorsal face indistinguishable from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 28:64:67. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate- punctate to punctate with dense interspersed coarse micropunctures where visible; sculpture sparser and micropunctures absent laterally and over integumental spots. T3–5 sculpture predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible; T6, except pygidial plate, virtually concealed by dense setation, densely foveolate-punctate where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 densely and coarsely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold present. S3–5 sculpture mostly concealed by dense setation, densely and finely foveolate- punctate with sparse micropunctures where visible; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose. MALE. Body length 14–16 mm. Head. Transversely subrectangular with posterolateral angles rounded in dorsal view; lateral margins of head convergent immediately behind eyes, but not contiguous with eye outline in dorsal view. Vertex width 0.8 × pronotal width. Eye almost circular. Ocelli small; OOD 5.3 × DLO, IOD 1.0 × DLO. Occipital carina distinct. Head surface sparsely and finely punctate; sculpture sparser and finer posterad. Gena ecarinate. Antennal scrobe concave to eye margin, with well-defined transverse dorsal scrobal carina. Clypeus concave laterally immediately below antennal insertion, conspicuously convex medially; densely and coarsely foveolate-punctate medially and along apical/ventral margin laterally; apical/ventral margin with a pair of medial short subacute free teeth medially. Scape bicarinate. Flagellomere 1 1.95 × pedicel length; flagellomere 2 2.6 × pedicel length. Mandible obliquely tridentate apically, medial tooth smaller than inner tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets well defined, sharply projected from anterior margin of pronotum, separated from well-defined humeral carina, anterolateral corners of pronotum not angulate. Anterior face of pronotum, with sparse fine punctures laterad with interspersed micropunctures, mostly unsculptured mediad; with medial longitudinal slightly concave smooth area. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures along inner and anterior margin. Dorsum of pronotum densely and coarsely foveolate-punctate to areolate-punctate with somewhat sharp intervals. Mesoscutum densely and finely foveolate-punctate, parapsis reduced to posterior half of mesoscutum, notaulus absent. Scutellum sloping throughout, somewhat depressed medially, without defined dorsal and posterior faces, densely and coarsely areolate-punctate to foveolate-punctate; anterior intervals somewhat aligned so as to form vestigial irregular longitudinal carina medially. Axilla produced posterolaterally as obliquely truncate projection, with inner margin slightly curved inward apicad in dorsal view; projection coarsely foveolate-punctate basad, unsculptured, smooth, shinning apicad. Metanotum slightly wider laterad, its surface obscured by dense setation. Propodeal dorsum convex, mostly concealed, densely areolate; sculpture of lateral face absent along most of anterior margin; dorsal face 215 indistinguishable from posterior face. Lateral face of pronotum sparsely and vestigially punctate with sparse interspersed micropunctures; mesopleura with conspicuous blunt projection on dorsal half; sculpture densely and coarsely areolate with interspersed micropunctures to simply micropunctate anterad. Metapleuron partially concealed by dense setation, micropunctate where visible, except basal third densely and coarsely areolate. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongated, roundly truncate apically; three submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; spurs finely serrate on margins. Metasoma. T1 0.5 × as wide as T2. T2 0.9 × as long as wide. Dorsal metasomal sculpture, except pygidial plate, partially concealed by dense setation, densely and finely punctate with sparse interspersed micropunctures where visible; sculpture sparser and less defined in apical segments; pygidial plate slightly broader than long, weakly defined by parallel carinae laterally; surface densely micropunctate throughout. S1 longitudinally elevated medially, terminating in slightly concave low longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate; sculpture sparser mediad; anteromedial crest-fold present, sternal pit absent. S3–7 sparsely and finely foveolate-punctate to punctate; S7 longer than broad, posterior margin projected medially into closely bidentate apex. Genitalia. Parapenial lobe slightly pronounced posteriorl, acute. Ratios of free length of paramere, cuspis and digitus, 83:58:18. Paramere virtually straight in dorsal view, upcurved posteriorly in lateral view, virtually asetose except for sparse scattered inconspicuous setae troughout, setae more evident on vetral surface. Cuspis slender, elongate, virtually straight and equally wide throughout in dorsal view, slightly upcurved posterad and somewhat subcaptitate on apex, with inconspicuous short setae on apex, scattered inconspicuous short setae elsewhere. Paracuspis poorly developed, sessile, lobe-like, wider than long with almost flate and asetose posterior margin. Digitus short, slightly curved inward in dorsal view, evenly upcurved and tapered posterad in lateral view, apex more abruptly curved and subcapitate, setose anterodorsally. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, with poorly defined lateral pocket on outer margin, apical distance between teeth 0.1 × length of valve, dense setae present along flat posterior margin and inconspicuous setae present at base of anterior tooth on outer surface, setae on posterior margin longer ventrad, posterior margin slightly sloping dorsad. Coloration and variations. FEMALE. Integument black to brownish black, except mandibles and antennal flagerllomeres partially reddish-brown, and T2 with four large orange integumental spots. Body setae predominantly golden varying in density, except the following areas with black setae varying in density: vertex, gena, malar space, and ventral surface of head; dorsum of propodeum, anterior half of mesoscutum, and propodeal dorsum medially; T1 medially, disc of T2 medially (except integumental spots), fringe of T2–3 sublaterally. MALE. Integument black. Body setae predominantly black varying in density, except the following areas with golden setae: antennal tubercles, lateral face of pronotum, mesosomal pleurae, metanotum, propodeum, and legs; T1, posterior two thirds of T2, fringe of T2–3 laterally, S1–4, and fringe of S2–4. Wings dark-brown throughout. Tibial spurs yellowish-white. Distribution. Brazil. Etymology. From the greek tetra “four” and trauma “wounded”, in reference to the four large orange spots on T2 of this species and the fact that this species has basically the same pattern as T. quasinotata but differs in structure. Material examined. Type material. Holotype, #f, BRAZIL, Espírito Santo, Linhares, IV.1973, P.C. Elias (MZSP). Paratypes. #1, BRAZIL: Pernambuco, Recife, Parque dos Dois Irmãos, 21.VII.2002, 08°00'33''S,34°56'31''W, STP Amarante e eq. [equipe]; 1#f, BRAZIL, Bahia, Padro, Cumuruxatiba, 20.III.1985, N.A.Menezez (MZSP); 1#f, BRAZIL, [Rio de Janeiro], Distrito Federal [Rio de Janeiro], IX.1928 (MZSP); 1#f, BRAZIL, Rio de Janeiro, Nova Iguaçu, Reserva Biológica do Tinguá, 9–12.III.2002, 22°34'43''S 43°26'08''W, S.T.P. Amarante e eq [equipe] (MZSP); 1#f, BRAZIL, Espírito Santo, Conc. [Conceição da] Barra, VII.1969, P.C.Elias (MZSP); 1#f, BRAZIL, Espírito Santo, Aracruz, Comboios, mata, VI.1993, M.M.A. de Oliveira (MZSP); 1#f, BRAZIL, Espírito Santo, Linhares, III.1973, P.C.Elias (MZSP); 1#f, BRAZIL, Ilha São Sebastião, 31.VIII.1963, H. Hzban (MZSP); 3#f, BRAZIL, São Paulo, Caraguatatuba, R. Flo [sic!], VII.1962, Exp.Dep.Zoo. [Expedição do Departamento de Zoologia] (MZSP); 1#f, BRAZIL, São Paulo, I.1938, E. schw. (MZSP). (CMNH). Allotypes. 2#m, BRAZIL, Espírito Santo, Linhares, X.1972, M. Alvarenga (CMNH). Remarks. Initially, T. quadrinotata appeared to be an easily recognizable, but variable species. It was originally known from females only and there seemed to be variation in the head setae color and mesosomal setal length. The male was also apparently easily recognized, based on its large body size, overlapping distribution, and parallel coloration with the female and other Atlantic Forest species. After careful examination of numerous suspected T. 216 quadrinotata males, however, we discovered that there were two species with identical coloration and remarkably similar external morphology, but distinct genitalia (Figs. 13A–F, 19A–F). Since there were two distinct species among the putative males of T. quadrinotata, we re-evaluated the species limits of the females. Those individuals with golden setae on the frons and uniformly short mesosomal setae were found to differ from the typical T. quadinotata by structural features as well, namely the occipital carina was swollen, the scutellar area was armed with distinct anterolateral transverse carinae, and the T1 was slightly broader and shorter. We therefore recognized these females as belonging to a distinct new species, named here as T. tetratrauma. The next step was to determine which of the males (short cuspis setae or long cuspis setae) belonged to each species. Both of these males and females overlap in distribution in the Atlantic Forest, so distribution couldn’t solve the association. Among the members of the species- group, only two males have short cuspis setae: T. incerta and one of these males. Therefore, the female of the short cuspis male should be similar to T. incerta. Traumatomutilla incerta and T. tetratrauma are the only species in this group with the occipital carina swollen and T1 comparatively short and broad. We therefore hypothesize that the male of T. tetratrauma is the one with short cuspis setae. The male with longer cuspis setae is associated with typical T. quadrinotata. Given the similarities between these two species, various collections will likely have numerous specimens of T. tetratrauma misidentified as T. quadrinotata. Perhaps when these specimens are re-evaluated, patterns about their distribution will become clearer. At this moment, it appears that T. tetratrauma is a Northern species, with most records focused in Espírito Santo state. Conversely, T. quadrinotata seems more widespread with many records from São Paulo and Santa Catarina states.
Traumatomutilla ursina (Gerstaecker, 1874) (Figs. 20A–C)
Mutilla ursina Gerstaecker, 1874: 74. holotype, #f, Brazil (ZMB), type examined. Ephuta (Traumatomutilla) ursina: André, 1902: p. 56. Traumatomutilla ursina: André, 1904: p. 40.
Diagnosis. FEMALE. In addition to the structural characters referenced in the species groups diagnosis, T. ursina can be defined by three unique features, body almost entirely covered by dense, long, brownish-black setae; lateral tubercles of mesonotum greatly reduced, as far apart as pronotal spiracles; and anterolateral corners of T2 virtually devoid of sculpture and setae, smooth, shinning. Description. FEMALE. Body length 16-17 mm. Head. Posterior margin virtually straight. Occipital carina evenly arched and equally wide throughout. Vertex width 0.8 × pronotal width. Eye almost circular, its length in frontal view 0.9 × distance from its ventral margin to mandibular condyle. Head densely and coarsely foveolate-punctate to areolate-punctate. Genal carina poorly defined, nearly vestigial. Mandible elongate, oblique, tapering slightly towards with small subapical tooth. Dorsal scrobal carina present, well-defined, disconnected from antennal tubercles and lateral scrobal carina. Antennal tubercle coarsely and irregularly rugose. Flagellomere 1 3.1 × pedicel length; flagellomere 2 2.0 × pedicel length. Mesosoma. Length 0.9 × width. Mesosomal dorsum mostly concealed by dense setation, densely and coarsely areolate-punctate with apparent scabrous intervals medially where visible. Anterior face of propodeum defined, short, slightly shorter than pronotal collar, coarsely striated longitudinally throughout with dense coarse punctures dorsomedially; dorsal face rounded into anterior face in lateral view. Humeral carina well- defined, projected dorsally, broadly separated from conspicuously projected angulate epaulet; anterolateral corners of pronotum sharply angulate in dorsal view. Pronotal spiracle slightly projected from lateral margin of pronotum, rounded, bulging. Lateral face of pronotum sparsely punctate with dense micropunctures except at smooth conspicuous subacute tubercle anterior to pronotal spiracle; mesopleuron densely micropunctate anteriorly, sparsely and vestigially punctate to foveolate-punctate to areolate-punctate ventrad on mesopleural ridge; dorsal half of mesopleural ridge simply micropunctate; metapleuron unsculptured smooth shinning on dorsal fourth, densely, coarsely and confusedly areolate on basal fourth, and with concealed by dense setation elsewhere. Lateral face of propodeum sparsely and coarsely foveolate-punctate posterad; virtually smooth and shinning anterad; with vestigially rugose intervals posterad. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 70:77:77:56:53. Lateral margin of mesonotum 217 conspicuously constricted anterior to propodeal spiracle, strongly diverging anterad, medially projected, into short, inconspicuous blunt process; with very small conspicuous tubercle posterior to lateral process. Propodeal spiracle strongly projected from lateral margin of mesosoma; post-spiracular area vestigial. Scutellar scale present, reduced, as narrow as surrounding sculpture; anterolateral carinae absent; scabrous intervals present on scutellar area. Propodeum slightly elongate, dorsal face longer than and poorly distinguished from posterior face. Metasoma. Ratios of width of T1, width of T2 and length of T2, 41:84:84. Disc of T2 mostly concealed by dense setation, densely and coarsely foveolate-punctate to punctate with dense interspersed micropunctures where visible; foveolations sparser and micropunctures absent laterally and over integumental spots; anterolateral corners of T2 virtually devoid of setae and sculpture, smooth, shinning. T3–6 sculpture, except pygidial plate, predominantly concealed by dense setation, densely and coarsely foveolate-punctate to simply punctate with interspersed micropunctures where visible. S1 sparsely, coarsely and confusedly foveolate-punctate, surface cuneiform, ending in short blunt longitudinal carina, slightly higher medially. S2 sparsely foveolate-punctate, sculpture conspicuously sparser posteromediad; anteromedial crest-fold vestigial. S3–4 sculpture mostly concealed by dense setation, densely and finely foveolate-punctate with sparse micropunctures where visible; S4 densely foveolate-punctate; S6 sparsely foveolate-punctate. Pygidial plate subpyriform, defined by lateral carinae at apical fourth of plate; surface irregularly rugose; interstice apparently granulose.
Coloration and variations. Integument black, except antennal flagellomeres and mandibles partially reddish-brown. Body setae predominantly and very densely black to brownish-black, except the following areas with silvery-white to silvery-golden setae varying in density: mesonotum, scutellar area, and propodeal dorsum laterally; mesopleuron, metapleuron, and lateral face of propodeum; T1 laterally, lateral felt lines of T2, lateral margins of T2, lateral areas of T2 disc, fringe of T2–4, fringe of T5 medially, S1–3, and fringe of S2–3. Distribution. Brazil. Material examined. (3#f) Type material. Holotype, #f, Brazil (ZMB); Additional material. BRAZIL, Minas Gerais: Corumbai [sic], 1#f, XI.1963 (DZUP); Araxá, 1#f, 5.V.1965, Elias, C. (DZUP); Matto Grosso [sic], Vaccaria, 1#f, XI.1922 (ZMB). Remarks. Traumatomutilla ursina is unusual, especially due to the conspicuously long brownish dorsal setae and the virtually asetose and unsculptured anterolateral corners of T2. Both these characters are unprecedented in South American Dasymutillini . Additionally, the head in frontal view seems to be wider than most Traumatomutilla species, especially related to how protuberant the eyes appear to be in frontal view. No structural or color variations were observed in the few specimens available for study which, save for the specimen from Araxá (DZUP) have no other indication of locality other than country and state. As mentioned before, T. ursina is one of four putative females that may be associated with either T. infernalis or T. pompiliformis.
Discussion
Species from the T. quadrinotata species-group were apparently defined by more consistent diagnostic color patterns than those seen in other groups, such as the T. indica and T. juvenilis species-groups. Apparently, the only species with conspicuous color variations are T. austera (mesosomal setal pattern) and T. quadripustulata (T2 spot color and mesosomal setal pattern). With the synonyms proposed herein, T. sancta and T. incerta also have color pattern variations in setal distribution and size of T2 integumental spots. Perhaps these relatively stable color forms and generally homogeneous structural characters are the main reason there are still four unassociated females and two unassociated males left in the group. On the other hand, the relatively small number of specimens examined may have prevented us from finding intermediate color forms or be able to confidently establish the minor structural differences as solid a basis for differentiating species. Therefore, even though it has been widely accepted that setae and color characters alone are inadequate for species delimitation (Williams et al. 2010), in this group, these characters seem to be more stable than in other groups. . Females of the T. quadrinotata species-group are unique their lateral mesonotal tubercles but, some of the included species bear traits that are apparently diagnostic for other species-groups.Some species have an indistinct medial longitudinal carina of the mesonotum, the main female charcters in the T. indica species-group. In the case of T. incerta and T. tetratrauma, the dorsolaterally slightly swollen occipital carina of the females is reminiscent of that 218 observed in females of the T. juvenilis species-group. In fact, the external morphology of the males of the T. quadrinotata species-group is nearly identical to that of the T. juvenilis species-group males, differing only by having an elongate hypopygium and lacking the setae filled pit on S2. The male genitalia of the T. quadrinotata species- group, however, is similar to those of the T. indica species-group males, except for the usually lobe-like paracuspis and convex posterior margin of the penis valve. Additionally, T. pompiliformis has the paracupsis slighlty node-like as in most species of the T. indica species-group, and there are several species within the T. indica species-group that have the posterior margin of the penis valve simply convex. These similarities between both species-groups are an indication that they more closely related to each other than to the remaining species-groups of Traumatomutilla and were further discussed by Bartholomay et al. (in prep).
Acknowledgements
We are grateful for the collection managers and curators that provided specimens for this study, including: Christine LeBeau (AMNH), Robert Zuparko (CASC), Andreas Köhler (CESC), John Rawlins (CMNH), Stephan Blank (DEI), Gabriel Melo (DZUP), James Pitts (EMUS), Mercedes Paris (MNCN), Agniéle Touret-Alby (MNHN), Felipe Vivallo (MNRJ), Orlando Silveira (MPEG), Kelli Ramos (MZSP), Fernando Silvestre (MuBio-UFGD), Gavin Broad (BMNH), Lynn Kimsey (UCDC), Fernando Silveira (UFMG), Robin Thomson (UMSP), Brian Harris (USNM), Michael Ohl, Viola Richter and Lukas Kirscher (ZMB), Lars Vilhelmsen (ZMUC), Denis Brothers (DJBC), Donald Manley (DGMC), Wouter Dekoninck (RBINS) and Karla Schneider (MLUH). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9).
References
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Figure legends.
Figures 1A–C. Traumatomutilla ameliae Casal, 1969, holotype, female, lines 5 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 2A–F. Traumatomutilla ameliae Casal, 1969, male, line 3 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved) ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale); lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 3A–G. A–C. Traumatomutilla austera (Gerstaecker, 1874), lectotype, female, lines 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla sigillata (Gerstaecker, 1874), holotype, female, lines 2 mm; D. Dorsal habitus; E. Lateral habitus; F. Type labels.
Figures 4A–B. Traumatomutilla chrysozona (Gerstaecker, 1874), female, lines 5 mm; A. Dorsal habitus; B. Lateral habitus.
Figures 5A–H. A. Traumatomutilla chrysozona (Gerstaecker, 1874), male, lectotype, line 2 mm; B. Vestigial coppery setae on female metasoma, lateral view; C. Type labels; D. Genitalia (halved), dorsal view; E. Genitalia (halved), ventral view; F. Genitalia (halved, penis valve removed), lateral/inner view; G. Cuspis (removed, not to scale), lateral/inner view; H. Penis valve (removed, not to scale), lateral/outer view.
Figures 6A–G. Traumatomutilla dives (André, 1906), lectotype, male, line 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia, (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view; G. Type labels.
Figures 7A–C. Traumatomutilla funebris (Gerstaecker, 1874), holotype, female, lines 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 8A–H. A–C. Traumatomutilla sodalicia (Kohl, 1882), holotype, female, lines 5 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–E. Traumatomutilla tabatinga Casal, 1969, holotype, female, line 2 mm; D. Dorsal habitus; E. Type labels; F–G. Traumatomutilla incerta (Spinola, 1841), female, most common color form; F. Dorsal habitus; G. Lateral habitus; H. Traumatomutilla incerta (Spinola, 1841), female, color form with reduced silvery- white setae on head and mesosoma, dorsal habitus.
Figures 9A–G. Traumatomutilla dentata (Smith, 1879), holotype, male, line 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia, (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view; G. Type labels.
Figures 10A–P. A–H. Traumatomutilla infernalis (Gerstaecker, 1874), holotype, male, lines 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Genitalia (halved), dorsal view; D. Genitalia, (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view; H. Type labels; I–P. Traumatomutilla floccosa (Gerstaecker, 1874), holotype, male, lines 5 mm; I. Dorsal habitus; J. Lateral habitus; K. Genitalia (halved), dorsal view; L. Genitalia, (halved), ventral view; M. Genitalia (halved, penis valve removed), lateral/inner view; N. Cuspis (removed, not to scale), lateral/inner view; O. Penis valve (removed, not to scale), lateral/outer view; P. Type labels.
221
Figures 11A–P. A–H. Traumatomutilla pompiliformis (Gerstaecker, 1874), lectotype, male, lines 5 mm; A. Dorsal habitus; B. Lateral habitus; C. Genitalia (halved), dorsal view; D. Genitalia, (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view; H. Type labels; I–P. Traumatomutilla serra (Cresson, 1902), holotype, male, lines 2 mm; I. Dorsal habitus; J. Lateral habitus; K. Genitalia (halved), dorsal view; L. Genitalia, (halved), ventral view; M. Genitalia (halved, penis valve removed), lateral/inner view; N. Cuspis (removed, not to scale), lateral/inner view; O. Penis valve (removed, not to scale), lateral/outer view; P. Type labels.
Figures 12A–C. Traumatomutilla quadrinotata (Klug, 1821), lectotype, female, lines 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels.
Figures 13A–F. Traumatomutilla quadrinotata (Klug, 1821), male, line 5 mm; A. Lateral habitus; B. Lateral habitus; C. Genitalia (halved), dorsal view; D. Genitalia, (halved), ventral view; E. Genitalia (halved, penis valve removed), lateral/inner view; F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 14A–C. Traumatomutilla quadripustulata (Klug, 1821), lectotype, female, lines 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus.
Figures 15A–D. A–B. Mutilla pruinosa (Smith, 1879), holotype, male, line 5 mm; A. Lateral habitus; B. Type labels; C–D. Traumatomutilla maraca (Cresson, 1902), holotype, male, line 3 mm; C. Lateral habitus; D. Type labels.
Figures 16A–G. A. Mutilla pruinosa (Smith, 1879), holotype, male, line 5 mm, dorsal habitus; B. Traumatomutilla maraca (Cresson, 1902), holotype, male, line 3 mm, dorsal habitus; C–G. Mutilla pruinosa (Smith, 1879), holotype, male genitalia; C. Dorsal view (halved); D. Ventral view (halved); E. Lateral view (halved, penis valve removed); F. Cuspis (removed, not to scale), lateral/inner view; G. Penis valve (removed, not to scale), lateral/outer view.
Figures 17A–F. A–C. Traumatomutilla sancta (Gerstaecker, 1874), holotype, female, lines 2 mm; A. Dorsal habitus; B. Lateral habitus; C. Type labels; D–F. Traumatomutilla solemnis (Cresson, 1902), holotype, female, lines 2 mm; D. Dorsal habiuts; E. Lateral habitus; F. Type labels.
Figures 18A–B. Traumatomutilla tetratrauma Bartholomay & Williams sp. nov., holotype, female, lines 2 mm; A. Dorsal habitus; B. Lateral habitus.
Figures 19A–F. Traumatomutilla tetratrauma Bartholomay & Williams sp. nov., male, line 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia, (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 20A–C. Traumatomutilla ursina (Gerstaecker, 1874), holotype, female, lines 2 mm; A. Dorsal habitus; B. Type labels; C. Lateral habitus. 222
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CAPÍTULO 4.11 – (formatado para submissão ao periódico ZOOTAXA)
Species groups of Traumatomutilla (Hymenoptera: Mutillidae): updates, keys, and redescriptions for the known males.
PEDRO R. BARTHOLOMAY1,*, KEVIN A. WILLIAMS2 ROBERTO A. CAMBRA3 & MARCIO L. OLIVEIRA1 1Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, Laboratório de Hymenoptera, Av. André Araújo, 2936, Manaus, Amazonas, Brazil. 2Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento CA 95832, USA. 3Museo de Invertebrados G. B. Fairchild, Estafeta Universitaria Apartado 00017, Universidad de Panamá, Panamá 0824, República de Panamá *Corresponding author, e-mail: [email protected]
Abstract
The species-groups of Traumatomutilla are updated according to their recent taxonomic treatments. All unplaced species known from males only are redescribed and illustrated. The male of Traumatomutilla virginalis (Gerstaecker, 1874) is described and illustrated.
Key words: Neotropical, Velvet ants, Sphaeropthalminae, Dasymutillini, Taxonomy
Introduction
Mutillidae, commonly known as velvet-ants, are a group of solitary aculeate wasps in which the females are always wingless, males are usually fully winged, and whose larvae act as ectoparasitoids of encapsulated immatures of other insects, especially other solitary Hymenoptera (Brothers, 2006; Williams, 2012). Traumatomutilla is one of the most diverse genera of Mutillidae in the Neotropical region, with over 120 species ranging from Panama to Argentina (Nonveiller, 1990, Bartholomay et al. in prep.). The extreme sexual dimorphism in Mutillidae has caused most species of this genus to be described based only on females or males with sexual associations depending on in situ observations, molecular data, or large numbers of specimens and distribution data (Bartholomay et al. 2019). The work of Williams et al. (2017) subdivided the species of Traumatomutilla known from females only into several species-groups based on exclusive characters or combinations of characters. This has allowed such a species rich genus to be addressed in smaller and more manageable units. As with the females, most males of Traumatomutilla were described prior to the establishment of the genus, based almost exclusively on variable color and setae characters, and without any mention of genitalia characters (Gerstaecker, 1874; Cresson, 1902; Mickel, 1952). A series of recent revisions of most species-groups of Traumatomutilla has addressed this issue and either associated and described males with their respective females or redescribed unassociated males and placed them within a revised species-group (Bartholomay et al. 2019, in prep.). Even though advances have been made regarding Traumatomutilla males, numerous species are still unapproachable by non-specialists (PRB & KAW pers. obs.). In this current study we address this issue by redescribing and fully illustrating the remaining species of Traumatomutilla known from males only. We then add them to a functional key along with the males treated in the recent species-groups reviews. Finally, we take this opportunity to update the species-group construct of Williams et al. (2017), considering the numerous advances made recently (Bartholomay et al. 2019, in prep).
Materials and terminology
During this study approximatelly 280 specimens have been examined and the following codens are used for institutions housing the material discussed:
AMNH - American Museum of Natural History, New York, New York, USA. 228
BMNH - British Museum of Natural History, London, England. CESC - Coleção Entomológica da Universidade de Santa Cruz do Sul, Santa Cruz do Sul, Rio Grande do Sul, Brazil. CISC - Essig Museum of Entomology, Department of Entomological Sciences, University of California, Berkeley, California, USA CMNH - Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA CSCA - California State Collection of Arthropods, Sacramento, California, USA CUIC - Cornell University Insect Collection, Department of Entomology, Ithaca, New York, USA CZMA - Coleção Zoológica do Maranhão, Caxias, Maranhão, Brazil. DEI - Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany DZUP - Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. EMUS - Department of Biology Insect Collection, Utah State University, Logan, Utah, USA. FSCA - Florida State Collection of Arthropods, Gainesville, Florida, USA INPA - Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil. MNHN - Muséum Nationale d’Histoire Naturelle, Paris, France MLUH - Martin Luther Universität Halle-Wittenberg, Halle, Germany MPEG - Museu Paraense Emílio Goeldi, Belém, Pará, Brazil PMAE - Royal Alberta Museum, Edmonton, Alberta, Canada. SEMC - Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA. UMSP - University of Minnesota Insect Collection, St. Paul, Minnesota, USA. USNM - United States National Museum of Natural History, Simthsonian Institution, Washington D.C., USA. UFES - Universidade Federal do Espírito Santo, Vitória, Espírito Santo, Brazil. UCDC - The Bohart Museum of Entomology, Universisty of California, Davis, California, USA ZMB - Museum für Naturkunde Berlin, Berlin, Germany ZMUC - Zoological Museum University of Copenhagen, Copenhagen, Denmark
General morphological terminology, definitions, abbreviations, and measurements followed that of Bartholomay et al. (2019). In the material examined section abbreviations, acronyms and additional or corrected data by the authors are given in brackets. The total number of males and females examined is provided in brackets at the beginning of the material examined section. The types of T. aethiops (Gerstaecker, 1874), T. mesoleuca (Gerstaecker, 1874), and T. vulnerata (Gersteacker, 1874), although selected and labeled lectrotypes by Mickel, were never published as such by the author. In this paper we maintain and officially designate those specimens selected by Mickel as the lectotypes for their respective species. Likewise, many of the species described by Cresson (1902) have more than one specimen in their type series, which is why Cresson (1916) provided a list of specimens with their respective catalog number in his collection, designating them as “types” and stating that those were the specimens used in the descriptions of all his previous studies. Though these specimens were not refered to nor labeled as lectotypes by the author, we consider them as the lectotypes of his species and Cresson (1916) as the paper that established them as such.
Taxonomy
The ungrouped males treated herein likely be associated with females of the T. trochanterata, T. inermis, T. integella, T. tabapaua, T. diabolica, or T. pilkingtoni species-groups. Such associations have not taken place yet because either the females of the species-group have not been revised yet (T. trochanterata and T. inermis species-groups) or the data examined at the time of their revisions was inconclusive regarding which males could potentially be associated (Bartholomay et al. 2018, in prep.).
Key to the known males of Traumatomutilla:
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1. Axilla acute (Fig. 1D) and/or mesopleuron tuberculate on dorsal half (Figs. 1A, 1E, 1G), if axilla appears obliquely truncate then cuspis laterally compressed, conspicuously wider medially in lateral view (“banana-shaped” - see couplet 5) ... 2 Axilla truncate (Figs. 1B, F, H) AND mesopleuron unarmed (Fig. 1C) … 10
2. Mesopleuron unarmed on dorsal half (Fig. 1C) … 3 Mesopleuron tuberculate on dorsal half (Figs. 1A, 1E, 1G) … 6
3. Cuspis sparsely setose, at most with conspicuous but sparse long setae posteriorly (Fig. 1I); if cuspis more densely setose, then always club-like and laterally compressed on posterior half (Figs. 1K–L); free length of cuspis ~0.9 × free paramere length and more than 2 × free digitus length (Fig. J) … T. indica species-group in part (see Bartholomay et al. in prep.) Cuspis densely and conspicuously setose throughout or at least along ventral surface; free length of cuspis ~0.6 × free paramere length and less than 2 × free digitus length (Fig. 1M, 5E–H, 14B–E) … 4
4. Apex of meso and metafemora rounded, at most slightly blunt; cuspis abruptly upcurved medially in lateral view, with long strongly sinuous setae on ventral margin (Fig. 1M) … T. gemella species-group (see Bartholomay et al. in prep.) Apex of meso and metafemora strongly truncate, almost sulcate; cuspis straight, at most slightly curved inward in dorsal view (Figs. 5E, 14B) … 5
5. Sculpture of mesonotum with intervals somewhat longitudinally aligned, parapsis more evident than notaulus (Fig. 1N); paramere more densely setose along outer and inner margins (Figs. 5E–F); cuspis laterally compressed, tapering apicad in lateral view, with dense simple setae throughout (Figs. 5G–H); penis valve with a pair of short acute posteroventral teeth, outer surface lacking lateral pocket at base of anterior tooth (Fig. 5I) … T. virginalis (Gerstaecker, 1874) Sculpture of mesonotum without aligned intervals, notaulus more evident than parapsis (Fig. 1O); paramere more densely setose on posterior half of ventral surface (Figs. 14B–D); cuspis laterally compressed, conspicuously wider medially in lateral view (“banana-shaped”), with conspicuous spine-like setae only apically on ventral margin, simply setose elsewhere (Fig. 14E); posteroventral teeth of penis valve longer, posteriormost tooth acute, anterior tooth truncate, with well-developed lateral pocket on outer surface (Fig. 14F) … T. rorida (Gerstaecker, 1874)
6. Frons with bifurcated tooth-like projection below median ocellus (Fig. 2A); T7 with elevated pygidial plate (Fig. B) … T. bifurca species-group (see Bartholomay et al. 2019b) Frons unarmed; T7 virtually flat, at most convex … 7
7. Hypopygium elongate, clearly longer than broad, defined laterally by parallel carinae (Fig. 2C); axillar projection usually acute in dorsal view (Fig. 1D) or oblique in posterior view … T. indica species-group in part (see Bartholomay et al. in prep.) Hypopygium usually as long as broad or broader basally than apically (Fig. 2D), if elongate, then never defined by lateral carinae; axillar projection always truncate (Figs. 1B, 1F) … 8
8. S2 without seta-filled pit … T. quadrinotata species-group (see Bartholomay et al. in prep.) S2 having seta-filled pit … 9
9. West of the Andes; paramere with medial tuft of setae ventrally and short inconspicuous scattered setae elsewhere (Fig. 2E); penis valve with teeth widely spaced (Fig. 2G) … T. vitelligera (Gerstaecker, 1874) East of the Andes; paramere with long setae throughout ventral surface (Fig. 2G); penis valve with teeth closely spaced (Fig. 2H) … T. juvenilis species-group (see Bartholomay et al. in prep.)
10. Clypeus unidentate on apical/ventral margin (Fig. 3A) … T. pomba (Cresson, 1902) 230
Clypeus with two closely spaced teeth or simply truncate on apical/ventral margin (Figs. 3B–C) … 11
11. T2-6 clothed with coppery-golden setae troughout (Fig. 3A) … T. americana (Linnaeus, 1758) Metasoma clothed predominantly with black or silvery-white setae (e.g. Fig. 7A) … 12
12. Cuspis parallel in dorsal view AND elongate (Figs. 3E–I, 7B–C) ... 13 Cuspis converging apicad in dorsal view OR short (e.g. Figs 8A, 15A) … 16
13. Cuspis in lateral view gently upcurving throughout length, with setae evenly distributed along ventral margin (Figs. E–G) … undescribed morphospecies: we have observed 12 apparently new morphospecies that key out here. Cuspis in lateral view either straight with extreme apex upcurved or sigmoidal, curving up near midlength and straightening out toward apex (Figs. 3I, 7E) … 14
14. S2 without setae filled pit; cuspis in lateral view abruptly narrowed in apical half, and conspicuously setose throughout (Fig. 3I) … T. ocellaris (Klug, 1821) S2 with setae filled pit; cuspis in lateral view equally wide throughout, at most slightly widened apically; cuspis setae greatly reduced, present mostly apically (Figs. 3H, 7E) … 15
15. Metasoma predominantly reddish (Fig. 3J); cuspis at most slightly widened apically (Fig. 3H) … T. quadrum (Klug, 1821) Metasoma entirely black to brownish-black (Fig. 7E, 12B); cuspis with conspicuous “finger-like” projection apicodorsally in lateral view (Figs. D–E) … T. barra (Cresson, 1902)
16 Metasoma either entirely orange/reddish or at least T2 marked with orange/reddish integumental spots (Figs. 3J, 18A, 19F) … 17 Metasoma integument all black to brownish-black (Figs. 12 A–F, 19A–E) … 18
17. T2 disc with a pair of dark reddish to orange integumental spots (Figs. 18A, 19F) and cuspis tapering and converging apicad in dorsal view (Figs. B–C) … T. vulnerata (Gerstaecker, 1874) Not matching the above characters, either metasoma entirely orange/reddish (similar to Fig. 3J) or genitalia differing … undescribed morphospecies: we have observed four apparently new morphospecies that key out here.
18. T2 disc finely and sparsely punctate, intervals smooth and shining, generally greater than three times wider than surrounding punctures; lacking micropunctures (Figs. 3K, 17A, 19A) … 19 T2 disc more densely punctate (e.g. Fig. 9A), if intervals smooth and shinning, then never more than two times wider than surrounding sculpture; micropunctures present … 20
19. T1 shape strongly petiolate, subnodose, sides slightly constricted before base of T2 in dorsal view (Fig. 3K); paramere virtually straight, slightly and evenly tapering apicad in dorsal view (Fig. 17B); cuspis equally wide throughout and virtually straight in lateral view (Fig. 17E) … T. tenuis (Smith, 1879) n. comb. T1 shape subpetiolate, sides parallel throughout in dorsal view (similar to Fig. 12A); genitalia differing … undescribed morphospecies: we have observed four apparently new morphospecies that key out here, including Santarém male in sabara’s type series
20. Cuspis subcylindrical, tapering and converging apicad in dorsal view (Figs. 6A, 9A, 10A, 11A, 15A, 18A) … 21 Cuspis broadly expanded and/or straight in dorsal view (Figs. 8A, 16A) … 25
21. Lateral face of propodeum and mesopleuron with conspicuous patches of dense silvery-golden setae (Fig. 6A) and pronotal setae completely black (Fig. 15A) … 22 231
Lateral face of propodeum and mesopleuron usually clothed with black setae only (Figs. 9A, 10A, 18A) if silvery setae present, then pronotum densely clothed with silvery-white setae (Fig. 11A) … 23
22. Fringes of T2–3 predominantly clothed with black setae, at most with scattered silvery-white setae laterally (Figs. 6A, 12A); pygidial plate irregularly and confusedly rugose … T. aethiops (Gerstaecker, 1874) Fringes of T2–3 completely clothed with silvery-white setae (Figs. 15A, 19C); pygidial plate with sparse coarse longitudinal rugae … T. soricina (Gerstaecker, 1874)
23. Pronotal dorsum clothed entirely with silvery-white setae (Figs. 11A, 12B); subapical tooth of penis valve broad, blunt, rounded, with conspicuous external pocket (Fig. 11F) … T. picada (Cresson, 1902) Pronotal dorsum clothed with black setae (Figs. 9A, 10A, 12E, 12D); subapical tooth narrow, acute to subacute, external pocket vestigial (Figs. 9F, 10F) … 24
24. Fringe of T3 entirely clothed with black setae, at most with silvery-white to silvery-golden setae laterally (Fig. 12D); anterior margin of scutellum with slightly projected vestigial longitudinal carina medially … T. mesoleuca (Gerstaecker, 1874) Fringe T3 entirely clothed with silvery-white to silvery-golden setae (Fig. 12E); anterior margin of scutellum straight, not projected medially … T. orbana (Cresson, 1902)
25. Paramere slender, evenly and slightly narrower apicad in dorsal view (Fig. 8A); cuspis with conspicuous lobe- like expansion basally in lateral view (Fig. 8E); penis valve with subapical tooth slightly longer than apical tooth, subapical, externolateral pocket reduced (Fig. 8F) … T. bartica Mickel, 1952 Paramere stout, subparallel medially, slightly and evenly narrower apicad in apical third (Figs. 16B–C); cuspis equally wide wide throughout in lateral view, dorsoventrally flattened (Figs. 16B–E); penis valve with subapical tooth much longer than apical tooth, broadly rounded, externolateral pocket well-developed (Fig. 16F) … T. taboca (Cresson, 1902)
Traumatomutilla bellica species-group
Diagnosis. FEMALES. Females of this species group can be defined by a unique combination of characters: vertex unarmed, apex of middle and hind femora truncate, scutellar scale lacking, and T2 with only two integumental spots. Additionally, the genal carina is weak, the mesosoma is not elongate, and the pygidial plate is broadly ovate. MALES. The males can be distinguished by the poorly defined mandibular teeth, intermandibular distance more than four times the length of the malar space, intervals of mesonotal sculpture longitudinally aligned, apex of meso and metafemora strongly truncate, and paramere densely setose throughout. MALES. Males of this species group can be defined by a unique combination of characters: mesopleuron simply swollen on dorsal half, axillar projections acute, intervals of mesonotal sculpture somewhat longitudinally aligned, apex of meso and metafemora strongly truncate, intermandibular distance more than 4 × the length of malar space. Included taxa. Traumatomutilla virginalis (Gerstaecker, 1874) and T. bellica (Cresson,1902). Distribution. Brazil and Paraguay. Remarks. Williams et al. (2017) mentioned the similarities between the females of this group with those of the T. inermis species-group. The hitherto unknown males of the T. inermis species-group will likely share some structural similarities with that of T. virginalis. Males of the T. bellica species-group have a wider intermandibular distance than other species-groups. They also have long and somewhat dense setae on the inner and outer surfaces of the parameres; this is rare in Traumatomutilla, but occurs in some unassociated males, such as T. bartica and T. taboca.
Traumatomutilla virginalis (Gerstaecker, 1874) (Figs. 1N, 4A–B, 5A–I)
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Mutilla virginalis Gerstaecker, 1874: 67 , holotype, #f, Brazil (ZMB). Ephuta (Traumatomutilla) virginalis: André, 1902, p. 56 (new combination). Traumatomutilla virginalis: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. See Bartholomay et al. (2019b). MALE. Poorly defined mandibular teeth, intermandibular distance more than four times the length of malar space, intervals of mesonotal sculpture longitudinally aligned, apex of meso and metafemora strongly truncate, and paramere densely setose throughout. Description. FEMALE. See Bartholomay et al. (2019b). MALE (hitherto unknown). Body length 12.5 mm. Head. Narrow rounded. Vertex width 0.9 × pronotal width. Eye almost circular. Ocelli small; OOD 4.4 × DLO, IOD 1.4 × DLO. Occipital carina sharp lamellate. Frons, vertex, and gena densely coarsely foveolate-areolatate, some intervals scabrous. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate medially; with shallow smooth transverse furrow on anteroventral margin. Scape bicarinate, dorsal carina largely obliterated. Flagellomere 1 2.2 × pedicel length; flagellomere 2 2.7 × pedicel length. Mandible elongate curved, oblique apically with any apparent teeth obliterated and undifferentiated; lacking dorsal or ventral projections. Mesosoma. Epaulets slightly projected from anterior margin of pronotum, connected with humeral carina. Anterior face of pronotum coarsely areolate dorsally, with scattered micropunctures ventrally. Tegula convex, mostly glabrous and impunctate except for dense coarse punctures on along inner and anterior margins. Mesoscutum densely and coarsely foveolate-areolate with many scabrous intervals, notaulus obliterated; parapsis present, reduced to posterior half of mesoscutum; longitudinal carina present medially. Mesoscutellum convex, areolate with scabrous intervals; dorsal and posterior faces poorly distinguished; intervals aligned forming longitudinal carina throughout mesoscutellum. Axilla with coarsely and densely foveolate-punctate dorsal surface and dorsally smooth posterolateral oblique incurved dentate projection. Metanotum surface obscured by dense setae. Propodeal dorsum convex to virtually flat basad, densely areolate; lateral face virtually unsculptured along anterior margin; dorsal face rounded into and poorly distinguished from posterior face. Lateral face of pronotum sparsely punctate with dense interspersed micropunctures; mesopleuron convex without tubercle on dorsal half; mesopleural sculpture densely areolate-punctate to foveolate-punctate with interspersed micropunctures. Metapleuron densely micropunctate throughout, except for indistinct areolations on basal fourth and indistinct rugosities on dorsal fourth. Wings. Forewing with elongate sclerotized pterostigma; marginal cell elongate, rounded-acute apically; two submarginal cells. Legs. Simply setose, no strong spines discernible dorsally; outer posterior margin of meso- and metafemur with sharp truncate furrow; spurs finely serrate on margins. Metasoma. T1 0.45 × as wide as T2. T2 length 0.9 × its width. Dorsal metasomal sculpture densely punctate with interspersed micropunctures where visible, except T2 disc with smooth intervals. Pygidial plate with indistinct rugae, intervals smooth, weakly defined by parallel carinae apicolaterally. S1 longitudinally elevated medially, terminating in slightly concave and blunt longitudinal carina. S2 sparsely and finely foveolate-punctate to punctate, without micropunctures; without seta-filled pit. S3–6 densely foveolate-punctate with sparse interspersed micropunctures; S7 ovate, longer than broad; foveolate-punctate, sculpture sparser medially; posterior margin projected medially into a rounded, apparently unidentate, tooth-like structure on posterior margin. Genitalia. Parapenial lobe not at all pronounced posteriorly, acute. Ratios of free length of paramere, cuspis and digitus, 101:51:13; paramere virtually straight in dorsal view, upcurved posteriorly in lateral view; sparsely setose on dorsal and ventral surfaces, densely setose on inner and outer surfaces; cuspis stout, equally wide throughout in dorsal view, laterally compressed, conspicuously broader at anterior third in lateral view, densely setose throughout; paracuspis well-developed, nearly sessile, node-like, longer than wide, with subtriangulate posterior margin, with strong conspicuous setae posterodorsally in lateral view, setae as long as paracuspis; digitus short, virtually straight in dorsal view and slightly upcurved in lateral view, sparsely setose anterodorsally, slightly narrower posterad in lateral view; penis valve strongly concave on inner surface, with well- defined pair of short teeth posteroventrally; both teeth acute, anterior tooth without lateral pocket on outer surface; apical distance between teeth 0.1 × length of valve; dense setae present along posterior margin and inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. FEMALE. See Bartholomay et al. (2019b). MALE. Integument black. Body setae predominantly black varying in density, except following areas with grayish-yellow setae varying in density and often interspersed with black setae: frons, legs, lateral portions of mesosoma, metanotum, dorsum of propodeum, T1, T2 233 anterolaterally, fringes of T2–3 laterally, and S1–3 partly or entirely. Tibial spurs whitish. Wings dark-brown with poorly discernible dark brown veins. Distribution. Brazil and Paraguay. Material examined. (1m#) PARAGUAY, San Pedro, Rio Ypane, Cororo, 1m#, XII.1983, M.A. Fritz (AMNH). Remarks. The sex association of T. virginalis was based firstly on a process of elimination, since most species known from males and undescribed male morphospecies were associated with their repsective female or species-group. Secondly there are clear conspicuous morphological characters that appears in both sexes of T. virginalis, namely the somewhat longitudinally aligned intervals of the mesonotal sculpture, the wide intermandibular distance, and the strongly truncate femoral apices. Finally the association of the male described herein to T. virginalis rather than T. bellica (Cresson, 1902) was based on the known distribution of both species, since T. bellica is only known from Chapada dos Guimarães, Mato Grosso, Brazil, and T. virginalis has also been recorded in Paraguay, including the Cororo area where the males were collected.
Unplaced species:
Traumatomutilla aethiops (Gerstaecker, 1874) (Figs. 6A–F, 12A, 12H)
Mutilla affinis Burmeister, 1854: 24, lectotype [designated here], #m, Brazil, [Rio de Janeiro], Nov. Frib. [Nova Friburgo] (MLUH), examined. Mutilla aethiops Gerstaecker, 1874: 317 (new name). Ephuta (Traumatomutilla) aethiops: André, 1902, p. 54 (new combination). Traumatomutilla aethiops: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections transversely truncate; S2 with small posteromedian pit filled with setae; free length of cuspis approximately 0.7 × free length of paramere; cuspis convergent posterad in dorsal view; mesopleuron, metapleuron and lateral face of propodeum with inconspicuous areas of appressed silvery-golden setae. Description. FEMALE. Unknown. MALE. Body length 11 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly dark fuscous, except hyaline brown basal third, veins blackish. Setae entirely black except with whitish setae on coxae, sparsely on meso and metatibia, sparsely meso and metabasitarsomeri, predominantly on propodeal dorsum, T1, T2 basally, fringe of T2–4 laterally (with scattered whitish setae medially), S1, S2 medially, and fringe of S2–3. Head. Transversely subrectangular, virtually straight on posterior margin. Vertex width 0.95 × pronotal width. Eye almost circular. Ocelli small; OOD 5.1 × DLO, IOD 1.5 × DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus depressed laterally below antennae insertion, convex medially; densely and coarsely punctate; with closely spaced pair of short subsessile tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 1.7 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, greatly reduced, restricted to posterior third of mesoscutum. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dense foveolate-punctate dorsally, except narrow posterior margin impunctate; projection narrowly separated from scutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum virtually equally wide throughout, its surface obscured by dense setae. Propodeum convex/gibbose, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations along posterior margin and scatterd punctures elsewhere. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of 234 mesopleuron at most slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, vestigially sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly narrow, rounded; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.9 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly longitudinally rugose throughout, lateral carinae obscure, short. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 densely and coarsely foveolate-punctate, with small shallow setae filled pit posteromedially; foveolations sparser and larger posterad. S3–6 densely and coarsely foveolate-punctate. S7 elongate sub-trapezoidal, densely foveolate-punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 97:66:31. Paramere slightly curved inward posterad, otherwise virtually straight throughout, with sparse short inconspicuous setae throughout. Cuspis tapering and strongly curved inward posterad in dorsal view, virtually equally wide throughout in lateral view, posterior third slightly laterally compressed, with strong dense setaet throughout ventral surface, setae conspicuously shorter anterad. Paracuspis well-developed, node-like, almost petiolate in shape, slightly posterior margin slightly subtriangulate, densely setose posterodorsally, setae as long as or shorter than paracuspis length. Digitus incurved in dorsal view and upcurved in lateral view, slightly tapering posterad in dorsal view, slightly narrower medially in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short teeth posteroventrally; posteriormost tooth acute, anterior tooth broadly sounded, blunt, slightly curved anteroventrally, with well-defined lateral pocket on outer surface; apical distance between teeth 0.1 × length of valve; strong setae present along posterior margin posterior margin; setae longer ventrad, posterior margin slightly sloping dorsad, inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish black. Body setae predominantly black varying in density, except the following areas with silvery-white to silvery-golden setae varying in density: mesopleuron posteroventrally, metapleuron, lateral face of propodeum, propodeal dorsum, coxae, femora, tibiae partially, basitarsomeri partially, T1, anterior fourth of T2, fringe of T2 laterally, S1, and S2–4 partially. Tibial spurs yellowish-white, wings dark brown infuscated, with strong violaceous and blueish reflections, veins poorly discernible. Distribution. Brazil. Material examined. (17m#) Type material. Lectotype: Mutilla aethiops, m#, BRAZIL, [Rio de Janeiro], Nov. Frib. [Nova Friburgo] (MLUH). Additional material. BRAZIL: Espírito Santo: Baixo Guandu, 1#m, 23–30.IX.1970, Elias, C.; Elias, CT. (DZUP); Faz. [Fazenda] Usina Paineiras, P1 [sic], 20°56'29''S 41°03'06''O, Itapemirim, 1#m,19– 26.XI.2010, M.T. Tavares e eq. [equipe] (UFES); Santa Teresa, 1#m, 19.II.1964, Elias, C., Elias, C.T. (DZUP); 1#m, 1–3.II.1968, Elias, C., Elias, C.T. (DZUP); 1#m, 16–28.VIII.1967, Elias, C., Elias, C.T. (DZUP); Minas Gerais: Caratinga, Faz. [Fazenda] Montes Claros, 1#m, 30–31.XII.1994, Melo, G.A.R. (DZUP); Serra do Caraça, S. [Santa] Barbara, 1600m [above sea level], 2#m, II.1969, F.M. Oliveira (EMUS); Rio de Janeiro: 1#m, 25.I.1919, R. Fischer (DEI); Mangaratiba, 2m#, Aug38–Feb39[sic!], YelSevFerv [sic!], M.E.S. Brazil, R.C. Shannon (EMUS); Represa Rio Grande, Guanabara, 2#m, XII.1969, M. Alvarenga (EMUS); Paraná: Quatro Barros, Morro Anhangava, 1#m, 10.I.2009, Ramos, K.S. (DZUP); Rio Capivari: 1#m, 22.I.2004, Melo, G.A.R. (DZUP); 1#m, 22.I.2004, Melo, G.A.R. (DZUP). Remarks. Burmeister (1854) described Mutilla affinis (Burmeister, 1854) based on a couple collected at Nova Friburgo, Rio de Janeiro, Brazil. Later, Gerstaecker (1874) determined that there was no evidence to associate the specimens in question and renamed the male as Mutilla aethiops. There is strong evidence that this species is in fact the male of T. inermis affinis, which is likely a junior synonym of T. inermis inermis (Klug, 1821). Most notably, the pattern of appressed silvery-golden setae on the mesopleuron, metapleuron, and lateral face of propodeum is typical of the Atlantic forest region and has repeatedly supplemented sex associations of different species in that region (PRB & KAW pers. obs.). The posteriorly convergent cuspis of T. aethiops is a character that also occurs in T. mesoleuca, T. orbana, T. picada, T. soricina, and T. vulnerata (refered to as the T. aethiops subgroup from now on). If the conspecificity of T. aethiops and T. inermis affinis is formalized, each of these species might be associated with females of the T. inermis species-group in the future.
235
Traumatomutilla barra (Cresson, 1902) (Figs. 7A–F, 12B, 12G)
Mutilla barra Cresson, 1902: 75, holotype [by monotypy], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães], Sept. [September] (CMNH), examined. Ephuta (Traumatomutilla) barra: André, 1902, p. 56 (new combination). Traumatomutilla barra: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections transversely truncate; cuspis slender, long, free length 0.9 × free paramer length, in lateral view with apex abruptly curved dorsad. Description. FEMALE. Unknown. MALE. Body length 8.5 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly light brown, veins dark brown. Setae entirely black except with whitish setae on head entirely; pronotum; mesoscutellum; metanotum; mesopleuron; metapleuron; propodeal dorsum; legs; T1; T2 basally, laterally, and apicolaterally; T3–4 entirely; T5–6 laterally; and S1–6. Head. Rounded, postero-medially swollen. Vertex width 1.1 × pronotal width. Eye almost circular. Ocelli small; OOD 5.1 × DLO, IOD 1.6 × DLO. Occipital carina distinct. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 1.8 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible oblique, apical teeth obliterated, apparently tridentate; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, restricted to posterior half of mesoscutum; with medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-areolate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex, areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly thickened, sub-truncate; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.5 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly transversely rugose throughout, lateral carinae strongest posteriorly, converging anteriorly. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, with seta-filled pit posteromedially. S3–6 foveolate-punctate. S7 elongate sub-trapezoidal, foveolate-punctate; apical margin projected medially into a single closely bidentate projection. Genitalia.: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 88:79:17. Paramere virtually straight throughout, with sparse strong setae on ventral surface medially and inconspicuous short setae elsewhere. Cuspis virtually straight throughout in dorsal view, posteriormost tip abruptly tapered in dorsal view and abruptly upcurved in lateral view, with sparse short setae slightly denser and more defined posteriorly. Paracuspis well-developed, node-like, not sessile, slightly longer than broad, posterior margin rounded, densely setose posterodorsally, setae as long as or shorter than paracuspis length, more defined than cuspis setae. Digitus virtually straight in dorsal view, slightly upcurved posteriorly in lateral view, tapering posterad in in lateral view. Penis valve strongly concave on inner surface, conspicuously narrower posteriorly than medially, with well-defined pair of short acute teeth posteroventrally, without 236 well-defined lateral pocket on outer surface; apical distance between teeth 0.15 × length of valve; strong setae present along convex posterior margin, inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. FEMALE. Unknwon. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: mesoscutum, axillar projections mesoscutellum partially, T2 (except anterior third and fringe laterally), fringe of T5–6 predominantly, T7, and S6–7. Tibial spurs yellowishw-white. Wings hyaline brown, slightly darkened apicad, without conspicuous reflections, veins easily discernible throughout. Distribution. Brazil. Bolivia and Paraguay Material examined. (10m#) Type material. Lectotype: Mutilla barra, f#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães], Sept. [September] (CMNH). Additional material. BRAZIL: Mato Grosso, Chapada dos Guimarães, 2m#, 13–18. XI.2013, Melo, G.A.R., Luz, D.R., Williams K.A. (DZUP); Mato Grosso do Sul, Baytaporã, 3m#, 03– 12.XII.2012, Salvaris, M. & Lampert, S. (DZUP); São Paulo, Fazenda Cambuy/Matão, 1m#, 27.XI.2005, P. Yamamoto (FSCA); BOLIVIA, Santa Cruz, 4km N [kilometers north of] Pedro Lorenzo, 27.II.1999, Irwin & Parker (EMUS); PARAGUAY: Cordillera, San Bernardino, 1m#, K. Fiebrig S.V. (ZMB); San Pedro: Rio Ypane, Cororo, 1m#, XI.1979, Fritz (AMNH); 1m#, II.1979, Fritz (AMNH). Remarks. The external morphology of T. barra is quite unremarkable and, apart from the conspicuously upcurved apex of the cuspis and the penis valve, the genitalia are reminiscent of species from the T. indica and T. quadrinotata species-group in length. At the moment, no putative females can be suggested as the counterpart for this male. Males with this particular color pattern (predominantly covered with silvery-white setae), have been associated with females that have extensive silvery-white markings and usually yellowish integumental spots on T2 (PRB & KAW pers. obs.).
Traumatomutilla bartica Mickel, 1952 (Figs. 8A–F, 12C)
Traumatomutilla bartica Mickel, 1952: 132, holotype, #m, Guiana, Bartica, Bartica district, Wm. [William] Beebe (UMSP).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate. apex of meso and metafemora truncate; cuspis very short, 0.5 × the length of paramere, with strong spine-like setae on ventral surface, inner margin expanded on basal third in ventral view. Description. FEMALE. Unknown. MALE (Hitherto undescribed). Body length 10 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly light-brown fuscous, slightly lighter basally, veins blackish. Setae entirely silvery-white except with black to brownish-black setae on ocelar area partially, pronotal dorsum predominantly, mesoscutum, mesoscutellum, axillar projections, posterior two thirds of T2, fringe of T2–6 mediall (interspersed with white on T3–5 and predominant in T6), T7, fringe of S5–6, and S6–7. Transversely subrectangular in dorsal view, slightly sinuous on posterior margin. Vertex width 0.9 × pronotal width. Eye almost circular. Ocelli small; OOD 4.3 × DLO, IOD virtually equal to DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate; sculpture sparser on vertex medially. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and finely punctate along anteroventral margin; with closely spaced pair of short blunt tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.3 × pedicel length; flagellomere 2 2.8 × pedicel length. Mandible vestigially bidentate, inner tooth smaller than outer tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum virtually entirely smooth, asetose, shinning, with longitudinal smooth furrow medially; sharply angulate around edges. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, restricted to posterior third of mesoscutum. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, densely foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dense foveolate-punctate, except at small basal area and posterior third impunctate; projection conspicuously separated from scutellum. Area 237 between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbose, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations to unsculptured and smooth anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron at most slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, vestigially sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly thickened, sub-truncate; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.7 × T2 width. T2 length 0.8 × T2 width. Dorsal metasomal sculpture sparsely and finely foveolate-punctate. T7 coarsely, confusedly and irregularly rugose to granulose throughout, lateral carinae obscure, short. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 sparsely and finely foveolate-punctate; foveolations sparser and smaller anterad. S3–6 sparsely and finely foveolate-punctate. S7 slightly elongate sub-rectangular, sparsely foveolate-punctate; apical margin projected medially into a single blunt projection. Genitalia.: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 75:38:18. Paramere virtually straight throughout, with sparse thin setae on throughout. Cuspis virtually straight and tapered posterad in dorsal view, slightly upcurved and basally swollen in lateral view, in ventral view with strong conspicuous node-like expansion on inner margin, with strong spaced and thick setae throughout ventral surface. Paracuspis well-developed, node-like, almost sessile, virtually as broad as long, posterior margin subtriangulate, sparsely and inconspicuously setose posterodorsally, setae longer than or shorter than paracuspis length. Digitus slightly curved inward and somewhat tapered posterad in dorsal view, upcurved posteriorly and constricted medially in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without well-defined lateral pocket on outer surface; apical distance between teeth 0.1 × length of valve; strong setae present along convex posterior margin, inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: vertex, dorsal half of front, dorsal half of gena, pronotal dorsum, mesoscutum, axillar projections, mesoscutellum, disc of T2 (except anterior third to anterior half), fringe of T2–6 medially, and S6–7 partially. Tibial spurs yellowish-white. Wings dark brown infuscated with strong violaceous and blueish reflections, except basal fourth to basal third hyaline, veins poorly discernible except at basa third to fourth. Distribution. Suriname, Guiana, Brazil, and Peru. Material examined. (30m#) Type material. Holotype: m#, GUIANA, Kartabo, Bartica District, 10.III.1922, Wm. [William] Beebe (UMSP). Additional material. SURINAME: Zanderij, Kreekbos, 1m#, 10–14.VII.1964, D.C. Geijskes (FSCA); BRAZIL: Pará: Marabá, 3m#, X.1974, J.F. Reinert (FSCA); Belém, Parque Ambiental: 2#m, 14– 18.VI.2004, A.L. Nunes e equipe (MPEG); 2#m, 09–13.VIII.2004, A.L. Nunes e equipe (MPEG); Ourém, Patauateua: 3#m, 20.VIII.1992, B. Mascarenhas (MPEG); 1#m, 23.VIII.1992, B. Mascarenhas (MPEG); 1#m, 15–17.VIII.1992, B. Mascarenhas (MPEG); Juruti, Área da Adutora, 02°27’9.04’’S 56°10’55.20’’W, 1#m, 21–24.VI.2008, E. Monteiro-Santos, Olivia Aguiar, R.L. Trindade, L.A. Quaresma (MPEG); Melgaço, Caxiuanã, ECFPn [Estação Científica Ferreira Pena], 1#m, 04–08.VIII.2000, A. Hook, C.K. Starr, W. Overal (MPEG); Tucuruí, Rio Tocantins, Saúde, 1#m, 03–05.VI.1984 (MPEG); Rondônia: Ouro Preto d’Oeste: 1m#, 18.X.1987, Elias, C. (DZUP); 1#m, 11– 13.XI.1984 (MPEG); Linha 62 km16, 3#m, 13–15.VI.1984 (MPEG); 62km SE [kilometers southeast of] Ariquemes: 1m#, 07.XI.1995, W.J. Hanson (EMUS); 1m#, 08.XI.1996, W.J. Hanson (EMUS); 1m#, 23.X.1997, W.J. Hanson (EMUS); 1m#, 07–15.XI.1995, W.J. Hanson (EMUS); 1m#, 22–31.X.1997, W.J. Hanson (EMUS); Nova Mamoré, Parque Estadual Guajará-Mirim, Rio Formoso, 1#m, 22.X.1995, Vidal, J. & Aquino, L. (INPA); Acre, Rio Branco, 1#m, 25.X–08.XI.1991, F. Ramos, A. Henriques, I. Gorayeb, N. Bittencourt (MPEG); Maranhão, Bom Jardim, REBIO [Reserva Biológica] Gurupi, 1#m, 17–27.I.2010, F. Limeira-de-Oliveira, J.T. Camara, A.T. Sousa (CZMA); PERU, Madre de Diós, CICRA Fld. Stn. [Centro de Investigacion y Capacitación Río Los Amigos Field Station] Garden; 12.56940°S 70.10100°W, 260m [meters above sea level], 1m#, 12–19.VIII.2010, M.J. Endara, ex. malaise trap PER10-08-MAT-011 / KUHNM-ENT SEMC1060205 [sic] (SEMC). Remarks. The cuspis of T. bartica is covered with strong spine-like setae along its ventral surface and also has the inner margin expanded basally in ventral view, which are unique characters within the genus. The remaining genitalia 238 characters and external morphology, however, are similar to some other known males, including T. taboca. There is not yet a clear candidate for the female of T. bartica, but distribution and the truncate apex of the femora indicate that it might belong in the T. inermis or T. integella species-group.
Traumatomutilla mesoleuca (Gerstaecker, 1874) (Figs. 9A–F, 12D, 12I)
Mutilla mesoleuca Gerstaecker, 1874: 317, lectotype [designated here], #m, Brazil, Sello. [Sellow] (ZMB), examined. Ephuta (Traumatomutilla) mesoleuca: André, 1902, 55 (new combination). Traumatomutilla mesoleuca: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate; apex of meso and metafemora rounded; cuspis short, 0.7 × the length of paramere, strongly convergent posterad in dorsal view, slightly sinuous and tapering posterad in lateral view. Description. FEMALE. Unknown. MALE. Body length 12 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly dark-brown, slightly lighter basally, veins blackish. Setae entirely black except with whitish setae on legs, mesopleuron predominantly, metapleuron, propodeal dorsum predominantly, T1, T2 basally, fringe of T2–5 laterally (greatly reduced on T5), S1–S2, and fringe of S2–4. Head. Transversely subrectangular, virtually straight on posterior margin. Vertex width 1.0 × pronotal width. Eye almost circular. Ocelli small; OOD 5.8 × DLO, IOD 1.8 × DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with closely spaced pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.1 × pedicel length; flagellomere 2 2.8 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, greatly reduced, restricted to posterior third of mesoscutum; with conspicuous medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection conspicuously separated from scutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbose, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron at most slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, vestigially sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly slightly thickened, rounded; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.8 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly longitudinally rugose throughout, lateral carinae obscure, short. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 densely and coarsely foveolate-punctate, with small shallow setae filled pit posteromedially; foveolations sparser and larger posterad. S3–6 densely and coarsely foveolate-punctate. S7 elongate sub-trapezoidal, densely foveolate- punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia.: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 78:56:20. Paramere slightly curved inward and curved outward posterad, with sparse short inconspicuous setae throughout outer and inner surfaces. Cuspis tapering and curved inward posterad in dorsal view, in lateral view 239 conspicuously wider on anterior half than posterior half, with strong dense setaet throughout, setae conspicuously shorter anterad. Paracuspis well-developed, node-like, not sessile, slightly longer than broad, posterior margin rounded and densely setose throughout, setae as long as or shorter than paracuspis length. Digitus slightly incurved in dorsal view and upcurved in lateral view, somewhat constricted before apex in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short teeth posteroventrally, posteriormost tooth acute, anterior tooth subacute, with poorly defined lateral pocket on outer surface, apical distance between teeth 0.1 × length of valve; strong setae present along posterior margin posterior margin, posterior margin somewhat truncate, slightly sloping dorsad, inconspicuous setae present at base of anterior tooth on outer surface. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except the following areas with silvery-whiite setae varying in density: mesopleuron partially, lateral face of propodeum partially, propodeal dorsum partially, legs predominantly, T1, basal third of T2, fringe of T2–5 laterally, and S1–5 (except fringe of S5). Tibial spurs yellowish-white. Wings light brown, without strong reflections or conspicuous darkened areas, veins easily discernible throughout. Distribution. Brazil, Bolivia, Paraguay and Argentina Material examined. (114m#) Type material. Lectotype: Traumatomutilla mesoleuca, m#, BRAZIL, Sello. [Sellow] (ZMB). Additional material. BRAZIL: Rio Grande do Sul, 3m#, Krüger, R.F. (DZUP); Minas Gerais: Berizal ,Fazenda Veredão, 18°39'54''S 41°39'56''W, 3#m, 12.XII.2012, J.A. Rafael & E.J. Grossi (INPA); Santana do Riacho, 1#m, 10.IV.1992, Melo,F.R. (DZUP); Bahia, Anagé, 1#m, 21.XI.1976, Elias, C., Enoque (DZUP); BOLIVIA: Beni,Rio Mamore, 10 km E [kilometers east of] San Antonio, 13#m, 13.VIII.1965, J.K. Bouseman (AMNH); approx. 10km E [approximately 10 kilometers east of] San Antonio, 1#m, 12.VIII.1965 (EMUS); Cochambamba: Villa Tunari: Hotel los Tucanes, 16°58’39’S 65°23.79’W, 320m [above sea level], 1#m, 04–06.IX.2000, M. Hause (EMUS); 16°54'S 65°22'W, 1#m, 15.II.2001, H. Helder (EMUS); 1#m, 4.IX.2000, M. Hauser (EMUS); 1#m, 15.VIII.2001, H. Heider (EMUS); Santa Cruz: 5km SE [kilometers southeast of] Buena Vista, Hotel Flaura Y Fauna: 17°30'S 63°39'W, FIT [flight interception trap] , 4#m, 15–24.XII.2003, S. & J. Peck (EMUS); Buena Vista:17°27.68'S 63°39.63W, Malaise, 2#m, 21.II.1999, F.D. Parker (EMUS); 3 km N [kilometers north of] Buena Vista, Hotel Flora y Fauna: 4#m, 15.XII.2003, S. & J. Peck (EMUS); 1#m, 31.X.1999,Porter & Stange,FSCA; Gral. [General] Saavedra Est. [Estación] Experimental, 2#m, XI.1973, Porter & Stange (FSCA); Caaguazu, Caaguazu: 1#m, XI.1977, Fritz (AMNH); 3#m, XII.1977, Fritz (AMNH); Paraguari, Cere [Cerro] Pelado, 1#m, 18.XII.1931, F. Sohade (UMSP); Cordillera, Inst. Agr. Nac. [Insituto Agricola Nacional] Caacupa, 1#m, 12.I.1981, E.D. Cave (USNM); Amambay, P.N. [Parque Nacional] Cerro Cera, 1#, 24.II.1981, E.D. Cave (USNM); San Pedro, Rio Ypane, Cororo, 30#m, XII.1983, Fritz (AMNH, EMUS); PARAGUAY, Mborero, 8#m, XI.1937 (CUIC); ARGENTINA: Catamarca, Santa Maria, 1#m, 04.I.1991, Fritz (AMNH); Corrientes:Ytuzaingo: 10#m, III.1982, Fritz (AMNH); 1#m, Fritz (EMUS); 1#m, I.1985, Fritz (EMUS); 1#m, III.1982, Fritz (EMUS); 1#m, XII.1981, Fritz (EMUS); Formosa, Est. Guaycolsec [sic!], 25 km N [kilometers north of] Formosa, 2#m, 10.III.1999, Heydon & Ledford (UCDC); Mendoza: 1#m (MNHN); Chacras de Coria, 2#m, 12.III.1980, C. Porter (FSCA); Missiones: P.N. [Parque Nacional] Iguazu Cent. Ecol. [Centro Ecológico], 6#m, 24.XII.1990, S & J Peck (PMAE); Punto Esperanza, 1#m, XII.1976, Fritz (AMNH); Salta: Pocitos, 1#m, XII.1981, Fritz (AMNH); Rosario de Lerma, 1#m, 16.XII.1983, M. Wasbauer (CSCA). Remarks. External morphology and genitalia place T. mesoleuca within the T aethiops subgroup. Given the evidence suggesting that T. aehtiops might be the male of T. inermis, T. mesoleuca could be associated with T. tetrastigma (Gerstaecker, 1874) or T. hemicycla (Gerstaecker, 1874) with which it shares distribution in southern South America. Also, T. hemicycla and T. tetrastigma are morphologically closer to T. inermis than to any other species within the T. inermis species-group. While T. mesoleuca is remarkably similar to T. vulnerata apart from minor genitalic differences (discussed in the remarks of T. vulnerata), it is unclear if these species are synonymous. Their shared distribution with each other and with T. tetrastigma and T. hemicycla suggests that these are either two different couples widespread in southern South America or all members of a single variable species.
Traumatomutilla orbana (Cresson, 1902) (Figs. 10A–F, 12E, 12J)
240
Mutilla orbana Cresson, 1902: 71, holotype [by monotypy], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães], April (CMNH), examined. Ephuta (Traumatomutilla) orbana: André, 1902, 56 (new combination). Traumatomutilla orbana: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate; apex of meso and metafemora rounded; cuspis short, 0.7 × the length of paramere, strongly convergent posterad in dorsal view, slightly sinuous and tapering posterad in lateral view; anterior margin of scutellum straight, not projected medially. Description. FEMALE. Unknown. MALE. Body length 12.5 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly dark-brown, slightly lighter basally, veins blackish. Setae entirely black except with gray-whitish setae on metapleuron sparsely; propodeal dorsum sparsely; legs; T1; T2 basally and apicolaterally; T3 entirely; T4 laterally; and S1–S4. Head. Rounded, postero-medially swollen. Vertex width 1.0 × pronotal width. Eye almost circular. Ocelli small; OOD 5.0 × DLO, IOD 1.3 × DLO. Occipital carina distinct. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.2 × pedicel length; flagellomere 2 2.5 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis obsolete. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate- punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbous, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate-rounded apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly narrow, rounded; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.5 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly longitudinally rugose throughout, lateral carinae strongest posteriorly. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, with seta-filled pit posteromedially. S3–6 densely and coarsely foveolate- punctate. S7 elongate sub-trapezoidal, densely foveolate-punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia.: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 84:60:25. Paramere slightly curved inward and curved outward posterad, with sparse short inconspicuous setae throughout. Cuspis tapering and curved inward posterad in dorsal view, in lateral view conspicuously wider on anterior half than posterior half, with strong dense setae throughout except at posterior third of inner surface, setae conspicuously shorter on anterior third. Paracuspis well- developed, node-like, nearly petiolate in shape, slightly longer than broad, posterior margin rounded and densely setose throughout, setae as long as or shorter than paracuspis length. Digitus incurved in dorsal view and upcurved in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, anterior tooth slightly curved anteroventrally in lateral view, with poorly defined lateral pocket on outer surface, apical distance between teeth 0.1 × length of valve; strong setae present along flat posterior margin, setae slightly longer ventrad. 241
Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly brownish-black varying in density, except the following areas with silvery-white setae varying in density: mesopleuron posteroventrally, metapleuron, lateral face of propodeum, posterior half of propodeal, dorsum femora and tibiae predominantly, T1, anterior fourth of T2, fringe of T2–3, fringe of T4–5 laterally, and S1–5 (except fringe of S5). Tibial spurs yellowish-white. Wings dark-brown with fain violaceous and blueish reflections, without conspicuous darkened areas, veins easily discernible throughout. Distribution. Brazil, and Paraguay. Material examined. (14m#) Type material. Lectotype: Traumatomutilla orbana, m#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães], April (CMNH). Additional material. BRAZIL: Mato Grosso, 9m#, various dates, Elias, C. (DZUP); São Paulo: Villa Americana [Americana], 1m#, II.1924, F.X. Williams (UMSP); Fazenda Cambuy/Matão, 1m#, 22.XI.2005, S. Halbert (FSCA); Campinas, 1m#, III.1921, F.X. Williams (UMSP); PARAGUAY, San Pedro, Rio Ypane, Cororo, 2m#, XII.1983, Fritz (AMNH). Remarks. External and genitalia morphology places T. orbana in the T. aethiops subgroup as another putative male for the T. inermis species-group. This species, however, might be synonymized with T. mesoleuca in the future, since they are separated by minor setal differences and an inconspicuous structural difference in the mesoscutellum. Additionally, while T. mesoleuca seems to be somewhat restricted or more common in the Pampa areas of South America, T. orbana occurs in the Cerrado and Chaco regions.
Traumatomutilla picada (Cresson, 1902) (Figs. 11A–F, 12F, 12K)
Mutilla picada Cresson, 1902: 74, holotype [by monotypy], #m, Brazil, [Pará], Santarém (CMNH), examined. Ephuta (Traumatomutilla) picada: André, 1902, p. 55 (new combination). Traumatomutilla picada: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen in dorsal half; axillar projections truncate; apex of meso and metafemora rounded; cuspis short, 0.7 × the length of paramere, slightly convergent posterad in dorsal view, slightly sinuous and conspicuously tapering posterad in lateral view; penis valve with anterior posteroventral tooth broad, rounded apically, and short strong nearly equally long setae on posterior margin. Description. FEMALE. Unknown. MALE. Body length 13 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly light brown, slightly lighter basally, veins blackish. Setae entirely black except with whitish setae on head entirely; pronotum; mesoscutellum; metanotum; mesopleuron; metapleuron; propodeal dorsum; legs; T1; T2 basally, laterally, and apicolaterally; T3–4 entirely; T5 laterally; and S1–6. Head. Rounded, postero-medially swollen. Vertex width 1.05 × pronotal width. Eye almost circular. Ocelli small; OOD 5.5 × DLO, IOD 1.4 × DLO. Occipital carina distinct. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 1.9 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis obscure, restricted to posterior half of mesoscutum; with conspicuous antero-medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbous, densely areolate, dorsal face shorter than and poorly distinguished from posterior 242 face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly narrow, rounded; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.5 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly longitudinally rugose throughout, lateral carinae strongest posteriorly. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, with seta-filled pit posteromedially. S3–6 densely and coarsely foveolate-punctate. S7 elongate sub-trapezoidal, densely foveolate-punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 97:68:33. Paramere slightly curved outward posterad, virtually straight elsewhere, with sparse short inconspicuous setae throughout. Cuspis tapering and curved inward posterad in dorsal view, in lateral view conspicuously wider on anterior half than posterior half, smoothly tapering posterad, with strong dense setae throughout except at posterior third of dorsal surface apparently asetose, setae conspicuously shorter on anterior third. Paracuspis well-developed, node-like, not sessile, slightly longer than broad, posterior margin rounded and densely setose throughout, setae as long as or shorter than paracuspis length. Digitus incurved in dorsal view and upcurved in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short teeth posteroventrally, posteriormost tooth acute, short than broad and blunt anterior tooth anterior tooth, with well-defined lateral pocket on outer surface, apical distance between teeth 0.15 × length of valve; strong setae present along flat posterior margin, setae slightly longer ventrad. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: mesoscutum, axillar projections, mesoscutellum partially, posterior half of T2 disc, fringe of T5 medially, T6–7, fringe of S5, and S6–7. Distribution. Brazil. Material examined. (1m#) Type material. Holotype: Traumatomutilla picada, m#, BRAZIL, [Pará], Santarém (CMNH). Additional material. BRAZIL, Pará, 1m#, 1846, Ghiliani (MNHN). Remarks. Traumatomutilla picada is another species that shares the morphology of the T. aethiops subgroup. The penis valve, however, is remarkably different than any other species within this subgroup. If indeed the species of the T. aethiops subgroup are the males of the T. inermis species-group, putative females for T. picada, based on distribution, are T. vagabunda (Smith, 1879) or T. crona Casal, 1969, especially the first since it was collected in the same type locality as T. picada. Nonveiller (1990) stated that this species occurred in Guyana and Brazil without providing the source for the Guyana record. We were unable to find such a record in the literature, neither did we find any specimens of T. picada outside Brazil.
Traumatomutilla pomba (Cresson, 1902) (Figs. 3A, 13A–F, 19A, 19G)
Mutilla pomba Cresson, 1902: 79, holotype (by monotypy), #m, Brazil, [Pará], Santarém (CMNH), examined. Ephuta (Traumatomutilla) pomba: André, 1902, p. 56 (new combination). Traumatomutilla pomba: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron unarmed on dorsal half; axillar projections truncate; apex of meso and metafemora rounded; hypopygium terminating in medial acute expansion on posterior margin; clypeus with medial anteroventral tooth-like projection; T1 shape strongly petiolate, almost nodose; cuspis straight and equally wide in dorsal and lateral view. Description. FEMALE. Unkown. MALE. Body length 8 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly light-brown fuscous, veins blackish. Setae entirely whitish except with black setae on mesoscutum, mesoscutellum, axillar projections, disc of T2 predominantly, 243 fringe of T2–5 medially (interspersed on T3–5), T6–7 (except pygidial plate), and S6–7. Head. Transversely subrectangular and with convex posterior margin in dorsal view; in frontal view, conspicuously subtriangular dorsally. Vertex width 1.05 × pronotal width. Eye almost circular. Ocelli small; OOD 3.1 × DLO, IOD 1.0 × DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate; sculpture coarser and denser on front. Gena ecarinate. Antennal scrobe concave to eye margin, without any scrobal carina. Clypeus virtually flat to slightly depressed laterally and convex medially; densely and coarsely punctate to micropunctate throughout; anteroventral margin projected medially into blunt tooth-like projection. Scape unicarinate. Flagellomere 1 1.8 × pedicel length; flagellomere 2 2.2 × pedicel length. Mandible weakly bidentate apically; lacking dorsal or ventral projections. Mesosoma. Epaulets conspicuous, sharp, broadly connected with humeral carina. Anterior face of pronotum virtually entirely smooth, asetose, shinning, virtually flat throughout. Dorsal face of pronotum with sharp upcurved edges on anterior margin. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate to areolate-punctate; notaulus and parapsis present, well-defined, restricted to posterior half of mesoscutum, with conspicuous medial longitudinal carina throughout. Mesoscutellum slightly convex, bearly flat, sloping posterad, without definable dorsal and posterior faces, densely foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, sparse foveolate-punctate, except posterior margin impunctate; projection conspicuously separated from scutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum virtually equally wide throughout, its surface obscured by dense setae. Propodeum convex, densely areolate, dorsal face indistinguishable from posterior face; lateral face mostly impunctate, with vestigial areolations to unsculptured and smooth anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures anteriorly; areolations vestigial anterad and posterad; dorsal half of mesopleuron at most slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, vestigially sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, subacute apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur narrow, rounded; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.5 × T2 width. T2 length 0.9 × T2 width. Dorsal metasomal sculpture sparsely and finely foveolate-punctate. T7 virtually unsculptured, smooth, at most with vestigial undefined sculpture apically; lateral carinae obscure, short. S1 longitudinally slightly elevated medially, forming low, shallowly concave carina. S2 densely and finely foveolate-punctate; foveolations sparser and smaller anterad. S3–6 sparsely and finely foveolate-punctate. S7 longitudinally subrectangular, sparsely foveolate-punctate; apical margin projected medially into a single acute projection. Genitalia.: Parapenial lobe not produced posteriorly, narrow, acute. Ratios of free length of paramere, cuspis and digitus, 85:56:19. Paramere virtually straight in dorsal view, slightly upcurved in lateral view, with dense long setae on anterior half and sparse short inconspicuous setae elsewhere. Cuspis straight in dorsal and lateral views, slightly tapered posteriorly in lateral view, weakly laterally compressed, with very short inconspicuous weak setae throughout. Paracuspis poorly developed, lobe-like, sessile, broader than long, posterior margin rounded with scarce weak inconspicuous setae, setae shorter than paracuspis length. Digitus incurved in dorsal view, slightly upcurved anteriorly and straight to downcurved posteriorly in lateral view, tapering posterad in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without lateral pocket on outer surface, apical distance between teeth 0.11 × length of valve; strong setae present along convex posterior margin. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with black to brownish-black setae varying in density: pronotal dorsum partially, mesoscutum, axillar projections, mesoscutellum partially, T2 disc (except anterior third), fringe of T2–6 medially, T7, fringe of S5, and S6. Tibial spurs yellowish-white. Wings hyaline brown on apical half, hyaline on basal half, without any conspicuous reflections, veins easily discernible throughout. Distribution. Colombia, French Guiana, Brazil, Ecuador and Peru. Material examined. (61m#) Type material. Holotype: Mutilla pomba, m#, BRAZIL, [Pará], Santarém (CMNH). Additional material COLOMBIA, 1#m (IAvH); FRENCH GUIANA, Saal [sic!], 1#m, 31.V.1997 (EMUS); BRAZIL: Vila Vera [sic!], 1#m, X.1973, M. Alavarenga (EMUS); Pará, 1#m, 13.II.1902, Ducke (MNHN); Óbidos, 1#m, 20.VII.1902, Ducke (MNHN); Tucuruí, 1#m, I.1979, M. Alvarenga (EMUS); Oriximiná, Porto Trombetas, Platô Garganta, 2#m, 14.VIII.2008, Freitas, H.M. (INPA); Vitória do Xingú, Margem esq. [esquerda] do rio Xingú: 2#m, 18–20.XI.2000, R. Santos & J. Dias (MPEG); 1#m, 10–12.XII.2000, C. Maciel & J. Dias (MPEG); 1#m, 14– 244
16.XII.2000, C. Maciel & J. Dias (MPEG); 1#m, 20–22.XI.2000, C. Maciel & J. Dias (MPEG); 1#m, 14–16.XII.2000, R. Santos & J. Dias (MPEG); Serra Norte, Manganês, 1#m, 12–13.IX.1985, M.F. Torres (MPEG); Benevides: Faz. [Fazenda] Morelândia, 1#m, 30.VI–02.VIII.1988, F.F. Ramos (MPEG); PA408, km06, 1#m, 26.VI.1981, I.S. Gorayeb (MPEG); Amazonas, EMBRAPA [Empresa Brasileira de Pesquisa Agropecuária, Guaraná cultivo convencional, Mata, 1#m, 05.I.2013, K. Schoeninger (INPA); Reserva Ducke: 3#m, 29.IX–13.X.2005, A. Aguiar (INPA); 3#m, 29.IX– 08.X.2005, A. Aguiar (INPA); 2#m, 08–14.X.2005, A. Aguiar (INPA); 1#m, 30.IX–08.X.2005, A. Aguiar (INPA); 1#m, 28.IX–10.X.2005, A. Aguiar (INPA); 1#m, X.2005, A. Aguiar (INPA); Presidente Figueiredo, Km24, AM240, 02°01’05’’S 59°49’59’’W, 1#m, 04– 08.IX.2008, Rondônia, 62km SE [kilometers southeast of] Ariquemes: 1#m, 12.XI.1991, E.M. Fischer (UCDC); 1#m, 01–14.xi.1997, Xavier-Filho, F.F. & Krolow, T. (INPA); B. Dozier (FSCA); 1#m, 8.XI.1994, W.J. Hanson (EMUS); 2#m ,22.X.1997, W.J. Hanson (EMUS); 1#m, 7.XI.1995,W.J. Hanson (EMUS); 1#m, 1.XI.1997, B. Dozier (EMUS); Itapuã do Oeste, FLONA [Floresta Nacional] do Jamari: 1#m, 4– 6.IX.2012, Melo, G.A.R., Rosa, B., Santos (DZUP); Ponto 3, 1#m, 15.VI.2013, Luz, D., Rosa, B., Williams, K.A. (DZUP); Ouro Preto d'Oeste, 1#m, 18.X.1987, Elias, C. (DZUP); Vilhena: 1#m, XI.1973, M. Alvarenga (EMUS); 2#m, 22.XII.1986, Elias, C. (DZUP); 1#m, 9.X.1986, Elias, C. (DZUP); 1#m, 15.X.1986, Elias, C. (DZUP); 1#m, 17.XII.1986, Elias, C. (DZUP); Mato Grosso: Sinop: 1#m, X.1976, M. Alvarenga (EMUS); 8#m, X.1970, M. Alvarenga (EMUS); ECUADOR, Napo, Yasuni Res. [Reserve] Station, 1#m, 04.V.2003, C. Bramer (EMUS); PERU, [Madre de Diós]: Pakitza, 1#m, 13.II.1992, B.V. Brown (USNM); CICRA Fld. Stn. [Centro de Investigación y Capacitación Río Los Amigos Field Station], Trail 6, research plot, 12.55207°S 70.10962°W, 295m [above sea level], 07–09.VI.2011, Chaboo team (SEMC); Puerto Maldonado, Los Amigos Biol. Stat. [Biological Station] CICRA [Centro de Investigación y Capacitación, Río Los Amigos], 1#m, 28.I.2014 (FSCA); Remarks. Traumatomutilla pomba is unique within the genus in external and genitalia morphology. The unidentate clypeus has not been observed in any other Traumatomutilla species. The cuspis being entirely straight and sparsely setose throughout is a character that can only be found in Frigitilla Williams in Bartholomay et al. (2015), within the South American Dasymutillini. In terms of putative females, the consistently small size of T. pomba (always less than 10mm) coupled with a strongly petiolate T1 shape parallels the morphology of females of the T. trochanterata and T. tabapua species-groups. The latter, however, is also apparently restricted to the Amazon and is the most likely candidate for T. pomba given the data available.
Traumatomutilla rorida (Gerstaecker, 1874) (Figs. 1O, 14A–F, 19B, 19J)
Mutilla rorida Gerstaecker, 1874: 318, holotype (by monotypy), #m, Brazil, [São Paulo], Salto Grande, Sello. [Sellow] S. [sic] (ZMB), examined. Mutilla sabara Cresson, 1902: 78, lectotype [designated by Cresson (1916)], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães], Jan. [January] (CMNH), type examined, [synonymized by Mickel, (1964)]. Ephuta (Traumatomutilla) rorida: André, 1902, p. 55 (new combination). Traumatomutilla rorida: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron unarmed at dorsasl half; axillar projections truncate; apex of meso and metafemora truncate; cuspis short, 0.6 × the length of paramere, laterally compressed, conspicuously wider medially in lateral view; digitus dorsoventrally flattened, obliquely truncate apically and conspicuously wider apicad in dorsal view; penis valve with basal posteroventral tooth truncate, with well-defined lateral pocket at base on outer margin. Description. FEMALE. Unknown. MALE. Body length 9 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly brown, veins dark brown. Setae entirely black except with whitish setae on scape; clypeus; gena ventrally; mesopleuron; propodeal dorsum; tibiae; tarsi; T1; T2 basally; and S1–3. Head. Rounded, postero-medially swollen. Vertex width 1.05 × pronotal width. Eye almost circular. Ocelli small; OOD 6.2 × DLO, IOD 2.0 × DLO. Occipital carina weak. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below 245 antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate, dorsal carina weak, interrupted by punctures. Flagellomere 1 1.8 × pedicel length; flagellomere 2 2.2 × pedicel length. Mandible oblique, teeth worn down, apparently tridentate apically; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, restricted to posterior half of mesoscutum; with conspicuous medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly thickened, sub-truncate; outer apical lobe of metafemur weakly thickened, sulcate, truncate; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 densely reticulate throughout, lateral carinae strongest posteriorly. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, without seta-filled pit. S3–6 foveolate-punctate. S7 sub-trapezoidal, foveolate-punctate; apical margin projected medially into an apparently unidentate projection. Genitalia.: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 80:49:25. Paramere slightly sinuous in dorsal view, slightly upcurved in lateral view, with dense conspicuous long setae throughout, except dorsal surface, setae denser and longer posterad. Cuspis straight and equally wide throughout in dorsal views, slightly upcurved and tapered posterad and anterad in lateral view, medioventrally expanded, somewhat “banana- shaped”, with strong thick setae on ventral surface at posterior half and inconspicuous short setae on anterior half. Paracuspis poorly developed, lobe-like, sessile, longer than broad, posterior margin rounded with conspicuous setae as long as paracuspis length. Digitus in dorsal view club-like, obliquely truncate posterad, inner-posterior marginrounded, conspicuously broader posteriorly than anteriorly, slihgtly incurved, in lateral view slightly upcurved at posterior extremity, strongly tapered posterad, densely setose throughout dorsal surface, setae shorter posterad. Penis valve strongly concave on inner surface, with well-defined pair of short teeth posteroventrally, posteriormost tooth acute, anterior tooth truncate apically, with well-defined lateral pocket on outer surface, apical distance between teeth 0.1 × length of valve; strong thick setae present along convex posterior margin. Coloration and variations. FEMALE. Unkown. MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: ventral half of gena, malar space, mesopleuron, metapleuron, latera face of propodeum, legs predominantly, propodeal dorsum, T1, basal fourth of T2, and S1–4 (except fringes of S3–4). Tibial spurs yellowish-white. Wings dark brown, without conspicuous reflections or darkened areas; veins easily discernible throughout. Distribution. Brazil. Material examined. (2m#) Type material. Holotype of Mutilla rorida, m#, BRAZIL, [São Paulo], Salto Grande, Sello S. (ZMB); Lectotype of Mutilla sabara, m#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães], Jan. [January] (CMNH). Additional material. BRAZIL: São Paulo: Campinas, 1m#, III.1921, F.X. Williams (UMSP). Remarks. The genitalia of T. rorida, especially the shape and setae of the paramere, cuspis, and penis valve, is remarkably similar to a male collected in copulo with a female of the T. inermis species-group from MNCN (possibly Traumatomutilla laida Casal, 1969). The fact that the T. aethiops subgroup is also likely to be associated with the T. inermis species-group suggests that males of the latter will have greater morphological diversity than what has been previously observed in other groups, such as the T. indica and T. quadrinotata species-groups. Additionally, one of the males in Cresson’s T. sabara type series was collected in Santarém, Pará state, in the Brazilian Amazon; it is 246 completely different from the lectotype and other described Traumatomutilla in external and genitalia morphology. The specimen has been labeled as an undescribed morphospecies.
Traumatomutilla soricina (Gerstaecker, 1874) (Figs. 15A–F, 19C, 19H)
Mutilla soricina Gerstaecker, 1874: 320, holotype [by monotypy], #m, Brazil, v. Winth. S. (ZMB), type examined. Ephuta (Traumatomutilla) soricina: André, 1902, p. 56 (new combination). Traumatomutilla soricina: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate; apex of meso and metafemora truncate; posterior half of mesopleuron, metapleuron and lateral face of propodeum with conspicuous areas of dense appressed silvery-goldens setae; cuspis short, 0.7 × the length of paramere, strongly convergent posterad in dorsal view, slightly sinuous and tapered posterad in lateral view. Description. FEMALE. Unknown. MALE. Body length 12.5 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly dark-brown, slightly lighter basally, veins blackish. Setae entirely black except with whitish setae on frons; gena; mesoscutellum; metanotum; mesopleuron; metapleuron; propodeal dorsum; legs; T1; T2 basally, laterally, and apicolaterally; T3 entirely; T4 basally; T5 laterally; and S1–S6. Head. Rounded, postero-medially swollen. Vertex width 0.95 × pronotal width. Eye almost circular. Ocelli small; OOD 4.7 × DLO, IOD 1.3 × DLO. Occipital carina distinct. Frons, vertex, and gena densely coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with pair of conspicuous tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 2.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, restricted to posterior half of mesoscutum; with conspicuous medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbous, densely areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate-rounded apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly narrow, rounded; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.8 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and mostly longitudinally rugose throughout, lateral carinae strong posterolaterally. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, with setae filled pit posteromedially. S3–6 densely and coarsely foveolate-punctate. S7 elongate sub-trapezoidal, densely foveolate-punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 85:62:25. Paramere virtually straight throughout in dorsal view, slightly leaning inward, upcurved in lateral view, virtually asetose apart from sparse short and barely visible setae. Cuspis tapering and strongly curved inward posterad in dorsal view, in lateral view conspicuously wider on anterior half than posterior half, smoothly tapering posterad, with strong dense setae 247 throughout except at dorsal surface with short sparse setae. Paracuspis well-developed, node-like, not sessile, longer than broad, posterior margin rounded and densely setose throughout, setae as long as or shorter than paracuspis length. Digitus strongly incurved in dorsal view and upcurved in lateral view. Penis valve strongly concave on inner surface, with well-defined pair of short teeth posteroventrally, posteriormost tooth acute, anterior tooth subacute, without apparent lateral pocket on outer surface, apical distance between teeth 0.15 × length of valve; strong setae present along flat posterior margin. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with black setae varying in density: dorsal half of front, dorsal half of gena, pronotal dorsum, mesoscutum, axillar projections, mesoscutellum partially, T2 disc (except anterior third), fringe of T2 medailly (in part), T5 partially, T6, T7 partially, fringe of S5–6, and S7. Tibial spurs yellowish-white. Wings dark brown, without conspicuous reflections, slightly lighter on basal fourth, veins discernible throughout. Distribution. Brazil and Paraguay. Material examined. (4m#) Type material. Holotype: Mutilla soricina, m#, BRAZIL, v. Winth. S. (ZMB). Additional material. BRAZIL: Mato Grosso, Chapada dos Parecis 1m#, 01.XII.2001, A. Foucart (EMUS); Minas Gerais, nr. [near] Timoteo, 12.II.1981, E.R. de Paula (EMUS); PARAGUAY, Cordillera, Inst. Agr. Nac. [Insituto de Agricultura Nacional] Caacupa, 12.II.1981, R.D. Cave (USNM). Remarks. Traumatomutilla soricina is another species within the T. aethiops subgroup and, therefore, also a likely candidate to be associated with the T. inermis species-group. The distribution of T. soricina in the Cerrado areas of Brazil and Chaco areas of Paraguay makes it impossible to choose a putative female at this point, since there are at least 10 likely candidates within the T. inermis species-group in those areas.
Traumatomutilla taboca (Cresson, 1902) (Figs. 16A–F, 19D, 19K)
Mutilla taboca Cresson, 1902: 78, lectotype [designated by Cresson (1916)], #m, Brazil, [Mato Grosso], Chapada [dos Guimarães], Campo, Sept. [September] (CMNH), examined. Ephuta (Traumatomutilla) taboca: André, 1902, p. 56 (new combination). Traumatomutilla taboca: André, 1904, p. 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate; apex of meso and metafemora truncate; paramere tapering only at apical third, with dense tuft of long setae on ventral surface at apical third; cuspis short 0.6 × the length of paramere, dorsoventrally flattened, greatly expanded along inner margin on ventral view, with strong spine-like setae on verntral surface and along inner margin; penis falve with basal tooth longer than apical tooth, broadly rounded, with conspicuous lateral pocket on outer surface. Description. FEMALE. Unknown. MALE (Hitherto undescribed). Body length 9 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly hyaline-brown, slightly darker on apical third, veins blackish. Setae entirely black except with whitish setae on legs, head predominantly, pronotum partially, mesopleuron partially, propodeum, metanotum predominantly, legs, T1, T2 basally, fringe of T2–4 laterally (vestigial on T4), and S1–5. Head. Transversely subrectangular, slightly convex on posterior margin. Vertex width 0.95 × pronotal width. Eye almost circular. Ocelli small; OOD 6.7 × DLO, IOD 2.8 × DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate along anteroventral margin, micropunctate to smooth elsewhere; with closely spaced pair of inconspicuous blunt tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 2.1 × pedicel length; flagellomere 2 2.5 × pedicel length. Mandible vestigially bidentate apically; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum virtually entirely smooth, asetose, shinning, virtually flat throughout; with rounded edges. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis 248 present, restricted to posterior third of mesoscutum, with conspicuous medial longitudinal carina. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, densely foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dense foveolate-punctate, except posterior margin impunctate; projection conspicuously separated from scutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum virtually equally wide throughout, its surface obscured by dense setae. Propodeum convex, densely areolate, dorsal face indistinguishable from posterior face; lateral face mostly impunctate, with vestigial areolations to unsculptured and smooth anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures anteriorly; areolations vestigial anterad and posterad; dorsal half of mesopleuron at most slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, vestigially sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly thickened, sulcate, truncate; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture sparsely and finely foveolate-punctate. T7 coarsely, confusedly and irregularly rugose to granulose throughout, lateral carinae obscure, short. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 sparsely and finely foveolate-punctate; foveolations sparser and smaller anterad. S3–6 sparsely and finely foveolate-punctate. S7 nearly subquadrate, sparsely foveolate-punctate; apical margin projected medially into a single blunt projection. Genitalia: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 70:41:18. Paramere in dorsal view virtually straight, slightly curved outward in posterior third, in lateral view slightly upcurved, with dense conspicuous tuft og long setae subapically, inconspicuous scattered short setae elsewhere. Cuspis broad in dorsal view, conspicuously tapered posterad ending in subacute apex, expanded anterad along inner margin, conspicuously flattened dorsoventrally and slightly constricted medially in lateral view, with stron thick setae throughout inner margin, setae conspicuously longer on posterior half. Paracuspis poorly developed, lobe-like, sessile, longer than broad, posterior margin rounded with conspicuous setae as long as or shorter than paracuspis length. Digitus in dorsal view club-like, obliquely truncate on posterior margin, slightly broader posteriorly than anteriorly, virtually straight, posterior half dorsoventrally compressed; slightly upcurved at posterior extremity and strongly tapered posterad in lateral view, densely setose throughout dorsal surface, setae shorter posterad, anterior half laterally compressed. Penis valve strongly concave on inner surface, with well-defined pair of teeth posteroventrally, posteriormost tooth acute, shorter than apically truncate anterior tooth, with well-defined lateral pocket on outer surface, apical distance between teeth 0.15 × length of valve; strong thick setae present along convex posterior margin. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: mesopleuron posteriorly, metapleuron, lateral face of propodeum, propodeal dorsum, metanotum medially, femora, T1, anterior third of T2, fringe of T2–5 laterally, and S1–5 (except fringe of S4–5). Tibial spurs yellowish-white. Wings dark brown, without conspicuous reflections, slightly lighter on basal fourth, veins discernible throughout. Distribution. Colombia, Brazil, Ecuador, Bolivia, and Peru. Material examined. (7m#) Type material. Holotype: Mutilla taboca, m#, BRAZIL, [Mato Grosso], Chapada [dos Guimarães], Campo, Sept [September] (CMNH). Additional material. COLOMBIA, Amazonas: Leticia, 1#m, 28.V.1979, R.C. Wilkerson, M.A. Tidwell (FSCA); BRAZIL: Mato Grosso do Sul, Chapada [dos Guimarães], 1#m, Feb. [February] (MNHN); Rondônia, Itapuã do Oeste, FLONA [Floresta Nacional] do Jamari, Trilha Pedra Grande, 09°11'39.4''S 63°04'55.3''W, 1#m, 08.X.2014, J.A. Rafael, F.F. Xavier-Filho, R.M. Vieira & R.H. Aquino (INPA); ECUADOR, Napo, Misahualli,nr. [near] Tena, 1#m, 26.VIII–02.IX.2000, Steven & Paul Keller (FSCA); PERU: Madre de Diós, CICRA Fld. Stn. [Centro de Investigación Científica Río Los Amigos Field Station] Garden, 12.56940°S 70.10100°W, 260m [above sea level], 1#m, 05–12.VIII.2010, M.J. Endara (SEMC); Cusco, Villa Carmen Field Station, ~ [approximately] 1.7km west cafeteria research transect, 12.89213°S 71.41920°W, 547m, 1#m, 28– 30.V.2011, D.J. Bennett & E. Razuri (SEMC); BOLIVIA, Santa Cruz: Hotel Flora y Fauna, 3 km N Buena Vista, 1#m, 6.XII.2003, S. & J. Peck (FSCA). Remarks. The external morphology, especially the truncate apex of the meso and metafemora in T. taboca, place this species as a likely candidate to be associated with the T. inermis species-group. The genitalia superficially resembles that of T. rorida in two main aspects: the digitus, virtually identical in both species and unlike what has been observed 249 for all the Traumatomutilla males; and the paramere which has a similar shape and setae distribution. That, coupled with the previously mentioned similarities between T. rorida and an undescribed male captured in copulo with a female from the T. inermis species, means that all three species are likely closely related. Additionally, a similar male morphospecies, also Amazonian in distribution, was been found that differs from T. taboca in the cuspis and penis valve shape (PRB & KAW pers. obs.).
Traumatomutilla tenuis (Smith, 1879), comb. nov. (Figs. 3K, 17A–F, 19E, 19I)
Mutilla tenuis Smith, 1879: 217, holotype (by monotypy), #m, [Brazil], [Amazonas], Ega [Tefé], B.M. Type Hym. 15.966 (BMNH), type examined. Mutilla tenuis: André, 1902, p.74 (incertae sedis)
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron unarmed on dorsal half; axillar projections truncate; apex of meso and metafemora rounded; T2 sculpture simply and finely punctate, with broad smooth and shinning intervals; cuspis very short, 0.5 × the length of paramere, slightly convergent but not curved in dorsal view, virtually equally wide throughout in dorsal and lateral views, with sparse short but conspicuous setae throughout. Description. FEMALE. Unknown. MALE. Body length 7.5 mm. Coloration. Head, mesosoma, metasoma, and appendages black. Tibial spurs yellowish-white. Wings predominantly light brown, slightly lighter basally, veins blackish. Setae sparse silvery except with blackish erect setae on mesoscutum; tegula; axilla; mesoscutellum; T2 medially; and T5–T7. Head. Rounded, postero-medially swollen. Vertex width 1.15 × pronotal width. Eye almost circular. Ocelli small; OOD 4.2 × DLO, IOD 1.2 × DLO. Occipital carina strong, sub-lamelliform. Frons, vertex, and gena densely moderately foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus slightly depressed laterally below antennae insertion, slightly convex medially; densely and coarsely punctate medially, micropunctate to smooth elsewhere; with pair of weak tooth-like projections medially on anteroventral margin, connected by obscure transverse lamella. Scape unicarinate. Flagellomere 1 2.0 × pedicel length; flagellomere 2 2.4 × pedicel length. Mandible oblique, apical teeth obliterated; lacking dorsal or ventral projections. Mesosoma. Epaulets elongate carinate, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus obliterated; parapsis present, restricted to posterior half of mesoscutum. Mesoscutellum slightly convex, sloping posterad, without definable dorsal and posterior faces, areolate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posterior faces, dorsal face dense foveolate-punctate at anterior half and impunctate at posterior half; projection separated from mesoscutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex, areolate, dorsal face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areolations anterad. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron slightly swollen. Metapleuron virtually smooth, and partially covered with microsetae, obscurely sculptured on dorsal fifth and ventral fourth. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, acute-rounded apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly thickened, sub-truncate; tibial spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.45 × T2 width. T2 length 0.9 × T2 width. Dorsal metasomal sculpture densely foveolate-punctate, sparser on T2. T7 mostly smooth with obscure punctation posteriorly, lateral carinae distinct, converging anteriorly. S1 longitudinally elevated medially, forming low, shallowly concave carina. S2 foveolate-punctate, with deep seta-filled pit posteromedially. S3–6 foveolate-punctate. S7 elongate sub-trapezoidal, foveolate-punctate; apical margin projected medially into a single closely bidentate projection. Genitalia: Parapenial lobe not produced posteriorly, acute. Ratios of free length of paramere, cuspis and digitus, 76:38:14. Paramere virtually straight in dorsal view, slightly upcurved in lateral view, with sparse scattered inconspicuous setae. Cuspis short, laterally compressed, slightly converging and tapering posterad in dorsal view, 250 slightly upcurved and equally wide throughout in lateral view. Paracuspis short, node-like, virtually as long as wide, posterior margin subtriangulate with conspicuously setose, setae sparse, as long as or shorter than paracuspis length. Digitus slightly incurved in dorsal view, upcurved in lateral view, conspicuously setose on dorsal margin. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without apparent lateral pocket on outer surface, apical distance between teeth 0.1 × length of valve; sparse setae present along convex posterior margin. Coloration and variations. FEMALE. Unknown. MALE. Integument black to brownish-black. Body setae predominantly silvery-white varying in density, except the following areas with brownish-black setae varying in density: mesoscutum, axillar projections, mesoscutellum partially, posterior half of T2 disc, T5 partially, T6–7, S5 partially, and S6–7. Tibial spurs yellowish-white. Wings hyaline-brown, without conspicuosu reflections, slightly darkened on apical third, veins easily discernible throughout. Distribution. Brazil. Material examined. (1m#) Type material. Holotype: Mutilla tenuis, m#, [BRAZIL], [Amazonas], Ega [Tefé], B.M. Type Hym. [Hymenoptera] 15.966 (BMNH). Remarks. Traumatomutilla tenuis is morphologically similar to the T. aethiops subgroup with the cuspis being slightly convergent posterad in dorsal view, more or less setose throughout, and the apex of the meso and metafemora truncate. Based on distribution, size, and the previously mentioned morphological characters, T. tenuis is a likely candidate to be associated with species of the T. integella species-group in the future.
Traumatomutilla vulnerata (Gerstaecker, 1874) (Figs. 18A–F, 19F, 19L)
Mutilla vulnerata Gerstaecker, 1874: 315, lectotype (designated here), #m, Brazil, [Rio Grande do Sul], St. Cruz [Santa Cruz do Sul], Hensel (ZMB), examined. Ephuta (Traumatomutilla) vulnerata: André, 1902: 56 (new combination). Traumatomutilla vulnerata: André, 1904: 40 (new combination).
Diagnosis. FEMALE. Unknown. MALE. Mesopleuron simply swollen on dorsal half; axillar projections truncate; apex of meso and metafemora truncate; T2 with a pair of dark red integumental spots; cuspis short, 0.7 × the length of paramere, strongly convergent posterad in dorsal view, strongly sinuous and slightly tapered posterad in lateral view. Description. FEMALE. Unknown. MALE. Body length 12 mm. Coloration. Head, mesosoma, metasoma, and appendages black, except T2 with a pair of medial dull orange to red integumental spots. Tibial spurs yellowish-white. Wings dark fuscous ath apical third, hyaline brown elsewhere, veins blackish. Setae entirely black except with whitish setae on meso and metatibia, meso and metatarsi, metanotum medially, propodeum dorsolaterally, T1, T2 basally, fringe of T2–4 laterally, S1, S2 medially, and fringe of S2–3. Head. Transversely subrectangular, virtually straight on posterior margin. Vertex width 0.93 × pronotal width. Eye almost circular. Ocelli small; OOD 5.7 × DLO, IOD 2.2 × DLO. Occipital carina distinct. Frons, vertex, and gena densely and coarsely foveolate-punctate. Gena ecarinate. Antennal scrobe concave to eye margin, with prominent transverse dorsal scrobal carina starting near eye margin and ending before antennal tubercle. Clypeus depressed laterally below antennae insertion, convex medially; densely and coarsely punctate; with closely spaced pair of tooth-like projections medially on anteroventral margin. Scape bicarinate. Flagellomere 1 1.5 × pedicel length; flagellomere 2 2.1 × pedicel length. Mandible obliquely tridentate apically, inner tooth slightly larger than middle tooth; lacking dorsal or ventral projections. Mesosoma. Epaulets small, not connected with humeral carina. Anterior face of pronotum punctate laterally, with longitudinal smooth furrow medially. Tegula convex, mostly smooth and impunctate with dense punctures along anterior and inner margins. Mesoscutum densely foveolate-punctate; notaulus and parapsis present, greatly reduced, restricted to posterior third of mesoscutum. Mesoscutellum simply convex, sloping, without definable dorsal and posterior faces, foveolate-punctate. Axilla produced posterolaterally as short truncate projection with conspicuous flat dorsal and posteiorior faces, dense foveolate-punctate dorsally, except narrow posterior margin impunctate; projection narrowly separated from scutellum. Area between mesoscutum and mesoscutellum shallowly concave, smooth, sloping posterad. Metanotum slightly narrower medially, its surface obscured by dense setae. Propodeum convex/gibbose, densely areolate, dorsal 251 face shorter than and poorly distinguished from posterior face; lateral face mostly impunctate, with vestigial areaolations along posterior margin and scatterd punctures elsewhere. Mesopleuron mostly areolate-punctate with interspersed micropunctures, areolations vestigial anterad and posterad; dorsal half of mesopleuron at most slightly swollen. Metapleuron virtually smooth, vestigially sculptured on dorsal and ventral fifths. Wings. Forewing with moderate elongate sclerotized pterostigma; marginal cell elongated, truncate apically; three submarginal cells. Legs. Setae simple, without any conspicuous spines; outer apical lobe of metafemur weakly narrow, rounded; spurs densely microsetose to microserrate laterally. Metasoma. T1 width 0.4 × T2 width. T2 length 0.85 × T2 width. Dorsal metasomal sculpture densely and coarsely foveolate-punctate. T7 coarsely and confusedly rugose throughout, lateral carinae obscure, short. S1 longitudinally elevated medially, forming pronounced concave carina. S2 densely and coarsely foveolate-punctate, with small shallow setae filled pit posteromedially; foveolations sparser and larger posterad. S3–6 densely and coarsely foveolate-punctate. S7 elongate sub-trapezoidal, sparsely foveolate-punctate except along smooth posterior third; apical margin projected medially into a single closely bidentate projection. Genitalia: Parapenial lobe not produced posteriorly, subacute. Ratios of free length of paramere, cuspis and digitus, 98:71:32. Paramere virtually straight throughout in dorsal view, slightly leaning inward, upcurved in lateral view, virtually asetose apart from sparse short and barely visible setae. Cuspis strongly curved inward and tapering posterad in dorsal view, in lateral view conspicuously wider on anterior half than posterior half, smoothly tapering posterad, weakly sigmoidal, with strong dense setae throughout except at dorsal surface with short sparse setae. Paracuspis well- developed, node-like, not sessile, slightly longer than broad, posterior margin rounded and densely setose throughout, setae as long as or shorter than paracuspis length. Digitus strongly incurved in dorsal view and upcurved in lateral view, apex subcapitate. Penis valve strongly concave on inner surface, with well-defined pair of short acute teeth posteroventrally, without apparent lateral pocket on outer surface, apical distance between teeth 0.1 × length of valve; strong setae present along shallowly convex posterior margin. Coloration and variations. Integument black to brownish-black, except for two dull reddish integumental spots on T2. Body setae predominantly black varying in density, except the following areas with silvery-white setae varying in density: metapleuron partiayll, lateral propodeal face, propodeal dorsum, metanotum medially, legs (except profemora), T1, anterior fourth of T2, fringe of T2 laterally, fringe of T3, fringe of T4–5 laterally, and S1–5 (except fringe of S5). Tibial spurs yellowish-white. Wings dark brown, without conspicuous reflections or darkened areas, veins discernible throughout. Distribution. Brazil, Argentina, and Uruguay. Material examined. (31m#) Type material. Lectotype (designated here): Mutilla vulnerata, m#, BRAZIL, [Rio Grande do Sul], St. [Santa] Cruz [do Sul], Hensel (ZMB). Additional material. BRAZIL: Rio Grande do Sul: 1#m, Krüger, R.F. (DZUP); Arroio do Sal, 1#m, 29.I.2008, A. Somavilla (CESC); Pelotas, 1#m, Biezanko (AMNH); Santa Cruz do Sul: JTI[Japanese Tobacco International]-Kannenberg: Malaise (137D) [sic!], 1#m, 01.I.2010, A. Köhler e equipe (CESC); Malaise (138D), 1#m, 25.XII.2009, A. Köhler e equipe (CESC); Malaise (139B), 1#m, 25.XII.2009, A. Köhler e equipe (CESC); Malaise (138D), 1#m, 01.I.2010, A. Köhler e equipe (CESC); Malaise (138B), 1#m, 25.XII.2009,A. Köhler e equipe (CESC); Malaise (138B), 1#m, 17.XII.2010, A. Köhler e equipe (CESC); JTI [Japanese Tobacco International] - Conv. [Convencional]: Malaise, Meio, 1#m, 27.XII.2013, A. Köhler e equipe (CESC); Malaise, Borda, 1#m, 19.XII.2013, A. Köhler e equipe (CESC); Malaise, Meio, 2#m, 19.XII.2013, A. Köhler e equipe (CESC); Malaise, Dentro, 2#m, 19.XII.2013, A. Köhler e equipe (CESC); Malaise, L2 [sic!], dentro, 1#m, 11.XII.2014, A. Köhler e equipe (CESC); Malaise, Borda, 1#m, 28.XII.2012, A. Köhler e equipe (CESC); Org. [Orgânico], Malaise L1 [sic!], borda, 1#m, 19.XII.2013, A. Köhler e equipe (CESC); Kannenberg, 1#m, 24.I.2012, A. Köhler (CESC); Xangri-lá, 1#m, 28.I.2008, J.T. Klein (CESC); ARGENTINA: [Buenos Aires], El Jabali, 1#m, 11.XI.1931, I.B. Anderson (BMNH); La Plata, 2#m (MNHN); Cordoba: Calamuchita, 1#m, Viana (AMNH); Miramar, 1#m, III.1992, Fritz (AMNH); Entre Rios: Palmar Colon, 1#m, XII.1974, Fritz (AMNH); P.N. [Parque Nacional] Colon, 1#m, I.1974 (AMNH); Pronunciamento, 1#m, Zelich (AMNH); Ubajay, 1#m, XII.1979, Fritz (AMNH); Santa Fé, Rosario, 1#m, Jos. Hubrich [sic!] (CISC); URUGUAY, Rio Negro, Arroyo Negro, 15km S [kilometers south og] Paysandu, 1#m, 27.XII.1963, R.G. van Gelder (AMNH). Remarks. Traumatomutilla vulnerata is the most distinct species of the T. aethiops subgroup due to T2 having two dark red integumental spots and its cuspis being conspicuously more sinuous than similar species. Like T. aehtiops, this will likely be associated with a species from the T. inermis species-group. A good candidate is T. tetrastigma, 252 which has been repeatedly collected in the same traps as males of T. vulnerata in tobacco fields in Santa Cruz do Sul, Rio Grande do Sul, Brazil, the type locality of this species (PRB pers. obs.). .
Updated Traumatomutilla species-groups construct.
Before the recent revisions of most Traumatomutilla species-groups (e.g. Bartholomay et al. 2018, 2019a, b) the genus was known from 183 species and subspecies, 133 of which were known from females only, 48 from males only, and only two known from both sexes (Williams et al. 2017). After recent new synonymies, new genera, new species, and new combinations, Traumatomutilla is currently comprised of 118 species, 71 known from females only, 15 from males only, and 32 known from both sexes (Table 1). This leaves the genus with 14 species-group, eight known from both sexes (references to each revision) and six known females only: T. inermis species-group with 31 species; T. trochanterata species-group with 19 species; T. integella and T. tabapua species-groups with 4 species each; and T. diabolica and T. pilkingtoni species-groups with only one species each. The 12 species treated above (known from males only) are not placed into a species-group at this time.
Table 1. Updated species-groups division for Traumatomutilla. “Females only” refers to taxa known only from females; “Males only” refers to taxa known only from males; “Both sexes” refers to taxa known from males and females. Species-group Species Females only Males only Both sexes americana T. americana (Linnaeus, 1758) X T. quadrum (Klug, 1821) X
T. ocellaris (Klug, 1821) X bellica T. bellica (Cresson, 1902) X T. virginalis (Gerstaecker, 1874) X bifurca T. bifurca (Klug, 1821) X T. oxira Casal, 1969 X diabolica T. diabolica (Gerstaecker, 1874) X gemella T. andrei (Cresson, 1902) X T. chuza Casal, 1969 X
T. diophtalma (Klug, 1821) X
T. gemella (André, 1906) X
T. peismatara Bartholomay & Cambra sp. nov. X indica T. aemulata (Cresson, 1902) X T. borba (Cresson, 1902) X
T. centralis (Burmeister, 1875) X
T. contempta André, 1908 X
T. geographica (Gerstaecker, 1874) X
T. grossa (Gerstaecker, 1874) X
T. guayaca Casal, 1969 X
T. impetuosa (Smith, 1879) X
T. indica (Linnaeus, 1758) X
T. ingens (André, 1903) X
T. mundula (Cresson, 1902) X
T. parallela (Klug, 1821) X
T. puella (Gerstaecker, 1874) X
T. protuberans (Gerstaecker, 1874) X
T. selligera (Gerstaekcer, 1874) X
T. spectabilis (Gerstaekcer, 1874) X
T. tristis (Klug, 1821) X
253
T. unimarginata (Cresson, 1902) X inermis T. aequinoctialis (Gerstaecker, 1874) X T. auxiliaris (Cresson, 1902) X
T. baguala Casal, 1969 X
T. caipira Casal, 1969 X
T. chapada (Cresson, 1902) X
T. chasca Casal, 1969 X
T. chilca Casal, 1969 X
T. confluens André, 1908 X
T. consimilis (André, 1895) X
T. coya Casal, 1969 X
T. crixa (Cresson, 1902) X
T. crona Casal, 1969 X
T. demissa (Cresson, 1902) X
T. dictynna (Cameron, 1895) X
T. gurisa Casal, 1969 X
T. hemicycla (Gerstaecker, 1874) X
T. inermis inermis (Klug, 1821) X
T. inermis affinis (Burmeister, 1854) X
T. ipanema (Cresson, 1902) X
T. ispiala Casal, 1969 X
T. laida Casal, 1969 X
T. lunigera (Gerstaecker, 1874) X
T. maipa Casal, 1969 X
T. moinga Casal, 1969 X
T. pereirai Suárez, 1960 X
T. pertela Casal, 1969 X
T. tetrastigma (Gerstaecker, 1874) X
T. trivirgata (Gerstaecker, 1874) X
T. vagabunda (Smith, 1879) X
T. vivax (Gerstaecker, 1874) X
T. zebrata (Gerstaecker, 1874) X integella T. barathra Bartholomay & Williams 2018 X T. integella (Cresson, 1902) X
T. pantherina Bartholomay & Williams 2018 X
T. poranga Bartholomay & Williams 2018 X juvenilis T. duplicata (Gerstaecker, 1874) X T. guarata Casal, 1969 X
T. juvenilis (Gerstaecker, 1874) X
T. juvenindica Bartholomay & Williams sp. nov. X
T. miniata (Gerstaecker, 1874) X pilkingtoni T. pilkingtoni Bartholomay & Williams, 2019b X quadrinotata T. ameliae Casal, 1969 X T. austera (Gerstaecker 1874) X
T. chrysozona (Gerstaecker, 1874) X
T. funebris (Gerstaecker, 1874) X
T. incerta (Spinola, 1841) X
T. quadrinotata (Klug, 1821) X
T. quadripustulata (Klug, 1821) X
254
T. pompiliformis (Gerstaecker, 1874) X
T. sancta (Gerstaecker, 1874) X
T. tetratrauma Bartholomay & Williams sp. nov. X
T. ursina (Gerstaecker, 1874) X
T. anhanga Bartholomay & Williams, 2018 X T. fratres Bartholomay & Williams, 2018 X tabapua T. luscoides André, 1908 X T. tabapua Casal, 1969 X trochanterata T. bellicosa (Cresson, 1902) X T. bellifera (Cresson, 1902) X
T. latona Mickel, 1952 X
T. lusca (Klug, 1821) X
T. moesta (Gerstaecker, 1874) X
T. ormena Casal, 1969 X
T. paraiba Casal, 1969 X
T. piasta Casal, 1969 X
T. preta Casal, 1969 X
T. rectilineata (André, 1898) X
T. tijuca Casal, 1969 X
T. triangulifera (André, 1906) X
T. trochanterata (Gerstaecker, 1874) X
T. trochanterata mirina Casal, 1969 X
T. trochanterata urupema Casal, 1969 X
T. tulumba Casal, 1969 X
T. valuta Casal, 1969 X
T. xiringa Casal, 1969 X
T. zayapa Casal, 1969 X
Unplaced species T. aethiops (Gerstaecker, 1874) X T. barra (Cresson, 1902) X
T. bartica Mickel, 1952 X
T. mesoleuca (Gerstaecker, 1874) X
T. orbana (Cresson, 1902) X
T. picada (Cresson, 1902) X
T. pomba (Cresso, 1902) X
T. rorida (Gerstaecker, 1874) X
T. soricina (Gerstaecker, 1874) X
T. taboca (Gerstaecker, 1874) X
T. tenuis (Smith, 1879) X
T. vulnerata (Gerstaecker, 1874) X
Totals 118 taxa 71 15 32
Discussion
Sex associations recently proposed for Traumatomutilla have provided solid evidence for the existence of morphological parallels between males and females. These characters can be effectively used to support sex association hypotheses, especially when coupled with distribution data and large series of specimens. For example, males of T. virginalis have conspicuously truncate femora which are one of the main characters of the females also. This particular character association was also observed between the putative couples involving the T. aethiops 255 subgroup and certain females of the T. inermis species-group. Another such character is the color of the meso- and metatibial spurs which aided in matching both sexes of T. centralis, T. puella, and T. ameliae. In terms of distribution data, existence of numerous males and females together in the same area is a useful tool in resolving sex associations. On the other hand, having samples from multiple locations is vital to ascertain if and how color and setae characters vary, intermediate color combinations serve as a basis for reconizing possible synonyms. Additionally, having at least one species known from both sexes can be the gateway to resolving most of, if not all, sex associations within a particular species-group. For example, provided that T. aethiops is in fact the male for T. inermis, it is likely that males morphologically similar to T. aethiops (i.e. T. mesoleuca, T. orbana, T. picada, T. soricina, and T. vulnerata) can be readily paired with females that share are morphologically similar to T. inermis. The T. inermis and T. trochanterata species-groups, with 31 and 19 species respectively, are still in need of revision and, as with the T. diabolica, T. integella, T. pilkingtoni, and T. tabapua species-groups, have no sex association. Evidence has already been found for different sex association hypotheses within the T. inermis species- group, including couples collected in copulo and the aforementioned distribution data for the T. aethiops subgroup. These hypotheses, however, can only be confirmed in the contex of a taxonomic revision for the species-group given its sheer number of species (now the largest in Traumatomutilla) and morphological diversity observed in the females, which suggests that male morphology will also vary. Putative males to be associated with the females of the T. integella and T. tabapua species-groups have been found in small numbers but, due to the rarity of both species- groups, none of these can be confidently associated with a specific female. Regarding the T. diabolica and T. pilkingtoni species-groups, even though both are somewhat limited in distribution to relatively well sampled areas (northern Argentina), no putative male has been found for either species. Finally, though there are putative males for the T. trochanterata species-group, none are described and as with the T. inermis species-groups preliminary data based on females suggests that male morphology may vary considerably. Additionally, the T. trochanterata species- group also contains certain species whose placement within the species-group and, in the case of T. valuta Casal, 1969, within the genus is uncertain. The species-group construct proposed by Williams et al. (2017) allowed for a compartmentalized study of Traumatomutilla, without which a revision of this complex and diverse genus would have been much more difficult and time consuming. In fact, a similar approach could prove to be equally helpful and effective with other large Neotropical genera of Mutillidae in need of revisionary work such as Hoplomutilla Ashmead, 1899, Pseudomethoca Ashmead, 1896, and Ephuta Say, 1836.
Acknowledgements
We are grateful for the collection managers and curators that provided specimens for this study, including: Christine LeBeau (AMNH), Andreas Köhler (CESC), John Rawlins (CMNH), Stephan Blank (DEI), Gabriel Melo (DZUP), James Pitts (EMUS), Agniéle Touret-Alby (MNHN), Orlando Silveira (MPEG), Gavin Broad (BMNH), Lynn Kimsey (UCDC), Robin Thomson (UMSP), Brian Harris (USNM), Michael Ohl, Viola Richter and Lukas Kirscher (ZMB). This project was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001, Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazônia (PPG-ENT), Project PRJ 12.10 Entomologia na Amazônia - Diversidade de Insetos of the Conselho Nacional de Pesquisas (CNPq). PRB was supported by the CNPq grant nº141158/2017-4 and CAPES PDSE grant nº88881.187031/2018-01. KAW was supported by CNPq’s Sciences Without Borders program (Complexos miméticos em vespas da família Mutillidae (Insecta, Hymenoptera): padrões de mimetismo e diversidade nos biomas brasileiros: Proceso 370106/2013-0). MLO is thankful for the CNPq productivity bursary, Brazil (306100/2016–9).
References
André. E. (1901) Descriptions de quelques espèces et variétés nouvelles de Mutilles d’Amerique appartenant au Museé Civique de Gênes. Zeitschrift für Hymenopterologie und Dipterologie. 1: 257–264. André, E. (1902) Fam. Mutillidae. In: Wytsman, P., Genera Insectorum, Fasc. 11. Bruxelles, 77 pp. André, E. (1904) Examen critique d’une nouvelle classification proposée par M. le Dr. W. H. Ashmead pour le famille des Mutillides. Revue d’Entomologie. 23(1): 27–41. 256
Ashmead, W.H. (1896) Description of new parasitic Hymenoptera. Transactions of the American Entomological Society, 23, 179–234. Ashmead, W.H. (1899) Superfamilis in the Hymenopter and generic synopses of the families Thynnidae, Myrmosidae, and Mutillidae. Journal of the New York Entomological Society, 7, 45–60. Bartholomay, P.R., Williams, K.A., Luz, D.R. & Morato, E.F. (2015) Frigitilla gen. nov., a new genus of Amazonian Mutillidae (Hymenotper). Zootaxa 3957 (1): 049–058. Bartholomay, P.R., Williams, K.A., Lopez, V.M. & Oliveira, M.L. (2019) Revsion of the Traumatomutilla americana species-group (Hymenopter, Mutillidae), Zootaxa, 4608 (1): 001–034. Brothers, D.J. (2006) Capítulo 14.4. Familia Mutillidae. In: Hanson, P.E. e Gauld, I.D. (Eds), Hymenoptera de la Región Neotropical. The American Entomological Institute, Gainesville, FL, pp. 586–593. Burmeister, H.C.C. (1854) Uebersicht der brasilianischen Mutillen. Abhandlungen der Naturforschenden Gesellschaft zu Halle, 2, 19–29. Casal, O.H. (1969) Sobre Traumatomutilla André (Hymenoptera, Mutillidae). Physis 28 (77): 279–298. Cresson, E.T. (1902) Descriptions of some Brasilian Mutilla. Transactions of the American Entomological Society 28: 1–82. Cresson, E.T. (1916) The Cresson types of Hymenoptera. Memoirs of the American Entomological Society 1: 79–85. Gerstaecker, A. (1874) Mutillarum Americae meridionalis indigenarum synopsis systematica et synonymica. Archiv für Naturgesichte 40: 41–77, 299–328. Klug, J.C.F. (1821) Entomologiae brasilianae specimen. – Nova Acta Academica Caesareae Leopoldino- Carolinae, 10 (2), 305-324. Mickel, C.E. (1952) The Mutillidae (wasps) of British Guiana. Zoologica, New York 37 (3): 105–150. Nonveiller G. (1990) Catalogue of the Mutillidae, Myrmosidae and Bradynobaenidae of the Neotropical Region including Mexico (Insecta, Hymenoptera). Hymenopterorum Catalogus (Nova Editio), 18. Den Haag, SPB Academic Publishing. 1–150. Say, T. (1836) Descriptions of new species of North American Hymenoptera, and observation on some already described. Boston Journal of Natural History, 1(3), 209–305. Smith, F. (1879) Descriptions of New Species of Hymenoptera in the Collection of the British Museum, London. 189– 227. Williams, K.A. (2012) Systematics of Mutillidae (Hymenoptera) with speciesl emphasis on Dasymutilla and their allies. All Graduate Theses and Dissertations. Paper 1200. Utah State University Press; Logan, UT, USA. 327 p. (Available at ~ http://digitalcommons.usu.edu/etd/1200/. Last accessed November 2018.) Williams, K.A., Bartholomay, P.R. & Oliveira, M.L. (2017) Species groups of Traumatomutilla André (Hymenoptera: Mutillidae). Insecta Mundi, 0533: 1-33.
Figure legends.
Figure 1 A–B. Traumatomutilla virginalis (Gerstaecker, 1874), male, line 2 mm; A. Lateral habitus; B. Dorsal habitus.
Figures 2 A–I. Traumatomutilla virginalis (Gerstaecker, 1874), male; A. Head, frontal view; B. Mesosoma, dorsal view; C. T7, posterodorsal view; D. S7, Ventral view; E. Genitalia (halved), dorsal view; F. Genitalia (halved), ventral view; G. Genitalia (halved, penis valve removed), lateral/inner view; H. Cuspis (removed, not to scale), lateral/inner view; I. Penis valve (removed, not to scale), lateral/outer view.
Figures 3 A–F. Traumatomutilla aethiops (Gerstaecker, 1874), lectotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
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Figures 4 A–F. Traumatomutilla barra (Cresson, 1902), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 5 A–F. Traumatomutilla bartica Mickel, 1952, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 6 A–F. Traumatomutilla mesoleuca (Gerstaecker, 1874), lectotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 7 A–F. Traumatomutilla orbana (Cresson, 1902), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 8 A–F. Traumatomutilla picada (Cresson, 1902), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 9 A–K. A. Traumatomutilla aethiops (Gerstaecker, 1874), lectotype, dorsal habitus, line 2 mm; B. Traumatomutilla barra (Cresson, 1902), holotype, dorsal habitus, line 2 mm; C. Traumatomutilla bartica (Mickel, 1952), dorsal habitus, line 2 mm; D. Traumatomutilla mesoleuca (Gerstaecker, 1874), lectotype, dorsal habitus, line 2 mm; E. Traumatomutilla orbana (Cresson, 1902), holotype, male, 2 mm; F. Traumatomutilla picada (Cresson, 1902), holotype, male, 2 mm; G–K. Type labels.
Figures 10 A–F. Traumatomutilla pomba (Cresson, 1902), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 11 A–F. Traumatomutilla rorida (Gerstaecker, 1874), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 12 A–F. Traumatomutilla soricina (Gerstaecker, 1874), holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 13 A–F. Traumatomutilla taboca (Cresson, 1902), lectotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 14 A–F. Traumatomutilla tenuis (Smith, 1879) comb. n., holotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 15 A–F. Traumatomutilla vulnerata (Gerstaecker, 1874), lectotype, male, 2 mm; A. Lateral habitus; B. Genitalia (halved), dorsal view; C. Genitalia (halved), ventral view; D. Genitalia (halved, penis valve removed), 258 lateral/inner view; E. Cuspis (removed, not to scale), lateral/inner view; F. Penis valve (removed, not to scale), lateral/outer view.
Figures 9 A–L. A. Traumatomutilla pomba (Cresson), holotype, dorsal habitus, line 2 mm; B. Traumatomutilla rorida (Gerstaecker, 1874), holotype, dorsal habitus, line 2 mm; C. Traumatomutilla soricina (Gerstaecker, 1874), dorsal habitus, line 2 mm; D. Traumatomutilla taboca (Cresso, 1902), lectotype, dorsal habitus, line 2 mm; E. Traumatomutilla tenuis (Smith, 1879), holotype, male, 2 mm; F. Traumatomutilla vulnerata (Gerstaecker, 1874), lectotype, male, 2 mm; G–L. Type labels. 259
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5. Considerações finais
Após as revisões da presente tese, apenas dois grupos de espécies de Traumatomutilla permanecem sem um estudo taxonômico aprofundado: o grupo de T. trochanterata (19 espécies) e o grupo de T. inermis (31 espécies). É importante ressaltar que todos os espécimes- tipo de ambos os grupos já foram estudados, fotografados e, em alguns casos, redescritos. Muitos casos de sinonímias também já foram encontrados para os dois grupos assim como importantes evidências para associações sexuais. Diversos gêneros neotropicais de Mutillidae, incluindo Traumatomutilla, possuem padrões morfológicos nos machos que de algum modo possuem contrapartidas nas fêmeas e vice-versa. Isso pôde ser observado ao longo das revisões de todos os grupos, sobretudo nos casos de T. miniata (Gerstaecker, 1874) onde ambos os sexos tem o mesosoma conspicuamente angulado em comparação às outras espécies do grupo de T. juvenilis; T. chrysozona (Gerstaecker, 1874) cujos machos e fêmeas possuem cerdas de tonalidade igual e exclusiva no metasoma em relação às demais espécies do grupo; T. virginalis (Gerstaecker, 1874) onde ambos os sexos possuem esculturação semelhante no dorso do mesosoma; e T. bifurca (Klug, 1821) cujas fêmeas possuem a carena dorsal da escroba projetada próximo ao tórulo e os machos possuem projeções dentiformes na fronte abaixo do ocello mediano. Tais paralelos morfológicos entre os sexos podem ser decisivos e, quando aliados a dados de distribuição, servir de base para associações sexuais a serem propostas na futura revisão dos grupos de T. inermis, T. trochanterata, assim como na descoberta dos machos dos grupos de T. diabolica, T. pilkingtoni, T. integella e T. tabapua. Infêrencias a respeito de possíveis associações sexuais também podem ser feitas com base em padrões de coloração de espécies simpátricas, pois os dados apresentados corroboram a existência de padrões de coloração típicos para ambos os sexos em determinadas regiões/biomas. Diversos casos foram observados em que machos e fêmeas de diferentes espécies (ou até mesmo diferentes gêneros) ocorrendo na mesma região possuem padrões de coloração virtualmente idênticos. Esses padrões são particularmente evidentes nos casos de T. juvenindica Bartholomay & Williams sp. nov., T. guayaca Casal, 1969, T. indica (Linnaeus, 1758) e T. parallela (Klug, 1821) na Floresta Amazônica; T. gemella André, 1906, T. guarata Casal, 1969, T. quadrinotata (Klug, 1821) e A. auriculata (Gertstaecker, 1874) na Mata Atlântica; e T. ameliae Casal, 1969, T. miniata (Gerstaecker, 1874), e T. ingens (André, 1903) nos ambientes xéricos do norte da Argentina. Tal relação de coloração entre os sexos provou-se ferramente importante para associação sexual entre espécimes coletados nas mesmas áreas além de contribuir como evidência para a existência de complexos de mimetismo Mulleriano envolvendo mutilídeos sul-americanos semelhantes aos casos observados na América do Norte e África (Wilson et al. 2013, 2018). 265
Os machos dos grupos de T. diabolica, T. pilkingtoni, T. integella e T. tabapua ainda não são conhecidos ou carecem de confirmação para uma associação definitiva. Devido à sua distribuição restrita a algumas áreas da Argentina, os machos de T. diabolica e T. pilkingtoni devem ser encontrados com relativa facilidade examinando material coletado nessas mesmas áreas. Tendo em vista os padrões estruturais e de coloração observados entre os sexos de diversas espécies de Dasymutillini na América do Sul e com base na morfologia das fêmeas, é plausível os machos de T. diabolica sejam estruturalmente semelhantes aos do grupo de T. indica, T. juvenilis ou T. quadrinotata, e possuam padrão de coloração semelhante ao de espécies como T. ameliae Casal, 1969 e T. miniata (Gerstaecker, 1874). No caso de T. pilkingtoni hipotetiza-se que os machos se assemelhem estruturalmente aos do grupo de T. americana com padrão de coloração semelhante ao de espécies como Cephalomutilla zelichi Casal, 1963 e Tobantilla ephemeros Williams, Brothers & Pitts, 2011. Outra relação morfológica observada entre os sexos diz respeito aos fêmures médios e posteriores. Apenas cinco grupos de espécie em Traumatomutilla possuem fêmeas com o ápice dos fêmures médios e posteriores truncados, sendo eles os grupos de T. inermis, T. integella, T. bellica, T. vitelligera e T. quadrinotata (vestigialmente truncados). Dentre estes, apenas os grupos de T. bellica, T. vitelligera e T. quadrinotata tem a morfologia dos machos conhecida e bem estabelecida. A ocorrência dessa característca nos grupos com ambos os sexos conhecidos indica que nem todas as fêmeas com fêmures truncados possuem machos com o a mesma característica. Entretanto, até o momento, todos os machos com fêmures truncados foram associados a fêmeas com o mesmo estado. Entre os machos não associados, mais da metade possui fêmures truncados e, presumivelmente, pertencem aos grupos de T. inermis ou T. integella. Com base nessa relação morfológica pode-se fazer inferências a respeito de associações entre tais machos e fêmeas dos grupos de T. inermis e T. integella. As espécies deste último — exceto a própria T. integella (Cresson, 1902) — são registradas até o momento apenas para a Amazônia Brasileira onde quatro machos não associados com fêmures truncados são conhecidos, T. bartica Mickel, 1952, T. orbana (Cresson, 1902), T. picada (Cresson, 1902) e T. tenuis (Smith, 1879). Hipoteticamente, T. picada e T. orbana pertencem ao grupo de T. inermis com base em características de genitália em comum com T. aethiops e T. vulnerata. Entre T. bartica e T. tenuis, esta última se assemelha superficialmente ao padrão morfológico das fêmeas do grupo de T. integella, possuindo o corpo de uma maneira geral mais esguio e com esculturação mais fina e homogênea, além de possuir o primeiro tergito metasomal mais evidentemente peciolado. Novamente valendo-se dos fêmures truncados em ambos os sexos, pode-se prever que os machos do grupo de T. inermis tendem a ser morfologicamente mais diversos do que nos demais grupos de espécie de Traumatomutilla. Isto porque o subgrupo de T. aethiops e os machos de T. bartica, T. rorida e T. taboca tendem a ser associados com espécies do grupo de 266
T. inermis com base nesse caráter observado em ambos os sexos. Enquanto que as espécies do subgrupo de T. aethiops são muito similares entre si inclusive em características de genitália, T. bartica, T. rorida e T. taboca possuem genitálias evidentemente distintas mesmo que sua morfologia externa seja semelhante. Ainda que a associação de T. bartica e T. taboca com fêmeas do grupo de T. inermis seja hipotetizada com base apenas na característica dos fêmures truncados, um casal coletado in copulo no estado do Mato Grosso, Brasil fornece fortes evidências para a associação de T. rorida com fêmeas deste grupo. A fêmea do casal em questão aparenta ser próxima a T. laida Casal, 1969 (provavelmente uma variaçãode coloração) e o macho é estruturalmente similar a T. rorida (Gerstaecker, 1874). Mais importante é o fato de que o macho possui a valva do penis, parâmeros e paracuspis virtualmente idênticos aos de T. rorida, diferindo na forma no dígito, e especialmente no aspecto lateral da cuspis, muito mais estreita do que em T. rorida. Essas similaridades e diferenças indicam que T. rorida provavelmente venha a ser reconhecida como o macho de alguma espécie do grupo de T. inermis, supostamente uma fêmea próxima à T. laida. Sendo este o primeiro estudo a examinar, descrever e ilustrar extensamente a genitália de Traumatomutilla, é importante ressaltar que aparentemente o isolamento reprodutivo entre espécies do gênero não se dá pelo sistema de chave/fechadura entre os aparelhos reprodutores de ambos os sexos. Os machos da maioria das espécies tratadas aqui não possuem características de genitália exclusivas, sobretudo dentro dos grupos de espécies. Em muitos casos, machos com morfologia externa conspicuamente distinta possuem características de genitália quase idênticas, tal qual os casos entre T. vidua e T. parallela, assim como T. ingens e T. grossa. Em contrapartida, também foram observados casos em que machos com características externas pouco distintas possuem morfologia externa conspicuamente diferente, como no caso de T. juvenindica e T. juvenilis. Situações similares também ocorrem ao se comparar a morfologia de ambos os sexos de diferentes espécies dentro do mesmo grupo de espécies. Frequentemente, espécies na quais as fêmeas são pouco distintas possuem machos marcadamente diferentes em várias características externas e de genitália. Isso pode ser observado com maior evidência no grupo de T. gemella como um todo no qual as fêmeas são distinguidas predominantemente com base em características de coloração e cerdas enquanto que os machos possuem diferenças claras em estruturas e esculturação. Ao mesmo tempo também foram observados casos em que machos virtualmente idênticos em estrutura são pareados com fêmeas com diferenças estruturais claras como observado entre T. ameliae, T. chrysozona e T. quadrinotata. Esse fenômeno é relativamente frequente em diversos outros gêneros de Mutillidae, ao ponto de que grupos como Odontophotopsis Schuster, Sphaeropthalma Blake e outros gêneros noturnos serem compostos por espécies cujos machos são facilmente separados enquanto que as fêmeas não possuem quaisquer diferenças observáveis (Pitts et al 2007; Williams, 2012). O contrário também já foi 267
observado em gêneros diurnos da região Neotropical como Pseudomethoca, no qual as fêmeas geralmente podem ser separadas por características estruturais distintas enquanto que os machos, quando distinguíveis, só podem ser separados com base em características de genitália (obs. pes.). Essa constante disparidade interspecífica entre machos e fêmeas dentro dos mesmos grupos de espécie também pode ser observada nos padrões de coloração. O melhor exemplo dessa situação é o que foi observado nos grupos de T. gemella e T. quadrinotata, nos quais a maioria das fêmeas possuem diferenças de coloração marcantes e aparentemente bem definidas, enquanto que os machos possuem modificações mínimas nos padrões de cerdas apenas. Por outro lado, em grupos como o de T. americana, os machos são facilmente separados por características de coloração e cerdas enquanto que as fêmeas de determinadas áreas possuem padrões semelhantes ou virtualmente idênticos. Tanto as dissimilaridaes quanto as semelhanças estruturais e/ou de coloração indicam a ocorrência do fenômeno de Dual Sex-Limited Mimicry (mimetismo duplo limitado pelo sexo) proposto or Evans (1968, 1969) no qual machos e fêmeas de uma mesma espécie participam de complexos miméticos com modelos distintos. Williams (2012) encontrou padrões semelhantes entre os sexos do grupo de Dasymutilla gloriosa nos desertos norte-americano. O mesmo autor sugeriu que a diferença no mimetismo de machos e fêmeas se devia a uma maior pressão de seleção sobre as fêmeas, supostamente por uma maior exposição das mesmas a predadores e variáveis climáticas por sua condição áptera. É possível que um mecanismo de seleção semelhante esteja atuando sobre as espécies de Traumatomutilla resultando nas diferenças e similaridades estruturais entre machos e fêmeas dentro dos mesmos grupos de espécie. Além das revisões dos grupos de T. trochanterata e T. inermis, juntamente e a procura pelos machos dos grupos de T. diabolica, T. pilkingtoni, T. integella e T. tabapua, futuros estudos com Traumatomutilla devem focar nas relaçãoes filogenéticas entre as espécies que compõem o gênero e sua relação com Dasymutilla Ashmead. Além dos novos gêneros aqui propostos, outras espécies ou grupos de espécie de Traumatomutilla podem vir a ser considerados como gêneros distintos, o que também já foi observado para Dasymutilla (obs. pes.). Até o momento foram encontrados outros dois supostos novos gêneros neotropicais cuja morfologia se confunde parcialmente com Traumatomutilla. A redução drástica no número de espécies de Traumatomutilla aliada à pouca diversidade morfológica encontrada nas espécies que compõem o gênero indica que a utlização de ferramentas moleculares juntamente com o estudo da morfologia será indispensável para elucidar as relações entre os Dasymutillini Sul- Americanos e seus congêneres na América do Norte.
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