Instituto Nacional De Pesquisas Da Amazônia Coordenação De Biodiversidade Divisão De Entomologia - Dient

INSTITUTO NACIONAL DE PESQUISAS DA AMAZÔNIA COORDENAÇÃO DE BIODIVERSIDADE DIVISÃO DE ENTOMOLOGIA - DIENT

SISTEMÁTICA DE Kapala Cameron, 1884 (HYMENOPTERA: CHALCIDOIDEA: EUCHARITIDAE)

KARINE SCHOENINGER

MANAUS, AMAZONAS ABRIL DE 2018

KARINE SCHOENINGER

SISTEMÁTICA DE Kapala Cameron, 1884 (HYMENOPTERA: CHALCIDOIDEA: EUCHARITIDAE)

Orientador: Dr. Marcio Luiz de Oliveira (INPA) Co-orientadores: Dr. John Heraty (University of California – Riverside, EUA) Dr. Javier Torréns (CRILAR – Anillaco, Argentina)

Tese apresentada ao Instituto Nacional de Pesquisas da Amazônia, como parte dos requisitos para obtenção do título de Doutora em Ciências Biológicas, área de concentração em Entomologia.

MANAUS, AMAZONAS ABRIL DE 2018 iii

Tese aprovada como requisito para a obtenção do título de Doutora em Ciências Biológicas, área de concentração em Entomologia, no Programa de Pós-Graduação em Entomologia do Instituto Nacional de Pesquisas da Amazonia, pela comissão formada pelos doutores (as):

Drª. Rosaly Ale Rocha Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular

Dr. Daniel Rodrigo R. Fernandes Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular

Dr. Danilo Pacheco Cordeiro Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular

Dr. Edinaldo Nelson dos S. Silva Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular

Dr. Ronildo Baiatone Alencar Instituto Nacional de Pesquisas da Amazônia (INPA) Membro titular

S365 Schoeninger, Karine

Sistemática de Kapala Cameron, 1884 (Hymenoptera: Chalcidoidea: Eucharitidae) / Karine Schoeninger. - Manaus: [sem editor], 2018. xxi, 280f.: il. color

Tese (Doutorado) - INPA, Manaus, 2018. Orientador: Marcio L. Oliveira. Coorientadores: John Heraty, Javier Torréns Área de concentração: Entomologia.

1. Eucharitinae. 2. Complexo furcata. 3. Parasitoides de formigas.. I. Título

CDD 595.79

Sinopse: Foi realizada a revisão taxonômica, bem como um estudo filogenético com base em dados morfológicos e moleculares para o gênero Kapala , com ênfase nos complexos de espécies iridicolor e furcata. Foram descritas 15 espécies novas , realizadas 4 sinonímias, designados três léctótipos e realizadas sete associações sexuais. Dois novos gêneros são propostos, Ecariatana n. gen. e Torquata n. gen.

Palavras-chave: Eucharitinae, complexo furcata, complexo iridicolor, Região Neotropical, parasitoides de formigas

AGRADECIMENTOS

Ao Instituto Nacional de Pesquisas da Amazônia (INPA) e ao Curso de Pós-Graduação em Entomologia, pela infraestrutura e oportunidade de realização do doutorado deste projeto. À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior pelo primeiro ano de concessão de bolsa de doutorado sanduíche (processo PDSE - 88881.133943/2016-01). Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), pelos três últimos anos de concessão de bolsa. Ao meu orientador Marcio L. Oliveira pela orientação e suporte ao longo destes últimos anos. Gostaria de agradecer imensamente aos Drs. John Heraty e Javier Torréns pela co- orietação e apoio ao longo destes quatro anos de doutorado. O zelo de ambos pelo mundo natural e o seu amor por Eucharitidae são aparentes em todos os seus trabalhos. Vocês me encorajaram a pensar de maneira mais criativa e a estabelecer metas elevadas. Eu aprendi muito nos últimos anos com vocês. A cada curador das 52 coleções que enviaram material de estudo e que proporcionaram a realização deste trabalho, em especial a curadora do The Natural History Museum, Dra. Natalie Dale-Skey, por toda a sua ajuda e compreensão. Aos amigos do INPA e colegas do Laboratório de Hymenoptera: Pedro, Breno, Alexandre, Diego, David, Sian, Bruno, Itanna, Thiago, Jeane, Patrik, Nikolas, Alexssandro, Ísis, Daniara, Thaís, Daniel, pelo companheirismo, discussões eternas sobre trabalho e, principalmente filogenia, e pelos momentos de descontração. Desde já, aos membros da banca pelas contribuições e sugestões. Um agradecimento especial aos meus pais, Artélio e Marilene, pelo amor contante e pelo apoio, tanto nos momentos felizes quanto nas dificuldades. Por terem compreendido a minha ausência durante todos esses anos (seis anos até o momento e sei que não foi fácil). Sem vocês eu não teria chegado até aqui. Amo vocês! Minha gratidão sincera a todo o pessoal que me incentivou nesta tarefa. E a todos os que, indiretamente colaboraram para que eu pudesse terminá-la. Obrigada.

RESUMO

Kapala Cameron, 1884 é amplamente difundido em toda a região Neotropical, com exceção do Chile e uma espécie disjunta que é encontrada na África Central e Madagascar. A necessidade de uma revisão das espécies de Kapala foi observada a partir de suas descrições originais, que são extremamente sucintas, o que resultou em muitas identificações errôneas. A grande maioria das descrições menciona a coloração, que é extremamente variável; a presença de dois longos espinhos, presença de antenas flabeladas e mesoscuto alto e convexo. Essas características são comuns a todas as espécies tornando a identificação de suas espécies quase impossível. Apesar disso, a maior dificuldade encontrada dentro do grupo é a sua alta variação instraespecífica, mas ao mesmo tempo altamente conservada. Esta é a primeira tentativa de revisar as espécies conhecidas, descrever novos táxons e estabelecer uma linha de base para a taxonomia do gênero. Além disso, uma nova proposta filogenética para Kapala é fornecida. No Capítulo I, com base no exame dos espécimes-tipo e material complementar, redescrevemos as espécies-tipo do gênero e estabelecemos os limites das espécies válidas; quatro sinônimos foram realizados, K. atrata (Walker, 1862) sin. nov. e K. surgens (Walker, 1862) sin. nov. foram sinonimizadas com K. flabellata (Fabricius, 1804), K. striaticeps (Cameron, 1913) sin. nov. foi sinonimizada com K. inexagens (Walker, 1862) e K. sulcifacies (Cameron, 1904) sin. nov. foi sinonimizado com K. romandii (Guérin-Meneville, 1845); três lectótipos foram designados: K. cynipsea (Walker, 1862) pres. desig., K. inexagens (Walker, 1862) pres. desig. e K. terminalis (Ashmead, 1892) pres. desig., e sete espécies tiveram machos associados a fêmeas. Sete novas espécies são descritas: K. confusa n. sp., K. corcovata n. sp., K. genistriata n. sp., K. gracilispina n. sp., K. haplospinosa n. sp., K. jalisca n. sp. e K. spinaepplanata, e uma chave de identificação para todas as espécies é fornecida. No capítulo II, um estudo filogenético com base em dados morfológicos e moleculares foi realizado com o objetivo de testar a monofilia de Kapala e as relações entre as suas espécies. Para este propósito, 61 caracteres relacionados à morfologia externa foram analisados. Para as análises moleculares, partes de cinco regiões de genes foram amplificadas: 18S, 28S-D2, 28S-D3-D5, COI-nj e COII. Como resultado, foi verificado em todas as análises, morfológica e molecular, que Kapala é parafilético, dividido em três complexos de espécies, que por sua vez, estão divididos por agrupamentos de gêneros distintos. No capítulo III, foi realizada a revisão do complexo de espécies iridicolor, bem como uma filogenia molecular vii

com base em 5 genes, 3 nucleares e dois ribossomais. Como resultados, foram reconhecidas seis espécies dentro do complexo iridicolor, incluindo K. iridicolor descrita por Cameron (1904) e cinco novas espécies, sendo elas: K. cavicornis n. sp., K. ceciliae n. sp., K. ipa n. sp., K. longicornis n. sp. e K. parairidicolor n. sp. Além disso, uma diagnose do complexo K. iridicolor é fornecida, bem como uma chave de identificação para as espécies. No capítulo IV, foi realizada a revisão do complexo de espécies furcata e, também, uma filogenia molecular, sendo reconhecidas três novas espécies: K. deltalis n. sp., K. parafurcata n. sp. e K. quasimodo n. sp. Além disso, uma diagnose para o complexo furcata é fornecida, bem como uma chave de identificação para as suas espécies. No capítulo V, como base na filogenia morfológica e molecular, dois novos gêneros são descritos para a família Eucharitidae, Ecarinata n. gen. e Torquata n. gen., sendo fornecida uma chave revisada e atualizada para os gêneros Neotropicais de Eucharitidae.

Palavras-chave. Eucharitinae, complexo furcata, complexo iridicolor, neotropical, parasitoides de formigas.

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ABSTRACT

Kapala Cameron, 1884 is widespread throughout the Neotropical region, except for Chile and a disjunct species is found in Central Africa and Madagascar. The need for a review of the Kapala species was observed from their original descriptions, which are extremely succinct, resulting in many misidentifications. The vast majority of descriptions mention coloration, which is extremely variable; the presence of two long spines, flabellate antennas in males and mesoscuto high and convex. These characteristics are common to all species making identification of their species almost impossible. Despite this, the greatest difficulty found within the group is its high intraspecific variation, but at the same time highly conserved. This is the first attempt to revise known species, to describe new taxa, and to establish a baseline for genus taxonomy. In addition, a new phylogenetic proposal for Kapala is provided. In Chapter I, based on the examination of the species-type and complementary material, we redescribe the type species of the genus and establish the limits of valid species; four synonyms were performed, K. atrata (Walker, 1862) n. syn. and K. surgens (Walker, 1862) n. syn. was synonimized with K. flabellata (Fabricius, 1804), K. striaticeps (Cameron, 1913) n. syn. was synonymized with K. inexagens (Walker, 1862) and K. sulcifacies (Cameron, 1904) n. syn. was synonymized with K. romandii (Guérin- Meneville, 1845); three lectotypes were designated: K. cynipsea (Walker, 1862) pres. desig., K. inexagens (Walker, 1862) pres. desig. and K. terminalis (Ashmead, 1892) pres. desig., and seven species had males associated with females. Seven new species are described: K. confusa n. sp., K. corcovata n. sp., K. genistriata n. sp., K. gracilispina n. sp., K. haplospinosa n. sp., K. jalisca n. sp. e K. spinaepplanata, and an identification key for all species is provided. In Chapter II, a phylogenetic study based on morphological and molecular data was performed with the objective of testing Kapala monophyly and the relationships among its species. For this purpose, 61 characters related to external morphology were analyzed. For the molecular analyzes, parts of five gene regions were amplified: 18S, 28S-D2, 28S-D3-D5, COI-nj and COII. As a result, it was verified in all morphological and molecular analyzes that Kapala is paraphyletic, divided into three species complexes, which in turn are divided by groups of different genera. In chapter III, the iridicolor species complex was reviewed, as well as a molecular phylogeny based on 5 genes, 3 nuclear and 2 ribosomal genes. Six species were recognized within the iridicolor complex, including K. iridicolor described by Cameron (1904) and five new species: K. cavicornis n. sp., ix

K. ceciliae n. sp., K. ipa n. sp., K. longicornis n. sp. and K. parairidicolor n. sp. In addition, a diagnosis of the K. iridicolor complex is provided, as well as an identification key for the species. In chapter IV, a review of the furcata species complex and a molecular phylogeny was performed, with three new species being recognized: K. deltalis n. sp., K. parafurcata n. sp. and K. quasimodo n. sp. In addition, a diagnosis for the furcata complex is provided, as well as a key identification for its species. In Chapter V, as a basis in morphological and molecular phylogeny, two new genera are described for the family Eucharitidae, Ecarinata n. gen. and Torquata n. gen. A revised and updated key is provided for the Neotropical genera of Eucharitidae.

Keywords. Ants parasitoids, Eucharitinae, furcata complex, iridicolor complex, neotropical.

SUMÁRIO

1 Introdução geral ...... 1 1.1 Aspectos biológicos de Eucharitidae ...... 1 1.2 Aspectos taxonômicos de Eucharitidae ...... 3 1.3 Kapala Cameron, 1884 ...... 5 Capítulo I ...... 8 Introduction ...... 10 Material and methods ...... 11 Morphological descriptions and redescriptions ...... 11 Measurements ...... 12 Specimen imaging ...... 14 Specimens repository ...... 14 Results ...... 15 Taxonomy ...... 15 Kapala Cameron, 1884 ...... 16 Key to Kapala species ...... 17 K. furcata species complex ...... 21 Kapala cuprea Cameron, 1913 ...... 22 Kapala furcata (Fabricius, 1804) ...... 26 Kapala splendens Ashmead, 1904 ...... 29 K. iridicolor species complex ...... 32 Kapala iridicolor (Cameron, 1904) ...... 32 K. romandii species complex ...... 41 Kapala argentina Gemignani, 1933 ...... 42 Kapala chacoensis Gemignani, 1947 ...... 46 Kapala cynipsea (Walker, 1862) ...... 48 Kapala flabellata (Fabricius, 1804) ...... 50 Kapala floridana (Ashmead, 1885) ...... 57 Kapala inexagens (Walker, 1862) ...... 63 xi

Kapala ivorensis Risbec, 1954 ...... 66 Kapala izapa Carmichael, 2006 ...... 70 Kapala romandii (Guérin-Meneville, 1845) ...... 74 Kapala terminalis Ashmead, 1892 ...... 81 New species descriptions ...... 84 Kapala corcovata Schoeninger n. sp...... 84 Kapala genistriata Schoeninger n. sp...... 87 Kapala gracilispina Schoeninger n. sp...... 89 Kapala haplospinosa Schoeninger n. sp...... 91 Kapala jalisca Schoeninger n. sp...... 94 Kapala spinaepplanata Schoeninger n. sp...... 96 Species with uncertain position ...... 98 Kapala dicerodera (Spinola, 1853) ...... 98 Kapala confusa Schoeninger n. sp...... 99 Acknowledgment ...... 101 References ...... 101 Capítulo II ...... 129 Introdução ...... 131 Material e métodos ...... 133 Dados morfológicos ...... 133 Escolha dos terminais ...... 133 Morfologia e análises dos caracteres ...... 133 Dados moleculares ...... 135 Análises moleculares ...... 136 Parcimônia ...... 136 Máxima Verossimilhança (ML) ...... 137 Resultados ...... 137 Análise de caracteres morfológicos ...... 137 Análise cladística morfológica ...... 158 Análise molecular: parcimônia ...... 161 Análise molecular: máxima verossimilhança ...... 162 xii

Discussão ...... 165 Conclusão ...... 167 Agradecimentos ...... 167 Referências bibliográficas ...... 168 Capítulo III ...... 180 Introduction ...... 182 Materials and methods ...... 183 Molecular phylogenetic analysis ...... 183 Taxonomy ...... 184 Morphological descriptions ...... 184 Measurements ...... 184 Specimen imaging ...... 185 Specimens repository ...... 185 Results and discussion ...... 186 Phylogeny ...... 186 Taxonomy ...... 187 Kapala Cameron, 1884 ...... 187 Kapala iridicolor species complex ...... 188 Key to the species of the K. iridicolor complex ...... 188 Kapala iridicolor (Cameron, 1904) ...... 189 Kapala cavicornis Schoeninger & Heraty, n. sp...... 195 Kapala ceciliae Schoeninger & Torréns, n. sp...... 197 Kapala ipa Schoeninger & Heraty, n. sp...... 200 Kapala longicornis Schoeninger & Oliveira, n. sp...... 202 Kapala parairidicolor Schoeninger & Torréns n. sp...... 204 Acknowledgment ...... 207 References ...... 207 Capítulo IV ...... 220 Introduction ...... 222 Material and methods ...... 222 Molecular phylogenetic analysis ...... 222 xiii

Taxonomy ...... 223 Morphological descriptions ...... 223 Measurements ...... 224 Specimen imaging ...... 224 Specimens repository ...... 225 Results and discussion ...... 226 Phylogeny ...... 226 Taxonomy ...... 226 Kapala furcata species complex ...... 227 Key of identification to Kapala furcata species complex ...... 228 Kapala cuprea Cameron, 1913 ...... 229 Kapala furcata (Fabricius, 1804) ...... 231 Kapala splendens Ashmead, 1904 ...... 233 New species descriptions ...... 234 Kapala deltalis Murray & Schoeninger, n. sp...... 234 Kapala parafurcata Murray & Schoeninger, n. sp...... 237 Kapala quasimodo Murray & Schoeninger, n. sp...... 241 Acknowledgment ...... 243 References ...... 244 Capítulo V ...... 253 Introduction ...... 254 Materials and methods ...... 256 Morphological descriptions ...... 256 Measurements ...... 256 Specimen imaging ...... 256 Specimens repository ...... 257 Results ...... 257 Revised key for the Neotropical genera of Eucharitidae ...... 257 Taxonomy ...... 267 Ecarinata Schoeninger & Torréns, n. gen...... 267 Torquata Schoeninger & Murray n. gen...... 269 xiv

Acknowledgment ...... 272 SÍNTESE ...... 276 Referências bibliográficas ...... 277

LISTA DE TABELAS

Capítulo II ...... 150

Tabela S1. Lista dos gêneros e espécies do grupo externo e interno utilizados na análise filogenética morfológica, região biogeográfica, país, museu depositário e observações (Obs.). AFRO = Afrotropical; OCE = Oceania; NEO = Neotropical; GE = Grupo externo; GI = Grupo interno; CF = complexo K. furcata; CI = complexo K. iridicolor; CS = complexo K. romadii 192 Tabela S2. Lista dos gêneros e espécies do grupo externo e interno utilizados na análise filogenética molecular, sexo, número de identificação de depósito (Numb. ID), DNA voucher, museu de deposito e localidade de coleta ...... 194 Tabela S3. Lista dos gêneros e espécies do grupo externo e interno utilizados na análise filogenética molecular, sexo, número de identificação de depósito (Numb. ID), DNA voucher, museu de deposito e localidade de coleta ...... 197 Tabela SI 4. Primer oligonucleotídeos utilizados neste estudo. Sequencias marcadas com “*” foram modificadas a partir das referências da publicação original ...... 200

Capítulo III ...... 201

Table S1. Locality, museum, identification number (ID) and DNA voucher number information for the 35 taxa of iridicolor complex and outgroup (furcata complex) used in the molecular analysis ...... 237 Table S2. Taxa, identification number, DNA voucher number information and the respective genes used for molecular analysis ...... 239

Capítulo IV ...... 241

Table S1. Locality, museum, identification number (ID) and DNA voucher number information for the 35 taxa of furcata complex and outgroup (iridicolor complex) used in the molecular analysis ...... 272 Table S2. Taxa, identification number, DNA voucher number information and the respective genes used for molecular analysis ...... 273

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LISTA DE FIGURAS

Introdução geral ...... 22

Figura 1. Associações taxonômicas hospedeiro-parasitoide. Tanglegrama simplificado de uma análise de 29 gêneros de Eucharitidae e 23 gêneros de Formicidae. Linhas de interação finas indicam a utilização de apenas um gênero como hospedeiro e linhas grossas. Fonte: adatado de Murray et al. (2013) ...... 26

Capítulo I ...... 29

Figure 1. Kapala spp. A. head ♀ (frontal view); antenna ♀ (lateral view); C. antenna ♂ (lateral view) ...... 126 Figure 2. Kapala flabellata ♂. D. mesosoma (lateal view); E. mesosoma (dorsal view); F. habitus; G. forewing ...... 127 Figure 3. Specimen measurements. A. Head (frontal view); B. antenna (lateral, ♂); C. antenna (lateral, ♀); D. habitus (lateral, ♀; E. habitus (dorsal, ♀); F. Forewing ...... 128 Figure 4. K. cuprea: A. Habitus (lateral, ♂); B. head (front, ♂); C. antenna (lateral, ♀); D. Habitus (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution ...... 129 Figure 5. K. furcata: A. habitus (lateral, ♀); B. front (head, ♀); C, antenna (lateral, ♀); D. front (head, ♂); E. habitus (lateral, ♂); G. habitus (dorsal, ♂); H. distribution ...... 130 Figure 6. K. splendens: A. habitus (lateral, ♀); B. antenna (lateral, ♀); C. head (frontal, ♀); D. habitus (dorsal, ♀); E. habitus (lateral, ♂); F. distribution ...... 131 Figure 7. K. iridicolor: A. habitus (lateral, ♀); B. head (front, ♀); C. antenna (lateral, ♀); D. habitus (lateral, ♂); E. habitus (dorsal, ♀); F. habitus (dorsal, ♂). G. distribution. Red setae = indicating the mesonotum smoothly curved ...... 132 Figure 8. K. argentina: A. habitus (lateral, ♀); B. head (front, ♀); C. mesosoma (lateral, ♀); D. head (front, ♂); E. habitus (lateral, ♂); F. habitus (dorsal, ♀); G. habitus (dorsal, ♂); H. distribution ...... 133 Figure 9. K. chacoensis: A. habitus (lateral, ♂); B. head (front, ♂); C. head (front, ♀); D. mesoscutum (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution ...... 134 Figure 10. K. cynipsea: A. habitus (lateral, ♀); B. head (front, ♀); C. distribution ...... 135 Figure 11. K. flabellata: A. habitus (lateral, ♂); B. head (front, ♂); C. head (front, ♀); D. mesosoma (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution ...... 136 Figure 12. K. floridana: A. habitus (lateral, ♂); head (front, ♂); C. mesosoma (lateral, ♂); D. antenna and mesosoma (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution. Red setae = Mesoscutellar disc not elevated above base of frenal process ...... 137 Figure 13. K. inexagens: A. habitus (lateral, ♀); B. head (front, ♀); C. habitus (dorsal, ♀); D. antenna (lateral, ♀); E. distribution ...... 138 Figure 14. K. ivorensis: A, habitus (lateral, ♀); B. head (front, ♀); C. antenna and mesosoma (lateral, ♀); D. habitus (lateral, ♂); E. habitus (dorsal, ♀); F. habitus (dorsal, ♂); G. distribution ...... 139 xvii

Figure 15. K. izapa: A. habitus (lateral, ♂); B. head (frontal, ♂); C. mesosoma (lateral, ♂); D. habitus (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution ...... 140 Figure 16. K. romandii: A. habitus (lateral, ♂); B. head (front, ♂); C. antenna (lateral, ♀); D. head (frontal, ♀); E. habitus (dorsal, ♀); F. mesosoma (lateral, ♂); G. habitus (dorsal, ♂); H. distribution ...... 141 Figure 17. K. terminalis: A. habitus (lateral, ♂); B. head (front, ♂); C. mesosoma (lateral, ♂); D. antenna (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution. Red setae = mesoscutellar disc not elevated above base off renal process ...... 142 Figure 18. K. corcovata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsolateral, ♂); D. mesosoma (front, ♀); E. habitus (lateral, ♀); F. distribution ...... 143 Figure 19. K. genistriata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma (lateral, ♂); E. distribution ...... 144 Figure 20. K. gracilispina n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma, ♂); E. distribution ...... 145 Figure 21. K. haplospinosa n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. head and mesosoma (laterofrontal, ♀); E. distribution ...... 146 Figure 22. K. jalisca n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma (lateral, ♂); E. distribution ...... 147 Figure 23. K. spinaepplanata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. frenal processes (dorsal, ♂); D. mesosoma (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution ...... 148 Figure 24. K. confusa n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesososma (lateral, ♂); E. distribution. Red setae = mesoscutellar disc elevated like a keel-shape ...... 149

Capítulo II ...... 150

Figura 1. Cerdas nos olhos compostos. (A) cerdas ausentes (Kapala flabellata); B. cerdas presentes (Lirata spinaelevis) ...... 159 Figura 2. Formato do olho composto. A. globular e convexo (Kapala flabellata); B. arredondado e pronunciado (Lirata spinaelevis n. sp.); C. com protuberância em forma de “mamilo” (Isomerala coronata) ...... 159 Figura 3. Depressão escrobal. A. depressão escrobal rasa (Colocharis hungi); B. profunda com as margens laterais abruptamente arredondadas (Isomerala coronata) ...... 160 Figura 4. Carena entre o ocelo mediano e ocelos laterais. A. carena ausente (Kapala iridicolor); B. carena presente (Neolirata daguerrei) ...... 161 Figura 5. Área supraclipeal (indicada). A. sulco lateral indistinto (Liratella sp.); B. sulco lateral crenulado (Colocharis hungi) ...... 161 Figura 6. Área supraclipeal (indicada), padrão do tegumento. A. liso (Torquata n.gen.); B. estriado (Kapala haplospinosa n. sp.); C. rugoso (Lirata pustula); D. pontuado (Dilocantha sp.) ...... 162 Figura 7. Padrão do tegumento da fronte. A. estrias grossas, as quais são separadas uma das outras (indicado); B. estrias finas, próximas umas das outras (indicado) ...... 163 Figura 8. Padrão das estrias da fronte. A. longitudinal. B. oblíquo (do triângulo ocelar em direção à margem inferior do olho); C. oblíquo (do ocelo lateral para a margem inferior do torulus) ...... 164 Figura 9. Fronte. A. inchada (Kapala longicornis n. sp.); B. achatada (Kapala flabellata) ..... 164 xviii

Figura 10. Margem posterolateral da gena. A. estreita, com a margem posterolateral arredondada (Kapala argentina); B. larga e angulada medialmente (indicado), sem carena na margem posterior (Isomerala coronata) ...... 165 Figura 11. Comprimento do escapo. A. não atingindo o ocelo médio (Parakapala decarloi); B. atingindo o ocelo médio (Lirata spinaelevis n. sp.) ...... 166 Figura 12. Formato de Fl2 da fêmea. A. flagelômeros simples/cilíndricos (Dilocantha sp.).; B. serreado (Ecarinata n. gen.); C. serreado com o ápice terminando em uma ponta afiada (Lirata diabla); D. ramificado, alongado (Lasiokapala serrata) ...... 167 Figura 13. Mesoescutelo em vista lateral. A. baixo, não projetado sobre a cabeça (Chalcura sp.); B. vertical, projetado sobre a cabeça e com a margem anterior suavemente curvada (Kapala ceciliae n. sp.); C. vertical, projetado sobre a cabeça e com margem anterior abruptamente curvada na metade dorsal (Kapala argentina) ...... 169 Figura 14. Lóbulos laterais do mesoescuto, em vista lateral. A. elevado a aproximadamente a altura do lóbulo médio; B. não elevado à altura do lóbulo médio ...... 169 Figura 15. Lóbulos laterais do mesoescuto em vista frontal. A. não projetado (Kapala parafurcata n. sp.); B. projetado (Kapala deltalis n. sp.) ...... 170 Figura 16. Flange do mesoescutelo. A. flange muito fina ao longo da margem da tégula (indicado) (Parakapala decarloi); B. flange triangular que se projeta posteriormente do canto posterolateral do mesoescuto sobre a tégula (indicado) (Lirata pustula) ...... 171 Figura 17. Formato do mesoescutelo em vista lateral. A. achatado (Kapala inexagens); B. inchado/convexo (Kapala terminalis) ...... 171 Figura 18. Margem medial posterior do disco mesoescutelar em vista lateral. A. arredondada (Chalcura sp.); B. terminando em um ápice elevado (Kapala haplospinosa n. sp.); C. medialmente deprimida (Kapala floridana) ...... 172 Figura 19. Projeção da margem posterior do mesoescutelo.A. curto (Colocharis hungi); B. longo (Kapala argentina) ...... 173 Figura 20. Ápice dos processos frenais. A. arredondado ou acuminado (Kapala iridicolor); B. emarginado (Kapala cavicornis n. sp.); C. emarginado com uma flange côncava na margem inferior dos processos frenais (Kapala haplospinosa n. sp.) ...... 174 Figura 21. Formato dos processos frenais das fêmeas (imagens superiores em vista lateral e inferiores em vista dorsal). A. cilíndricos e paralelos (Kapala flabellata); B. amplos e achatados dorsoventralmente (Dicoelothorax sp.); C. formando uma carapaça arqueada sobre o metassoma (Galearia latreillei) ...... 174 Figura 22. Processos frenais, padrão de esculturação do tegumento (aspecto dorsal). fêmea. A. liso ou aparentemente quase liso (Lasiokapala serrata). B. com carenas longitudinais ao longo de todo o comprimento ou tornando-se oblíquas no ápice (Kapala cuprea); C. carenas transversais ou circulares (Lirata luteogaster); D. rugoso (Lirata maranhensis) ...... 176 Figura 23. Disco propodeal, em vista lateral. A. convexo (Lirata striatissima); B. achatado (Liratella nigra) ...... 176 Figura 24. Carena delimitando o disco propodeal. A. ausente (Ecarinata n. gen.); B. presente (Kapala iridicolor, indicado) ...... 177 Figura 25. Espiráculo propodeal. A. circular (indicado); B. circular com uma incisão ventral (indicado). Fonte: adaptado de Heraty, 2002 ...... 177 Figura 26. Cladograma obtido com pesagem implícita (k = 7.1875) dos caracteres através de busca tradional pelo TNT (L = 300); IC = 35; IR = 60). Os círculos pretos representam mudanças únicas, os brancos representam paralelismos ou convergências (homoplasia). As linhas coloridas xix

representam as espécies do gênero Kapala. Valores de bootstrap são mostrados nos retâncgulos lilases. À direta, a filogenia proposta por Heraty (2002) para o clado Kapala ...... 181 Figura 27. Análise de parcimônia com buscas heurísticas através do New Technology Search, com algoritmos Sectorial Search, Ratchet, Drift e Tree fusing (10 rounds) pelo TNT. Valores de bootstrap são mostrados abaixo dos ramos. Ramos em azul representam o complexo de espécies K. romandii; em vermelho o complexo K. furcata; em verde o complexo K. iridicolor ...... 184 Figura 28. Análise de máxima verossimilhança de 89 táxons terminais pelo RaxML. Suporte de bootstrap mostrado acima dos ramos ...... 185

Capítulo III ...... 201

Figure 1. Maximum Likelihood analysis of iridicolor complex species. The analyses include 35 taxa and alignment of 18S (804BP), 28S-D2 (585 pb); 28S D3-D5 (538 pb); COI-nj (382 pb) and COII (279 pb). Support values is maximum likelihooh bootstrap, all values are shown ...... 231 Figure 2. Front head of Kapala iridicolor complex. A. Kapala longicornis n. sp. (front smooth); B. Kapala parairidicolor n. sp. (presence of striae on the frons) ...... 232 Figure 3. Antenna of females of Kapala iridicolor complex, in lateral view. A. Kapala iridicolor; B. Kapala cavicornis n. sp.; C. Kapala ceciliae n. sp.; D. Kapala ipa n. sp.; E. Kapala longicornis n. sp.; F. Kapala parairidicolor n. sp...... 232 Figure 4. Dorsal view of Kapala iridicolor complex (females). A. K. iridicolor; B. K. cavicornis n. sp.; C. K. ceciliae n. sp.; D. K. ipa n. sp.; E. K. longicornis n. sp.; F. K. parairidicolor n. sp.; Red seta indicating the apex of frenal processes emarginated with an elongate flange like spoon- shaped ...... 233 Figure 5. Lateral view of Kapala iridicolor complex A‒B. K. iridicolor (♀ and ♂, respectively); C‒D. K. cavicornis n. sp. (♀ and ♂); E‒F. K. ceciliae n. sp. (♀ and ♂). Red seta indicating the mesoscutum low and smoothly curved anteriorly in lateral view ...... 234 Figure 6. Lateral view of Kapala iridicolor complex. A‒B. K. ipa n. sp. (♀ and ♂, respectively); C‒D. K. longicornis n. sp. (♀ and ♂); E‒F. K. parairidicolor n. sp. (♀ and ♂) ...... 235 Figure 7. Distribution of the species of K. iridicolor complex. A. K. iridicolor; B. K. cavicornis n. sp.; C. K. ceciliae n. sp.; D. K. ipa n. sp.; E. K. longicornis n. sp.; F. K. parairidicolor n. sp...... 236

Capítulo IV ...... 241

Figure 1. Maximum Likelihood analysis of furcata complex species. The analysis includes 24 taxa and alignment of 18S (204 pb), 28S-D2 (587 pb), 28S D3-D5 (543 pb), COI-nj (756 pb) and COII (251 pb). Support values is maximum likelihood bootstrap, all values are shown ...... 267 Figure 2. Kapala furcata complex faces. A. K. furcata (♀); B. K. splendens (♀); C. K. cuprea (♀); D. K. parafurcata n. sp.; E. K. quasimodo n. sp. (♂); F. K. deltalis n. sp. (♀) ...... 268 Figure 3. Head and mesososma of K. furcata complex. A. K. quasimodo n. sp. (♂); B. K. parafurcata n. sp. (♀); C. K. delatlis n. sp. (♀); K. cuprea (♀) ...... 268 Figure 4. Kapala furcata complex lateral habitus of mesosoma. A. K. furcata (♀); B. K. splendens (♀); C. K. cuprea (♀); D. K. deltalis n. sp. (♀); E. K. quasimodo n. sp. (♂); F. K. parafurcata n. sp. (♀) ...... 269 xx

Figure 5. Kapala furcata complex dorsal view of mesosoma. A. K. furcata (♀); B. K. splendens (♀); C. K. cuprea (♂); D. K. parafurcata n. sp. (♀); E. K. deltalis n. sp. (♀); F. K. quasimodo n. sp. (♂) ...... 270 Figure 6. Species distribution map, from holotype and paratype material. Points mapped in SimpleMappr ...... 271

Capítulo V ...... 275

Figure 1. Mesosoma in profile; A. Prepectus separated from pronotum by a suture; B. Prepectus fused to pronotum ...... 279 Figure 2. Mesosoma in dorsal view. A. Scutellum rounded or with a rounded flange; B. Scutellum with prominent process ...... 279 Figure 3. Mesosoma in dorsal view. A. Scutellum with a flattened flange; B. Scutellum rounded ...... 280 Figure 4. Mesosoma in profile. A. Frenum broad; B. Frenum with short bifurcating spines ... 280 Figure 5. Mesosoma in profile. A. Prepectus reaching tegula; B. Prepectus not reaching tegula ...... 281 Figure 6. Antenna in profile. A. Funicular segments pectinate; B. Funicular segments simple...... 282 Figure 7. Mesosoma in dorsal view. A. Frenal processes short and blunt; B. Frenal processes long...... 282 Figure 8. Mesosoma in dorsal view. A. Frenal processes flat; B. Frenal processes cylindrical. 283 Figure 9. Specimen in profile. A. Galearia; B Dicoelothorax...... 283 Figure 10. Mesosoma in dorsal view. A. Frenal processes dorsoventrally flat; B. Frenal processes triangular...... 284 Figure 11. Head in full-face view. A. Isomerala; B. Generalized form, several genera...... 284 Figure 12. Mesosoma in dorsal view. A. Frenal processes cylindrical and smooth; B. Frenal processes not smooth...... 285 Figure 13. Head in full-face view. A. Eyes with erect bristles; B. Eyes with short setae...... 285 Figure 14. Mesosoma in dorsal view. A. Scutellum smooth and shining; B. Scutellum longitudinally ribbed...... 286 Figure 15. Metasoma in dorsal view. A. Two medial incisions over the posterior margin of the first gastral tergite; B. One medial incision over the posterior margin of the first gastral tergite...... 287 Figure 16. Mesosoma in dorsal view. A. Axillular groove deep and curved medially; B. Axillular groove shallow and linear...... 287 Figure 17. Ecarinata sp.1 n. sp. A. Habitus (lateral, ♀); B. anetnna (lateral, ♀; Fl2 indicating); C. propodeum (carina absent); D. head (front, ♂); E. habitus (dorsal, ♀); F. habitus (lateral, ♂). 295 Figure 18. Torquata sp.1 n. sp. A. habitus (lateral view); B. head (front); C. Maxillary and labial palpi (lateral view); D. Mesososma (dorsal view) ...... 296

xxi

REJEIÇÃO PARA FINS DE NOMENCLATURA ZOOLÓGICA

Este trabalho, na forma em que se apresenta (tese de doutorado), não deve ser considerado como publicação válida para finas de nomenclatura zoológica. Este é o disclaim e dénégation mencionado no Código Internacional de Nomenclatura Zoológica (edição 1999), capítulos três, artigos 8.2 e 8.3.

REJECTION FOR ZOOLOGICAL NOMENCLATURE PURPOSES

This work, in the form in which it is presented (doctoral thesis), should not be considered as a valid publication for zoological nomenclature purposes. This is the disclaim and dénégation mentioned in the International Code of Zoological Nomenclature (edition 1999), chapters three, articles 8.2 and 8.3.

1 Introdução geral

Os insetos da ordem Hymenoptera compreendem uma das formas de vida dominantes no planeta, tanto em termos de números de espécies, quanto na diversidade de estilos de vida que se desenvolveram neste grupo (Austin & Downton, 2000). Existem mais de 115.000 espécies reconhecidas, o que a coloca como a quarta maior ordem de insetos, e esse número deverá aumentar com uma análise taxonômica mais aprofundada (Sharkey, 2007; Heraty et al., 2012). Muitos dos membros (vespas, abelhas e formigas) são economicamente importantes por uma variedade de razões, incluindo a polinização e o controle biológico (Sharkey, 2007). Dentro de Hymenoptera, espécies de Chalcidoidea possuem uma incrível variedade de formas e comportamentos. Muitos (> 22.500) dos Chalcidoidea reconhecidos são insetos diminutos que parasitam uma grande variedade de hospedeiros (Munro et al., 2011; Heraty et al., 2012).

1.1 Aspectos biológicos de Eucharitidae

Eucharitidae forma um dos grupos menos característicos de Chalcidoidea, morfológica e biologicamente (Melo et al., 2012), sendo considerado o mais numeroso e diversificado dentre os himenópteros parasitoides de insetos sociais. Eucaritídeos possuem um ciclo de vida altamente especializado e, diferente da maioria das espécies de himenópteros parasitoides, eles não colocam seus ovos diretamente no hospedeiro, mas em um substrato externo, a partir do qual as larvas emergirão e, posteriormente sairão em busca do seu hospedeiro (Heraty, 2002; Lachaud & Perez- Lachaud, 2012, 2015). As espécies de Eucharitidae são todas parasitoides externos (principalmente) ou internos de larvas e/ou pupas de formigas (Clausen, 1941). As fêmeas adultas depositam seus ovos sobre ou dentro do tecido da planta, podendo ser depositados também em brotos e nectários florais. Ao contrário da maioria dos himenópteros parasitoides, a larva de primeiro instar de Eucharitidae, denominada “planídia”, é ativa, podendo mover-se mediante saltos ou curvando-se e, conseguindo assim chegar até o ninho das formigas hospedeiras utilizando comportamentos foréticos (Clausen, 1941; Heraty, 1985, 2002; Lachaud et al., 2012; Lachaud et al., 2015; Pérez- Lachaud et al., 2006; Torréns, 2013). Esses comportamentos foréticos envolvem uma ligação 2

direta com formigas operárias forrageadoras ou, através de hospedeiros intermediários, por exemplo tisanópteros, os quais carregam a planídia para dentro da colônia. Dentro dela, a planídia se adere externa ou internamente à larva de alguma formiga, permanecendo inativa até esta chegar à fase de pré-pupa ou pupa. Quando isso acontece, a planídia migra para a região ventral do tórax da formiga, iniciando sua alimentação com vistas a continuar seu desenvolvimento que possui mais dois estádios adicionais (Clausen 1940; Heraty, 2002; Pérez-Lachaud et al., 2006; Lachaud et al., 2015; Torréns, 2013). Posteriormente, os parasitoides adultos emergem e deixam o ninho por conta própria ou, esse processo pode ser realizado pelas formigas que os depositam entre os resíduos da colônia (Clausen, 1941; Pérez-Lachaud et al., 2006; Buys et al., 2010; Torréns, 2013). A ausência de agressão em relação aos eucaritídeos adultos contrasta com o comportamento violento demonstrado pelas formigas operárias quando reconhecem indivíduos de outras espécies ou presas potenciais (Wheeler, 1910; Dejean et al., 1999), sugerindo a existência de algum tipo de mimetismo químico ou morfológico pelos eucaritídeos parasitoides (Perez- Lachaud et al., 2015). O reconhecimento de Nestmate em insetos sociais baseia-se principalmente em hidrocarbonetos cuticulares (CHC) e, aparentemente, existem perfis CHC específicos que definem as relações entre os membros do grupo (Howard & Blomquist, 2005; d'Ettorre & Lenoir, 2010). No caso de Eucharitidae, as vespas emergentes têm um perfil de CHC semelhante ao de seu hospedeiro (Vander Meer, 1989; Howard et al., 2001; Lachaud-Perez et al., 2015), mas essa semelhança não é suficiente para que os parasitoides consigam suprimir inteiramente o reconhecimento como intrusos na colônia (Howard et al., 2001). Alguns autores compararam os perfis de CHC entre vespas e formigas operárias e concluíram que elas contêm os mesmos compostos, mas em proporções diferentes ─ elas não diferiram significativamente (Howard et al., 2001; Perez-Lachaud et al., 2015). Esses resultados revelam apenas uma imitação parcial do perfil químico do hospedeiro e isso pode ser interpretado como uma estratégia evolutiva para o parasitoide sair da colônia mais rapidamente. A vida de um Eucharitidae adulto é efêmera (5-7 dias) (Clausen 1941; Perez-Lachaud et al., 2010) e um ninho de formigas é tipicamente um complexo de galerias e câmaras que são sempre guardados pelas formigas operárias (Hölldobler & Wilson, 1990). Assim, ser carregado para fora do ninho por 3

uma destas formigas pode ser uma maneira de economizar tempo e energia (Perez-Lachaud et al., 2015).

1.2 Aspectos taxonômicos de Eucharitidae

Eucharitidae foi tratada pela primeira vez como família por Walker (1862) e, cujo nome deve-se ao gênero Eucharis descrito por Latreille em 1805. Essa família contém 54 gêneros e aproximadamente 500 espécies válidas (Heraty, 1985, 2002; Torréns, 2013). Na literatura, o número de subfamílias pertencentes a Eucharitidae é de três ou quatro, sendo elas, Oraseminae e Eucharitinae propostas por Burks (1979) e Gollumiellinae por Heraty et al. (2004). Contudo, Bouček (1988) incluiu também Akapalinae (hospedeiro desconhecido). Porém, utilizando dados morfológicos, Akapalinae foi recuperada como grupo-irmão de Eucharitidae com 85% de suporte de bootstrap (Heraty, 2002) e através da utilização de dois genes nucleares ribossomais, 18S e 28S, Jambyia Heraty & Darling, 2007 (Incertae sedis) foi recuperado com grupo-irmão de Eucharitidae, sem suporte de ramo (Munro et al., 2011). Além disso, através de uma análise combinada utilizando 232 caracteres morfológicos e três genes, foi verificado que Akapalinae é grupo-irmão de Eucharitidae sensu stricto (Heraty et al., 2012). A monofilia da família está embasada em sinapormorfias tanto de adultos como de larvas. Em adultos, o pronoto é reduzido, ventral ao mesonoto e não visível em vista dorsal (Heraty & Darling, 1984; Heraty, 2002). Não há sulco malar, as mandíbulas são falcadas e em forma de foice e o labro possui dígitos, embora estas duas últimas características tenham sido secundariamente perdidas em alguns gêneros (Heraty, 1994). Contudo, o suporte mais forte para a monofilia do grupo é obtido a partir de características dos imaturos, as quais são: ovos pedunculados e planídia com redução das setas dorsais nos tergitos VII e IX (de um total de 12 tergitos). Eucaritídeos são encontrados em todas regiões zoogeográficas, com exceção da Nova Zelândia, regiões polares e algumas ilhas oceânicas mais isoladas e, com uma maior diversificação nos trópicos (Heraty, 1985, 2002; Torréns, 2013). Heraty (2002) propôs que a família teve origem no final do Cretáceo ou início do Eoceno, devido sua distribuição disjunta no hemisfério sul. Baseado na descrição de duas espécies a partir de âmbar do mar Báltico, 4

Perilampus pisticus Heraty & Darling, 2009 (Perilampinae) e Paleocharis rex Heraty & Darling, 2009 (Eucharitinae), uma idade mínima de 40 milhões foi proposta Eucharitinae (Heraty & Darling, 2009). Gollumiellinae é composta por dois gêneros, Gollumiella Hedqvist, 1978 com distribuição por toda a região Indo-pacífica, incluindo o Japão e nordeste da Austrália; e Anorasema Bouček, 1988, presente nas sub-regiões da Indo-China, Malásia e Filipinas (Heraty et al., 2004). Já Oraseminae, grupo irmão de Eucharitinae, é considerado monofilético (Heraty, 1994, 2000), sendo composto por quatro gêneros; Indosema Husain & Agarwal, 1983, Orasemorpha Bouček, 1988 e Timioderus Waterston, 1916, os quais ocorrem nos trópicos do Velho Mundo e Orasema Cameron, 1884 com distribuição circumtropical e neártica, a qual é fragmentada em uma série discreta de grupos de espécies geograficamente restritas (Heraty, 1994, 2000). Eucharitinae compreende duas tribos: Psilocharitini, que inclui dois gêneros do Velho Mundo, Neolosbanus Heraty, 1994 e Psilocharis Heraty, 1994 (Heraty, 1994, 2002), e Eucharitini, um grupo morfologicamente diverso com 44 gêneros. A monofilia de Eucharitini é bem suportada pela fusão anterior do prepecto e do pronoto, com invólucro completo dos espiráculos internos do mesotórax; perda do segmento anelar antenal e o primeiro fragma formando um forte rebordo acima da margem dorsal do pronoto (Bouček, 1988; Heraty, 1985, 1989, 2002). Os gêneros dessa tribo foram revisados somente para a região da Australásia (Bouček, 1988) e para região Neártica (Heraty, 1985), porém até o momento não houveram tratamentos abrangentes para os gêneros neotropicais (Torréns, 2013, 2016; Torréns et al., 2007; Torréns et al. 2013). Dentro de Eucharitini (Eucharitinae), há um grupo monofilético que ataca formigas das subfamílias Ponerinae, Ectatominae e Myrmeciinae (clado PEM) (Fig. 2) (Muray et al., 2013). O clado PEM é suportado por análises moleculares e algumas morfológicas (Heraty, 2002; Muray et al. 2013) e é constituído por três grupos: os clados Chalcura e Schizaspidia do Velho Mundo e o clado Kapala do Novo Mundo.

O clado Kapala está fortemente apoiado em todas as análises com bons caracteres e análises de bootstrap (88%) (Heraty, 2002; Murray et al., 2013). Baseado em evidências moleculares, estima-se que o clado Kapala divergiu dos táxons do Velho Mundo, a aproximadamente 35,6 m.a.a (25,7-46,4) (Murray et al., 2013). Possui gêneros com múltiplas modificações morfológicas relacionadas à forma da cabeça, morfologia da antena, padrões de escultura e morfologia dos processos frenais, os quais são particularmente variáveis dentro do clado (Murray, 2014). O dimorfismo sexual está particularmente confinado a diferenças na morfologia da antena, do metasoma e nas diferenças da morfologia dos espinhos frenais, os quais são muito mais reduzidos e finos nos machos do que nas fêmeas. O clado Kapala é composto por 13 gêneros, sendo Kapala o mais abundante e diverso.

1.3 Kapala Cameron, 1884

Kapala Cameron, 1884 é especialmente comum e especioso na região Neotropical. Ocorre desde o sul dos Estados Unidos (Arizona, Texas e Flórida) até a Argentina, não ocorrendo 6

no Chile. Apenas Kapala ivorensis Risbec, 1954, possui ocorrência restrita para a região Afrotropical e Madagascar (Heraty, 1985, 2002; Murray & Heraty, 2016). Até o momento, sabe- se que as espécies de Kapala atacam três gêneros de Ponerinae e três de Ectatominae (Lachaud & Pérez-Lachaud, 2012; Murray et al., 2013). No total, existem 18 espécies válidas (Heraty, 2002). Kapala foi proposto por Cameron (1884) como um nome substituto para Chirocerus Brullè, 1846, o qual era pré-ocupado por Chirocerus Latreille, 1825 (pág. 447). Portanto, Chirocerus Brullè, 1846 passou a ser homônimo-júnior de Kapala. O gênero, inicialmente, incluía apenas uma espécie Kapala furcata (Fabricius, 1804) descrita a partir de uma única fêmea coletada na América do Sul. Muitos exemplares presentes em coleções científicas foram determinados sob o nome da espécie-tipo, no entanto, ela não é muito comum (Heraty, 2002), consequentemente, muitas determinações estão erradas. As espécies desse gênero exibem uma grande quantidade de variações morfológicas externas, o que dificulta o estabelecimento dos limites da maioria das espécies (Heraty & Wooley, 1993; Heraty, 2002) tornando complicada a identificação destas e até mesmo fatigante. Dados moleculares sugerem que Kapala é parafilético em relação à maioria dos outros gêneros do Clado Kapala (Heraty, 2002; Murray, 2014), sendo o gênero dividido em três complexos de espécies: furcata, iridicolor e sulcifacies (Heraty & Wooley, 1993; Murray et al., 2013; Murray, 2014). Esses três complexos são separados filogeneticamente por dois agrupamentos de gêneros morfologicamente distintos e pelo gênero Isomerala Shipp,1894 (Murray, 2014.). Contudo, de acordo com Heraty (2002) através de dados morfológicos, foi sugerido que Kapala é monofilético, não existindo, porém, sinapomorfias que sustentem sua monofilia. O gênero é definido por um conjunto de simplesiomorfias que incluem espinhos frenais lirifomes com estrias longitudinais ou em espiral, olhos nus, ramos antenais do macho maiores que a altura da cabeça, flagelômero basal da fêmea raramente mais que duas vezes mais longo que largo e margem posterior do disco escutelar elevada a uma crista proeminente entre as bases dos espinhos frenais (Heraty, 2002). Em contraste, os outros gêneros reconhecidos podem ser altamente divergentes na sua morfologia (Torréns, 2013). No entanto, Kapala é fenotipicamente distinto dos demais gêneros, e pesquisas estão em andamento utilizando a próxima geração de sequenciamento para ver se esse é um artefato de amostragem de genes limitados ou um fenômeno real (Heraty, em prep.). 7

O estudo sistemático de Kapala constitui um importante alvo de pesquisa uma vez que o elevado grau de variação morfológica evidente dentro das espécies e a falta de caracteres confiáveis para separá-las vêm dificultando a sua taxonomia e definição. Baseado em trabalhos anteriores (Heraty & Woolley; Heraty, 2002; Torréns et al., 2007; Torréns & Heraty, 2013; Murray et al., 2013; Murray & Heraty, in prep.) há um quadro existente para o agrupamento de fenótipos e desenvolvimento de limites mais concretos para as espécies propostas. Para tanto, no capítulo I, uma revisão taxonômica das espécies previamente descritas de Kapala é fornecida, e cujas redescrições contemplam novos caracteres indispensáveis para o estabelecimento dos limites específicos, bem como para associação sexual. Pela primeira vez, uma chave de identificação para as espécies do gênero é fornecida, bem como a descrição de sete novas espécies, K. confusa n. sp., K. corcovata n. sp., K. gracilispina n. sp., K. haplospinosa n. sp., K. jalisca n. sp., K. spinaepplanata and K. genistriata n. sp. No capítulo II, é realizada uma filogenia com limites moleculares e, morfológicos, o qual irá fornecer um sólido conhecimento para futuros estudos sobre o gênero, bem como táxons relacionados. A análise crítica morfológica extensiva do gênero permitiu explorar novas características morfológicas, tanto para a identificação das espécies de Kapala e para a definição do gênero; já as análises moleculares além de auxiliar a definir o gênero também proporcionaram estabelecer as relações internas entre as espécies. Nos Capítulos III e IV, a revisão taxonômica dos complexos de espécies iridicolor e furcata, são fornecidas, respectivamente. No Capítulo V, como base na filogenia morfológica e molecular, dois novos gêneros são descritos para a família Eucharitidae, Ecarinata n. gen. e Torquata n. gen. Além disso, uma chave revisada e atualizada para os gêneros Neotropicais de Eucharitidae é fornecida.

Capítulo I

Schoeninger, K., Torréns, J., Heraty, J.M, Oliveira, M.L. (2018) Taxonomy of Kapala (Hymenoptera: Eucharitidae): New species, redescriptions and identification key for the species of Kapala

Taxonomy of Kapala (Hymenoptera: Eucharitidae): new species, redescriptions and an identification key

KARINE SCHOENINGER1,4, JAVIER TORRÉNS2, JOHN M. HERATY3 & MARCIO L. OLIVEIRA1

1Coordenação de Biodiversidade, Programa de Pós-Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia, CEP 69067-375, Manaus, AM, Brazil. E-mails: [email protected]; [email protected] 2CRILAR-CONICET, Entre Ríos y Mendoza, 5301 Anillaco, La Rioja, Argentina. E-mail: jtorrens@crilar- conicet.gob.ar 3Department of Entomology, University of California, Riverside, CA 92521, United States. E-mail: [email protected] 4Corresponding author. [email protected]

Abstract. Kapala Cameron 1884, is a diverse and speciose genus within Eucharitidae, of which the species often exhibit a large amount of intraspecific morphological variation. Based on examination of the type species and complementary material, we review the current knowledge of the Neotropical species of Kapala. We redescribe the type species of the genus and establish the limits of valid species; four synonyms were performed, K. atrata n. syn. and K. surgens n. syn. was synonimized with K. flabellata (Fabricius, 1804), K. striaticeps n. syn. was synonymized with K. inexagens (Walker, 1862) and K. sulcifacies (Cameron, 1904) n. syn. was synonymized with K. romandii (Guérin-Meneville, 1845); three lectotypes were designated: K. cynipsea (Walker, 1862) pres. desig., K. inexagens (Walker, 1862) pres. desig. and K. terminalis Ashmead, 1892 pres. desig., and seven species had males associated with females. Seven new species are described: K. confusa n. sp., K. corcovata n. sp., K. genistriata n. sp., K. gracilispina n. sp., K. haplospinosa n. sp., K. jalisca n. sp. e K. spinaepplanata n. sp., and an identification key for all species is provided.

Keywords. Ant parasitoids, Chalcidoidea, identification key, Neotropical region, taxonomy. 10

Introduction

Eucharitidae is the only family of insetcs where all member attack Formicidae, more specifically Ectatominae, Formicinae, Myrmeciinae, Myrmicinae and Ponerinae (Clausen 1940; Heraty 2002; Lachaud & Pérez-Lachaud 2012; Murray et al. 2013). Eucharitidae includes some of the most distinct members and whithin this family Kapala Cameron, is one of the most common, diverse and specious, which species are distributed across the Neotropical region, with exception of Kapala ivorensis Risbec, found in the Afrotropical region and Madagascar (Heraty 2002, Murray & Heraty, 2016). Kapala was proposed by Cameron (1884) as a replacement name for Chirocerus Bruleè, 1846, who was pre-occupied by Chirocerus Latreille, 1825. Therefore, Chirocerus Bruleè, 1846 became the junior synonyms of Kapala. The genus initially included only one species Kapala furcata Fabricius, 1804 described from a single female collected in South America. The need for a review of the already described species of Kapala was observed from their original descriptions, which are extremely brief, which resulted in a many erroneous identifications. The vast majority of the descriptions mention coloration, which is extremely variable; the presence of two long spines, presence of flabelled antennae and high and convex mesoscutum. These characteristics are common to all Kapala species, and accurate determination of the species is almost impossible based on the original descriptions. Recently, Pérez-Lachaud et al. (2006) described a new species, Kapala izapa Carmichael, 2006 in which they inserted new characters as striae on the frons and in enterily body and measures to try to delimit the species. This genus currently is defined by a suite of symplesiomorphies including frenal spines lyre-shaped with longitudinal or spiral carina, eyes bare, male antennal rami longer than height of head, female basal flagellomeres rarely more than twice as long as broad and posterior margin of the scutellar disc raised to a prominent ridge between the bases of the frenal spines (Heraty 2002). However, Kapala are phenotypically distinct from these other genera, but there is still no synapomorphy that supports Kapala as monophyletic. Species of Kapala can be divided into three clades based on both morphological and molecular data: furcata, iridicolor and sulcifacies (Heraty & Woolley 1993; Murray, 2014; Schoeninger et al., in prep). Based on molecular data 11

only, these clades are separated phylogenetically by two clusters of morphologically distinct genera and by genus Isomerala Shipp, 1894 (Murray,2014; Schoeninger et al., in prep). In synthesis, a revison of Kapala is an important starting point for a more general overview in the genus and the entire Kapala clade (includes 13 genera). There is a critical need for a broad morphological examination of the genus to explore new morphological characters, both for identification of the species and for the definition of the genus within the Kapala clade. Several characters remain unexplored within the genus, but this requires examination of a broad range of species and specimens to sort out character differences from character variation. This is the first attempt to revise the known species, describe new taxa and establish a baseline for the taxonomy of the genus. Therefore, in this study, we rediscribed the species types of the genus, established well defined limits to the valid species through the insertion of new characters; four synonyms were performed; three lectotypes were designated, as well as the sexual association of seven species was characterised. In addition, seven new species were described and an identification key for all the species was provided.

Material and methods

Morphological descriptions and redescriptions

The descriptive format follows Heraty (2002), with details on sculpture from Harris (1979) and Gibson et al. (1998). Morphological terms and abbreviations are from the Hymenoptera Anatomy Ontology (HAO; Yoder et al. 2010), Heraty (2002) and Gibson et al. (1998). The basal flagellomere is counted as Fl2 following Heraty (2002). The funiculars include Fl2 and the following unfused flagellomeres before the clava, which is defined by 1–3 fused flagellomeres. The paired spines of the mesoscutellum arise from the frenum and are refered here as frenal process (character 50; Heraty 2002). Morphological abbreviations (see Figs 1 and 2): acl, anteclypeus; acr, acropleuron; ao, anterior ocellus; ax, axilla; axg, axillular groove; axl, axillula; cal, callus; clv, clava; cly, clypeus; cx, coxa; es2, mesepisternum; Fl2, flagellomere basal; flg, flagellum; fmd, femoral depression; frl, frenal line, fp, frenal processes; fub, funicle; gen, gena;

Gt1, first gastral tergite; iod, interocular distance; lep2, lower mesepimeron; llm, lateral lobe of 12

mesoscutum; mlm, midlobe of mesoscutum; msp, malar space; mv, marginal vein; not, notaulus; pdl, pedicel; pet, petiole; pl3, metapleuron; pmv, postmarginal vein; po, posterior ocellus; ppd, propodeum; pre, prepectus; sca, supraclypeal area; scd, scrobal depression, scp; scape, sct, mesoscutellar disc; smv, submarginal vein; sss, scutoscutellar suclcus; stv, stigmal vein; tgl, tegula; tor, torulus; tsa, transscutal articulation; upe2, upper mesipimeron.

Measurements

Measurements were made through a Zeiss Stemi SV6® with a 1.5X objective lens. The measures and ratios between structures are given as the value of the holotype [in brackets], followed by the range. The stigmal vein on the fore wing is often irregular in shape and so was measured at the widest point and the longest distance from the wing margin. All measurements are given in millimeters, and the abbreviations used are presented in Fig. 3. The following measurements were recorded.

Axillar length: dorsal length from transscutal articulation (TSA) to scutoscutellar AXL sulcus (SSS). Body length: the length of mesosoma from anterior margin to the apex of frenal BDL process. C3L Metacoxa length: length from basal to apical margin. C3W Metacoxa width: width at widest point. EYH Eye height: vertical height of eye. FLG Flagellum length of female: length of all flagellomeres from pedicel to apex of clava. Flagellomere basal length: length of basal flagellomere (female) or basal Fl2 flagellomere including ramus (male).

Fl2W Flagellomere basal width: width of flagellomere at apex, female only.

Fl3 Flagellomere 3 length; dorsal length of Fl3 from basal margin to apex, female only. F3L Metafemur length: lateral length of femur F3W Metafemur width: width in center from lateral view. FPH Frenal process height: lateral height of process at base. 13

Frenal process length: dorsal length of frenal process from medial point of process FPL base on frenal line to apex of the frenal process. Forewing length: maximum length of the forewing in a straight-line spanning from FWL the humeral plate to the maximum wing apex. Forewing width; the maximum height of the forewing, measured in lateral view, in a FWW straight-line spamming from the costal margin to the posterior margin. Head height; height measured from dorsal margin of median ocellus to median apex HDH of clypeus. HDW Head width; distance across face at widest point from outer margin of eyes. IOD Interocular distance; distance between eyes measured across dorsal margin of eyes. Mesoscutum height: lateral height from anterolateral ventral margin at anterior MSH pronotal junction to dorsal margin at transscutal articulation. Mesonotum length: lateral length of mesonotum not including the frenal process, MSL from midlobe to apex of mesoscutellum at frenal line. Malar space length: length measured from ventral margin of eye to base of MSP mandibles. Mesoscutum width: lateral diagonal from anterior lower margin of mesoscutum to MSW dorsal margin at transscutal articulation (TSA). MVL Marginal vein length: length from apex of costal cell to stigmal vein. PTL Petiole length: lateral length of petiole from flange at the base to apex. PTW Petiole width; width of petiole across medial. Scutellum apex height; lateral height of crest from dorsal base of frenal process to SAH dorsal margin. SCL Mesoscutellar length: dorsal length of mesoscutellum from SSS to apex medial to scutellar processes. Mesoscutellar width: width of mesoscutellum at base of frenal process and between SCW axillular carinae. SMVL Submarginal vein length: length from base of costal cell to base of parastigmal vein. SPL Scape length: measured from the base to the apical margin. SPW Scape width: width measured at apex. 14

Specimen imaging

Specimens were photographed using a Leica M205A stereomicroscope coupled with a Leica DMC4500, and processed using Leica Application Suite V4.1 Interactive Measurements, Montage. Images were stacked and combined using Helicon Focus (©HeliconSoft) using the C method, 100% full resolution and saved at 600 DPI. The final images and plates were edited using Adobe Photoshop and Adobe Illustrator.

Specimens repository

The following intitutions served as source of material and type depositories for specimens examined for this study: Archbold Biological Station, Lake Placid, Florida, USA (ABS); American Museum of Natural History, New York, USA (AMNH); The Natural History Museum, London, England (BMNH); California Academy of Sciences, San Francisco, California, USA (CAS); Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA (CMNH); Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada (CNC); Cornell University Insect Collection, Ithaca, New York, USA (CUIC); Coleção Zoológica do Maranhão, Universidade Estadual do Maranhão, Caxias, Brazil (CZMA); University of Guelph Insect Collection, Guelph, Ontario, Canada (DEBU); Escuela Agricola Panamericana, Tegucigalpa, Honduras (EAPZ); Essig Museum of Entomology, University of California, Berkeley, California, USA (EMEC); Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, Florida, USA (FSCA); Hymenoptera Institute Collection, University of Kentucky, Lexington, USA (HIC); Instituto Fundación Miguel Lillo, Tucuman, Argentina (IFML); Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (INPA); Los Angeles County Museum of Natural History, Los Angeles, California, USA (LACM); APTA, Laboratório Regional de Ribeirão Preto, São Paulo, Brasil (LRRP); Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (MACN); Milwaukee City Public Museum, Milwaukee, Wisconsin, USA (MCPM); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ); Museu de Ciências Naturais da Fundação Zoo-Botânica do Rio Grande do Sul, Porto Alegre, 15

Brazil (MCNZ); Muséum d’Histoire Naturelle, Geneva, Switzerland (MHNG); Museo de La Plata, Universidad Nacional de La Plata, Division Entomologia, La Plata, Argentina (MLP); Mississippi Museum of Natural Science, Mississipi, USA (MNCN); Museu Nacional da Costa Rica (MNCR); Museum National d’Histoire Naturelle, Paris, France (MNHN); Museu Paraense Emílio Goeldi, Belém, Pará, Brazil (MPEG); Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brazil (MZFS); Lund University, Lund, Sweden (MZLU); Museum and Institute of Zoology, Polish Academy of Science, Warsaw, Poland (MZPW); Museu de Zoologia de São Paulo, São Paulo, Brazil (MZSP); National Museum of Natural History, Washington D.C., USA (NMNH);, Prague, Czech Republic (NMPC); Royal Belgium Institute of Natural Science, Brussels, Belgium (RBINS); Royal Ontario Museum, Toronto, Ontario, Canada (ROME); Iziko Museum of Capetown, Cape Town, South Africa (SAMC); Staatliches Museum für Naturkunde, Stuttgart, Germany (SMNS); Universidad Autónoma Nuevo León, Facultad de Ciencias Forestales, Linares, Mexico (UANL); University of California, Riverside, California, USA (UCRC); University of California, Davis, California, USA (UCDC); Coleção Entomológica, Departamento de Biologia, Universidade Federal do Espírito Santo, Brazil (UFES); University of Michigan, Museum of Zoology, Ann Arbor, Michigan, USA (UMMZ); Universidad Nacional Autonoma de Mexico, Mexico, D.F., Mexico (UNAM); Zoologisk Museum, Copenhagen, Denmark (ZMUC); Zoologisches Institute und Zoologisches Museum, Universität von Hamburg, Hamburg, Germany (ZMUH). Specimens are identified with unique UCRC (University of California – Riverside Collection) specimen identifiers that indicate the museum of deposition. All specimens were databased in a FileMaker v.11 ® database maintained in the Heraty lab. Localities were georeferenced using Google Earth (coordinates in italics) or from the original label data. The distribution map was constructed in Simple Mappr (Shorthouse, 2010).

Results

Taxonomy

Kapala Cameron, 1884

Chirocerus Brullè, 1846: 571, pl. 38, fig. 5. Type species: Eucharis furcatus Fabricius, by monotypy. Preoccupied by Chirocerus Latreille, 1825: 447, homonym discovered by Cameron, 1884: 102-103. Chirocera Desmarest, 1860: 161, fig. 141. Unjustified emendation of Chirocerus Brullè, with reference to Chirocera furcatus. Kapala Cameron, 1884: 102-103. Replacement name. Kapala; Kirby, 1886: 32. [Kirby questions the identity of Brullè’s drawing and suggests that it looks closer to Thoracantha atrata Walker than to either of the Fabricius species. Indeed, the drawing appears closer to Schizaspidia than to what is now recognized as Kapala. The figure legend in Brullè (1846: pl. 38) refers to Chirocerus furcatus Westwood, not Fabricius, and may not, unfortunately, refers to the Fabricius species. Kapala; Ashmead, 1888[45]: 187 (key). Kapala; Ashmead, 1904[238]: 269,270 (key), 472. Capala Schulz, 1906: 150. Unjustified emendation. Holcokapala Cameron, 1913: 117. Heraty, 2002: 167. Kapala; Gemignani, 1933: 485. Subsequent description. Kapala; Heraty, 1985: 84. Subsequent description. Kapala; Heraty, 2002: 167-170. Subsequent description.

Diagnosis. Genus currently defined by frenal processes slightly lyriform in dorsal view, with longitudinal or spiral carina (except in K. chacoensis Gemignani, 1947, whose process is soft without defined carina) and usually emarginated apically (in lateral view); eyes bare; maxillary and labial palpi with 3/3 or 3/2 formula; posterior margin of the scutellar disc raised to a prominent ridge between the base of the frenal process (however, this is considered to have been lost in a few species, for example, in K. floridana (Ashmead, 1885) and K. terminalis Ashmead, 1892); propodeal spiracles with a strongly excised ventral margin. Both sexes with short basal flagellomeres; the flagellum of the female simple or serrate with 8, 9 or rarely 10 flagellomeres, length of flagellum 0.8–1.7x head height and basal flagellomere (Fl2) 1.2‒3.3x as long as apical 17

width; the 10-segmented flagellum of the male pectinate with the rami thin and long (0.8‒1.8X longer than height of head).

Key to Kapala species

(K. dicerodera (Spinola, 1853) not included in this key; holotype lost and questionably placed in Kapala)

1 Smooth face or with faint striae on the frons (Fig. 7B); mesocutum low in lateral view (Fig. 7C); lateral lobes of mesoscutum usually densely pilose and weakly striated … 2 - Striated face (Fig. 8B); mesoscutum strongly elevated in lateral view (Fig. 11D); lateral lobes of mesoscutum with sparsely setose and strongly striated … 3

2(1) Frons flat and with faint longitudinal striae (Fig. 24B). Mesosoma with fine and closely longitudinal striae (Fig. 24C); medial apex of mesoscutellar disc elevated like a keel-shape (Fig. 24D, indicated) and wings brown infuscate … K. confusa n. sp. - Frons swollen and smooth (Fig. 7B). Mesosoma with wide spaced longitudinal striae (Figs. 7C, E); medial apex of mesoscutellar disc not elevated and wings hyaline … K. iridicolor Cameron (K. iridicolor complex)

3(1) Females with 8 flagellomeres (Fig. 16C); frons with striae widely spaced and pattern of striae longitudinal; lower face with transverse striation (Figs. 11B, C). Males with frenal processes usually with roughly sculptured carinae, giving them a serrated appearance (Figs. 11A, E; 14E) … 4 (K. romandii complex) - Females with 9 or 10 flagellomeres (Fig. 5C); frons with striae closely to each other and pattern of striae oblique; lower face with striation usually oblique below torulus (Fig. 5B), if it is transversely striated then females with 9 flagellomeres. Males with frenal processes with longitudinal carina (Fig. 5G) … 19 (K. furcata complex)

4(2) Frenal processes flattened laterally with longitudinal striae and apex straight (Figs. 23A, C); front head densely pilose (Fig. 23B) … K. spinaepplanata n. sp. - Frenal processes cylindrical with longitudinal or oblique striae (Fig. 11E; 22C) and apex rounded or emarginate (Fig. 12A, F; 22C); front head with sparse setae … 5

5(4) Medial apex of mesoscutellar disc not elevated above base of frenal processes (Figs. 12C; 17C, indicated) … 6 - Medial apex of mesoscutellar disc elevated above base of frenal processes (Figs. 8E; 21A) …8

6(5) Mesocutellar disc relatively flat and not rounded in lateral view (Fig. 12C). Frons with wide longitudinal striae, which are bifurcate giving a rugose appearance (Fig. 12B). Male with rami of

Fl2 1.0–1.1x head height. Femurs brown with distal apex yellow (Figs. 12A, E) and females with

Gt1 reddish-orange (Fig. 12E) … K. floridana (Ashmead) - Mesoscutellar disc swollen and rounded in profile view (Fig. 17C; 22A). Frons with longitudinal striae without a rugose appearance (22B). Male with rami of Fl2 > 1.2x head height.

Femurs yellow or entirely brown in some species and females with Gt1 brown to dark-brown … 7

7(6) Male with ramus of Fl2 brown and long, 1.6‒2.2x head height. Midlobe of mesoscutum with fine transversal carinae (Figs. 17F, G). Frenal processes long, 1.9‒2.0x as long as length of axilla and mesoscutellar disc, curved in lateral and dorsal view and, longitudinal striated … K. terminalis Ashmead

- Male with rami of Fl2 yellow, 1.2‒1.4x head height. Midlobe of mesoscutum with tranversal and longitudinal carinae, giving a rough appearance (Figs. 22B, C). Frenal processes short, 1.4‒1.5x as long as length of axilla and mesoscutellar disc, slightly curved in lateral view and U- shape or parallel in dorsal view (Fig. 22C), oblique striated … K. jalisca n. sp.

8(5) Mesocutellar disc smooth or with some faint lateral striae (Fig. 13C) and, medial apex raised 0.5–0.6x height of frenal processes in lateral view. Female antenna short, 0.8x head height (Fig. 13D). Male unknown … K. inexagens (Walker) 19

- Mesoscutellar disc with well-marked longitudinal or slightly oblique striae (Fig. 8F; 19C) and, medial apex raised >0.7x height of frenal processes in lateral view. Female antenna > 0.9x head height … 9

9(8) Mesoscutum elevated and projected over the head (Fig. 9A, 18A) … 10 - Mesoscutum elevated but not projected over the head (Fig. 8A; 15C) … 11

10(9) Midlobe elevated with apex pronounced and bilobed (Fig. 18C, D); frenal processes thin with apex rounded … K. corcovata n. sp. - Midlobe elevated, but apex rounded and not pronounced (Fig. 9A, E); frenal processes soft with fine curviform striae giving a rough and irregular surface appearance (9F, G) … K. chacoensis Gemignani

11(9) Frenal processes short and stout, 1.1x as long as length of axilla and mesoscutellar disc (Fig. 10A). Petiole short 1.4x longer than broad … K. cynipsea (Walker) - Frenal processes long >1.4x as long as length of axilla and mesoscutellar disc. Petiole > 1.8x longer than broad … 12

12(11) Medial apex of mesoscutellar disc raised 1.7–2.0x height of frenal processes; frenal processes light-brown to even reddish-brown (Fig. 21C). Supraclypeal area of males striated (Fig. 21B) and females with supraclypeal area usually smooth … K. haplospinosa n. sp. - Medial apex of mesoscutellar disc raised < 1.7x height of frenal processes; frenal processes black. Both sexes with supraclypeal area smooth … 13

13(12) Frenal processes very long, > 1.7x as long as length of axilla and mesoscutellar disc, exceeding the metasoma (Fig. 11A; 19A) … 14 - Frenal processes long, < 1.7x as long as length of axilla and mesoscutellar disc, but not exceeding the metasoma … 18

14(13) Basal flagellar rami of males >1.6x height of head (Figs. 15A; 19A) and flagellum of females >1.2x height of head … 15 - Basal flagellar rami of males <1.6x height of head (Fig. 20A) and flagellum of females <1.2x height of head (Fig. 16C) … 16

15(14) Frons with oblique wide striae from the inferior margin of torulus to the lateral ocelli (Fig. 19B). Frenal processes 2.3–2.4x as long as length of axillae and mesoscutellar disc, with base black and apical half reddish orange (Fig. 19C). Gt1 orange with some spots brown … K. genistriata n. sp. - Frons with longitudinal striae (Fig. 15B). Frenal processes 1.8–2.0x as long as length of axillae and mesoscutellar disc, entirely brown (some females) or apically brown and black basally (Figs.

15E, F). Gt1 basally dark brown and apically light brown (Fig. 15A) … K. izapa Carmichael

16(14) Medial apex of mesoscutellum with carina, more pronounced in males (Fig. 11E). Both sexes with flagellomeres reddish-brown to deep-black (Figs. 11A, D) … K. flabellata (Fabricius) - Medial apex of mesoscutellum with carina. Both sexes with flagellomeres light brown to brown (Figs. 8C, D) … 17

17(16) Face with well-marked striae (Fig. 8B). Malar space 1.0–1.1x eye height. Body black … K. argentina Gemignani - Face with faint striae (Fig. 14B). Malar space 0.8–0.9x eye height. Body metallic-green … K. ivorensis Risbec

18(13) Lower face smooth to even transversal striate (Fig. 20B); length of rami of Fl2 0.9‒1.1x head height; mesoscutellar disc and axilla with a medial longitudinal depression (Fig. 20C); frenal processes very thin with apex rounded … K. gracilispina n. sp.

- Lower face transversal striate (Fig. 16B); length of rami of Fl2 1.1‒1.2x head height; mesoscutellar disc and axilla without a medial longitudinal depression; frenal processes not thin with apex emarginated … K. romandii (Cameron) 21

19(2) Frons and face with oblique striations (Fig. 4B). Frenal processes short and stout with longitudinal striae in basal half, which becoming oblique in apical half (Fig. 4E) … K. cuprea Cameron - Frons with oblique striae from the torulus to the lateral ocelli, and lower face with transverse striae (Fig. 5B; 6C). Frenal process longitudinal striate … 20

20(19) Gena with margin broad and angulate (Fig. 6C). Flagellum of females 8‒9 segmented

(Fig. 6B); length of flagellum 1.0–1.1x head height; males with length of rami of Fl2 1.1X head height. Frenal processes 1.0–1.8x longer than the basal width of mesoscutellum. Body color metallic green to blue (Fig. 6A) … K. splendens Ashmead - Gena with margin not broad and angulate (Fig. 5B). Flagellum of females 9–10 segmented (Fig.

5C); length of flagellum 0.8–1.0x head height; males with length of rami of Fl2 1.2–1.3x head height. Frenal processes 1.9–2.3x longer than the basal width of scutellum. Body color black (Fig. 5A) … K. furcata (Fabricius)

K. furcata species complex

Diagnosis. The K. furcata species complex can be defined by a combination of the following characters: face with fine striae; facial setae spars, more prominent on upper frons, with a row of short transverse setae on anteclypeus; labial palpi 2-3 segmented and females with 9-10 flagellomeres. When there are 10 flagellomeres present, F8-F10 are generally broader than long. There is a variation in facial striae; most species have the frons adjacent to scrobes with oblique striae, traversing between middle ocellar triangle medially to the lower eye margin and lower face with oblique striae from inferior margin of torulus to the posterior genal margin. Two species, K. furcata and K. splendens, however, has striae oblique from the inferior margin torulus to the lateral ocelli and the lower face with transverse striae adjacent to the clypeus. Many of the specimens in this complex have thick frenal processes and robust bodies.

Included taxa. K. cuprea Cameron, 1913; K. furcata (Fabricius, 1804); K. splendens Ashmead, 1904. 22

Kapala cuprea Cameron, 1913 (Figs 4A‒G)

Kapala cuprea Cameron, 1913: 116‒117. Type data: Guiana. Holotype ♂, [by original designation]. Type depository: BMNH; type n° 5.390, examined. [Description of the male]. [Holotype relatively complete: exemplar pinned on a quadrangular carboard; region of the axilla and apex of the midlobe dirty with a green gum (like); digits broken and lost; right hind leg broken after coxa, right medial leg broken after femur]. Thoracantha cynipsea Walker, 1862. Type data: Brazil: Villa Nova. Syntype male. Type depository: BMNH; type n° 5.636b. (Withdraw K. cynipsea syntypes (in part., only one male). New syn. Kapala cynipsea; Ashmead, 1904a: 403 [new combination].

Diagnosis. Recognized by a combination of the following characters: frons and face with oblique striations (Fig. 4B); frenal processes short and stout, 1.1‒1.4x as long as length of axillae and mesoscutellar disc; basal half of frenal processes longitudinaly striated and with strong oblique striae in apical half (Fig. 4E).

Redescription. Male. Body length [4.3] 3.8‒4.6 mm; length of mesosoma excluding the frenal processes [2.6] 2.0‒2.6 mm. Head, mesosoma, coxae, frenal process and petiole black; scapes, pedicels, flagellomeres, rami of flagellum and legs yellow; wings hyaline, venation brown; Gt1 black with light brown apex. Head. [1.5] 1.4‒1.5x as broad as high, with few sparse setae more prominent in frons (Fig. 4B). Scrobal depression with longitudinal striae. Frons with wide oblique striae, start in the ocellar triangle towards the inferior margin of the eyes; lower face with transversal striae below torulus. Eyes separated by [2.0] 1.8‒2.0x their height. Malar space [0.9] 0.7‒0.9x eye height with oblique striae. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape [2.2] 2.2‒3.0x longer than broad, [0.2] 0.2‒0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 [1.0] 1.0‒1.2x head height and [1.0] 0.9‒1.0x as long as following rami. 23

Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and [1.3] 1.2‒1.5x as high as broad. Midlobe with transversal striae in frontal view and, dorsally slightly bilobed in fontal view; lateral lobes with faint longitudinal striae in lateral view. Axilla and scutellar disc with wide longitudinal striae. Mesoscutellar disc [1.5] 1.3‒1.8x as long as axilla with medial apex raised [1.4] 1.0‒1.5 height of frenal processes in lateral view. Axillula longitudinal striated. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with faint longitudinal striae; lower mesepimeron rugose; femoral depression crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad, and delimited by carina, reticulate. Frenal processes cylindrical, short and stout, [1.7] 1.6‒2.2x longer than the basal width of mesoscutellum, [1.1] 1.1‒1.4x as long as length of axillae and mesoscutellar disc, slightly arched lateral view and convex with the apex next to each other, in dorsal view; frenal processes longitudinal striated in basal half and with strong oblique striae in apical half; apex emarginated. Forewing 2.4‒2.6x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.2‒1.4x longer than marginal vein; stigmal vein 1.5‒2.5x longer than broad; postmarginal absent. Hindwing fringed. Metacoxa semiglobose [1.5] 1.5‒2.0x longer than broad with a medial row of semi-erect setae. Metafemur [5.3] 5.1‒6.7x longer than broad, with faint striae and sparse setae. Metasoma. Petiole cylindrical, reticulate with faint longitudinal carina, [3.6] 3.5‒4.0x longer than broad and [1.6] 1.6‒2.0x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 3.1‒5.4 mm; length of mesosoma excluding the frenal processes 1.7‒3.2 mm (Figs. 4D, F). Similar to male except for the following: Antenna with 11 segments; scape 3.3‒3.5x longer than broad; flagellum 9 segmented, length of flagellum 0.9‒1.0x head height (Fig. 4C); Fl2 1.2‒2.0x as long as apical width and 1.2‒1.3x as long as following flagellomere. Mesoscutellar disc 1.2‒1.6x as long as axilla with medial apex raised 0.8‒1.0x height of frenal processes in lateral view. Petiole 3.0‒3.5x longer than broad and 1.1‒1.5x as long as metacoxa. Wings hyaline to eventually infuscate.

Comments. In the present study it was performed the synonymy of the male of K. cynipsea (Walker, 1862) with K. cuprea. After analyzing the syntype of K. cynipsea (BMNH: type n°. 5.636b), it was verified that it has the short and robust frenal process with oblique striae in apical half, as well as oblique striation on the frons and lower face, typical characters of K. cuprea. Also, it was described for the first time the female of K. cuprea.

Biological notes. Associate with Ponerinae (Pachycondyla crassinoda Latreille, 1802) (Meyers, 1931).

Distribution. Bolivia, Brazil, Ecuador, French Guiana, Guyana, Peru, West Indies (Fig. 4G). Holotype. GUIANA [probably British Guiana]: P. Cameron Coll., 1914‒110, [1♂, Type Hym. 5.390, BMNH: UCRCENT00310005]. Examined.

Additional material examined: BOLIVIA: Santa Cruz Dept.: 5km SSE Buena Vista, Hotel Fauna y Flora, 440m, 17°29'56"S, 63°39'08"W, 6-15.x.2003, forest, flight intercept trap, S. & J. Peck, [2♂, CNC: UCRCENT00300676, UCRCENT00300677]. Buena Vista, 17°27'32"S, 63°39'33"W, 8.vii.1973, C. Potter, L. Styange & E. Demares, [2♂, IMLA: UCRCENT00242080, UCRCENT00242081]. Estac. Experimental General Saavedra, 430m, 17°47'38"S, 63°11'00"W, 9.vii.1972, C. Porter & L. Stange, [1♂, IMLA: UCRCENT00313142]. BRAZIL: Amapá: [Pedra Branca do] Amapari – Tucano 2, 0°46'10''N, 51°57'13'', 6-7.xi.1993, Malaise, W. França, [3♂, MPEG]. Idem, 11-15.xi.1993, J. Pena, [5♂, MPEG]. Idem, but serraria, 10-14.xi.1995, equipe, [3♂, MPEG]. Bahia: Porto Seguro, PARNA Monte Pascoal, 16°53'33.65"S, 39°13'0.96"W, 12.viii-2.ix.2013, Malaise, [1♂, UFES]. Mato Grosso: [Campo Novo do Parecis], [Salto] Utiariti, Rio Papagaio, 13°40'30''S,57°53'31''W, x.1966, Lenko & Pereira, [1♂, n° 07549, MZUSP]. Mato Grosso do Sul: [Dois Irmãos do] Buriti, 20°41'23''S, 55°16'43''W, 8.ii.1961, J. & B. Bechyné, [1♂, MPEG]. Pará: Aldeia Coraci 12 km W Caninde, Rio Gurupi, 1°48'38"S, 46°24'03"W, 16- 26.iv.1963, B. Malkin, [1♂, AMNH: UCRCENT00237855]. Benevides, PA 408 - Km 06, 01°21'30''S, 48°14'42''W, 23.iv.1981, Fligth trap, I.S. Gorayeb, [1♂, MPEG]. Idem, 24.iv.1981, [1♂, MPEG]. Idem, 25.iv.1981, [1♂, MPEG]. Rio Araguaia, 19-31.1983, Malaise, J.A. Rafael, [1♂, INPA]. Santarém, 0°57'27"S, 46°59'38"W, [1♀, CMNH: UCRCENT00172445]. Turcurui, 25

3°42'00"S, 49°42'00"W, i.1978, M. Alvarenga, [1♀, UCRC: UCRCENT00305549]. Pernambuco: Caruaru, Serra dos Cavalos, 800 m, 8°22'23''S, 36°01'04''W, 7.viii.2015, Varredura, J.A. Rafael & P.C. Grossi, [1♂, INPA]. Roraima: Ilha de Maracá, Rio Uraricoera, 3°24'N, 61°42'W, 19-24.vii.1987, Malaise, J.A. Rafael & L.S. Aquino, [1♂, INPA]. ECUADOR: Napo: Sacha, 1°04'59.3"S, 77°37'05"W, 9.iii.1983, L. Huggert, [1♂, MZLU: UCRCENT00242568]. Orellana: 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 216m, 0°39'25.7"S, 76°27'10.8"W, 2.vii.1995, terre firme forest, fogging, T.L. Erwin et al., [1♀, UCRC: UCRCENT00091807]. Idem, but 9.vii.1995 [1♀, ????: UCRCENT00092227]. Rio Piraña Bridge, Reserva Etnica Waorani, Onkone Gare Camp, 216.3m, 0°39'25.7"S, 76°27'10.8"W, 17.x.2005, terre firme forest, Fogging, T.L. Erwin, M.C. Pimienta et al., [1♂ 1♀, USNM: UCRCENT00247778, UCRCENT00247775]. Sucumbíos: Napo River, Sacha Lodge, 230m, 0°30'00"S, 76°30'00"W, 13-23.iv.1994, malaise trap, P. Hibbs, [1♂, CNC: UCRCENT00320767]. Idem, but 290m, 3-13.vi.1994, [1♂, CNC: UCRCENT00320768]. Idem, but 4-14.v.1994, [1♂, CNC: UCRCENT00320853]. GUYANA: Bartica, Kartabo, 0-30m, 5°45'15"N, 57°42'16"W, 24.iii.1924 [1♀, AMNH: UCRCENT00238159]. PERU: Madre de Dios: Los Amigos Bio. St., 300m, 12°33'44.4"S, 70°05'47.1"W, 25.xii.2010, trail 10, swp, J. Heraty, [1♂, UCRC: UCRCENT00301932]. Rio Tambopata Res., 30 km air SW Pto. Maldonado, 290m, 12°50'00"S, 69°20'00"W, 6.xii.1982, J.J. Anderson, [1♂, FSCA: UCRCENT00411912]. TRINIDAD: 10°41'26"N, 61°13'16"W, v.2004, [1♂, UCRC: UCRCENT00172511]. Brasso Seco, Rd to Paria Bay, 148m, 10°44'57"N, 61°15'53"W, 25 Jul 2013, forest, swp, Heraty & Baker, [2♂ 1♀, UCRC: UCRCENT00412122‒24]. Curepe, 10°38'48"N, 61°24'56"W, 21.vii.1978, malaise trap, [1♂, UCRC: UCRCENT00305559]. nr. Chaguaramas, 400m, 10°43'01"N, 61°11'52"W, 24.xi.1977, W. & E. Mason, [1♂, CNC: UCRCENT00425741]. Simla Field Station, Arima Valley, 10°41'49"N, 61°17'06"W, 8- 9.iii.1977, tropical rain forest, malaise trap, P. Feinsinger, [1♀, FSCA: UCRCENT00306377]. Simla Res. St., 250m, 10°41'34"N, 61°17'23"W, 22.vii.2013, station, Malaise, Heraty & Baker, [1♂, UCRC: UCRCENT00412133]. Tunapuna, Mt. St. Benedict, 500m, 10°40'20"N, 61°23'51"W, 21-8.vii.1993, Mt. Tabor rainforest, Malaise, S&J Peck, [15♂, UCRC: UCRCENT00320102‒03, UCRCENT00320105‒17]. U.S. VIRGIN ISLANDS: St. Thomas, 1500m, 18°20'17"N, 64°53'39"W, 1859, Thoreg, [1♂, NMW: UCRCENT00317112]. 26

Kapala furcata (Fabricius, 1804) (Fig. 5A‒H)

Eucharis furcata Fabricius, 1804: 158. Type locality: Am.: mer. [=America Meridionali, South America]. Holotype ♀, by monotypy. Lectotype designated by Heraty, 2002:171 [invalid Lectotype designation, ICZN Art. 74.2]. Type depository: ZMUC; Type n° ZMUC: 00241187 (image examined). [Holotype not complete: specimen pinned with body dirty; right wings broken and lost; apical half of left frenal process broken and lost; anterior legs broken and lost]. Thoracantha furcata; Walker, 1839: 22. Chirocerus furcatus; Brullé, 1846: 571. [Kirby (1886) questions the identity of Brullé’s drawing and suggests that it looks closer to Thoracantha atrata Walker than to either the Frabricius species. According Heraty (2002), the drawing appears closer to Kapala sulcifacies (Cameron). The figure legend in Brullé (1846: pl. 38) refers to Chirocerus furcatus Westwood, not Fabricius, and probably refers to a species identified by Westwood. Chirocera furcata; Chenu [Desmarest], 1860: 161, illustrated. Schizaspidia furcata; Walker, 1871: 66. Schisaspidia furcata; Walker, 1872: 65, [misspelling of generic epithet]. Kapala furcata Cameron, 1884: 103, pl. 5, fig. 17. [Change of combination and designation by monotypy as type of Kapala].

Diagnosis. Recognized by a combination of the following characters: frons with oblique wide striae from the torulus to the lateral ocelli, and with transverse striae adjacent to clypeus, often deeply impressed across ventral half of face (Fig. 5B). Female with 9-10 flagellomeres (Fig. 5B). The medial apex of mesoscutellar disc is lower than other species of the furcata complex, and in lateral view is < 1x height of the base of the frenal process (Fig. 5A)

Redescription. Female. Body length 3.9‒5.0 mm; length of mesosoma excluding the frenal processes 2.4‒3.1 mm. Head, mesosoma, coxae, frenal processes and petiole black to 27

green-black; scapes and pedicels brown, flagellum brown to dark-brown; legs pale yellow; wings hyaline to an even light infuscation, venation brown; Gt1 dark-brown. Head. 1.4‒1.5x as broad as high, with few sparse setae more prominent in frons. Scrobal depression with faint irregular striae giving a rugose aspect. Frons with oblique wide striae from the torulus to the lateral ocelli; lower face with transverse striae adjacent to clypeus, often deeply impressed across ventral half of face (Fig. 5B). Eyes separated by 2.1‒2.3x their height. Malar space 0.7‒0.9x eye height, transversally striated. Supraclypeal area and clypeus smooth; anteclypeus with a row of short transverse setae. Maxillary/labial palpi variously segmented 3/3 or 3/2. Antenna with 11 or 12 segments; scape 2.7‒3.5x longer than broad, 0.2‒0.3x head height, smooth surface with sparse setae; flagellum 9‒10 segmented, basal flagellomere cylindrical or slightly serrate, clava rounded (Fig. 5C); length of flagellum 0.8‒1.0x head height; Fl2 1.7‒2.0x as long as apical width and 1.2‒1.7x as long as following flagellomere. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.4‒1.6x as high as broad. Midlobe with transversal striae and dorsally rounded in frontal view; lateral lobes with longitudinal striae. Axilla with longitudinal striae and mesoscutellar disc with slightly oblique striae to the medial apex (Fig. 5F). Mesoscutellar disc 1.5‒1.9x as long as axilla with medial apex raised 0.6‒0.8x height of frenal processes in lateral view. Axillula with longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with weak transverse striae, lower mesepimeron fine reticulate; femoral depression crenulate. Callus slightly swollen, carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat and broad, weakly reticulate and may have some weak longitudinal carinae. Frenal processes cylindrical, 1.9‒2.3x longer than the basal width of mesoscutellum, 1.05‒1.2x as long as length of axilla and mesoscutellar disc; frenal processes, arched in lateral view and convex in dorsal view; frenal processes with longitudinal striae and with apex rounded or emarginated (Figs. 5A, F). Forewing 2.5‒3.4x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.1‒1.4x longer than marginal vein; stigmal vein 2.0‒3.0x longer than broad; postmarginal vein absent. Hindwing fringed. Metacoxa semiglobose 1.7‒1.9x longer than broad, with medial row of setae. Metafemur 6.2‒6.8x longer than broad, smooth with sparse setae. 28

Metasoma. Petiole slightly flattened dorsally with fine parallel longitudinal carinae,

2.8‒3.8x longer than broad and 1.2‒1.5x as long as metacoxa. Gt1 bare except for sparse minute setae; tergal scar present or absent. Male. Body length 3.7‒4.5 mm; length of mesosoma excluding the frenal processes 2.1‒2.8 mm. Similar to female except for the following: Antenna with 12 segments; scape 2.3‒2.7x longer than broad; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 1.2‒1.3x head height and 0.9‒1.0x as long as following rami. Mesoscutellar disc elongated (Fig. 5G). Frenal processes short and thick (Fig. 5G). Metafemur 5.4‒6.3x longer than broad. Petiole 5.3‒6.3 longer than broad and 1.9‒2.5x as long as metacoxa.

Comments. This species has been misidentified in most collections. The type is a very robust individual, at least twice the size of the most Kapala. There is a variation on the females regarding in the number of flagellomeres being 9‒10 and the number of maxillary palpomeres being 2 or 3. Also, some populations of males found in Paraná and São Paulo (Brazil) have thick frenal projections and welmarked longitudinal carinae, besides having irregular striae in the frons giving a rough appearance.

Biological notes. Reported as ovipositing into flower buds of Mikania sp. (Asteraceae) (Clausen, 1940b), although K. furcata is not known to occur in Panamá, where the observation took place. The same species was observed on foliage infested with aphids and were feeding on the honeydew (Clausen, 1941).

Distribution. Argentina, Brazil and Uruguay (Fig. 5H).

Holotype. Am.: mer. [=America Meridionali, South America]: Schmidt, [1♀, ZMUC 00241187, ZMUC].

Additional material examined: ARGENTINA: Misiones: Santa Ana, near Loreto, 84m, 27°20'11"S, 55°31'51"W, 27.iii.2003, humid forest, J. Heraty, [1♀, UCRC: 29

UCRCENT00092093]. Salta: Finca El Rey, 1000m, 24°42'00"S, 64°38'00"W, ii.1953, N. Kusnezov, [1♀, IMLA: UCRCENT00313148]. Tucuman: Siambon, 26°42'00"S, 65°27'00"W, i.1945, D. Olea, [1♂, IMLA: UCRCENT00242082]. Tucuman, 26°48'30"S, 65°13'03"W, i.1947, Cordoba, [1♀, IMLA: UCRCENT00313138]. BRAZIL: Rio Grande do Sul: -, 29°32'04"S, 53°23'26"W, Stieglmayr, [4♂ 6♀, NMW: UCRCENT00242559‒62, UCRCENT00242564, UCRCENT00242566, UCRCENT00317114‒15, UCRCENT00317118‒19]. CPCN Pro-Mata, 325m, 29°30'00"S, 50°10'00"W, 4.iv.1997, J. Ketterl, [1♂, SMNS: UCRCENT00318556]. Idem, but 30.iv.1997, [1♂, SMNS: UCRCENT00318557]. Santa Catarina: Corupa (Hansa Humbolt), 26°25'34"S, 49°14'36"W, xii.1940, A. Maller, [1♀, AMNH: UCRCENT00238157]. Pinhal, 27°14'00"S, 51°55'00"W, iv.1947, A. Maller, [1♂, AMNH: UCRCENT00238160]. Rio Natal, 26°21'00"S, 49°18'00"W, i.1945, A. Maller, [3♀, AMNH: UCRCENT00238151, UCRCENT00238153‒54]. Rio Vermelho, 27°29'28"S, 48°24'32"W, ii.1945, A. Maller, [2♀, AMNH: UCRCENT00238150, AMNH: UCRCENT00238152]. Idem, but vi.1958, Dirings, [1♀, MZUSP n° 07555, MZUSP]. São Paulo: Ribeirão Grande, Parque Estadual Intervales, 24°16’23.6S, 48°25’21.08’’W, 22.i.2010, Malaise - Ponto 1, N.W. Perioto e eq., [1♂ 1♀, LRRP]. Idem, but Malaise - Ponto 5, [1♂, LRRP]. Idem, but 21.i.2011, Malaise - Ponto 5, [1♂, LRRP]. Idem, 21.iii.2011, [2♂, LRRP]. Idem, 22.iv.2011, [1♂, LRRP]. URUGUAY: Tacuarembo: 40 km NW Tacuarembo, 200-300m, 31°29'45"S, 56°18'08"W, 2-9.ii.1963, J.K. Bouseman, [2♂, AMNH: UCRCENT00237807, AMNH: UCRCENT00238155].

Kapala splendens Ashmead, 1904 (Figs. 6A‒F)

Kapala splendens Ashmead, 1904a: 473. Type data: Brazil: Chapada. Holotype ♀, by original designation. Type depository: U.S. National Entomological Collection, United States National Museum (USNM), Washington, DC, USA; type nº 8084, (examined). [Description of female, illustrated. Holotype relatively complete: apex of antennae broken and lost; left middle leg broken and lost; coxa and hind legs broken and lost; apex of right and left forewings broken; right and left hindwing with apical half broken; abdomen and petiole broken and glued erroneously below the frenal process]. 30

Diagnose. Recognized by a combination of the following characters: body very robust than the others Kapala species; margin of gena broad and angulate (Fig. 6C); frons with fine longitudinal striae curving towards the supraclypeal area and face with fine transversal striae; flagellum 8 or 9 segmented; head and mesosoma brilliant metallic green and metasoma blue with green reflections.

Redescription. Female. Body length [8.3] 4.4‒8.3 mm; length of mesosoma excluding the frenal processes [4.7] 2.3‒4.7 mm. Head, mesosoma, coxae and petiole green to bright metallic blue; frenal process metallic blue; scapes, pedicels and legs yellow, flagellomeres light brown; wings slightly infuscate, venation brown; Gt1 blue with green reflections. Head. [1.5] 1.4‒1.5x as broad as high, with few sparse setae. Scrobal depression rugose with some striae. Frons with fine longitudinal striae curving towards the supraclypeal area; face with fine transversal striae (Fig. 6C). Eyes separated by [2.2] 1.9‒2.2x their height. Malar space [0.9] 0.8‒0.9x eye height with oblique striae. Gena with margin broad and angulate (Fig. 6C). Supraclypeal area with upper half with transverse striae and basal half smooth. Clypeus smooth, laterally marked by a shallow groove. Maxillary/labial palpi 3/3 segmented. Antenna with 10 or 11 segments; scape [2.8] 2.8‒4.0x longer than broad, [0.3] 0.2‒0.3x head height, smooth surface with sparse setae; flagellum 8‒9 segmented (Fig. 6B), length of flagellum [1.1] 1.0‒1.1x head height; Fl2 [1.6] 1.4‒1.6x as long as apical width and [1.2] 1.0‒1.2x as long as following flagellomere. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, in lateral view, and [1.3] 1.3‒1.8x as high as broad. Midlobe with fine transverse striae and dorsally slightly bilobed in frontal view; lateral lobes with fine longitudinal striae. Axilla with fine longitudinal striae and mesoscutellar disc with fine striae slightly oblique (Fig. 6D). Mesoscutellar disc [1.6] 1.5‒1.7x as long as axilla with medial apex raised [0.6] 0.6‒0.7x height of frenal processes in lateral view. Axillula with fine longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with some longitudinal striae, lower mesepimeron fine reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and delimited by carina, smooth. Frenal processes cylindrical, [1.6] 1.0‒1.8x longer than 31

the basal width of mesoscutellum, [1.2] 1.2‒1.3x as long as length of axilla and mesoscutellar disc; frenal processes, arched in lateral view and slightly convex in dorsal view and with longitudinal striae and with apex rounded (Figs. 6A, D). Forewing [2.8] 2.5‒2.8x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.6] 1.5‒1.6x longer than marginal vein; stigmal vein [1.2] 1.2‒1.6x longer than broad; postmarginal vein short. Hindwing fringed, 7 hamulli curved hook-shaped. Metacoxa semiglobose [1.6] 1.5‒1.7x longer than broad. Metafemurr [4.7] 2.8‒4.7x longer than broad, smooth with sparse setae. Metasoma. Petiole slightly flattened dorsally, fine reticulate and with weak longitudinal striae, [3.1] 3.1‒3.4x longer than broad and [1.5] 1.5‒1.6x as long as metacoxa. Gt1 smooth and bright with tergal scar present.

Male. Body length 6.0 mm (Fig. 6E); length of mesosoma excluding the frenal process 3.5 mm. Similar to female except for the following: supraclypeal area completely covered by transverse striae; upper region of the clypeus with soft transverse striae. Antenna with 12 segments, rami of Fl2 1.1x head height. Mesoscutellar disc 2.0x as long as axilla (Fig. 6E). Frenal processes 2.0x longer than the basal width of mesoscutellum. Petiole 7.7x longer than broad and 2.6x as long as metacoxa.

Biological notes. Unknown.

Distribution. Argentina (Misiones), Brazil (Goiás, Mato Grosso and Rio de Janeiro) (Fig. 6F). Holotype. BRAZIL: Chapada, in April and November, H.H. Smith coll., [1♀, USNMENT 00809391, USNM type n° 8084].

Additional material examined: ARGENTINA: 38°30'42.01"S, 63°38'23.89"W, [1♀, MLPA]. Misiones: San José, 27°46'12.18"S, 55°46'49.33"W, xii.1942, Bridarolli, S.J., [1♂, MACN]. BRAZIL: Goiás: Alto Paraíso de Goiás, BR 010, 18°56'48''S, 47°25'53''W, 1400m, 28.xii.2011, V. Kanamura & K. Ramos, [1♀, MZUSP]. Goiás, 15°56'02''S, 50°08'24''W, 26.iii.1978, A. Raw, [1♀, UCRC_ENT 297983, BMNH]. Mato Grosso: Chapada, 15°27'29.90"S, 55°45'8.85"W, H.H. 32

Smith, [2♀, Acc.2966, UCRC_ENT 00240757, CMNH; Acc. 2966, UCRC_ENT 00240756, CMNH]. Rio de Janeiro: 22°54'54.90"S, 43°10'7.27"W, [1♀, UCRC_ENT 00416661, USNM].

K. iridicolor species complex

Diagnosis. The iridicolor species complex can be defined by a combination of the following characters: frons swollen, smooth (wide longitudinal striated in specimens from Mexico and Honduras) and with sparse setae, more prominent on upper frons and vertex; lower face smooth; females with flagellum 9 segmented. Mesoscutum low and smoothly curved anteriorly in lateral view; lateral lobe of mesoscutum smooth and pilose. Mesoscutellum elongated. The dorsolateral projection of prepectus is an elongate triangle, broadly rounded at the apex.

Included taxa. K. iridicolor (Cameron, 1904).

Kapala iridicolor (Cameron, 1904) (Figs. 7A‒G)

Lirata iridicolor Cameron, 1904: 60. Type data: Nicaragua: Chinandega. Holotype ♀, by monotype. Type depository: BMNH, Type nº 5.385, examined. [Holotype relatively complete: right medial leg broken after tibia; left hind leg broken after the coxa and glued to the side of the body; metasoma broken and lost]. Kapala iridicolor; Heraty & Woolley, 1993: 522. [description of both sexes, illustrated.].

Diagnosis. Recognized by a combination of the following characters: frons swollen and smooth (Fig. 7B); mesoscutum smoothly curved anteriorly (Fig. 7C, indicated); frenal processes cylindrical and very thin with apex acuminate (Figs. 7E, F); female with mesoscutellar disc smooth except for some longitudinal weak striae laterally and a short flagellum 0.8–0.9x head height (Fig. 7C).

Redescription. Female. Body length [3.3] 1.8‒3.3 mm; length of mesosoma excluding the frenal processes [1.7] 1.1‒1.9 mm. Head e mesosoma black; frenal processes dark brown at the base and light brown at the apical half; scapes and pedicels yellow; flagellum brown; coxae dark brown; legs light yellow; wings hyaline, venation brown; Gt1 black with apex light brown. Head. [1.4] 1.3‒1.4x as broad as high, smooth surface with sparse setae, more prominent on upper frons and vertex (Fig. 7B). Frons swollen. Scrobal depression with smooth surface. Eyes separated by [1.9] 1.9‒2.2x their height. Malar space [0.8] 0.8‒1.1x eye height, smooth and shiny except for some faint striae. Supraclypeal area and clypeus smooth, clypeus laterally marked by a shallow groove. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 segments; scape [4.1] 3.3‒4.2x longer than broad, 0.3x head height, smooth surface with sparse bristles; flagellum 9 segmented, cylindrical and slightly serrated, clava rounded (Fig. 7C); length of flagellum [0.9] 0.8‒0.9x head height; Fl2 [2.0] 2.0‒3.3x as long as apical width and [1.3] 1.1‒1.5x as long as following flagellomere. Mesosoma. Mesocutum anteriorly smoothly curved in lateral view and [1.3] 1.1‒1.3x as high as broad (Fig. 7C). Midlobe with transverse striae and dorsally rounded in frontal view; lateral lobes smooth. Axilla with longitudinal striae. Mesoscutellar disc with smooth surface except for some longitudinal weak striae laterally, mesoscutellar disc [1.2] 1.2‒1.6x as long as axilla with medial apex raised [0.9] 0.8‒1.2x height of frenal processes in lateral view (Fig. 7A). Axillula smooth to with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth (Fig. 7C); lower mesepimeron weakly reticulate; femoral depression weakly crenulate. Callus carinate and weakly reticulate, covered by semi-erect setae. Metapleuron carinate and slightly reticulate. Propodeum flat, broad, smooth and delimited by carina. Frenal processes cylindrical and very thin, [1.7] 1.5‒2.6x longer than the basal width of mesoscutellum, [1.5] 1.3‒1.5x as long as length of axillae and mesoscutellar disc, arched in lateral view and convex in dorsal view; frenal processes with longitudinal striae and with apex acuminate (Figs. 7A, E). Forewing [2.5] 2.2‒2.7x longer than broad; covered by small bristles, except for the base bare; submarginal vein [1.3] 1.2‒1.4x longer than marginal vein; stigmal vein oblique rectangular, [2.7] 2.0‒2.7x longer than broad and 2.7X longer than postmarginal vein. Hindwing fringed, 4 hamulli curved hook-shaped. Metacoxa 34

semiglobose with sparse setae, [1.5] 1.5‒2.0x longer than broad. Metafemur [7.2] 5.2‒7.2x longer than broad, weakly striated with sparse setae. Metasoma. Petiole cylindrical, weakly reticulate with some fine striae, 3.0‒3.5x longer than broad and 1.4‒1.8x as long as metacoxa. Gt1 smooth; tergal scar on Gt1 present or absent.

Male. Body length 1.8‒3.6 mm, length of mesosoma excluding the frenal processes 1.1‒1.9 mm (Fig. 7D). Similar to female except for the following: malar space 0.7‒1.1x eye height; antenna with 12 segments; scape 2.0‒2.7x longer than broad, 0.2‒0.3x head height; length of rami of Fl2 1.0‒1.2x head height and 0.9‒1.0x as long as following flagellomere (Fig. 7D). Mesoscutellar disc with longitudinal striae (Fig. 7F); mesoscutellar disc 1.1‒1.9x as long as axilla with medial apex raised 1.0‒1.5x height of frenal processes in lateral view. Frenal processes 1.5‒2.7x longer than the basal width of mesoscutellum, 1.0‒1.5x as long as length of axillae and mesoscutellar disc. Petiole 4.8‒7.5x longer than broad and 2.1‒2.8x as long as metacoxa.

Comments. This species, according to Heraty & Woolley (1993) and Murray et al. (2013), is inserted within the K. iridicolor complex species, whose individuals share some characteristics like, face smooth and all females having 9-10 flagellomeres and a low profile of the mesocutum. Some specimens from Mexico and Honduras have faint striae on the frons, but not on the face, probably a new species within the complex. Much of the diversity and species of the K. iridicolor complex are found in northen South America and Central America.

Biological notes. Known to oviposit into closed flower buds of Cordia macrostachya, Cordia cana (Boraginaceae) and Gossypium hirsutum (Malvaceae) (Heraty & Woolley, 1993). Also collected on Gossypium hirsutum L., Hamelia patens Jacq., Melampodium divaricatum L., Onoseris onoseroides (Kunth.) B.L. Rob., Panicum sp., Phoradendron sp.; frutification of Carica papaya L., Passiflora edulis L., Terminalia catappa L.; flowering cactus-like Asclepiad.

Distribution. Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Honduras, Mexico, Nicaragua, Panama, Suriname, Peru and Venezuela (Fig. 7G). Usually occurs in sympatry with K. romandii. 35

Holotype. NICARAGUA: Chinandega, Baker coll., [♀, BMNH type n°5.385, BMNH: UCRCENT 310008].

Additional material examined. BRAZIL: Roraima: Ilha de Maracá, 3°24'N 61°42'W, 02- 13.v.1987, Malaise, J.A Rafael & equipe, [1♂, INPA]. COLOMBIA: Magdalena: PNN Tayrona Zaino, 50m, 11°20'01"S, 74°02'01"W, 14-29.vi.2000, Malaise trap, R. Henriques, [1♂, UCRC: UCRCENT00172493]. Boyacá: Muzo, 5°32'00"N, 74°06'00"W, Westwood, [1♂, ZSMC: UCRCENT00245267]. Cauca Valley: near Palmira, 1000m, 3°35'02"N, 76°15'01"W, v.1986, A. Braun, [1♀, UCDC: UCRCENT00243536]. 11 km W. Santa Marta, Pozo Colorado, 11°10'00"N, 74°14'00"W, 25-30.iv.1968, Borys Malkin, [1♀, EMEC: UCRCENT00236312]. PNN Tayrona Zaino, 50m, 11°20'00"N, 74°02'00"W, 13-30.v.2000, Malaise trap, R. Henriquez, [2♂, UCRC: -, UCRCENT00092217]. Idem, but 17-28.vii.2000, [1♂, UCRC: UCRCENT00092131]. Idem, but 14-29.vi.2000, [3♂, UCRC: UCRCENT00091815, UCRCENT00091822, UCRCENT00091850]. Idem, but 14-29.vii.2000, [1♂, UCRC: UCRCENT00092101]. Idem, but 28.iv-3.v.2000, [4♂, UCRC: UCRCENT00092119, UCRCENT00091816, UCRCENT00306546, UCRCENT00306555]. Idem, but 28.vi-17.vii.2000, [1♂, UCRC: UCRCENT00306543]. Idem, but 29.v-14.vi.2000, [1♂, UCRC: UCRCENT00306544]. Valle del Cauca: Cadelaira, Finca San Luis, 1010m, 3°24'00"N, 76°23'00"W, 4.vii.1975, tropical dry forest, Malaise trap, R.C. Wilkerson, [1♀, FSCA: UCRCENT00306477]. Central de Anchicaya, 3°52'56"N, 76°48'02"W, 28.i.1975, Malaise trap, R. Wilkerson, [1♂, FSCA: UCRCENT00306443]. Valle Rio Anchicaya: -, 400m, 10.ii.1977, Breed & C.D. Michener, [1♀, UCRC: UCRCENT00446413]. COSTA RICA: Curubande, 10°43'00"N, 85°25'00"W, xi.1981, forest edge, sweep, S.G. Compton, [1♀, SAMC: UCRCENT00246690]. Idem, but woodland, [1♀, SAMC: UCRCENT00246689]. Guardia, 40m, 10°33'54"N, 85°35'18"W, 21.vii.1967, A.D. Banegas, [1♀, EMEC: UCRCENT00236307]. Alajuela: Chiles de Aguas Zarcas, café, 300m, 10°23'56"N, 84°20'20"W, i.1990, R. Cespedes, [1♂, UCRC: UCRCENT00274871]. Idem, but xii.1989, [1♂ 1♀, UCRC: UCRCENT00302321‒22]. Idem, but iii.1990, P. Hanson & R. Cespedes, [1♂, UCRC: UCRCENT00320244]. San Pedro de la Tigra, Cacao, 200m, 10°22'07"N, 84°34'55"W, i-ii.1990, R. Cespedes, [2♂, UCRC: UCRCENT00274879, UCRCENT00302325]. Idem, but iii-iv.1990, [1♀ 5♂, UCDC: UCRCENT00243537, UCRC: UCRCENT00274876‒78, UCRCENT00302324, 36

UCRCENT00302326]. Idem, but xii.1990, [1♂, UCRC: UCRCENT00274870]. Guanacaste: ACG, Santa Rosa, 300m, 10°50'22"N, 85°37'06"W, 8-18.vi.1995, S. Dadelahi, L. Price & N. Zitani, [4♀, UCRC: UCRCENT00318838‒40, UCRCENT00320089]. Cerro el Hacha, NW Volcan Orosi, 300m, 10°59'57"N, 85°33'03"W, 1988, [5♂, UCRC: UCRCENT0027487‒75, UCRCENT00302323]. La Pacifica, 4 km. N.W. Canas, 10°27'16"N, 85°07'54"W, 8.ix.1971, Cordia cana, [1♀, EMEC: UCRCENT00236290]. Santa Rosa NP, 300m, 10°53'33"N, 85°45'59"W, 16.xi-7.xii.1985, D.H. Janzen & I.D. Gauld [1♂, UCRC: UCRCENT00320091]. Idem, but 27.iv-11.v.1985, [1♂, UCRC: UCRCENT00320090]. Santa Rosa NP Bosq. Hum. (11O), 10°53'33"S, 85°45'59"W, [1♂, UCRC: UCRCENT00274882]. Santa Rosa NP Sn. Emilio (6C), 10°52'15"N, 85°43'38"W, [1♂, UCRC: UCRCENT00274880]. Heredia: Est. Biol. La Selva, 50-150m, 10°20'00"N, 84°01'00"W, 1.vi.1993, [3?, UCRC: UCRCENT00446451, UCRCENT00446462, UCRCENT00446463]. Idem, but 1.iii.1994, [1?, UCRC: UCRCENT00446464]. F. La Selva 3km S Pto. Viejo, 80m, 10°26'00"N, 84°01'00"W, 29- 31.vii.1976, E.M. Fisher, [1♂, UCRC: UCRCENT00320316]. La Selva Biol. Sta., 64m, 10°25'49"N, 84°00'26"W, 14.viii.2010, Hamelia patens and nearby bushes, sweep, J. Heraty, [1♀, UCRC: UCRCENT00282474]. Puntarenas: Manuel Antonio National Park, 9°23'45"N, 84°08'16"W, 23-28.viii.1986, costal rainforest, screen sweep, L. Masner, [1♂, CNC: UCRCENT00315911]. Parque Nacional Corcovado, Est. Sirena, 8°30'00"N, 83°36'00"W, 22.iii.1981, H.A. Hespenheide, [1♂, UCRC: UCRCENT00305602]. ECUADOR: Bucay, 304.8m, 2°12'17"S, 79°10'40"W, 4.x.1922, F.X. Williams, [2♀, EMEC: UCRCENT00236291‒92]. Microwave tower, 1°49'52"S, 78°11'00"W, 6-7.iii.1976, Malaise trap, [1♂, CNC: UCRCENT00315880]. Rio Palengue, 150m, 0°35'11"S, 79°21'53"W, 4.ii.1983, forest, L. Huggert, [1?, MZLU: UCRCENT00242617]. Carchi: Chical, 1250m, 1°01'32"N, 78°14'28"W, viii.1983, YPT, J.E. Rawlins & M. Smyers, [1?, UCRC: UCRCENT00416377]. Esmeraldas: Bilsa Biol. Sta., 500m, 0°20'24"N, 79°42'36"W, 10.v-4.vi.1996, malaise trap, P. Hibbs, [1♂, UCRC: UCRCENT00092085]. Borbon, 1°05'21"N, 78°59'24"W, 1.ii.1988, Mike Huybensz, [1♀, MCZ: UCRCENT00242352]. Zapallo Grande, 0°47'00"N, 78°59'00"W, 25- 30.x.1987, Mike Huybensz [1♂, MCZ: UCRCENT00242385]. Idem, but 4.ii.1988, [2♂, MCZ: UCRCENT00242383‒84]. Guayas: Isla Puna, 2°46'35"S, 80°08'34"W, 22.iii.1988, Mike Huybensz, [1♂, SAMC: UCRCENT00246691]. Napo: Misahualli, 417m, 1°01'57"S, 37

77°40'13"W, 18.ii.1983, L. Huggert, [3?, MNCN: UCRCENT00322500, MZLU: UCRCENT00324495‒96]. Orellana: 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 216m, 0°39'26"S, 76°27'11"W, 4.ii.1996, terre firme forest, fogging, T.L. Erwin et al., [1♂, UCRC: UCRCENT00320096]. Idem, but 8.x.1995, [1♂, UCRC: UCRCENT00320097]. Tiputini Biodiversity Sta. nr. Yasuni National Park, Trans. Ent., 220-250m, 0°37'55"S, 76°08'39"W, 2.ii.1999, terra firme forest, Fogging, T.L. Erwin et al., [1♀, UCRC: UCRCENT00320093]. Pichincha: 16km SE Santo Domingo, Tinalandia, 680m, 0°18'17"S, 78°59'00"W, 15-30.vi.1975, S. & J. Peck, [2♂, CNC: UCRCENT00241041‒42]. 45 km S. Santo Domingo, Rio Palenque, 0°39'05"S, 79°19'33"W, 5.v-25.vii.1985, rain forest, FIT, Peck, [3♂, CNC: UCRCENT00316114‒16]. 47 km S Santo Domingo, Rio Palenque Stn., 160-180m, 0°35'11"S, 79°21'53"W, 29.iv-5.v.1987, rainforest, MT, B .Brown & L. Coote, [1?#, CNC: UCRCENT00315883]. Idem, but 250m, 0°39'05"S, 79°19'33"W, 17-25.ii.1979, S.A. Marshall, [11♂, CNC: UCRCENT00316001, UCRCENT00316024, UCRCENT00316052‒54, DEBU: UCRCENT00251183‒86, UCRCENT00251194‒95]. Idem but 14.vii.1975, A. Forsythe, [33♂, CNC: UCRCENT00315869‒70, UCRCENT00315872, UCRCENT00315874, UCRCENT00316006‒23, UCRCENT00316026‒36]. Idem, but 15.vi.1975, S. & J. Peck, [5♂ 1♀, CNC: UCRCENT00241009‒13, UCRCENT00313592]. Idem, but 18-30.v.1975, [77♂, CNC: UCRCENT00241015, UCRCENT00241017‒37, UCRCENT00300682, UCRCENT00313601‒04, UCRCENT00313606‒11, UCRCENT00315905‒06, UCRCENT00315916, UCRCENT00316071‒85, UCRCENT00316087‒94, UCRCENT00316096‒6113]. Idem, but 22-31.iii.1976, [10♂, CNC: UCRCENT00316037‒46]. Idem, but 28.v.1975, [1♂, CNC: UCRCENT00241014]. Idem, but 29.v.1975, [3♂ 1♀, CNC: UCRCENT00241038‒40, UCRCENT00313652]. 47km S. Santo Domingo, Rio Palenque Station, 200m, 0°39'05"S, 79°19'33"W, [1♂, CNC: UCRCENT00315875]. Rio Palenque, 200m, 0°36'00"S, 79°21'00"W, 6.iii-1.iv.1996, malaise trap, P. Hibbs, [2♂, UCRC: UCRCENT00172491‒92]. Rio Palenque, 0°08'48"S, 78°50'18"W, 25.iii.1983, L. Masner, [2♂ 1?, UCRC: UCRCENT00092037, UCRCENT00092096, UCRCENT00446422]. Rio Palenque, 0°39'05"S, 79°19'33"W, 20.ii.1979, S.A. Marshall, [1♂, DEBU: UCRCENT00251196]. Idem, but ii.1976, G. Shewel, [4♂, CNC: UCRCENT00316048‒51]. Idem, but Res. Sta., 0°39'05"S, 79°19'33"W, vi-viii.1985, S & J Peck, [1♂, CNC: UCRCENT00316118]. Idem, but ii.1983, MT, 38

Sharkey & Masner, [100♂ 4♀ 11?, CNC: UCRCENT00240971‒73, UCRCENT00240975‒79, UCRCENT00240981‒82, UCRCENT00313612‒16, UCRCENT00313618‒29, UCRCENT00313631‒37, UCRCENT00313639‒40, UCRCENT00313642, UCRCENT00313644‒48, UCRCENT00313651, UCRCENT00313653‒54, UCRCENT00313656‒66, UCRCENT00315812‒13, UCRCENT00315815, UCRCENT00315818‒29, UCRCENT00315831, UCRCENT00315833‒35, UCRCENT00315838‒39, UCRCENT00315841‒44, UCRCENT00315847‒50, UCRCENT00315854, UCRCENT00315861‒64, UCRCENT00316055‒58, UCRCENT00316060‒67, UCRCENT00316069‒70; UCRC: UCRCENT00305605, UCRCENT00446428, UCRCENT00446430‒38]. Idem, but 23-31.vii.1976, S. & J. Peck, [1♂, CNC: UCRCENT00316047]. Idem, but 22-26.ii.1976, malaise trap, G. Shewel, [2♂, CNC: UCRCENT00315878‒79]. Rio Palenque Stn., 0°39'05"S, 79°19'33"W, Malaise trap, [4♂, CNC: UCRCENT00240962‒65]. Idem, but 190m, 1-5.v.1987, lowland rainforest, sweep, L. Coote, [4♂, CNC: UCRCENT00240966‒68, UCRCENT00240970]. Idem, but 200m, 11.iv.1983, Sharkey & Masner, [33♂ 1♀, CNC: UCRCENT00240983‒93, UCRCENT00240995‒98, UCRCENT00241000‒01, UCRCENT00313593‒600, UCRCENT00315855‒60, UCRCENT00316117, UCRCENT00316231‒32]. Idem, but 200m, 0°08'48"S, 78°50'18"W, 6.iii- 1.iv.1996, malaise trap/pan trap, P. Hibbs, [2♂ 1?, UCRC: UCRCENT00092125, UCRCENT00446420, ????: UCRCENT00175171]. Rio Palenque, 7-25.ii.1979, S. Marshall, [2?, UCRC: UCRCENT00446414, UCRCENT00446510]. Tinalandia, 800m, 0°16'47"S, 78°57'48"W, 8.ii.1983, Masner & Sharkey, [1♀, CNC: UCRCENT00315866]. Idem, but Science Ctr, 200m, 0°36'00"S, 79°21'00"W, 25.iv-6.vii.1996, flight intercept trap/malaise trap, P. Hibbs, [1♂, UCRC: UCRCENT00091809]. Zamora Chinchipe: Rio Bombuscaro, 1100m, 4°07'12"S, 78°58'48"W, 26.iv-4.vii.1996, P. Hibbs, malaise trap/pan trap, [1♂, UCRC: UCRCENT00091806]. EL SALVADOR: Quezaltepeque, 13°50'00"N, 89°16'00"W, 1.vii.1963, D. Cavagnaro & M.E. Irwin, [1♂ 1♀, EMEC: UCRCENT00236305, UCDC: UCRCENT00279932]. Idem, but 17.vi.1963, [1♂, UCDC: UCRCENT00279933]. Idem, but 18.vi.1963, [1♂ 1♀, UCDC: UCRCENT00279931, UCRC: UCRCENT00306553]. Idem, but 22.vi.1963, [1♀, UCDC: UCRCENT00279930]. Idem, but 21.vi.1961, M.E. Irwin [2M#, UCDC: UCRCENT00279928‒29]. HONDURAS: C.A., EAP, 14°01'02"N, 87°01'39"W, 13.vi.1982, 39

R.W. Jones, [1♂, CNC: UCRCENT00241044]. Idem, but 24.iv.1982, [1♀, CNC: UCRCENT00241043]. Comayagua: 3 km S Comayagua, 600m, 14°24'47"N, 87°37'25"W, 23.vii.1989, R. Cave, [2♀, EAPZ: UCRCENT00282629, UCRCENT00282635]. Comayagua, 14°27'15"N, 87°38'35"W, 10.vii.1964, G.A. Axtell [1♂, UCDC: UCRCENT00279927]. El Paraiso: Moroceli, El Mesias, 14°07'12"N, 86°46'48"W, 20.iii.1991, Carica papaya fructification, M. Avila, [1♀, EAPZ: UCRCENT00316462]. Rapaco, 630m, 14°04'56"N, 86°54'26"W, 17.v.1993, Pimenta vegetative, B. Castro, [1♀, EAPZ: UCRCENT00316463]. Francisco Morazan Dept.: 32 Km E. Tegucigalpa, El Zamorano, 14°01'02"N, 87°01'39"W, 24.ix.1985, R. Caballero, [1♂, EAPZ: UCRCENT00282632]. San Antonio de Oriente, El Zamorano, 14°01'02"N, 87°01'39"W, 19.x.1988, P1assiflora edulis fructificacion, R. Cave, [1♂, EAPZ: UCRCENT00282636]. El Zamorano, 14°01'02"N, 87°01'39"W, 22-29.iii.1990, coffee plantation ‘baja Inga’, malaise, R. Cave, [1♂, EAPZ: UCRCENT00316472]. Idem, but 4- 10.v.1990, [1♂, EAPZ: UCRCENT00316473]. El Zamorano, 14°01'02"N, 87°01'39"W, 28.vi.1988, R. Cave, [1♀, EAPZ: UCRCENT00282633]. Est. Agr. Pan., Zamorano, 14°00'29"N, 87°00'35"W, 19.vii.1948, in monte, T.H. Hubbell, [1♀, FSCA: UCRCENT00306467]. Idem, but, 792.48m, 4.vii.1948, on gardenia, [1♀, UMMZ: UCRCENT00243582]. Olancho: 22 km. S La Union, 30.xi.1995, R. Turnbow, [1?#, FSCA: UCRCENT00424509]. El Boquerón Nat. Mon., 14°47'06"N, 86°00'42"W, 2.vii.2002, D. Yanega, [2♂, UCRC: UCRCENT00092077, UCRCENT00172479]. Tulin, 14°43'00"N, 86°11'00"W, 7.v.1979, C. Henriquez, [1♂, UCRC: UCRCENT00305998]. Catacamas, Catacamas, 1000m, 14°50'51"N, 85°53'32"W, 28.iv.1993, Phoradendron sp., R. Cave, [1♀, EAPZ: UCRCENT00316471]. San Francisco de la Paz, El Ocote, 15°26'11"N, 86°35'53"W, 16.vii.1989, Onoseris onoseroides, G. Marquez, [1♂, EAPZ: UCRCENT00282634]. Silca, El Carbonal, 14°47'36"N, 86°26'28"W, 15.x.1988, R. Cave, [1♂ 1♀, EAPZ: UCRCENT00282630‒31]. MEXICO: Chiapas: Rosario Izapa, 14°58'00"N, 92°09'00"W, 11.viii.1997, P. Lachaud, [2♀, ????: UCRCENT00002411, UCRCENT00002416]. Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 16.viii.2004, E. ruidum nest, G. Perez, [3♂ 3♀, UCRC: UCRCENT00235924, UCRCENT00318828‒31, ????: UCRCENT00002417]. Idem, but 19.vii.2002, Melampodium divaricatum, [1♀, ????: UCRCENT00002410]. Idem, but 5.vii.2004, on Panicum sp., [1♀, UCRC: UCRCENT00318837]. Tapachula, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 16.vii.2004, Melampodium divaricatum, G. Perez, [1♂ 1♀, 40

UCRC: UCRCENT00318835‒36]. Nuevo Leon: Salinas Victoria, 20 km N. Carr. Colombia, 450m, 26°02'03"N, 100°18'18"W, 13.vii.1983, A. Gonzalez, [3♂, UANL: UCRCENT00320609‒11]. Quintana Roo: Lazaro Cardena, 25 km NNE Leona Vicario Reserva Ecologia El Eden, 21°13'00"N, 87°11'00"W, 10.viii.1998, secondary growth near greenhouse, sweep, R. Rodriguez, [1♂, UCRC: UCRCENT00320099]. 25 km NNE Leona Vicario Reserva Ecologica El Eden, 21°13'00"N, 87°11'00"W, 19.viii.1998, savannah de cabana, sweep, R. Rodriguez, [1♂, UCRC: UCRCENT00320100]. Tamualipas: Gomez Farias, 23°01'49"N, 99°08'53"W, [1♂, UCRC: UCRCENT00320098]. NICARAGUA: near Comalapa, 12°16'55"N, 85°30'44"W, 1.xii.1976, from dooryard Gossypium hirsutum, swept, W.H. Cross, [1♀, MMNS: UCRCENT00242467]. Idem, but 1.xii.1976, on flowering cactus-like Asclepiad, [1♀, MMNS: UCRCENT00242468]. Chinandega: Chinandega, 12°37'28"N, 87°07'47"W, Baker, [2♂ 1♀, CUIC: UCRCENT00241291, UCRCENT00241293, UCRCENT00300485]. Managua: Tipitapa, 50m, 12°11'47"N, 86°05'44"W, 24.ix.1991, Terminalia catapa fructification, R. Caballero, [9♂, EAPZ: UCRCENT00316464‒70, UCRCENT00316474‒75]. PANAMA: Cerro Galera, 8°56'00"N, 79°37'00"W, 9.ix.1983, Neutusij, [1♂, ZMUH: UCRCENT00245182]. David, 40m, 8°25'40"N, 82°25'50"W, 20.vi.1978, G.J. Umphrey, [1♀, UCRC: UCRCENT00306355]. Canal Zone: 1mi. W Frijoles, 20-50m, 9°10'34"N, 79°48'21"W, 18.vii.1977, R.B. & L.S. Kimsey, [3♂, UCDC: UCRCENT00243502‒04]. Barro Colorado Island, 20-180m, 9°09'20"N, 79°50'46"W, 1.vii.1976, R.B. & L.S. Kimsey, [1♂, UCDC: UCRCENT00243517]. Idem, but 10.viii.1981, [1♂, UCDC: UCRCENT00243514]. Idem, but 12.viii.1978, R.B. & L.S. Kimsey, [2?#, UCDC: UCRCENT00243507, UCRCENT00243511]. Idem, but 13.vii.1976, [1♂, UCDC: UCRCENT00243512]. Idem, but 15.ix.1976, [2♂, UCDC: UCRCENT00243508, UCRCENT00243510]. Idem, but 15.vii.1976, [2♂, UCDC: UCRCENT00243493‒94]. Idem, but 17.vi.1981, [1♂, UCDC: UCRCENT00243525]. Idem, 19.vii.1976, [1♂, UCDC: UCRCENT00243518]. Idem, but 2.vii.1976, [1♂, UCDC: UCRCENT00243506]. Idem, but 20.ix.1978, [1♂, UCDC: UCRCENT00243534]. Idem, but 20.vi.1981, [1♂, UCDC: UCRCENT00243524]. Idem, but 20.vii.1976, [7♂ 1♀, UCDC: UCRCENT00243476‒79, UCRCENT00243481‒84]. Idem, but 22.vii.1976, [2♂, UCDC: UCRCENT00243474‒75]. Idem, but 26.vii.1976, [1♂, UCDC: UCRCENT00243520]. Idem, but 28.vi.1982, [1♂, UCDC: UCRCENT00243523]. Idem, but 3.viii.1976, R.B. & L.S. Kimsey, [1♂, UCDC: 41

UCRCENT00243505]. Idem, but 31.vii.1977, [1♂, UCDC: UCRCENT00243535]. Idem, but 6.vii.1976, [1♂, UCDC: UCRCENT00243486]. Idem, but 9.vii.1976, [1♂, UCDC: UCRCENT00243492]. Idem, but 13.ix.1978, R.B. Rimsey, [1♂, UCDC: UCRCENT00243533]. Idem, but 3-13.vi.1983, malaise trap, B. Gill, [1♂, CNC: UCRCENT00315886]. Idem, but 1- 8.vi.1994, MT, J. Pickering, [2?#, UCRC: UCRCENT00446423‒24]. Idem, but 9°10'00"N, 79°50'00"W, 23.vii.1977, H.A. Hespenheide, [1♂, UCRC: UCRCENT00305599]. Fort Clayton, 40m, 9°00'19"N, 79°34'50"W, 6.viii.1978, H.J. Harlan, [1♂, UCRC: UCRCENT00306002]. Puma Island, 20-70m, 9°14'10"N, 79°54'12"W, 24.vii.1982, R.B. Kimsey, [2♂, UCDC: UCRCENT00243529‒30]. Tigre Island, 20-60m, 9°13'52"N, 79°54'39"W, 10.vi.1982, R.B. Kimsey, [1♂, UCDC: UCRCENT00243515]. Idem, but 21.vi.1982, [4♂, UCDC: UCRCENT00243465‒66, UCRCENT00243470, UCRCENT00243472]. Idem, but 24.vii.1982, [2♂, UCDC: UCRCENT00243526‒27]. Idem, but 28.vi.1982, [5♂, UCDC: UCRCENT00243496‒97, UCRCENT00243499‒501]. Idem, but 31.v.1982, [1♂, UCDC: UCRCENT00243532]. PERU: Lima: Callanga, 1200m, 12°34'00"S, 76°19'00"W, Staudinger K., [1♂, Unknown: UCRCENT00320348]. SURINAME: Paramaribo, 5°49'24"N, 55°10'04"W, 25.viii.1911 [1♀, EMEC: UCRCENT00236313]. VENEZUELA: Coje des El Baul, 8°57'16"N, 68°18'04"W, 19.v.1967, J&B Bechyne, [1♂, BMNH: UCRCENT00239429]. Idem, but 20.vii.1967, [1♂ 1♀, BMNH: UCRCENT00239422, UCRCENT00239428]. El Pao, 8°02'26"N, 62°18'04"W, 19.v.1967, J&B Bechyne, [1♂, BMNH: UCRCENT00239430]. Aragua: Rancho Grande N. P., 1100m, 10°22'53"N, 67°37'08"W, 18.viii-3.ix.1992, cloud forest, maxinet, L. Masner, [3?#, ROME: UCRCENT00418062, UCRC: UCRCENT00418062, UCRC: UCRCENT00446455]. Lara: 1 km E. Barquisineto, 500m, 10°04'38"N, 69°15'18"W, 27.xii.1985, savannah, R. Jones & P. Kovarik, [6♂ 1♀, CNC: UCRCENT00241002‒08].

K. romandii species complex

Diagnosis. The K. romandii species complex can be defined by a combination of the following characters: frons with striae widely spaced and pattern of striae longitudinal; lower face with transverse striation; frons flat lateral to scrobes cavity; mesoscutum with strongly elevated lateral profile; lateral lobes sparsely hairy, scutellum less produced. Females and males share the same 42

distinguishing characteristics, except that females have a more robust mesosoma, and the facial striae are usually less pronounced. Males with frenal processes usually with roughly sculptured carinae, giving them a serrated appearance.

Included taxa. K. argentina Gemignani, 1933; K. chacoensis Gemignani, 1947; K. cynipsea (Walker, 1862); K. flabellata (Fabricius, 1804); K. floridana (Ashmead, 1885); K. inexagens (Walker, 1862); K. ivorensis Risbec, 1954; Kapala izapa Carmichael, 2006; K. romandii (Guérin-Meneville, 1845); K. terminalis Ashmead, 1892; K. corcovata n. sp.; K. genistriata n. sp.; K. gracilispina n. sp.; K. haplospinosa n. sp.; K. jalisca n. sp.; K. spinaepplanata n. sp.

Kapala argentina Gemignani, 1933 (Fig. 8A‒H)

Kapala argentina Gemignani, 1933: 485‒487. Ilust. Type data: Argentina: Salta. Holotype ♀, [by original designation]. Type depository: MACN; type n° 7.116 (examined). [Holotype relatively complete: glued to card; antennae broken and lost (illustrated in original description); fore legs, including coxae, broken and lost; wings glued together]. Kapala argentina; Gemignani, 1937: 164. Male description. Illust.

Diagnosis. Pattern of striation on the frons with striae near to scrobal depression, curved to the ocellar triangle towards the lower margin of the torulus and longitudinal striae near the eyes (Fig. 8B). Female with clava yellow (Fig. 8C); males with scapes, pedicels and flagellomeres yellow and rami brown (Figs. 8D, E).

Redescription. Female. Body length [3.8 mm, holotype] 3.1–3.8 mm; length of mesosoma excluding the frenal processes [1.7] 1.3–1.7 mm. Head, mesosoma and petiole black; frenal processes with basal half black and apical half black to reddish-brown; legs yellow; coxae and Gt1 reddish-brown; wings hyaline, venation brown. Head. [1.4] 1.4‒1.5x as broad as high, with few sparse setae, more prominent in frons. Scrobal depression with some irregular striae. Frons with wide striae near to scrobal depression 43

curved to the ocellar triangle towards the lower margin of the torulus, longitudinal striae near the eyes (Fig. 8B); lower face with striae curved from the lower margin of the torulus towards the malar space. Eyes separated by [2.1] 2.0‒2.1x their height. Malar space [1.0] 1.0‒1.1x eye height with oblique striae. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/3 segmented. Scape [-] 3.2‒3.8x longer than broad, [-] 0.2‒0.3x head height (Fig. 8C); length of flagellum [-

]1.1‒1.2x head height; Fl2 [-] 2.9‒3.3x as long as apical width and [-] 0.8‒1.3x as long as following flagellomere. Mesosoma. Mesoscutum with dorsal half abruptly curved anteriorly in lateral view, and [1.5] 1.5‒1.7x as high as broad. Midlobe slightly bilobed in frontal view and with transverse striae, which extend to notauli; lateral lobes with longitudinal striae. Axilla with wide longitudinal striae and mesoscutellar disc with striae slightly oblique, converging towards the posterior medial apex (Fig. 6F). Mesoscutellar disc [1.5] 1.3‒1.5x as long as axilla with medial apex raised [1.0] 1.0‒1.4x height of frenal processes in lateral view (Fig. 8C). Axillula with longitudinal striae widely separated from each other. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with faint longitudinal striae; lower mesepimeron weakly reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semierect setae. Metapleuron reticulate. Propodeum flat, broad, and delimited by carina, reticulate. Frenal processes cylindrical, [2.5] 2.3‒2.8x longer than the basal width of mesoscutellum, [1.7] 1.7‒2.1x as long as length of axillae and mesoscutellar disc, slightly arched in lateral view and, in dorsal view, curved with the apex next to each other; frenal processes with well-marked longitudinal striae, apex excavated, with ventral region like a flange-shape more elongated (Fig. 8F). Forewing [-] 2.4‒2.6x longer than broad covered by small bristles, except for the base bare; submarginal vein 0.9‒1.1x as long as marginal vein; stigmal vein 1.8‒2.2x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa semiglobose [1.6] 1.6‒2.0x longer than broad. Metafemur [6.0] 6.0‒6.6x longer than broad, smooth with sparse setae. Metasoma. Petiole slightly flattened dorsally, fine reticulate with weak longitudinal striae,

3.4‒3.7x longer than broad and 1.2‒1.6x as long as metacoxa. Gt1 smooth and tergal scar absent.

Male. Body length 3.5‒3.8 mm; length of mesosoma excluding the frenal processes 1.4‒1.8 mm. Similar to female except for the following: scapes, pedicels and base of flagellomeres yellow, rami dark brown (Fig. 8D); lower face with transversal striae. Eyes separated by 1.9‒2.3x their height. Malar space 0.8‒1.0x eye height. Antennae with 12 segments, scape 2.0‒2.3x longer than broad, flagellum 10 segmented, each flagellomere with a single long rami; length of rami of Fl2 1.3‒1.4x head height and 0.9x as long as following rami. Medial apex of mesoscutellar disc raised 1.0‒1.4x height of frenal processes in lateral view. Frenal processes 2.5‒3.3x longer than the basal width of mesoscutellum, 1.8‒2.3x as long as length of axillae and mesoscutellar disc, with irregular longitudinal striae, giving a serrated appearance (Fig. 8G). Petiole 5.3‒8.3x longer than broad and 2.3‒2.9x as long as metacoxa.

Comments. Gemignani (1937) based on the analysis of a batch of insects coming from R. Saenz Peña (Chaco, Argentina) concluded that they should be males of K. argentina. Therefore, he made a brief description of the males only mentioning that they differed from the female because of the antenna with rami, elongated petiole and small metasoma (characters typical of males of the genus Kapala). Thus, here were inserted new characters for the males, as well registered for the first time K. argentina for Brazil (Fig. 8H), expanding its geographical distribution.

Biology. Unknown.

Distribution. Argentina, Bolivia and Brazil (Fig. 8H).

Holotype. [ARGENTINA]: Província Salta, [Spegazzini, Carlos leg.], [1♀, MACN: type n° 7116].

Additional material examined: ARGENTINA: Buenos Aires: Zelaya, 34°22'10"S, 58°52'11"W, 9.iii.1939, J. Daguerre, [1♀, MACN: UCRCENT00242251]. Chaco: -, 26°37'10"S, 60°57'41"W, [1♀, MACN: UCRCENT00242218]. Colonia Benitez, 50-60m, 7°20'00"S, 58°56'00"W, 10.xii.1948, R. Golbach, [3♂, IMLA: UCRCENT00242077‒79]. Salta: Aguaray, 22°14'22"S, 63°43'59"W, 14.ii.1950, R. Golbach, [1♀, IMLA: UCRCENT00313150]. Idem, but 45

19.ii.1950 [1♀, IMLA: UCRCENT00313151]. Cobos, 24°44'00"S, 65°05'00"W, Fritz, [1♀, AMNH: UCRCENT00172425]. Salta, 1200m, 24°46'59"S, 65°24'44"W, [1♀, MACN: UCRCENT00242255]. Tartagal, 22°31'13"S, 63°47'57"W, A. Martinez, [1♀, AMNH: UCRCENT00238149]. Orán, Abra Grande, 24°18'00"S, 65°57'00"W, 18.iv.-5.v.1969, C. Porter, [11♂ 5♀, MCNZ: UCRCENT00242312‒13, UCRCENT00242315‒17, MCZ: UCRCENT00242319‒53]. Abra Grande, 24°18'00"S, 65°57'00"W, 18-25.x.1968, C. Porter, [2♂, MCZ: UCRCENT00242314, UCRCENT00242318]. Tucuman: Burruyacu, Villa Padre Monti, 1100m, 26°29'00"S, 64°58'00"W, 7.ii.1948, R. Golbach, [1♂, IMLA: UCRCENT00313149]. BOLIVIA: Rio Yacuma, Espiritu, 250m, 14°12'00"S, 66°37'00"W, 23.vii.1950, W. Forster, [1♀, ZSMC: UCRCENT00245259]. BRAZIL: Amazonas: Manaus, INPA I - Bosque da Ciência, 03°05'40.35''S, 59°59'18.02''W, 13.vi.2015, Varredura, D.D.D., [1♀, INPA]. Idem, but 5.v.2015, N.T.B Antunes, [1♀, INPA]. Bahia: Ilhéus, Mata Esperança, 14°46'S, 39°04'W, 17.v.2002, Varredura 14:40 - 14:45, A.M. Penteado-Dias & eq., [1♀, MZUSP: n° 07629]. Espírito Santo: Colatina, 19°32'20''S, 40°37'51''W, xii.1969, E.M. Oliveira, [1♀, MZUSP: n° 07551]. Vitória, EC 63, 20°19'08''S, 40°20'16''W, 03.xii.1996, Armadilha Luminosa, [1F#, INPA]. Idem, but 18.xi.1996, [1♀, INPA]. Mato Grosso: Corumbá - SISBIOTA Pantanal, 19°00'32''S, 57°39''10''W, 1.i.2012, Malaise 02, C. Araújo, [1♂, MZSP]. Idem, but 17.xii.2011, [2♂, MZSP]. Piauí: Piracuruca, P.N. de Sete Cidades, Posto do ICMBio, 04°05'57''S, 41°42'34''W, 15- 30.vi.2013, Malaise, J.A. Rafael, F. Limeira-de-Oliveira, T.T.A. Silva, [1♂, CZMA]. Idem, but 19-28.ii.2013, [1♂, CZMA]. Idem, but, 12-27.iii.2013, [1♂, CZMA]. Idem, but 1-10.iii.2013, Limeira-de-Oliveira, T.T.A. Silva, [1♂, CZMA]. Guaribas, Parque Nacional Serra das Confusões, Andorinha, 09°08'27.8''S, 43°33'42.1''W, 515 m, 1-10.vii.2014, Malaise, J.A. Rafael, F. Limeira-de-Oliveira & G.A. Reis, [2♂, CZMA]. Idem, but, Riacho dos Bois, 09°13'11.9''S, 43°29'26.2''W, 575 m, 20-31.v.2014, [1♂, CZMA]. Rio Grande do Sul: -, 29°32'04"S, 53°23'26"W, Stieglmayr [3♂, NMW: UCRCENT00242563, UCRCENT00242565, UCRCENT00317117].

Kapala chacoensis Gemignani, 1947 (Figs. 9A‒H)

Kapala chacoensis Gemignani, 1947: 1‒2. Type data: Argentina: Chaco, Resistencia. Holotype ♂, [by original designation]. Type depository: MACN; Type n° 38.347 (examined). [Description of male, illustrated]. [Holotype relatively complete: the head and two legs separated from the body but glued in the same cardboard; the last flagellomeres of both antennae are missing].

Diagnosis. Recognized by a combination of the following characters: mesoscutum elevated and slightly projected over the head (Figs. 9A, D); frenal processes soft with fine curviform striae giving to the surface a rough and irregular appearance (Figs. 9F, G). Male frons with wide longitudinal striae, which extend to the supraclypeal area (Fig. 9B); females with supraclypeal area smooth (Fig. 9C). General coloration of the body greenish black and frenal process reddish- brown.

Redescription. Male. Body length [5.0] 3.1‒5.0 mm; length of mesosoma excluding the frenal processes [2.2] 1.4‒2.2mm. Head, mesosoma, coxae and petiole shiny black-green; scapes, pedicels, flagellomeres and legs yellow, except for the base of femora brown; rami of flagellomeres brown; frenal processes reddish-brown; wings hyaline, venation brown; Gt1 reddish-brown. Head. [1.6] 1.4‒1.6x as broad as high, with few sparse setae (Fig. 9B). Scrobal depression with wide transversal striae. Frons wide longitudinal striated, the striae near to scrobal depression extending toward the supraclipeal area; lower face with wide transversal striae (Fig. 9B). Eyes separated by [2.1] 1.9‒2.1x their height. Malar space [1.1] 0.9‒1.1x eye height with transversal striae. Supraclypeal area striated. Clypeus smooth. Antenna with 12 segments; scape [1.2] 1.2‒1.8x longer than broad, [0.1] 0.1‒0.2x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 [1.2] 1.1‒1.2x head height and [1.0] 1.0‒1.05x as long as following rami. 47

Mesosoma. Mesoscutum with dorsal half abruptly curved anteriorly in lateral view and [1.6] 1.4‒1.6x as high as broad, slightly projected over the head (Figs. 9A, D). Midlobe with transverse striae, extending to the lateral lobe and with a soft central depression in dorsal view; lateral lobes with faint longitudinal striae. Axilla and mesoscutellar disc slightly swollen, weakly longitudinal striated and with a slight depression, crossing them longitudinally in the center and becoming more pronounced between the frenal processes (Fig. 9F). Mesoscutellar disc [1.6] 1.5‒1.9x as long as axilla, with medial apex raised [1.0] 0.9‒1.0x height of frenal processes in lateral view. Axillula with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with faint longitudinal striae; lower mesepimeron reticulate; femoral depression weakly crenulate (Fig. 9D). Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate and delimited by carina. Propodeum flat and broad, reticulate. Frenal processes cylindrical and soft, [3.0] 2.6‒3.1x longer than the basal width of mesoscutellum, [1.7] 1.6‒2.0x as long as length of axillae and mesoscutellar disc; frenal processes with fine curviform striae giving to surface a rough and irregular appearance, process slightly curved in dorsal view, and slightly straight in lateral view (Figs. 9A, F), apex excavated (Fig. 9F). Forewing [2.6] 2.6‒2.9x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.1] 1.1‒1.4x longer than marginal vein; stigmal vein [2.0] 2.0‒2.5x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa semiglobose [1.6] 1.5‒2.0x longer than broad. Metafemur [5.6] 5.0‒6.0x longer than broad, with irregular striae and sparse setae. Metasoma. Petiole cylindrical, reticulate and with longitudinal striae, [3.9] 3.1‒4.0x alonger than broad and [1.6] 1.5‒1.6x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 4.1 mm; length of mesosoma excluding the frenal processes 1.9 mm (Fig. 9G). Similar to male except for the following: supraclypeal area and clypeus smooth (Fig. 9C). Antenna with 10 segments (Fig. 9D); scape 3.8x longer than broad, 0.3x head height; flagellum 8 segmented, length of flagellum 1.0x head height; Fl2 1.3x as long as apical width and 1.0x as long as following flagellomere. Petiole 2.8x as longer than broad and 1.1x as long as metacoxa.

Comments. The specimens collected in Maranhão (Brazil) have the axilla and the mesoscutellar disc almost smooth, with only a few weak striae (Fig. 9F), in addition, they are smaller than the specimens from Argentina. In the present study, is reported the first record of K. chacoensis for Brazil and a description of the female.

Biological notes. Unknown.

Distribution. Argentina and Brazil (Fig. 9H).

Holotype. ARGENTINA: Dep.[artamento] Resistência, 10.xii.1935, J.B. Daguerre [coletor], [1♂, n° 38.347, MACN].

Additional material examined: ARGENTINA: Vila E. Lord., 15.xi.-,[1♀, MLPA]. Santa Fé: Vera, 29°27'57"S, 60°12'55"W, x.1950, Duret, [2♂, AMNH: UCRCENT00172426, UCRCENT00237843]. Chaco: Colonia Benítez, 27º20'56''S, 58º58'21''W, 08.ix.2007, J. Torréns, [1♀, MACN; 1♀, UCRC]. BRAZIL: Maranhão: Carolina, P.N. Chapada das Mesas, Cachoeira da Prata, 7°19'S, 47°20'06''W, 25-30.vi.2009, Malaise, A.L. Costa, M.B. Aguiar-Neto, P.A.M. Moraes, [3♂, CZMA].

Kapala cynipsea (Walker, 1862) (Figs 10A‒C)

Thoracantha cynipsea Walker, 1862: 379. Type data: Brazil: Santarém. Syntypes (♀ and ♂, type n° 5.636a and 5.636b, respectively) (examined). Lectotype ♀ (n° 5.636a) present designation. Type depository: BMNH; [type n° 5.636b belongs now to K. cuprea]. [Lectotype relatively complete: right antenna broken after the fourth funicular segment; left antenna broken after the third funicular segment; anterior legs broken and lost, including coxae]. Schizaspidia cynipsea; Walker, 1871: 66. 49

Kapala cynipsea; Ashmead, 1904: 473. Additional citation: De Santis, 1980: 208 (catalog).

Diagnosis. Recognized by a combination of the following characters: mesosoma robust; frenal processes short and stout, 1.1x as long as length of axilla and mesoscutellar disc (Fig. 10A); frons with longitudinal striae curving to the supraclypeal area (Fig. 10B).

Female. Body length 4.2 mm; length of mesosoma excluding the frenal process 2.5 mm. Head, mesosoma, coxae, frenal processes and petiole black; scapes, pedicels and legs yellow; flagellomeres brown; wings hyaline, venation brown; Gt1 black with apex light-brown. Head. 1.4x as broad as high, with few sparse setae more prominent in frons (Fig. 10B). Scrobal depression reticulate with some faint longitudinal striae laterally. Frons with wide longitudinal striae curving to the supraclypeal area; lower face with some faint transversal striae (Fig. 10B). Eyes separated by 2.1x their height. Malar space 0.8x eye height. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2 segmented. Scape 3.0x longer than broad, 0.3x head height, surface smooth with sparse setae; Fl2 1.4x as long as apical width and 1.0x as long as following flagellomere. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.4x as high as broad. Midlobe with transverse striae and dorsally rounded in frontal view; lateral lobes with fine longitudinal striae. Axillar and mesoscutellar disc longitudinal striated. Mesoscutellar disc 1.1x as long as axilla with medial apex raised 1.2x height of frenal processes in lateral view. Axillula with longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with some longitudinal striae; lower mesepimeron fine reticulate; femoral depression smooth. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, smooth, broad and delimited by carina. Frenal processes cylindrical, short and stout, 1.5x longer than the basal width of mesoscutellum, 1.1x as long as length of axillae and mesoscutellar disc; slightly arched in lateral view and slightly convex medially in dorsal view; frenal processes with longitudinal striae well marked and apex excavated (Fig. 10A). Forewing 2.5x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.2x longer than marginal vein; stigmal vein 50

1.6x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa semiglobose 2.8x longer than broad, with a row of setae semi-erect medially. Metafemur 4.6x longer than broad, smooth with sparse setae. Metasoma. Petiole slightly flattened dorsally, 1.4x longer than broad and 1.0x as long as metacoxa. Gt1 smooth and bright with tergal scar absent.

Male. Unknown.

Biological notes. Unknown.

Distribution. Brazil (Fig. 10C). Lectotype. [BRAZIL]: [Pará]: Santarém [1♀, UCRC_ENT 310055, BMNH: type n° 5.636a].

Additional material examined: BRAZIL: Santa Catarina: Nova Teutonia, 27°00'00"S, 52°23'00"W, 26 Jan 1939, Fritz Plaumann, [1♀, BMNH: UCRCENT00239460]. Amazonas: Macura, 200m, 7°29'00"S, 72°28'00"W, Apr, [1♀, CMNH: UCRCENT00240753]. Pará: Santarém, 30m, 2°26'22"S, 54°41'55"W, [2♀, CMNH: UCRCENT00240749, UCRCENT00240751]. Idem, but vii.1919, S.M. Klages, [1♀, CMNH: UCRCENT00240750].

Kapala flabellata (Fabricius, 1804) (Fig. 11A‒G)

Eucharis flabellatus Fabricius, 1804: 158. Type data: “America meridional Dom. Smidt. Mus. Dom. Lund.” [Original description]. Syntypes ♂, labelled “E. flabellate ex Am. mer. Schmidt”. Lectotype designated by Heraty, 2002: 171 (examined). Type depository: ZMUC. [Description of the male. Lectotype in good conditions: individual pinned between axilla and head]. Kapala furcata; Kirby, 1886: 32. Kapala flabellata; Heraty, 2002: 171 [new combination]. 51

Thoracantha atrata Walker, 1862: 383. Type data: Brazil [?? no label on specimen]. Holotype ♂, by original designation. Type depository: BMNH; Type n° 5.635 (examined). [Description of the male. Holotype relatively complete: palpi labial broken and lost; left side of the mesopleura broken; right anterior leg broken after femur; left anterior leg broken and lost]. New syn. Kapala atrata; Ashmead, 1904: 473. [taxonomic position]. Thoracantha surgens Walker, 1862: 384. Type data: Brazil: Santarém, Villa Nova. Holotype ♂, by original designation. Type depository: BMNH; Type n° 5.632 (image examined). Description of the male. Holotype in good conditions: complete. New syn.

Diagnosis. Recognized by a combination of the following characters: frenal processes long and thin with apex emarginated and exceeding the metasoma (Figs. 11A, E, F); medial apex of mesoscutellum with carina and raised 1.0‒1.5x height of frenal processes in lateral view (Fig. 11E). Male with antennae reddish-brown to deep-black.

Redescription. Male. Body length 2.6‒5.1 mm; length of mesosoma excluding the frenal processes 1.2‒2.2 mm. Head, mesosoma, petiole and frenal processes black; scapes, pedicels and flagellomeres and rami brown; legs yellow; coxae and Gt1 reddish brown; wings hyaline, venation brown. Head. 1.5‒1.6x as broad as high, with few sparse setae (Fig. 11B). Scrobal depression punctate and striated. Frons with striae near to scrobal depression curved to the ocellar triangle towards the lower margin of the torulus, longitudinal striae near the eyes; lower face with transversal striae. Eyes separated by 1.8‒2.1x their height. Malar space 0.8‒1.0x eye height with oblique striae. Supraclypeal area with upper region punctate, the rest of supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/3 or 3/2 segmented. Antenna with 12 segments; scape 2.0‒2.7x longer than broad, 0.2‒0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 1.1‒1.4x head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.7‒2.0x as high as broad. Midlobe with wide transversal striae and dorsally slightly bilobed in 52

frontal view; lateral lobes with fine longitudinal striae. Axilla with wide longitudinal striae and mesoscutellar disc with striae slightly oblique converging towards the posterior medial apex. Mesoscutellar disc 1.1‒1.5x as long as axilla and medial apex with carina and raised 1.0‒1.5x height of frenal processes in lateral view (Fig. 11E). Axillula with longitudinal striae widely separated from each other. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with wide longitudinal striae; lower mesepimeron weakly reticulate; femoral depression weakly crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad, reticulate and delimited by carina. Frenal processes cylindrical and long with apex exceeding metasoma (Fig. 11E), 3.0‒4.3x longer than the basal width of mesoscutellum, 1.8‒2.9x as long as length of axillae and mesoscutellar disc, straight in lateral view and, in dorsal view, curved with the apex near to each other; frenal processes with well-marked irregular longitudinal striae, giving a serrated appearance; apex excavated. Forewing 2.4‒2.7x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.05‒1.3x longer than marginal vein; stigmal vein 1.5‒2.6x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa semiglobose 1.6‒2.0x longer than broad. Metafemur 5.2‒6.7x longer than broad, with faint striae and sparse setae. Metasoma. Petiole cylindrical, reticulate, with two longitudinal carinae, 6.0‒9.6x longer than broad and 2.4‒3.2x as long as metacoxa. Gt1 smooth and tergal scar present.

Female. Body length 2.9‒5.3mm; length of mesosoma excluding the frenal processes 1.3‒2.5 mm. Similar to male except for the following: Frons longitudinal striated to slightly oblique (Fig. 11C). Eyes separated by 2.0‒2.2x their height. Scape and pedicel yellow; scape

3.0‒4.5x longer than broad; length of flagellum 0.9‒1.1x head height; Fl2 1.7‒3.0 as long as apical width and 1.0‒1.5x as long as the following flagellomere (Figs. 11C, D). Mesoscutellar disc with medial apex raised 0.7‒1.3 height of frenal processes in lateral view (Fig. 11D). Frenal processes 1.7–2.6x as long as length of axilla and mesoscutellar disc, with longitudinal striae (Fig. 11F). Metacoxa 1.8‒2.2x longer than broad. Metafemur 5.5‒7.6x longer broad. Petiole short, 3.0‒4.6x longer than broad and 1.3‒1.7x as long as metacoxa. Tergal scar present or absent. 53

Comments. The population of K. flabellata from Costa Rica and Guadeloupe have more robust bodies and deep black color, as well as, scape and antenna entirely black. Populations from Bahia and Espírito Santo (Brazil) are smaller in size, with antenna, coxae and frenal processes reddish- brown. In the present study we performed the synonimy of K. flabellata with K. atrata (= Thoracantha atrata) and K. surgens (= Thoracantha surgens).

Biological notes. Unknown.

Distribution. Brazil, Costa Rica, French Guiana, Guadeloupe, Panamá, Peru, Trinidad, Tobago and Venezuela (Fig. 11G).

Lectotype. “America meridional Dom. Smidt. Mus. Dom. Lund.” [ZMUC 00241188, image analyzed].

Additional material examined: BRAZIL: [?? no label on specimen], [Type of Thoracantha atrata, BMNH type n° 5.635, BMNH: UCRC_ENT: 310054]. Santarém, Villa Nova, [Type of Thoracantha surgens, BMNH type n° 5.632, BMNH: UCRC_ENT 310051]. Amazonas: AM 010, Km 26, Reserva Ducke, 3°0'13.68"S, 59°55'7.37"W, 2.iv.1978, Jorge Arias, [1♂ 1♀, INPA]. Idem, but 7.ii.1978, [1♂, INPA]. Idem, but 20.ix.1978, [1♀, INPA]. Manaus, ZF 03 Km 23 - Fazenda Esteio 1208, 02°22'34''S, 59°52'39''W, 14.viii.1984, Arm. Malaise, [1♂, INPA]. Manaus, Reserva Ducke, Platô leste/oeste - dossel, 3°0'13.68"S, 59°55'7.37"W, 14.ii-06.iii.2007, Armadilha suspensa, G. Freitas & M. Feitosa, [1♂, INPA]. Manaus, Reserva Ducke, 26 Km NE Manaus, 3°0'13.68"S, 59°55'7.37"W, 16.v.1978, J. Arias & N. Penny, [1♂, INPA]. Idem, but 23.v.1978, [1♂, INPA]. Idem, but 27.ix.1978, [1♀, INPA]. Manaus, Reserva Ducke - Igapó Tinga, 3°0'13.68"S, 59°55'7.37"W, 03.iii.2003, Armadilha Malaise, J.M.F. Ribeiro, [1♀, INPA]. Marãa - Rio Japurá, Maguari, 01°51'21S, 65°34'51''W, 27-31. X.1988, Armadilha Shanon, J. Dias, [1♀, MPEG]. Presidente Figueiredo, AM 240, Km 24, 02°00'46''S, 59°49'43''W, Armadilha Malaise, Henriques, A., Xavier-Filho, F., Carmo, D., [1♂, INPA]. Bahia: Camacã, Serra Bonita, 15°25'13.74"S, 39°29'46.58"W, 03.ii.2009, Armadilha Luminosa, F.R. Fernandes, [1♂, MZUSP]. Igrapiúna - RESEC Michelin, Seringueira, Guibó Mata, 100 m, 13°50'S, 39°15'W, 4-5.ii.2010, 54

Silva, M.O., Prates, P. Silva L.J., Rosa, J.F., Queiroz, M.V., Santos, F., Ramalho, M., Coelho, J.F., [1♀, UEFS]. Porto Seguro - PARNA Monte Pascoal, 16°54'14.72"S, 39°13'2.78"W, 12.viii- 02.ix.2013, Armadilha Malaise, [2♂, UFES, amostra 1986 and amostra 1980]. Ceará: Guaramiranga, 4°15'37.50''S, 38°56'4.17''W, 17-19.vi.2015, Malaise, David Nogueira, [1♀, INPA]. Espírito Santo: Atílio Vivácqua - Fazenda Oriente, área 2, 20°54'50''S, 41°11'52''W, 20- 22.ii.2003, Armadilha Malaise, [1♂, UFES, amostra 1047]. Cariacica - R.B. Duas Bocas, 20°16'23.42"S, 40°28'40.19"W, 06.ii.1997, varredura, [2♂, UFES, amostra 244]. Idem, but 10.xii.1996, [2♂, UFES, amostra 328]. Guarapari, Pq. Est Paulo Cesar Vinha, 20°35'07"S, 40°24'37"W, 18.xi.2006, Varredura, M.T. Tavares et al., [1♂, UFES: UFES00095386]. Idem, but 08-15.v.2006, Möericke, [2♂, UFES, amostra 166]. Sooretama - Reserva Biológica de Sooretama, 19°0'32.79"S, 40°7'2.14"W, 5-7.vi.2011, Möericke, [1♂, UFES, amostra 1880]. Idem, but 30.x.-06.xi.2002, Malaise, [1♀, UFES, amostra 1339]. Maranhão: Carolina, PARNA Chapada das Mesas, Riacho Cancela, 225 m, 07°06'44.2''S, 47°17'56.8''W, 1-10.viii.2013, Armadilha Malaise, J.A. Rafael, F. Limeira-de-Oliveira & T.T.A. Silva, [1♂, CZMA]. Idem, but, 11-14.vi.2013, Varredura, J.A. Rafael, F. Limeira-de-Oliveira, A.A. Santos, [1♀, CZMA]. Idem, but Riacho Sucuruiu, 240 m, 07°07'05.6''S, 47°18'31.6''W, 11-14.v.2013, Armadilha Malaise, J.A. Rafael, F. Limeira-de-Oliveira, T. Câmara & A.A. Santos, [1♂, CZMA]. São Pedro da Água Branca, Fazenda Esplana, 04°59'05''S, 48°08'03''W, 7.xii.2001, Malaise, J.A. Rafael & J. Vidal, [1♂, MZSP n°51493, MZUSP]. Minas Gerais: Cássia, Região do Lajeado, Sítio Genoveva, 20°34'58''S, 46°55'19''W, 30.xii.2015-12.i.2016, Armadilha Malaise, Pádua D.G. & Pádua, A.G., [1♂ 1♀, INPA]. Pará: Belém, Tenoné - Parque Crocodilo Zoo, 01°27'21''S, 48°30'14''W, 04.viii.1999, J.M.F. Ribeiro, [1♂, MPEG]. Idem, but Utinga, 25.iv.1961, Bechyné, [2♂, MPEG]. Primavera, 00°56'34''S, 47°06'57''W, 14.ii.1987, Marcio Zanufe, [1♂, MPEG]. Rio Nhamundá, 25 m, 01°35'11''S, 57°37'32''W, 17-20.v.2008, Armadilha Malaise, J.A. Rafael e equipe, [1M#, INPA]. Pernambuco: Recife, 8°03'14"S, 34°52'52"W, -.viii.1976, F. D. Benton, [1♂, CNC: UCRCENT00172314]. Idem, but Mata do Bananal, 26.i.2015, Grossi P.C., [1♂, INPA]. Idem, but Parque dos Dois Irmãos, 08°03'14''S, 34°52'52''W, 17-20.vii.2002, Armadilha Malaise Bosque pt. 02, S.T.P.A. Amarante & eq., [2♂, MZSP n°07654 and MZSP n°07655, MZUSP]. Idem, but 20-23.vii.2002, Malaise trilha, [1♀, MZSP n°07667, MZUSP]. Rondonia: Vilhena, 14°46'55''S, 60°22'18''W, 26.iv.2006, J.A. Rafael & F.F. Xavier-Filho, [3♂, INPA]. Roraima: 55

Amajari, Serra do Tepequem, 03°39'07''N, 61°22'15''W, 1-15.iv.2016, Armadilha Malaise, Rafael et al., [1♀, INPA]. Caracarai - Parque Nacional de Viruá, 1°29'23.3''N, 61°00'08.7''W, 19.iv.2015, Armadilha Malaise, J.A. Rafael, R.A. Heleodoro, D.M. Mendes, D. Marques & C. Maldaner, [4♂, INPA]. São Paulo: Iguape, 24°42'29"S, 47°33'19"W, 1991, Wettstein, [1♀, NMW: UCRCENT00317113]. Iguape, Est. Ecol. Juréia-Itatins, 24°31'17.8''S, 47°12'1.6''W, 19.vi.2010, Malaise ponto 5, N.W. Perioto & eq., [1♂, LRRP]. Ribeirão Grande, Parque Estadual Intervales, 24°16'23.6''S, 48°25'21.8''W, 22.i.2010, Malaise ponto 2, N.W. Perioto & eq., [1♀, LRRP]. São Luis do Paraitinga, P.E.S.M. - Nucl. Sta. Virginea, 23°20'0.8''S, 44°49'57.2''W, 20.iv.2011, Malaise ponto 6 teste, N.W. Perioto & eq., [1♂, LRRP]. Teodoro Sampaio, Parq. Estação Morro do Diabo, 22°36'15.8''S, 52°18'02.5''W, 15.vii.2010, Malaise ponto 1, N.W. Perioto & eq., [1♀ 1♂, LRRP]. Idem, but 17.v.2010, [1♀, LRRP]. Idem, but 18.ii.2010, [1♂, LRRP]. Ubatuba, P.E.S.M. - Nucl. Picinguaba, 23°20'0.8''S, 44°49'57''W, 18.i.2010, Malaise ponto 5, N.W. Perioto & eq., [2♂, LRRP]. Idem, but 200 m, 23°19'08.4''S, 44°49'04.8''W, 18- 21.i.2016, E.F. Santos & C.P. Scott-Santos, [1♀, MZSP n°07781, MZUSP]. COSTA RICA: Alajuela Prov.: Est. San Ramon Oeste, 3-19.iv.1994, C. Cano, [1♂, INBIO: INBIOCRI01769292]. Finca La Selva NE Dos Rios, 400m, 10°26'15"N, 84°00'01"W, 27.iii.1988, Hanson [1♂, UCRC: UCRCENT00172497]. La Fortuna, Sector Catarata, 500m, vi- vii.1998, Malaise, G. Carballo, [2♂#, INBIO: INBIOCRI03018281, INBIOCRI03018389]. P.N. Volcan Tenorio, Sector El Pilon, Send. El Mirador, 700-800mm, 22.vi.2003, J. Azofeifa, Light Trap, [1♂, INBIO: INBIOCRI03732257]. Sector Cerro Chato, 1100m, 2.ix-10.x.1997, G. Carballo, Malaise, [1♂, INBIO: INBIOCRI02574408]. Guanacaste Prov.: Cerro El Hacha, 12 km SE La Cruz, 300m, -.v.1988, M. Espinoza, [1♂, INBIO: INBIOCRI00644961]. Idem, but, NW Volcan Oros, 300m, 10°59'57"N, 85°32'59"W, 1988, [1♂, UCRC: UCRCENT00172505]. Est. Patilla, 9 km S Sta Cecilia, 700m, 10°59'28"N, 85°25'39"W, -.iv.1991, C. Moraga, [2♂, INBIO: INBIOCRI00687085, INBIOCRI00687127]. Idem, but -.ii.1995, [1♂, INBIO: INBIOCRI02134401]. Idem, but -.ii.1993, [1♂, INBIO: INBIOCRI01806134]. Idem, but - .ix.1989, C. Moraga, P. Riox, [1♂, INBIO: INBIOCRI01089256]. Idem, but 21.iii.-21.vi.1989, GNP Biod. Sur., [1♂, INBIO: INBIOCRI00089700]. Idem, but -.ix.1988, I. Gauld, [3♂, UCRC: UCRCENT00172499, UCRCENT00172500, UCRCENT00235915]. Idem, but 10°59'22"N, 85°25'33"W, -.vi. 1988, P. Hanson, [1♂, UCRC: UCRCENT00305913]. Idem, but -.x.1988, [1♂, 56

UCRC: UCRCENT00172496]. Idem, but -.ix.1988, [1♂, UCRC: UCRCENT00172501]. Idem, but -.iv.1988, [2♂, UCRC: UCRCENT00172502‒03]. Idem, but P.N. Guanacaste, 22.viii.1993, C. Moraga [1♂, INBIO: INBIOCRI01638544]. Idem, but 3-18.x.1991, [1♂, INBIO: INBIOCRI00548741]. Idem, but 31.iii.-15.iv.1992, [1♂, INBIO: INBIOCRI00416444]. Idem, but -.iii.1991, [2♂, INBIO: INBIOCRI00701431, INB0IOCRI00701433]. Idem, but -.i.1990, P. Rios, [1♂, INBIO: INBIOCRI00213978]. Idem, but -.viii.1991, [1♂, INBIO: INBIOCRI03032932]. Idem, but .ii-iii.1993, [1♂, INBIO: INBIOCRI01904524]. Idem, but - .ii.1995, [1♂, INBIO: INBIOCRI02135067]. Idem, but -.ix.1989, [1♂, INBIO: INBIOCRI00035575]. Idem, but 2-19.iii.1992, [3♂, INBIO: INBIOCRI00420475, INBIOCRI00420563, INBIOCRI00420774]. Idem, but 21.iii.-7.iv.1993, [1♂, INBIO: INBIOCRI01386416]. Idem, but 3-18.x.1991, [3♂, INBIO: INBIOCRI00340398, INBIOCRI00340409, INBIOCRI00340411]. Idem, but -.iv.1994, [1♂, INBIO: INBIOCRI01793719]. Idem, but -.vii.1994, [2♂, INBIO: INBIOCRI02049020, INBIOCRI02049046]. Idem, but -.iii.1995, [2♂, INBIO: INBIOCRI0224202‒21]. Idem, but - .v.1991, [1♂, INBIO: INBIOCRI00651151]. Idem, but -.iii.1995, [2♂, INBIO: INBIOCRI02252332‒33]. Idem, but -.v.1990, II curso Parataxon. [1♂, INBIO: INBIOCRI00261545]. Idem, but -.v.1992, F. Araya, [1♂, INBIO: INBIOCRI00805140]. Finca Loaiciga, 6km S Sta. Cecilia, P.N. Guanacaste, 500m, 23.ix-14.x.1992, P. Rios, [1♂, INBIO: INBIOCRI00811547]. Sta. Rosa NP Hacienda 1-0, 10°53'33"N, 85°45'59"W, xii.1986-i.1987, [1♂, UCRC: UCRCENT00172504]. Idem, but 200m, -.i.1987, I. Gauld, [1♂, UCRC: UCRCENT00305914]. Heredia: La Selva Biol. Sta. 3km S Pto. Viejo, 10°26'00"N, 84°01'00"W, 20.iv.1989, H.A. Hespenheide, [1♀, UCRC: UCRCENT00279856]. Limón: 16 km W. Guapiles, 400m, 10°12'45"N, 83°56'06"W, -.xii.1989, Paul Hanson, [1♀, UCRC: UCRCENT00305912]. Idem, but, -.ii.1989, [1♀, UCRC: UCRCENT00172498]. 7 km SW Bribri, 50m, 9°35'28"N, 82°52'54"W, -.ix.1989, Paul Hanson, [1♂, UCRC: UCRCENT00172506]. Puntarenas Prov.: Cerro Mueller, 744m, 30.xi.1995, L. Ângulo, [1♂, INBIO: INBIOCRI02484924]. Golfito, P.N. Corcovado, Estacion Agujas, Cerro Rincon, 745m, 20.vi.2000, J. Azofeifa, Manual, [1♂, INBIO: INBIOCRI03133318]. P.N. Corcovado, Cerro Rincon, La Tigrilla, 600m, 24.vi-31.vii.2002, J. Azofeifa Zuniga, MT, [2♂, INBIO: INBIOCRI03983117‒18]. Idem, but Estac. Agujas, 745m, 24.vi.2002, Esteban Leon, #69885, [1♂, INBIO: INBIOCRI03495180]. Idem, but 17.vii.2001, J. 57

Azofeifa, Manual, [1♂, INBIO: INBIOCRI03329238]. San Jose: Ciudad Colón, 800m, 9°54'33"N, 84°14'31"W, -.iv-.v.1990, Luis Fournier, [1♂, UCRC: UCRCENT00172636]. Idem, but .xii.1989-.i.1990, [1♂, UCRC: UCRCENT00172638]. Idem, but .ii.1990, [1♂ 1♀, UCRC: UCRCENT00172635, UCRCENT00172637]. Idem, but .iii.-.iv.1990, [3♂, UCRC: UCRCENT00172495, UCRCENT00172639‒40]. FRENCH GUIANA: 21 Km SE Roura on Kaw Rd., 04°33'57.0''N, 52°12'43.3''W, 6-7.ii.2010, MV Ligth, J.E. Eger, [2♂ 2♀, FSCA]. Idem, but 16.ii.2010, [1♂, FSCA]. Idem, but 21.iv.2007, [1♀, FSCA]. Amazone Nature Lodge, 30 Km SE Roura on Kaw Rd., 04°33'57.0''N, 52°12'43.3''W, 10-18.iv.2007, D.G. Hall & J.E. Eger, [1♂ 1♀, FSCA]. GUADELOUPE: Rte. Forestiere de grosse montagne, 16°16'59"N, 61°44'59"W, 20.iv.1979, J. J. Menier, light trap, [1♀, MNHN: UCRCENT00172274]. PANAMA: Coiba: Estacion Biol., 7°28'06"N, 81°45'24"W, 19.i.1994, Malaise, J.L. Nieves, [1♂, MNCN: UCRCENT00301016]. Idem, but 21-23.i.1994, [1♂, MNCN: UCRCENT00301015]. Idem, but 27-29.i.1994, [1♂, MNCN: UCRCENT00301012]. PERU: Huannes, Tingo Maria, 9°17'43"S, 75°59'51"W, 29.i.1984, L. Huggert, [1♂, MZLU: UCRCENT00172399]. Madre de Dios, Puerto Maldonado, 12°36'12"S, 69°11'30"W, 1.i.1984, L. Huggert, [1♀, MZLU: UCRCENT00172406]. Idem, but 3.i.1984, [2♀, MZLU: UCRCENT00172400 - 01]. Idem, but 6-11.i.1984, [1♂, MZLU: UCRCENT00172396]. SURINAME: Saramacca, 6 m, 05°81'77.5''N, 55°59'0.64''W, 15- 25.ix.2005, Malaise, G.J. Steck, [1♂, FSCA]. TRINIDAD: St. Augustine, Pax Guest House, 3- 10.xi.2000, blackligth, R.E. Woodruff, [1♀, FSCA]. TOBAGO: 1 1/8mi ESE Adelphi, 150m, 11°12'12"N, 60°42'24"W, 15.ii.1977, secondary scrub along pasture edge, sweep, P. Feinsinger, [1♂, FSCA: UCRCENT00241438]. VENEZUELA: Merida: Tabay Mucuy, Send. Lag. Suero, 1900m, 18.vi-2.viii.1989, S. & J. Peck, [1♂, CNC: UCRCENT00426061].

Kapala floridana (Ashmead, 1885) (Fig. 12A‒H)

Thoracantha floridana Ashmead, 1885: 95‒96. Type data: USA: east Florida. Holotype ♂, by monotypy. Type depository: USNM; type n° 2827 (examined). [Description of male]. [Holotype relative complete: anterior legs broken and lost; coxa medially not visible 58

(covered by a brown resin, glue); left middle leg broken, but it is glued in the triangle; left wing apex missing]. Kapala floridana; Ashmead, 1892: 357. Additional citations: Wheeler, 1907: 17, plate IV, figs. 56 and 57 [biology and illustration]; Burks, 1979: 877 (catalog); Heraty, 1985: 86- 87 [subsequent description of both sexes, illustrated].

Diagnosis. Recognized by a combination of the following characters: mesoscutellar disc not elevated above base of frenal processes (Figs. 12A, C); frons with longitudinal striae, some of striae are bifurcate giving a rugose appearance (Fig. 12B); femurs brown with distal apex yellow (Figs. 12A, E). Females with metasoma reddish orange (Fig. 12E).

Redescription. Male. Body length [2.9] 2.5‒3.3 mm; length of mesosoma excluding the frenal processes [1.7] 1.1‒1.7 mm. Head, mesosoma, coxae and petiole black; frenal processes black with apex reddish-brown; scapes dark-brown, pedicels, flagellum and rami brown; femurs brown with distal apex yellow, tibia and tarsi yellow; wings hyaline to infuscate, venation brown;

Gt1 dark-brown with apex yellow. Head. [1.5] 1.4‒1.5x as broad as high, with few sparse setae. Scrobal depression rugose. Frons with wide longitudinal striae, some of striae are bifurcate giving a rugose appearance (anastomosed) (Fig. 12B); lower face transversally striated. Eyes separated by [2.3] 2.1‒2.3x their height. Malar space [1.0] 0.9‒1.0x eye height. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape [3.1] 3.0‒3.5x longer than broad, [0.2] 0.2x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 [1.0] 1.0‒1.1 head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.5‒1.6x as high as broad. Midlobe with transversal and longitudinal striae, in frontal view and dorsally rounded in frontal view; lateral lobes with longitudinal striae. Axilla longitudinaly striated with a longitudinal groove, extending to apex of mesoscutellum (Fig. 12F). Mesoscutellar disc slightly swollen, in lateral view, with longitudinal and irregular striae, [1.6] 1.1‒1.7x as long as axilla and not elevated above base of frenal processes (Fig. 12C). Axillula with irregular striae giving a rugose appearance. Prepectus triangular with elongated posterior region and rounded 59

apex. Mesepisternum and upper mesepimeron with longitudinal striae; lower mesepimeron reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and rugose, its upper region with longitudinal carina. Frenal processes cylindrical, [2.0] 2.0‒2.6x longer than the basal width of mesoscutellum, [1.4] 1.4‒1.8x as long as length of axilla and mesoscutellar disc, straight in lateral view and, in dorsal view, parallel; frenal processes with longitudinal irregular striae, giving a serrated appearance; apex of frenal processes excavated, with ventral region like a flange-shape more elongated (Fig. 12F). Forewing [2.1] 2.1‒2.5x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.2] 1.2‒1.4x longer than marginal vein; stigmal vein rectangular [2.0] 1.2‒2.0x longer than broad; postmarginal vein indistinct. Hindwing fringed. Metacoxa semiglobose [1.5] 1.5‒1.9x longer than broad. Metafemur [6.0] 5.7‒6.5x longer than, with faint striae and sparse setae. Metasoma. Petiole cylindrical and rugose with longitudinal striae dorsally, [5.0] 5.0‒7.2x longer than broad and [2.3] 2.0‒2.4x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 2.6‒3.1 mm; length of mesosoma excluding the frenal processes 1.2‒1.0 mm (Fig. 12G). Similar to male except for the following: scapes and pedicels yellow to brown; flagellum brown; Gt1 reddish-orange (Fig. 12E). Antenna with 11 segments; scape 4.2‒4.7x longer than broad and 0.3‒0.4 head height; flagellum 8 segmented, cylindrical and slightly serrated (Fig. 12D), length of flagellum 1.1‒1.2x head height; Fl2 2.2‒2.7x as long as apical width and 1.1‒1.4x as long as following flagellomere. Petiole 3.2‒3.6x longer than broad and 1.1‒1.3x as long as metacoxa.

Biological notes. The specimen type was taken while feeding (= ovipositing?) on flowers of Ilex glabra (L.) Gray (Aquifoliaceae). Also, this species is associate with Pogonomyrmex badius Latreille (Wheeler, 1907). This association is based only on the opinion of Ashmead and is probably incorrect. Other specimens also were collected on Hypericum cistifolium Lam., Hypericum reductum Lam., Chamaecrista fasciculata (Michx.) Greene (= Cassia fasciculata Michx.); on flowers of Pterocaulon pycnostachyum (Michx.) Elliott; flower buds of Eupatorium 60

perfoliatum L., Eupatorium pilosum Walter, Eupatorium mohrri Greene (= Eupatorium recurvans) and Sericocarpus tortifolius (Michx.) Ness (= Aster bifoliatus (Walter) H.E. Ahles).

Distribution. United States of America (Florida) (Fig. 12H).

Holotype. [United States of America]: east Florida, [1♂, USNMENT 00802962, USNM type n° 2827].

Additional material examined: USA: Florida: 5 miles E. of Lake Butler T5S R20E section 35, 30°02'01"N, 82°16'27"W, 10.vii.1986, Lloyd R. Davis. Jr., [1♂, UCRC: UCRCENT00300867]. Idem, but 2.x.1986, [1♂, UCRC: UCRCENT00301022]. Idem, but 26.vi.1986, [4♂ 3♀, FSCA: UCRCENT00241437, UCRC: UCRCENT00300803‒08]. Idem, but 27.vi.1986, [4♂ 2?, UCRC: UCRCENT00172645, UCRCENT00300796‒97, UCRCENT00300800‒01, UCRCENT00301027]. Idem, but 31.vii.1986, [12M#, UCRC: UCRCENT00282952‒61, UCRCENT00300795, UCRCENT00301218]. Idem, but 6.vii.1986, [1♀, UCRC: UCRCENT00172481]. Idem, but 7.viii.1986, [3♀, UCRC: UCRCENT00300862‒64]. Idem, but collected on flowers of Pterocaulon pycnostachyum, [1♀, UCRC: UCRCENT00300868]. Idem, but 2.x.1986, ex: flower buds of Eupatorium perfoliatum, [4♀, UCRC: UCRCENT00301023‒26]. Idem, but, 27.vi.1986, ex: flower buds of Eupatorium pilosum, [3♀, UCRC: UCRCENT00300798‒99, UCRCENT00300802]. Idem, but 30.vii.1986, ex: flower buds of Eupatorium pilosum, [42♀, UCRC: UCRCENT00282967‒71, UCRCENT00301018‒21, UCRCENT00301219‒51]. Idem, but 30.vii.1986, ex: flower buds of Eupatorium recurvans, [3♀, UCRC: UCRCENT00282964‒66]. Idem, but 30.vii.1986, Hypericum cistifolium, [1♀, UCRC: UCRCENT00282973]. Idem, but 6.viii.1986, ex: flower buds of Eupatorium pilosum, [39♀, UCRC: UCRCENT00282963, UCRCENT00300824‒33, UCRCENT00300835‒61, UCRCENT00301216]. Idem, but 6.viii.1986, ex: flower buds of Eupatorium recurvans, [15♀, UCRC: UCRCENT00300810‒23, UCRCENT00301217]. Idem, but 9.vii.1986, ex: flower buds of Eupatorium recurvans, [2♀, UCRC: UCRCENT00282972, UCRCENT00301017]. Idem, but 9vii.1986, ex: flower buds of Eupatorium pilosum, [12♀ 1?, UCRC: UCRCENT00172646, UCRCENT00300784‒94, UCRCENT00300809]. 8-10mi E Naples S. of Corkscrew Sw., 2m, 61

26°08'28"N, 81°38'36"W, 13.xii.1985, roadside weeds, G. Otis, [1♂, UCRC: UCRCENT00172487]. Idem, but, J. Heraty, [1♀, UCRC: UCRCENT00172483]. 9 mi SSW Ocala (KCE), 20-30m, 29°03'55"N, 82°11'36"W, 13.x.-5.xi.1975, Turkey oak, Mal. Trap, J. Wiley, [20♂ 9♀, ABS: UCRCENT00305294, FSCA: UCRCENT00306374, UCRC: UCRCENT00300875‒79, CRCENT00300881‒95, UCRCENT00300923‒27, UCRCENT00301028‒31]. Idem, but 4-10.ix.1975, Mal. Trap, J. Wiley, [20♂, UCRC: UCRCENT00300979‒92, UCRCENT00301004‒07, UCRCENT00172482, UCRCENT00172488]. Idem, but [2♂ 3♀, FSCA: UCRCENT00241400‒02, UCRCENT00241426, UCRCENT00306375]. Idem, but 19.ix.-2.x.1975, [2♂, UCRC: UCRCENT00302312‒13]. Alachua Co., 20-50m, 29°14'02"N, 82°21'01"W, 11.xi.1956, weeds, sweep, R. A. Morse, [1♂, FSCA: UCRCENT00241399]. Archbold Bio. Research Station, 27°10'54"N, 81°21'06"W, 11.x.1986, collected on buds of Aster bifoliatus (Walt.) Ahles “white- topped aster” flowers, malaise trap, Lloyd R. Davis. Jr., [1♀, UCRC: UCRCENT00301030]. Idem, but 11-17.viii.1987, D.B. Wahl, [1♂ 1♀, UCRC: UCRCENT00301035‒36]. Idem, but 18.ix-5.x.1988, malaise trap, D.B. Wahl, [1♀, UCRC: UCRCENT00300871]. Idem, but 27.x- 30.xi.1988, malaise trap, D.B. Wahl, [1♀, Heraty: UCRCENT00235923]. Idem, but 28.v.2005, from Hypericum reductum, road 224, swept, M. Deyrup, [1♂, ABS: UCRCENT00305300]. Idem, but 30.vi-27.vii.1987, D.B. Wahl, [2♀, UCRC: UCRCENT00301033‒34]. Idem, but 31.viii.2001, from seasonal pond, north bayhead E. of SR. 8, sweep, M. & L. Deyrup, [2♂ 2♀, ABS: UCRCENT00305290, UCRCENT00305308, UCRCENT00305285, UCRCENT00305288]. Charlotte Co., 5mi S. of Punta Gorda, 26°50'45"N, 82°01'59"W, 10.x.1981, G.F. & J.F. Hevel, [4♂ 2♀, USNM: UCRCENT00245041‒42, UCRCENT00245035‒36, UCRCENT00245038‒39]. Citrus Co., Homosassa Tract, 28°45'47"N, 82°24'37"W, 19.x.2002, sandhill community, Jackson Mosley, [1♂ 1♀, ABS: UCRCENT00305307, UCRCENT00305299]. Clay Co., Goldhead Branch S.P., 29°50'08"N, 81°57'03"W, 28.v.1991, G. Zolnerowich, [2♂ 2♀, UCRC: UCRCENT00301008‒11]. Dade Co., Miami Matheson Hammock, 25°40'36"N, 80°15'46"W, 15.x.1990, R.S. Anderson, [1♀, UCRC: UCRCENT00301038]. Duval Co., St. Johns Bluff, 30°23'06"N, 81°30'02"W, 30.viii.1976, E.E. Grissell, [1♀, FSCA: UCRCENT00241411]. Gainesville, Sta. 40, 29°39'06"N, 82°19'29"W, 30.v.1967, moist oak-pine flat-woods, open, Vacc. Myrcinites-grass, F. W. Mead, [1♀, FSCA: 62

UCRCENT00306376]. Gainesville, 29°39'06"N, 82°19'29"W, 28.iii.1976, decid. for., E.E. Grissell, [12♂ 2♀, FSCA: UCRCENT00241412‒25]. Idem, but A.E.I, 29°35'53.6"N, 82°21'54.8"W, 15.vii.-2.viii.1987, D.B. Wahl, [1♀, UCRC: UCRCENT00301032]. Goldhead Branch S.P., 29°50'57"N, 81°57'39"W, 28.v.1991, G. Zolnerowich, [1♂, UCRC: UCRCENT00172461]. Gould Rd. Preserve, 27°08'01"N, 81°19'36"W, 18 May 2009, fl. scrub, site 1, flight trap w. pan, M. Deyrup, H. Otte & A. May, [1♂, ABS: UCRCENT00305304]. Highlands Co., Archbold Bio. Research Station, 27°10'54"N, 81°21'06"W, 1.vii.2008, in flight “lousy ten acres”, M. Deyrup, [1♀ ABS: UCRCENT00305301]. Hilliard, 30°41'18"N, 81°55'21"W, 19.viii.1930, [1♀, CMNH: UCRCENT00172490]. Idem, R. H. Beamer, [1♂, CMNH: UCRCENT00172486]. Myakka River St. Pk, 27°14'21"N, 82°19'00"W, 23.vi.1965, C.W. O'Brien, [1♂, LACM: UCRCENT00172489]. St. Augustine, 29°53'39"N, 81°18'48"W, 17.vi.1938, G. P. Englehardt, [2♂, UCRC: UCRCENT00301040-a and b]. St. Petersburg, 27°46'23"N, 82°38'24"W, 28.iv.1908, Van Duzee, [1♀, USNM: UCRCENT00245040]. Juniper Spr Rd., 41m, 29°13'33"N, 81°43'35"W, 19.ix.2001, sand/oak scrub, J. Heraty, [1?#, UCRC: UCRCENT00092111]. Lake placid, Archbold Bio. Research Station, 27°10'54"N, 81°21'06"W, 16.xi.1987, malaise trap, M. Deyrup, [1♂, UCRC: UCRCENT00300870]. Idem, but 17.v.1989, malaise trap, M. Deyrup, by Lake Annie, [1♂, ABS: UCRCENT00305298]. Idem, but 21.xi.1986, trail 2 SSo, malaise trap, M. Deyrup, [1♀, ABS: UCRCENT00305291]. Idem, but 24.vii.1985, [1♀, UCRC: UCRCENT00300939]. Idem, but 25.xi.1983, trail 1, [1♂, ABS: UCRCENT00305297]. Idem, 25.x.1984, [1♂, UCRC: UCRCENT00300938]. Idem, but, 8.viii.1987, SSo, [1♀, ABS: UCRCENT00305292]. Idem, but 8.x.1986, [1♂, UCRC: UCRCENT00301029]. Idem, but 9.ix.1993, cage road, [1♂ 1♀, ABS: UCRCENT00305289, UCRCENT00305305]. Lake Co., Green Swamp WMA, 28°19'38"N, 81°54'14"W, 30.vii.1987, malaise trap, V. Gupta, [2?#, UCRC: UCRCENT00092024‒25]. Lake Eaton, 29°15'32"N, 81°52'03"W, 10.ix.-2.x.1975, malaise trap, J. Wiley, [1♀, UCRC: UCRCENT00300901]. Idem, but 13.x.-5.xi.1975, [1♂ 1♀, FSCA: UCRCENT00241427, UCRC: UCRCENT00302311]. Idem, but 8-13.x.1975, Wiley & Holler, [12♀ 29♂ ABS: UCRCENT00305293‒96, UCRC: UCRCENT00300880‒96, UCRCENT00300900‒22, UCRCENT00300928‒37, UCRCENT00302305]. Levy Co., Sea Horse Key, 29°05'51"N, 83°04'01"W, 22.vi.1957, H.V. Weems Jr., [1♂, FSCA: UCRCENT00241384]. Liberty Co., Torreya State Park, 30°34'10"N, 63

84°56'59"W, 22.vii.1974, H.V. Weems, Jr., [1♂, FSCA: UCRCENT00172480]. Torreya State Park, 30°34'10"N, 84°56'59"W, 5.vii.1965, malaise trap, H.V. Weems Jr., [1♂, FSCA: UCRCENT00241398]. Marion Co., 8.0 km N Hwy 40 on 65 nr. Juniper Springs Rec. Area, 30m, 29°14'58"N, 81°43'38"W, 30.v.1987, pine/palmetto forest, J. Heraty, [14♂ 1♀, UCRC: UCRCENT00305717‒24, UCRCENT00305726‒32]. Ocala Ntl Fst, 29°10'31"N, 81°45'22"W, 22.viii.2014, sweep Cassia fasciculata, A. Baker & L. Davis [2♂ 2♀, UCRC: UCRCENT00436501‒02, UCRCENT00439131‒32]. Okeechobee Co., Kissimmee Prairie State Pk., 27°32'46"N, 81°00'52"W, 7.xi.2007, dry prairie south of headquarters, main road, M. Deyrup, [1♂ 5♀, ABS: UCRCENT00305286, UCRCENT00305287, UCRCENT00305302, UCRCENT00305306, UCRCENT00305309, UCRCENT00305310]. Polk Co., TNC Tiger Creek Pres., 27°49'18"N, 81°28'38"W, 6.vii.2009, scrub, flight trap, M. Deyrup, H. Otte & A. May, [1♀, ABS: UCRCENT00305303]. Putnam Co., 2mi W Interlachen, 29°38'14"N, 82°00'17"W, 30.v.1991, G. Zolnerwich, [1♂, UCRC: UCRCENT00172465]. Union Co., 5 miles E. of Lake Butler T5S R20E section 35, 30°02'01"N, 82°16'27"W, 10 Jul 1986, Lloyd R. Davis. Jr., ex: flower buds of Eupatorium pilosum, [1F# 1?#, UCRC: UCRCENT00300865, UCRCENT00300866].

Kapala inexagens (Walker, 1862) (Fig. 13A‒E)

Thoracantha inexagens Walker, 1862: 381. Type data: Brazil: Santarem. Syntypes (2♀, type n° 5.637a and 5.637b). Lectotype ♀ (n°5.637b) present designation. Type depository: BMNH; types n° 5.637a and 5.637b (examined). Description of the female. [Lectotype in good conditions: right antenna broken and lost, and left antenna broken after pedicel (rest of flagellomeres lost); left frenal process with apex broken and lost]. Schizaspidia inexagens; Walker, 1871: 66. Kapala inexagens; Ashmead, 1904: 472. Additional citation: De Santis, 1980: 209 (catalog). Holcokapala striaticeps Cameron, 1913: 117. Type data: Guyana [probably British Guyana]. Holotype ♀, by monotypy. Type depository: Natural History Museum, London, 64

England (BMNH), type n° 5.391 (examined). Description of female. [Holotype relatively complete: apex of frenal process broken and lost; metasoma broken and lost]. New syn. Kapala striaticeps; Heraty, 2002: 173. [taxonomic position].

Diagnosis. Recognized by a combination of the following characters: mesoscutellar disc flat and smooth or with some faint lateral striae (Fig. 13C); medial apex of mesoscutellar disc raised 0.5‒0.6x height of frenal processes in lateral view; antenna short, 0.8x head height (Fig. 13D); frons with faint slightly oblique striae and lower face with some faint transversal striae (Fig. 13B).

Redescription. Female. Body length [3.2] 3.2‒4.0 mm; length of mesosoma excluding the frenal processes [1.8] 1.6‒2.1 mm. Head, mesosoma, coxae and petiole black; frenal processes dark brown; scapes, pedicels and legs yellow; wings membrane hyaline, venation light brown; Gt1 dark brown. Head. [1.5] 1.4‒1.5x as broad as high, with few sparse setae. Scrobal depression reticulate with some faint irregular striae. Frons with faint slightly oblique striae; lower face with some faint transversal striae (Fig. 13B). Eyes separated by [2.1] 2.1‒2.2x their height. Malar space [0.9] 0.8‒0.9x eye height with faint oblique striae. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2 segmented. Antenna with 10 segments; scape [2.2] 2.2‒3.5x longer than broad, [0.2] 0.2‒0.3x head height; flagellum 8 segmented, length of flagellum 0.8x head height; Fl2 1.3‒1.7x as long as apical width and 1.1x as long as following flagellomeres (Fig. 11D). Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and [1.5] 1.5‒1.7x as high as broad. Midlobe with transversal striae and dorsally rounded in frontal view; lateral lobes with faint longitudinal striae. Axilla with faint longitudinal striae and mesoscutellar disc smooth (Fig. 13C). Mesoscutellar disc [1.4] 1.4‒1.5x as long as axilla with medial apex raised [0.5] 0.5‒0.6x height of frenal processes in lateral view (Fig. 13A). Axillula with longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with faint longitudinal striae; lower mesepimeron weakly reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semi- 65

erect setae. Metapleuron reticulate. Propodeum flat, broad, and delimited by carina, smooth and bright. Frenal processes cylindrical and thin, [1.7] 1.7‒2.0x longer than the basal width of mesoscutellum [1.2] 1.2‒1.4x as long as length of axillae and mesoscutellar disc, slightly arched in lateral view and convex in dorsal view; frenal processes longitudinal striated and apex rounded (Figs. 13A, C). Forewing [2.4] 2.4‒2.5x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.3] 1.0‒1.4x longer than marginal vein; stigmal vein [1.5] 1.2‒1.7x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa semiglobose [1.4] 1.4‒1.7x longer than broad, with a row of setae semi-erect medially. Metafemur [4.4] 4.2‒4.5x longer than broad, with faint striae and sparse setae. Metasoma. Petiole slightly flattened dorsally, reticulate and with weak longitudinal striae,

[2.9] 1.8‒2.0x longer than broad and [1.6] 1.0‒1.6x as long as metacoxa. Gt1 smooth and tergal scar absent.

Male. Unknown

Biological notes. Unknown.

Distribution. Brazil (Fig. 13E).

Lectotype. BRAZIL: [Pará], Santarém, [♀, UCRC_ENT 309824, BMNH: type n° 5.637a and UCRC_ENT 310056, BMNH: type n° 5.637b].

Additional material examined. BRAZIL: Pará: Vigia [de Nazaré], 00°51'28''S, 48°08'31'', 6.iv.1961, Bechiné, [1♀, MPEG]. Belterra, Floresta Nacional Tapajós, C. Jamabaquá, 2°49'24''S, 55°0'52''W, 11.ix.2010, Varredura, J. Almeida, L. Cézar, F. Fernandes, R. Kawada & S. Moraes, [1♀, MZUSP]. Goiás: Rib. Vaozinho, 16°40'00''S, 49°15'00''W, 12.ii.1962, J. Bechyné, [1♀, MZUSP 7557]. GUIANA: [probably British Guiana], P. Cameron Coll., [1♀, BMNH type n° 5.391, BMNH: UCRCENT310014].

Kapala ivorensis Risbec, 1954 (Fig. 14A‒G)

Kapala ivorensis Risbec, 1954: 1086‒1090. Type data: Ivory Coast, Adiopodoumé. Holotype ♀, by monotypy. Type depository: MNHN, (examined). Description of female, illustrated. [Holotype relatively complete: the specimen is conditioning on excavated lamina; head e mesosoma separated from each other. Antennae broken and lost, eyes cracked; mesosoma with fungus].

Diagnosis. Recognized by a combination of the following characters: frons with fine oblique striae since the ocellar triangle toward the lower margin of the torulus, longitudinal striae near to eyes and lower face with faint transversal striae (Fig. 14B); antenna relatively long, 1.1‒1.2x head height and basal flagellomere smaller than following flagellomere (Fig. 14C); frenal processes long (Figs. 14A, E). It occcurs only in the Old World (Fig. 14G).

Redescription. Female. Body length 2.8‒3.6 mm; length of mesosoma excluding the frenal processes 1.4‒1.8 mm. Head, mesosoma and petiole black; frenal processes and coxae reddish-brown; scapes, pedicels and legs yellow; flagellomeres brown; wings membrane hyaline, venation brown; Gt1 reddish-brown. Head. 1.4‒1.5x as broad as high, with few sparse setae. Scrobal depression reticulate with some faint striae. Frons with fine oblique striae, since the ocellar triangle toward the lower margin of the torulus, longitudinal striae near to eyes; lower face with faint transversal striae (Fig. 14B). Eyes separated by 1.9‒2.1x their height. Malar space 0.8‒0.9x eye height with faint oblique striae. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/3 segmented. Antenna with 10 segments; scape smooth with sparse setae, 3.0‒4.0x longer than broad, 0.3x head height; flagellum 8 segmented (Fig. 14C), slightly serrated, length of flagellum 1.1‒1.2x head height; Fl2 1.7‒2.0x as long as apical width and 0.7‒1.0x as long as following flagellomere. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.5‒1.6x as high as broad. Midlobe with transversal striae and dorsally slightly bilobed in frontal view; lateral lobes with faint longitudinal striae. Axilla and mesoscutellar disc longitudinal 67

striated with a median longitudinal groove. Mesoscutellar disc 1.0‒1.2x as long as axilla with medial apex raised 0.7‒1.3x height of frenal processes in lateral view. Axillula with wide longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with some fine longitudinal striae and punctuations (Fig. 14C); lower mesepimeron reticulate; femoral depression crenulate. Callus slightly swollen and covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad, reticulate and delimited by carina. Frenal processes cylindrical, 2.5‒3.1x longer than the basal width of mesoscutellum, 1.7‒2.0x as long as length of axillae; frenal processes, arched in lateral view and slightly convex in dorsal view; with longitudinal striae and apex excavated and with ventral region like a flange-shape more elongated (Fig. 14E). Forewing 2.6‒2.7x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.0‒1.1x longer than marginal vein; stigmal vein 1.3‒2.0x longer than broad; postmarginal vein indistinct. Hindwing fringed, 4 hamulli curved hook-shaped. Metacoxa semiglobose 1.4‒2.0x longer than broad. Metafemur 5.0‒6.2x longer than broad, striated with sparse setae. Metasoma. Petiole slightly flattened dorsally, reticulate and with faint longitudinal striae,

3.0‒4.0x longer than broad and 1.2‒1.3x as long as metacoxa. Gt1 smooth and bright with tergal scar absent. Male. Body length 2.9‒3.6 mm; length of mesosoma excluding the frenal processes 1.1‒1.7 mm (Figs. 14D, E). Similar to female except for the following: antenna with 12 segments, flagellum 10 segmented, each flagellomere with a single long rami; length of rami of

Fl2 1.1‒1.2x head height and 0.9x as long as following flagellomere. Petiole 5.0‒7.0x longer than broad and 1.7‒2.1x as long as metacoxa.

Comments. K. ivorensis is an exception, being the only species within Kapala occupying a range extending across the Afrotropics. It was described by Risbec (1954) from a sigle female specimen from Ivory Coast, though it was misidentified as a male. According Heraty (2002) no males had been collected for this species even though several collections of females had been made, and therefore proposed that K. ivorensis might be a thelytokous species. Therefore, we describe for the first time the male of K. ivorensis. Also, the specimens used to redescribed K. ivorensis was come from Ivory Coast (UCRC_ENT 302315, UCRC_ENT 242391 and 242392). 68

Biological notes. Unknown.

Distribution. Ethiopian (Cameroon, Ivory Coast, Kenya, Nigeria, Sierra Leone [Walker, 1846; mistakenly identified as Thoracantha furcata]; Uganda, Democratic Republic of Congo). Malagasy (Madagascar) (Fig. 14G).

Holotype. IVORY COAST: Lagunes: Adiopodoumé, Abidjan, 5°20'08"N, 4°07'45"W, [1♀, MNHN: UCRCENT00298983].

Additional material examined: CAMEROON: Musone, Mboandony, 4°26'54"N, 9°16'03"E, 11.xii.1981, sweep, S.G. Compton, [1♀, UCRC: UCRCENT00302317]. Victoria, 0-80m, 4°00'34"N, 9°13'13"E, 5-18.xi.1975, W. Schacht, [1♀, ZSMC: UCRCENT00245264]. Centre: Messamendongo, 3°48'35"N, 11°31'17"E, 1-13.iv.2003, malaise trap, T. Marc, [1♂ 1♀, UCRC: UCRCENT00235916‒17]. DEMOCRATIC REPUBLIC OF THE CONGO: Libenge, Savane Liki-Bembe, 3°39'00"N, 18°38'00"E, 26.ii.1948, R. Cremer & M. Neuman, [2♀, ISNB: UCRCENT00242087‒88]. Libenge, Vallee Liki Bembe, Bavula, 4°29'10"N, 19°07'10"E, 26.ii.1948, R. Cremer & M. Neuman, [3♀, ISNB: UCRCENT00242089, UCRCENT00242093, UCRCENT00242094]. Idem, but 28.ii.1948, [3♀, ISNB: UCRCENT00242090‒92]. Tanganyika Territory, S. of Bukavu, 6°13'23"S, 27°54'30"E, 28.viii.1931, J. Ogilvie, [1♀, BMNH: UCRCENT00309830]. University of Kisangani, maise field, 0°30'51"N, 25°10'34"E, 18.i.2013, [1♀, ????: UCRCENT00320312]. Yangambi, D. for. Arboretum parc. 2B, 0°45'47"N, 24°26'20"E, 21.xi.1946, Lele, [1♂ 2♀, ISNB: UCRCENT00242095‒97]. GAMBIA: Western Div.: Lamin, Abuku Nature Reserve, 12°23'29"N, 16°39'09"W, 27.i.1978, L. Huggert, [2♀, BMNH: UCRCENT00309828, UCRCENT00309832]. Guinea: Mt. Nimba, 514-740m, 7°41'00"N, 8°23'00"W, xii.1990-iii.1991, rainforest, FIT, L. Leblanc, [1♀, UCRC: UCRCENT00292763]. IVORY COAST: Reserve do Banco, 5°23'00"N, 4°03'00"W, R Paulian & C Delamare, [2♀, MHNG: UCRCENT00242391‒92]. Bouake: -, 7°41'17"N, 5°01'50"W, i- x.1981, rice fields, P. Cochereau, [1♀, UCRC: UCRCENT00302315]. KENYA: 0°01'25"S, 37°54'22"E, vi.1968, Storch, [1♀, ZMUH: UCRCENT00245181]. Coast: Arabuko-Sokoke Forest, 3°25'13"S, 39°53'49"E, 8-9.i.2000, sweep, R. Copeland, [1♀, UCRC: 69

UCRCENT00302068]. Nyanza: LK. Victoria Ungoye; ICIPE res. site, 1145m, 0°36'54"S, 34°05'31"E, 22.viii.-8.ix.1998, MT in 2nd, S. Miller & P. Otieno, [1♀, UCRC: UCRCENT00302069]. MADAGASCAR: 11 km SE Ampasimanotra (-Brickaville), 5m, 18°54'00"S, 49°08'00"E, 15.iv.1989, on low vegetation, littoral rainforest, P. S. Ward, [1♀, UCRC: UCRCENT00302316]. Analanjirofo Region: Tampolo, 15°43'36"S, 49°57'19"E, 1.viii.1949, Guérin-Méney, [1♀, MNHN: UCRCENT00298978]. Tamatave: 11 km. N. Tamatave, 0-30m, 18°03'25"S, 49°22'59"E, 15.i.1985, John Wenzel, [1♀, UCRC: UCRCENT00302070]. Toamasina: Ankadirano, Rte East Fénérive, 24°13'00"S, 44°16'59"E, viii.1958, Randimby, [2♀, MNHN: UCRCENT00298886, UCRCENT00298890]. Mobot site, Analalava 7 km SW of Foulpointe, 18m, 17°41'36"S, 49°27'37"E, 28.ix.-5.x.2007, on sand in low altitude dense humid forest, malaise trap, M. Irwin, R. Harin’Hala, [1♀, CASC: UCRENT00018900]. Idem, but 3-11.i.2008, [1♀, CASC: UCRENT00018899]. NIGERIA: Ibadan, 7°23'47"N, 3°55'00"E, 15.ii.1963, D.C. Eidt, [1♀, CNC: UCRCENT00300633]. Ikoyi Lagos, 6°27'00"N, 3°25'59"E, ii.1973, M.A. Cornes, [1?#, NMPC: UCRCENT00416489]. Olokemeji Ibadan, 7°25'00"N, 3°32'00"E, [1?#, USNM: UCRCENT00416827]. Ondo: 1.6 km E Owena, 268m, 7°11'54"N, 5°01'50"E, 19.vii.2008, cacao plantation, sweep, J. Mottern, [1♀, UCRC: UCRENT00000313]. Osun: Ile-Ife, W State, 7°31'16"N, 4°31'20"E, viii.1973, J.T. Medler, [1?#, NMPC: UCRCENT00416513]. REPUBLIC OF CONGO: Dept. Pool: Lesio- Louna Pk, Abio, 330m, 3°06'01"S, 15°31'26"E, 16-2.xi.2008, malaise trap, Sharkey & Braet, [2♀, USA: UCRCENT00241582, UCRCENT00241605]. Idem, but 16-23.ix.2008, [1♀, HIC: UCRCENT00241575]. Lesio-Louna Pk, Iboubikro, 340m, 3°16'12"S, 15°28'16"E, 19-25.xi.2008, Sharkey & Braet, malaise trap, [1♀, USA: UCRCENT00241573]. Idem, but 330m, ix.2008, [1♀, HIC: UCRCENT00241571]. Pool Dept.: Abio, Lesio-Louna Pk., 330m, 3°06'01"S, 15°31'26"E, 11-18.ix.2008, Sharkey & Braet, [1♀, HIC: UCRCENT00241581]. Idem, but 30.ix.-7.x.2008, [1♂, HIC: UCRCENT00241604]. Sao Tome: Poto CIAT compound, 0°14'33"N, 6°36'34"E, 7- 12.vi. 1999, malaise trap, A. Polaszek, [2♀, UCRC: UCRCENT00092139‒40]. SIERRA LEONE: Freetown, Cape Sierra Hotel area, 8°31'00"N, 13°17'00"W, 23.xi.1993, along roadside, sweep, Mac Donald, [2♀, MZLU: UCRCENT00172295‒96]. S.L. Peninsula, 8°16'00"N, 13°04'00"W, 25.xii.1999, along roadside, sweep, Mac Donald, [1♀, MZLU: UCRCENT00172294]. Moyamba District: Njala, 8°09'36"N, 12°23'02"W, 17.vi.1931, 70

solanaceous plant, E. Hargreaves, [1♀, BMNH: UCRCENT00309827]. SOUTH AFRICA: KwaZulu-Natal: Durban, 29°53'00"S, 31°03'00"E, iv.1942, Marley, [1♀, SAMC: UCRCENT00242751]. Montclair, 29°55'00"S, 30°58'00"E, 16.v.1945, Marley, [1♀, SAMC: UCRCENT00242750]. Mpumalanga: Stridjum tunnel area, 730m, 24°27'47"S, 30°36'31"E, 31.i.2006, J. Heraty, [1♀, UCRC: UCRCENT00278291]. Transvaal: Zoutpansberg, 22°58'00"S, 29°45'00"E, [1♀, MZPW: UCRCENT00242647]. UGANDA: Kawanda, 0°50'00"N, 31°55'00"E, vii.1941, T.H.C. Taylor, [1♂, BMNH: UCRCENT00309829]. Idem, but v.1941, [1♀, BMNH: UCRCENT00309825]. Idem, but v.1943, [1♀, BMNH: UCRCENT00309826]. Bwamba Co., Semuliki Nat. Pk., north +/- savannah part inside Nat’l Park, 0°50'00"N, 30°03'00"E, 16.iii.2013, A. Gumovsky, [1♀ 1?, ????: UCRCENT00320309, ????: UCRCENT00320310]. Ankole Dist.: Kalinzu Forest, 0°25'00"S, 30°05'00"E, 6-16.ii.1973, H. Gonget, [4♀, ZMUC: UCRCENT00245098‒99, UCRCENT00245104‒05]. Buganda: Kampala, 0°18'49"N, 32°34'53"E, 4.ii.1930, on citrus, H. Hargreaves, [1♂, BMNH: UCRCENT00309831]. Kampala: Muyenga Hill, 0°17'49"N, 32°36'54"E, 10.i.1973, H. Gonget, [5♀, ZMUC: UCRCENT00245095‒96, UCRCENT00245103, UCRCENT00245106‒07]. Idem, but 13.iii.1973, [1♀, ZMUC: UCRCENT00245102]. Idem, but 26.iii.1973, [1♀, ZMUC: UCRCENT00245108]. Idem, but 4.i.1973, [2♀, ZMUC: UCRCENT00245097, UCRCENT00245101]. Idem, but ii.1973, [1♀, ZMUC: UCRCENT00245100].

Kapala izapa Carmichael, 2006 (Figs. 15A‒G)

Kapala izapa Carmichael, 2006: 568‒571. Type data: Mexico: Chiapas, Rosario Izapa. Holotype ♂, by original designation. Type depository: USNM, (examined). [Both sexes described and illustrated. Holotype in good conditions].

Diagnosis. Recognized by a combination of the following characters: basal flagellar rami of males 1.7‒1.8x height of head (Fig. 15A) and flagellum of females 1.2‒1.3x height of head; flagellar segments light brown and with slightly lighter scapes; frenal processes arching laterally 71

and dorsally (Figs. 15A); frenal processes entirely brown (some females) or apically brown and black basally.

Redescription. Male. Body length [3.2] 3.0‒3.2 mm; length of mesosoma excluding the frenal processes [1.5] 1.3‒1.5 mm. Head, mesosoma, coxae, and petiole black with a dark green sheen; frenal processes black basally, becoming brown apically; legs yellow; antenna, and wings venation light brown; wings membrane hyaline; Gt1 basally dark brown and apically light brown. Head. [1.6] 1.6‒1.7x as broad as high, with few sparse setae more prominent in frons. Scrobal depression weakly rugose. Frons longitudinaly striated; lower face with some faint transversal striae (Fig. 15B). Eyes separated by [2.2] 2.2‒2.3x their height. Malar space [0.8] 0.8‒1.2x eye height. Supraclypeal area and clypeus smooth and bare. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape [1.6] 1.6‒2.0x longer than broad, [0.2] 0.2‒0.3x head height; flagellomeres 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 [1.6] 1.6‒1.7x head height and [0.9] 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.4‒1.6x as high as broad. Midlobe with transversal striae and dorsally rounded in frontal view; lateral lobes with faint longitudinal striae. Axilla with longitudinal striae; mesoscutellar disc with slightly oblique striae converging to midline (Fig. 15E). Mesoscutellar disc [1.4] 1.2‒1.4x as long as axilla with medial apex raised [1.6] 1.3‒1.6x height of frenal processes in lateral view. Axillula with longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum with some longitudinal faint striae; upper mesepimeron smooth (Fig. 15C); lower mesepimeron weakly reticulate; femoral depression weakly crenulated. Callus carinate and weakly reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and weakly reticulate with a longitudinal carina. Frenal processes cylindrical, [3.1] 3.0‒3.3x longer than the basal width of mesoscutellum, [1.8] 1.8‒2.0x as long as length of axillae and mesoscutellar disc, arched in side view and slightly convex in dorsal view (Fig. 15E); frenal processes with irregular longitudinal striae, giving a serrated appearance; apex of frenal processes emarginated, with ventral region like a flange-shape more elongated. Forewing [2.3] 2.2‒2.3x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.2] 1.0‒1.2x longer than marginal vein; stigmal vein rectangular 2.0x as longer than broad; 72

postmarginal vein absent. Hindwing fringed. Metacoxa semiglobose [1.8] 1.8‒2.0x longer than broad, with a row of semi-erect setae medially. Metafemur [6.0] 5.7‒6.0x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical and rugose with fine parallel carinae, [7.3] 7.3‒8.0x longer than broad and [2.4] 2.0‒2.4x as long as metacoxa. Gt1 smooth with faint sparse punctuations, tergal scar absent.

Female. Body length 2.8‒2.9 mm; length of mesosoma excluding the frenal processes 1.5‒1.6 mm (Fig. 15D). Similar to male except for the following: scapes and pedicels yellow; flagellum light brown; frenal processes black or brown basally. Head 1.4‒1.5x as broad as high. Antenna with 10 segments; scape 4.0x longer than broad; flagellum 8 segmented, weakly serrate and terminal clava with three fused segments and slightly expanded; length of flagellum 1.2‒1.3x head height; Fl2 1.5x as long as apical width and 0.9‒1.0x as long as following flagellomere. Mesoscutellar disc 1.4‒1.7x as long as axilla. Frenal processes 2.3‒2.6x longer than the basal width of mesoscutellum, 1.4‒1.5x as long as length of axilla, frenal processes tapering to a single flattened point (Fig. 15F). Petiole 3.0‒4.0x longer than broad and 1.3‒1.5x as long as metacoxa.

Comments. K. izapa is very similar to K. roamndii, but can be distinguished by differences in the antennae: rami of Fl2 of males 1.6‒1.7x head height of head (1.1‒1.2 in K. romandii); flagellum of females 1.2‒1.3 height of head (0.9‒1.1x in K. romandii); frenal processes arching both laterally and dorsally (relatively level and straight in K. romandii); and frenal processes are entirely brown (some females) or apically brown and black basally (entirely black in K. romandii).

Biological notes. Host association: Ectatomma ruidum Roger (Formicidae: Ectatominae) [Perez et al. 2006, Howard et al. 2001, Lachaud & Perez-Lachaud 2009].

Distribution. Costa Rica and Mexico (Fig. 15C).

Holotype: MEXICO: Chiapas: Rosario Izapa, 14°58'00"N, 92°09'00"W, iv.1997, G. Perez & J.P Lachaud, ex: Ectatomma ruidum, [1♂, UCRC: UCRCENT00305918]. Paratypes. COSTA RICA: Alajuela: 5-10 mi. west of La Fortuna, 300m, 10°29'13"N, 84°43'53"W, 25.ix.1972, J. Helava, [1♂, UCRC: UCRCENT00320183]. Heredia: F. La Selva 3km S Pto. Viejo, 10°26'00"N, 84°01'00"W, 2.iv.1987, Hamelia sp., H.A. Hespenheide, [1♂, UCRC: UCRCENT00320184]. Idem, but 28.vi.1986, [3♂, UCRC: UCRCENT00320177‒79]. Idem, but 29.iii.1987, [1♂, UCRC: UCRCENT00320180]. 4.iv. 1987, [1♂, UCRC: UCRCENT00320182]. Idem, but 19.vii.1982, at foliar nectaries of Solanum, [1♂, UCRC: UCRCENT00320181]. Idem, but 25.x.1978, MT #48, T.W. Sherry, [2♂, UCRC: UCRCENT00305915, UCRCENT00320176]. La Selva Biol. Sta. 3km S Pto. Viejo, 10°26'00"N, 84°01'00"W, [2♂, UCRC: UCRCENT00235900, UCRCENT00235901]. MEXICO: Chiapas: Rosario Izapa, 14°58'00"N, 92°09'00"W, iv.1997, G. Perez & J.P Lachaud, [3♂ 1♀, UCRC: UCRCENT00235899, UCRCENT00320139‒40, UCRCENT00320148]. Idem, but iv.1997, ex: Ectatomma ruidum, [4♂ 12♀, UCRC: UCRCENT00305916, UCRCENT00320137‒38, UCRCENT00320146‒47, UCRCENT00320149‒59].

Additional material examined: COSTA RICA: Heredia: La Selva Biol. Sta., 15m, 10°24'00"N, 84°00'00"W, 15-17.ii.1988, rainforest, screen sweep, B. Hubley & D.C. Darling, [12♂ 2♀, ROME: UCRCENT00242730‒36, UCRC: UCRCENT00320192‒98]. MEXICO: Chiapas: Rosario izapa, 14°58'00"N, 92°09'00"W, 1.viii.1997, H.S. As, [1♂, UCRC: UCRCENT00320175]. Idem, but 25.ii.1997, C.O. Azwedo, [1♂, UCRC: UCRCENT00305734]. Idem, but 25.iii.1997, [1♂, UCRC: UCRCENT00305735]. Idem, but 26.xii.1997, [1♂, UCRC: UCRCENT00305917]. Idem, but 27.v.1996, [1♂, UCRC: UCRCENT00320174]. Idem, but iv.1997, ex: Ectatomma ruidum, G. Perez & J. P Lachaud, [13?#, UCRC: UCRCENT00320160‒72]. Idem, but ex. O. laticeps, P. Lachaud, [1♂, UCRC: UCRCENT00320173]. Tapachula Canton Leoncilla, 25m, 14°45'59"N, 92°24'12"W, 16.viii.2004, [1♀, UCRC: UCRCENT00172520]. Veracruz: Veracruz, 19°10'22"N, 96°07'60"W, x.1962, N.L.H. Krauss, [1♂, BMNH: UCRCENT00297969].

Kapala romandii (Guérin-Meneville, 1845) (Fig. 16A‒H)

Thoracantha romandii Guérin-Meneville, 1845: 415: Type data: Colombie [Colombia]. Holotype ♀, type designation unknown. Type depository: MNHN, (examined). Description of female. Ashmead, 1904a: 473. [Change of combination]. Kapala romandii; Ashmead: 1904:473. [new combination]. Lirata sulcifacies Cameron, 1904: 61. Type data: Nicaragua: Chinandega. Holotype ♂, by monotypy. Type depository: BMNH; type n° 5.387, (examined). [Description of male. Additional material is in CUIC. Holotype relatively complete: anterior and posterior left leg broken after femur; anterior right leg broken and lost; right medial leg broken after femur]. New syn. Kapala sulcifacies; Heraty & Woolley, 1993: 522. [Subsequent description of both sexes, illustrated. Change of combination]. Lirata fulvicornis Cameron, 1904: 61. Type data: Nicaragua: Managua. Holotype ♂, by monotypy. Type depository: BMNH; type n° 5.386 (image examined). [Description of male. Synonymy by Heraty & Wolley, 1993: 522. Additional material is in CUIC]. Lirata nigriventris Cameron, 1904: 61‒62. Type data: Nicaragua: Chinandega. Holotype ♀, by monotypy. Type depository: BMNH; type n° 5.388 (image examined). [Description of female. Synonymy by Heraty & Wolley, 1993: 522. Additional material is in CUIC].

Diagnosis. Recognized by a combination of the following characters: frons with wide longitudinal striae (Fig. 16B), less pronounced in females (Fig. 16E); lower face with faint transversal striae to smooth; antennae short; frenal processes short with apex emarginated (Fig. 16F) and medial apex of mesoscutellar disc raised 1.2‒1.7x height of frenal processes in lateral view (Fig. 16G).

Redescription. Male. Body length [2.8] 2.6‒3.1 mm; length of mesosoma excluding the frenal processes [1.3] 1.2‒1.7 mm. Head, mesosoma, coxae and petiole black; scapes, pedicels, 75

flagellomeres and rami brown; frenal processes reddish-brown to black; legs yellow; wings hyaline, venation brown; Gt1 black with apex ferruginous (Fig. 16A). Head. [1.5] 1.4‒1.5x as broad as high, with few sparse setae more prominent in frons. Scrobal depression weakly rugose. Frons with wide longitudinal striae, which curve to the lower margin of torulus; face with some faint transversal striae (Fig. 16B). Eyes separated by [2.4] 1.8‒2.4x their height. Malar space [0.9] 0.8‒0.9x eye height with slightly oblique striae. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/3. Antenna with 12 segments; scape [2.5] 2.5‒3.5x longer than broad, [0.2] 0.2‒0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 [1.2] 1.1‒1.2x head height and 0.9x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, in lateral view, and [1.4] 1.4‒1.8x as high as broad. Midlobe with transversal striae and dorsally rounded in frontal view; lateral lobes with faint longitudinal striae. Axilla and mesoscutellar disc with longitudinal striae (Fig. 16F). Mesoscutellar disc [1.2] 1.2‒1.4x as long as axilla with medial apex raised [1.4] 1.2‒1.7x height of frenal processes in lateral view (Fig. 16G). Axillula with wide longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum with some longitudinal faint striae (Fig. 16G); upper mesepimeron and lower mesepimeron smooth; femoral depression smooth. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and rugose. Frenal processes cylindrical, [2.8] 2.5‒2.8x longer than the basal width of mesoscutellum, [1.8] 1.4‒1.7x as long as length of axillae and mesoscutellar disc, arched in side view and convex in dorsal view (Figs. 16A, F); frenal processes with irregular longitudinal striae, giving a serrated appearance (Fig. 16F); apex of frenal processes emarginated. Forewing 2.3‒2.4x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.0‒1.2x longer than marginal vein; stigmal vein rectangular oblique 1.2‒1.5x longer than broad; postmarginal vein absent. Hindwing fringed. Metacoxa semiglobose [1.3] 1.8‒2.2x longer than broad, with a row of setae semi-erect. Metafemur [5.2] 5.2‒6.3x longer than broad, with faint striae and sparse setae. Metasoma. Petiole cylindrical and rugose, [5.6] 5.2‒7.4x longer than broad and [1.9]

1.9‒2.1x as long as metacoxa. Gt1 smooth with faint sparse punctuations, tergal scar absent. 76

Female. Body length 2.5‒3.0 mm (Fig. 16D); length of mesosoma excluding the frenal processes 1.4‒1.7 mm. Similar to male except for the following: face with striae less pronounce (Fig. 16E). Antenna with 10 segments; scape 3.0‒3.3x longer than broad; flagellum 8 segmented, the first three segments long and slightly serrated (Fig. 16C), length of flagellum 1.0x head height; Fl2 1.2‒1.5x as long as apical width and 1.0x as long as following flagellomere. Mesoscutellar disc with medial apex raised 1.0‒1.3x height of frenal processes in lateral view. Frenal processes 1.7‒2.3x longer than the basal width of mesoscutellum, 1.2‒1.4x as long as length of axillae and mesoscutellar disc. Metacoxa 1.5‒1.6x longer than broad. Metafemur 4.7‒6.0x longer than broad. Petiole slightly flattened dorsoventrally, 2.7‒3.0x longer than broad and 1.3‒1.5x as long as metacoxa.

Comments. K. ramandii is inserted within the complex of K. romandii complex (with 16 species here rediscribed and described), which is one of the most specious clades exhibiting low supports across differents analyzes previously made by Murray et al. (2013). This species shares some characteristics like, face striated, high profile of mesoscutum and female have 8 flagellomeres. Besides, the amount of intraspecific variation and the species boundries will require a large revision of morphological characters and molecular data for accurate determination. Up to now, are more than 20 morphotypes classified within this complex in addition to the species described above, but which have a high degree of intraspecific hybridization and variation.

Biological notes. Known to oviposit into closed flower buds of Cordia macrostachya (Jacq.) (Boraginaceae), Grossypium hirsutum (L.) (Malvaceae) and flowering asclepiad (Heraty & Woolley, 1993). Eggs laid in clusters of 200-300 eggs (Heraty & Darling, 1984). Usually occurs in sympatry with K. iridicolor.

Distribution. Brazil, British Virgin Islands, Colombia, Costa Rica, Ecuador, El Salvador, Guiana, Honduras, Mexico, Nicaragua, Panama, Peru, Suriname, Trinidad & Tobago, Venezuela (Fig. 16H).

Holotype. NICARAGUA: Chinandega, Cameron coll., [1♂, BMNH: UCRC_ENT 310003, BMNH type n°5.387].

Additional material examined: BRAZIL, Rio Grande do Norte: Baixa Verde, 150m, 5°33'00"S, 37°57'00"W, W.M. Mann, [5♂ 4♀, LACM: UCRCENT00242200‒03, UCRCENT00242213, UCRCENT00305104, UCRCENT00305106‒08]. BRITISH VIRGIN ISLANDS: Virgin Gorda, Gorda Peak Nat. Pk., Gorda Pk trail, 18°29'04"N, 64°24'14"W, 6.i.1993, beating, M. Ivie & D. Chadwick, [1♀, CNC: UCRCENT00300702]. COLOMBIA: Cauca: Rio Micay Lopez, 90m, 2°56'00"N, 77°12'00"W, 20.v.1977, MT, Mac A. Tidwell, [1♂, FSCA: UCRCENT00306401]. Chocó: 50-60 km W. Bolivar, 500m, 5°45'00"N, 76°30'00"W, 11- 13.iv.1973, montane forest, J. Helava, [5♂ 2♀, CNC: UCRCENT00316139‒40, UCRCENT00316309‒11, UCRCENT00316314‒15]. Cundinamarca: Fusagasunga, 1700m, 4°20'18"N, 74°21'50"W, E. Pehike S. [1♀, MZPW: UCRCENT00242648]. Huila: Gallego, 1300m, 2°15'50"N, 75°58'07"W, 1937 [1♂, UCRC: UCRCENT00306363]. Magdalena: Curumani, 60 km. S. Becerril, 150m, 9°15'21"N, 73°31'07"W, 22.vii.1968, Borys Malkin, [1♂ 2♀, AMNH: UCRCENT00237894, UCRC: UCRCENT00301047‒48]. PNN Tayrona Zaino, 50m, 11°20'01"N, 74°02'01"W, 14-29.vi.2000, malaise trap, M.240, R. Henriquez, [3♂, UCRC: UCRCENT00091931‒32, UCRCENT00092130]. Meta: Carimagua, 370m, 4°34'00"N, 71°20'00"W, i.1973, M. Corn, [1♀, UCRC: UCRCENT00306564]. Tolima: Armero, 350m, 4°58'00"N, 74°54'00"W, 30.i-5.ii.1977, E.L. Peyton, [1♀, UCRC: UCRCENT00301044]. Valle del Cauca: 15 km. S. of Cali, 1000-1500m, 3°17'10"N, 76°35'31"W, 27.x.1971, Eberhard & Garcia, [2♂, CNC: UCRCENT00316303, UCRCENT00316305]. 17 km. S. of Cali, 1000m, 3°14'03"N, 76°33'33"W, 10.iv.1971, Eberhard & Garcia, [3♂ 3♀, CNC: UCRCENT00316126‒28, UCRCENT00316304, UCRCENT00316307‒08]. Buenaventura, 3°53'12"N, 77°04'13"W, 6.iv.1965, A.E. Michelbacher, [2♀, EMEC: UCRCENT00236262, UCRC: UCRCENT00306359]. Cadelaira, Finca San Luis (Lago), 1010m, 3°24'00"N, 76°23'00"W, 21-23.iii.1975, tropical dry forest, malaise trap, R.C. Wilkerson, [1♂, FSCA: UCRCENT00318601]. Cali, 1000m, 3°25'14"N, 76°31'20"W, iv.1971, [2♂, CNC: UCRCENT00320626‒27]. Central de Anchicaya 30 km. E. Buenaventura, 560m, 3°52'56"N, 76°48'02"W, 10.vi.1975, tropical very wet forest, MT, R.C. Wilkerson, [1♂, FSCA: 78

UCRCENT00306404]. Rio Zabaletas, 4 km. E. Zabaletas, 1000m, 3°42'25"N, 76°17'06"W, 3- 5.vi.1975, tropical rain forest, malaise trap R.C. Wilkerson, [3♂, FSCA: UCRCENT00306400, UCRCENT00306414‒15]. Triana, km. 43, Buenaventura Street, 3°51'25"N, 76°48'25"W, M.A. Tidwell, [1M#, FSCA: UCRCENT00306402]. COSTA RICA: Alajuela: Chiles de Aguas Zarcas, 300m, 10°23'56"N, 84°20'20"W, i.1990, café, R. Cespedes, [1♂, UCRC: UCRCENT00301174]. La Fortuna, 600m, 10°28'14"N, 84°38'43"W, 18.ii.1964, H.E. Evans, [9♂ 2♀, MCZ: UCRCENT00242361, UCRCENT00242363‒69, UCRCENT00242378, UCRCENT00306341‒42]. San Pedro de la Tigra, 200m, 10°22'07"N, 84°34'55"W, iii-iv.1990, Cacao, R. Cespedes, [2♂, UCRC: UCRCENT00305476‒77]. Guanacaste: Est. Pitilla, 9km S Santa Cecilia, 700m, 10°58'43"N, 85°25'00"W, ix.1989, P. Hanson, [2♂ 1♀, UCRC: UCRCENT00172305, UCRCENT00302399, UCRCENT00305465]. Santa Rosa NP 300m, 10°53'33"N, 85°45'59"W, 11.v-1.vi.1985, D.H. Janzen, I.D. Gauld, [1♂, UCRC: UCRCENT00320119]. Santa Rosa NP, 300m, 10°53'33"N, 85°45'59"W, 22.vi-13.vii.1985, D.H. Janzen, I.D. Gauldm [2♂, UCRC: UCRCENT00320120, UCRCENT00320122]. Heredia: Bio. La Selva, 50-150 mm, 10°26'00"N, 84°01'00"W, vii.1999, CRI002-623422, [2♂, INBIO: UCRCENT00282421‒22]. Chilamate, 100m, 10°26'60"N, 84°05'55"W, 4.ii.1989, P. Hanson, [4♂, UCRC: UCRCENT00302417, UCRCENT00305473‒75]. Limon: Guapiles, 10°12'21"N, 83°47'21"W, 17.vii.1964, R.E. Woodruff, [1♂ 2♀, FSCA: UCRCENT00241439‒40, UCRCENT00306406]. Los Diamantes, 10°12'21"N, 83°47'21"W, 7-12.iv.1964, F.S. Truxal, [1♂ 3♀, LACM: UCRCENT00242208, UCRCENT00242210, UCRCENT00242215, UCRCENT00305103]. Pandora, 20-200m, 9°44'30"N, 82°57'60"W, 29-30.iv.1964, F.S. Truxal [1♀, LACM: UCRCENT00242216]. ECUADOR: El Oro: Puyango, 450m, 3°52'36"S, 80°04'39"W, 27.ii.1988, Mike Huybensz, [3♂, MCZ: UCRCENT00242354‒55, UCRCENT00242360]. Guayas: Isla Puna, 2°46'35"S, 80°08'34"W, 22.iii.1988, Mike Huybensz, [13♂ 7♀, MCZ: UCRCENT00242341‒50, UCRCENT00242375‒76, UCRC: UCRCENT00305981‒85, UCRCENT00305992‒94]. Imbabura: Carolina, 900m, 0°47'32"N, 78°17'37"W, 18-20.xii.1975, W. Schacht [1♀, ZSMC: UCRCENT00245261]. Manabi: Chone, 0°41'42"S, 80°05'19"W, v.1976, Fritz, [4♀, AMNH: UCRCENT00237811, UCRCENT00237814, UCRCENT00237824‒25]. Napo: Reserva Ethinca Waorani, 1km S Onkone Gare Camp, 220m, 0°39'10"S, 76°26'00"W, 27.x.1998, fogging terra firme forest, T.L. 79

Erwin et al, [1♀, UCRC: UCRCENT00306556]. Orellana: Tiputini Biodiversity Sta. nr. Yasuni National Park, 220-250m, 0°37'55"S, 76°08'39"W, 29.vi.1998, Fogging, Lot 1803, T.L. Erwin et al., [1♀, UCRC: UCRCENT00305969]. Pichincha: Puerto Quito, 0°07'00"N, 79°16'00"W, 3.vi.1987, Martin Cooper [1♀, UCRC: UCRCENT00306565]. Rio Palenque Res. Stn., 0°39'05"S, 79°19'33"W, ii.1983, MT, Sharkey & Masner, [44♂ 2♀, CNC: UCRCENT00313259, UCRCENT00313438, UCRCENT00313447‒75, UCRCENT00313551, UCRCENT00313554‒61, UCRCENT00313570, UCRCENT00313574, UCRCENT00313579, UCRCENT00316134, UCRCENT00316148, UCRCENT00316150]. EL SALVADOR: La Libertad, 19 mi W, 500-750m, 13°45'02"N, 89°35'37"W, 13.viii.1965, A. Raske & C. Slobodehikoff, [1♂, EMEC: UCRCENT00236261]. GUIANA: Blairmont, 6°16'47"N, 57°34'34"W, viii.1923, F.X. Williams, [1♂, BMNH: UCRCENT00239421]. New Amsterdam, 6°13'50"N, 57°30'21"W, viii.1923, F.X. Williams, [2♀, BMNH: UCRCENT00239417, BMNH: UCRCENT00239418]. HONDURAS: Comayagua: 3 km S Comayagua, 600m, 14°24'47"N, 87°37'25"W, 23.vii.1989, R. Cave [1♂, EAPZ: UCRCENT00282620]. Olancho: Montana del Malacate, 15°08'04"N, 85°35'36"W, 3.vii.2002, D. Yanegam [2♀, UCRC: UCRCENT00092082, UCRCENT00092088]. Yoro: El Negrito, Nuevo San Antonio, 15°19'00"N, 87°42'00"W, 10.xi.1989, R. Cave, Artocarpus altilis [1♀, EAPZ: UCRCENT00282626]. MEXICO: Chiapas: 12 mi NW Huixtla, 50m, 15°14'28"N, 92°35'31"W, 28.vi.1965, C. Slobodchikoff & A. Raske, [3♀, UCRC: UCRCENT00306159‒60, UCRCENT00306162]. Jalisco: Est. Biologia Chamela, 100m, 19°30'00"N, 105°02'00"W, 18.xii.1987, P.S. Ward, on low vegetation tropic. dry forest, #9258, [1♂, UCRC: UCRCENT00105489]. Sinaloa: Magistral rt. 40, 23°23'36"N, 105°56'37"W, 18.viii.1985, G. Ekis [1♂, UCDC: UCRCENT00280080]. NICARAGUA: Chinandega: Chinandega, 12°37'28"N, 87°07'47"W, Baker, [8♂ 6♀, CUIC: UCRCENT00241276, UCRCENT00241280‒90, UCRCENT00300486‒87]. Managua: Managua, 12°08'11"N, 86°15'05"W, Baker [3♂, CUIC: UCRCENT00241277‒79]. Masachapa, 11°47'15"N, 86°30'51"W, 25.vii.1970, L. H. Bolston, [1♀, MMNS: UCRCENT00172297]. PANAMA: Bocas del Toro: Bocas del Toro, 5m, 9°20'09"N, 82°15'16"W, vii.1908, W. Robinson, [1♂, USNM: UCRCENT00245051]. Changuinola, 10m, 9°25'48"N, 82°31'12"W, 1.viii.1924, G.C. Wheeler, [1♀, LACM: UCRCENT00242207]. Rio La Gloria, 8 km W Rambaia, 35m, 8°59'04"N, 82°13'57"W, 8.i.2001, forest, sweep, L. Masner, [1♂, UCRC: UCRCENT00102760]. Canal 80

Zone: Barro Colorado Island, 20-180m, 9°09'20"N, 79°50'46"W, 10.vii.1976, R.B. & L.S. Kimsey, [1♂, UCRC: UCRCENT00302342]. Coiba: Estacion Biologica, 7°28'06"N, 81°45'24"W, 12-19.ii.1994, Malaise, J.L. Nieves, [1♂, MNCN: UCRCENT00322477]. Herrera: Chitre, 7°58'00"N, 80°26'00"W, vii.1981, N.L.H. Krauss, [1♂, AMNH: UCRCENT00237819]. PERU: Chiclayo, 30m, 6°46'36"S, 79°50'47"W, 10.vii.1944, Berry, [1♀, MACN: UCRCENT00242253]. La Libertad: Trujillo, Simbal Ca., 7°58'23"S, 78°48'42"W, 7.iv.1974, C. Potter & L. Stange, [1♀, IMLA: UCRCENT00313143]. Madre de Dios: Rio Tambopata Res., 30km (air) SW Pto. Maldonado, 290m, 12°50'00"S, 69°20'00"W, 4-19.ix.1984, D. H. Kavanaugh, [1♂, CASC: UCRCENT00292309]. SURINAME: Sipaliwini: Raleigh Vallen Voltzberg Res Foengoe, 53m, 4°43'00"N, 56°12'00"W, 26.i-15.ii.1982, J. Carpenter & D. Trail, [1♀, ROME: UCRCENT00242712]. TRINIDAD & TOBAGO: Curepe, Sta. Margarita, 10°38'48"N, 61°24'56"W, 4.xii.1977, Mason [1♂ 1♀, CNC: UCRCENT00313332, UCRENT00001369]. Dept. Agriculture grounds, Port of Spain, 10°39'34"N, 61°28'44"W, 2.xi.1918, Harold Morrison, [1♀, CNC: UCRCENT00300704]. Las Cuevas, 10°47'02"N, 61°23'20"W, 16.iii.1995, Along Roadside, R.L. Manuel, [1♂, UCRC: UCRCENT00091808]. Maracas Vlly., 10°41'49"N, 61°24'19"W, 22.v-7.vi.1974, Special Traps, Yaseen Stickum, [2♂ 4♀, CNC: UCRCENT00300728‒31, UCRCENT00313383‒84]. Maracas, 10°40'48"N, 61°24'41"W, 16.v.1961, N. Gopaul, [1♂ 1♀, CNC: UCRCENT00300733, UCRCENT00313382]. Idem, but xii.1977, W. & E. Mason, [1F#, CNC: UCRCENT00313588]. VENEZUELA: Aragua: Pto. de Cata, 10m, 10°29'21"N, 67°44'22"W, 25.xii.1985, P. Kovarik & R. Jones, [1♂, UCRC: UCRCENT00306370]. Bolivar: 7.6km SE Guasipati, 175m, 7°25'03"N, 61°51'27"W, 22.iii.1982, G.F. & J.F. Hevel, [1♂ 1♀, UCRC: UCRCENT00172307, UCRCENT00306372]. Guarico: 57 km East of Santa Maria de Ipire, 240m, 8°47'30"N, 64°50'24"W, 19.iii.1982, G.F. & J.F. Hevel [1♂, UCRC: UCRCENT00302388]. Lara: 1 km E. Barquisineto, 500m, 10°04'38"N, 69°15'18"W, 27.xii.1985, savannah, R. Jones & P. Kovarik, [1♂ 1♀, TAMU: UCRCENT00242862, UCRC: UCRCENT00306156]. Miranda: 2.5 km E. Carenero, 10m, 10°32'29"N, 66°05'43"W, 26.iii.1982, G.F. & J.F. Hevel, [1♂, UCRC: UCRCENT00306373]. Zulia: El Tucuco, 700m, 9°50'44"N, 72°48'44"W, 24.vi.1979, Schuster & Grigarick [1♀, UCDC: UCRCENT00280079]. Los Angeles del Tucuco, 265m, 9°50'56"N, 72°48'32"W, 15-16.iv.1981, edge of disturbed rain forest, E.E. Grissell, [1♂ 1♀, UCRC: UCRCENT00172306, 81

UCRCENT00302383]. Maracaibo, Caño Colorado, 9°54'58"N, 62°54'48"W, 27.vi.1979, R.W. Brooks, R.P. Kimsey & L.S. Kimsey [1♂ 2♀, UCDC: UCRCENT00280081‒83].

Kapala terminalis Ashmead, 1892 (Figs. 17A‒H)

Kapala terminalis Ashmead, 1892: 358. Type data: Cuba. Syntypes 4♂. Lectotype ♂ (type n° 27.301) present designation. Type depository: USNM, type n° 27.301 (examined). [Description of male. Lectotype relatively complete: pedicel and flagellomeres of the left antennae broken and lost; second antennal rami of the right antenna with the apex broken and lost; left middle leg broken and lost; tarsomeres (3, 4 and 5) of the right hind leg broken and lost].

Diagnosis. Recognized by a combination of the following characters: mesoscutellar disc rounded in profile view and not elevated above base of frenal processes in both sexes (Fig. 17C, indicated); male with rami of Fl2 long, 1.6‒2.2x head height (Fig. 17A); female with flagellum long 1.4‒1.7x head height (Fig. 17D), the first three segments long and slightly serrated; general color metallic green to violaceus.

Redescription. Male. Body length [3.5] 2.4‒3.5 mm; length of mesosoma excluding the frenal processes [1.5] 1.2‒1.5 mm. Head and mesosoma metallic bronze-green; scapes and legs yellow; flagellum and rami brown; frenal processes with basal half metallic green and apical half purplish-blue; petiole and coxae dark-blue; wings membranes hyaline, venation brown; Gt1 dark- brown with apex yellowish. Head. [1.6] 1.4‒1.6x as broad as high, with few sparse setae more prominent in frons. Scrobal depression rugose. Frons with longitudinal striae; lower face with transversal striae (Fig. 17B). Eyes separated by [2.2] 2.1‒2.3x their height. Malar space [1.0] 0.9‒1.0x eye height with slightly oblique striae. Supraclypeal area and clypeus with faint striae. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape [2.5] 2.5‒3.5x longer than broad, [0.4] 0.3‒0.4x head height; 82

flagellum 10 segmented, transversal and each flagellomere with a single long rami (Fig. 17A); length of rami of Fl2 [1.8] 1.6‒2.2 head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, in lateral view, and 1.3‒1.5x as high as broad. Midlobe with fine transversal striae and dorsally rounded in frontal view; lateral lobes with faint longitudinal striae. Axilla with fine longitudinal striae. Mesoscutellar disc rounded, in lateral view, with slightly oblique striae and not elevated above base of frenal processes (Figs. 17C, F); mesoscutellar disc [1.4] 1.2‒1.5x as long as axilla. Axillula with fine longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with fine longitudinal striae (Fig. 17C); lower mesepimeron reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and rugose. Frenal processes cylindrical, [2.8] 2.5‒2.8x longer than the basal width of mesoscutellum, [1.9] 1.9‒2.0x as long as length of axilla and mesoscutellar disc, straight in lateral view and, in dorsal view, parallel; frenal processes with fine longitudinal striae and apex rounded (Fig. 17F). Forewing [2.8] 2.6‒2.8x longer than broad covered by small bristles, except for the base bare; submarginal vein [1.0] 1.0‒1.2x longer than marginal vein; stigmal vein rectangular oblique [1.9] 1.9‒2.0x longer than broad; postmarginal vein long. Hindwing fringed. Metacoxa elongated [2.1] 1.7‒2.1x longer than broad. Metafemur [6.5] 6.2‒6.7x longer than broad, with faint striae and sparse setae. Metasoma. Petiole cylindrical and rugose with two weak parallel carinae dorsally, [7.1]

7.3‒8.0x longer than broad and [2.1] 2.0‒2.4x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 2.8‒3.1 mm (Fig. 17E); length of mesosoma excluding the frenal processes 1.5‒1.6 mm. Similar to male except for the following: head 1.3‒1.4x as broad as high. Antenna with 10 segments; scape 3.5‒5.3x longer than broad; flagellum 8 segmented, the first three segments are long and slightly serrated, length of flagellum 1.4‒1.7x head height (Fig.

17D); Fl2 3.0‒3.3x as long as apical width and 0.8‒1.0x as long as following flagellomere. Frenal processes thin, 2.1‒2.5x longer than the basal width of mesoscutellum, 1.5‒2.0x as long as length of axillae and mesoscutellar (Fig. 17G). Metafemur 5.8‒7.2x longer than broad. Petiole 4.2‒5.0x longer than broad and 1.3‒1.7x as long as metacoxa. 83

Biological notes. Association with Odontomachus insularis, Guérin-Menéville (reported as O. haematodus insularis pallens Wheeler) (Clausen, 1941).

Distribution. Bahamas, Cuba, Dominican Republic, Grand Cayman Islands, Haiti, Puerto Rico and U.S. Virgin Islands (Fig. 17H).

Lectotype. CUBA: [1♂, USNMENT 00809412, USNM Type n° 27301].

Additional material examined. BAHAMAS: Eleuthera: Rainbow Bay, 25°26'43"N, 76°41'26"W, xi.1986, malaise trap, J.R. Wiley, [2♂ 1♀, FSCA: UCRCENT00306380‒81, UCRENT00000756]. West Indies: South Bimini Isl., 25°42'33"N, 79°16'27"W, vi.1951, M. Cazier C.&P. Vaurie, [1♂, USNM: UCRCENT00172524]. Idem, but v.1951, [2♂, AMNH: UCRENT00001338‒39]. CUBA: Havana, 23°07'00"N, 82°23'19"W, 6.v.1922, [1♀, UCRC: UCRCENT00306557]. Hoyo, Colorado, 22°58'00"N, 82°32'00"W, Jul 1931, C.P. Clausen, [1♂ 1♀, CMNH: UCRENT00001345‒46]. Manacas Sta. Clara, 22°35'49"N, 80°20'18"W, 7.vi.1930, S.C. Bruner & L. Bouole, [1♂, USNM: UCRCENT00172528]. Sierra Range1, P. del Rio, Cuba, 457.2m, 19°59'47"N, 76°44'55"W, 29.viii.1927, J. Aculia. & Y.S.C. Brumer, [1♂, USNM: UCRCENT00172522]. DOMINICAN REPUBLIC: 18°44'00"N, 70°09'36"W, 13.vi.1974, [1♂ 1♀, CNC: UCRCENT00172525, UCRENT00001360]. Km. 6, W. of Ciudad Trujillo R. D., 18°28'44"N, 69°53'27"W, 22.xii.1955, J. Maldonado & Capriles, [1♂, USNM: UCRCENT00091498]. Cabrera, 19°38'16"N, 69°54'16"W, 22.vii.1978, R.O. Schuster & R.S. Rominger, [1♂, UCDC: UCRENT00001752]. La Altagracia, Boca de Yuma, P.N. del Este, 18°18'02"N, 68°41'30"W, 29.ix.1992, Grimaldi, [4♂, AMNH: UCRCENT00237846, UCRCENT00237850‒51, UCRCENT00237853]. Punta Cana, Reserva, 18°30'40"N, 68°22'38"W, 11-14.xi.2005, forest, screen sweep, L. Masner, [2♂ 2?, ????: UCRCENT00161507, ????: UCRCENT00161509‒11]. Rio San Juan, 19°38'53"N, 70°04'31"W, 1.viii.1978, [1♂, UCDC: UCRENT00001753]. Seashore W. of Ciudad Trujillo, 18°25'28"N, 69°59'49"W, 21.xii.1955, J. Maldonado Capriles, [1♂, USNM: UCRCENT00172527]. Samana: Honduras, 19°12'00"N, 69°22'00"W, 26.vii.1978, R.O. Schuster [1♀, UCDC: UCRENT00001357]. Las Galeras, 19°17'20"N, 69°12'05"W, 29.vii.1978, R.O. Schuster, [3♂ 1♀, 84

UCDC: UCRENT00000755, UCRENT00001358, UCRENT00001750‒51]. Samana, 19°12'42"N, 69°19'56"W, 29.vii.1978, R.O. Schuster, [1♀, UCDC: UCRCENT00243540]. Idem, but R.O. Schuster & R.S. Rominger, [3♂ 1♀, UCDC: UCRCENT00243541, UCRENT00001361‒63]. San Cristobal: Cambita Garabitos, 18°27'15"N, 70°11'52"W, 28.viii.1967, J. C. Schaffner, [1?, UCRC: UCRCENT00305919]. Villa Altagracia, 18°40'27"N, 70°10'20"W, 8.viii.1967, J. C. Schaffner, [1♀, USNM: UCRCENT00091499]. GRAND CAYMAN ISLANDS: West Iindies: 19°18'00"N, 81°13'00"W, 19.viii.1685, R. R. Askew, [1♂, UCRC: UCRCENT00091502]. HAITI: 18°57'50"N, 72°16'57"W, [1♂, ZSM: UCRENT00001344]. Diquini, 18°31'53"N, 72°23'32"W, W.M. Mann., [2♂, MCZ: UCRCENT00242340, UCRENT00001349]. Grand Anse, 18°32'14"N, 74°00'09"W, P.R. Uhler, [2♂, MCZ: UCRENT00000757, UCRENT00001337]. Kenscoff, 18°27'01"N, 72°17'00"W, 1- 6.viii.1961, J. Maldonado C., [1♂, USNM: UCRCENT00091495]. Port au Prince, 18°32'21"N, 72°19'44"W, 2.x.1934, Darlington, [1♂ 1♀, MCZ: UCRENT00001347‒48]. Idem, but ii.1925, G. N. Wolcott, [1♂, USNM: UCRCENT00091500]. Pt. au Prince, 18°32'21"N, 72°19'44"W, [2♂, USNM: UCRCENT00091496, UCRCENT00091497]. PUERTO RICO: Aquadilla: Smtih- Perez, 18°26'09"N, 67°09'20"W, 21.vii.1952, [1♂, RNC: UCRENT00001341]. Cerezos, circle 38, 28.ii.2004, Malaise, L Yunes, [1♂, UCRC: UCRCENT00397249]. U.S. VIRGIN ISLANDS: St. John: Lameshur Bay, got at VIERS, 0-60m, 18°19'20"N, 64°43'23"W, -.iii.1984, M. Ivie, [1♀, MAIC: UCRCENT00242271].

New species descriptions

Kapala corcovata Schoeninger n. sp. (Fig. 18 A‒F)

Diagnosis. Recognized by a combination of the following characters: midlobe elevated with pronounced, rounded and bilobed apex (Figs. 18 C, D, E); lateral lobes not elevated, approximately the height of midlobe; axilla swollen (Fig. 18A); mesoscutellar disc and axilla with a medial longitudinal depression; frenal processes thin with rounded apex.

Description. Male. Body length 3.0‒3.5 mm; length of mesosoma excluding the frenal processes 1.4‒1.8 mm. Head, mesosoma, coxae, petiole and frenal processes black; scapes brown to dark-brown, pedicels, flagellomeres and rami brown; legs pale-yellow, base of anterior femora brown; wings hyaline, venation brown; Gt1 dark-brow, subsequent tergites light brown. Head. 1.5‒1.6x as broad as high, with sparse setae. Scrobal depression punctate. Face with a pattern of fine uniform striations, frons adjacent to scrobes with longitudinal striae and lower face with transversal striae (Fig. 18B). Eyes separated by 2.1‒2.3x their height. Malar space 1.1‒1.3x eye height. Supraclypeal area and clypeus smooth; anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 2.2‒2.5x longer than broad, 0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 1.1‒1.2x head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.5‒1.8x as high as broad. Midlobe elevated with apex pronounced and rounded, in lateral view and; bilobed in frontal view with regular transverse striae (Figs. 18C, D); lateral lobes not elevated, approximately the height of midlobe with faint longitudinal striae (Figs. 18A, E). Axilla swollen. Axilla and mesoscutellar disc longitudinal striated and with a depression crossing them longitudinally in the center. Mesoscutellar disc 1.0‒1.1x as long as axilla with medial apex raised 1.2‒1.5x height of frenal processes in lateral view. Axillula with fine longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum smooth to some even longitudinal striae; mesepimeron slightly reticulate; femoral depression weakly crenulate. Callus carinate, reticulate and pilose. Metapleuron carinate and reticulate. Propodeal disc flat and broad, with shallow depression on dorsal margin. Frenal processes cylindrical and thin (Fig. 18C), 2.8‒3.8x longer than the basal width of mesoscutellum, 1.5‒1.8x as long as length of axillae and mesoscutellar disc, weakly curved in lateral and dorsal view, frenal processes with well-marked irregular longitudinal striae, giving a serrated appearance; apex rounded. Forewing 2.3‒2.4x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.0‒1.3x longer than marginal vein; stigmal vein rectangular, 1.2‒2.0x longer than broad and postmarginal vein absent. Hindwing fringed. Metacoxa semiglobose with a 86

medium line of setae, 1.7‒2.1x longer than broad. Metafemur 5.3‒7.0x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical and colliculate, 5.7‒8.0x as longer than broad, 2.1‒2.5x longer than metacoxa. Tergal scar on Gt1 present or absent.

Female. Body length 2.8 mm (Fig. 18E); length of mesosoma excluding the frenal processes 1.3 mm. Similar to male except for the following: Antenna light brown, with 10 segments (Fig. 18D); scape 3.3x longer than broad, length of flagellum 1.1x head height; Fl2 1.7x as long as apical width and 1.2x as long as following flagellomere. Maxillary/labial palpi 3/2 segmented. Apex of midlobe more pronounced (Fig. 18E). Frenal processes 3.2x longer than the basal width of mesoscutellum and with longitudinal striae. Metacoxa 2.3x longer than broad. Petiole 3.6x longer than broad, and 1.3x as long as metacoxa. Etymology. The specific name refers to the elevated mesocutum whose shape resembles the Morro do Corcovado shape, located in the city of Rio de Janeiro (Brazil) and which is also one of the main tourist attractions of the city.

Biological notes. Unknown.

Distribution. Brazil (Fig. 18F).

Type material. Holotype. BRAZIL, Pará: Conceição do Araguaia, 23-25.v.1983, 08°15'28"S, 49°15'54"W, Malaise trap, J.A. Rafael, [1♂, INPA]. Paratypes (7♂ 1♀). Ceará: Meruoca, 3°31'54.69"S, 40°27'49.41"W, 20-21.vi.2015, Malaise, David Nogueira [1♂, INPA]. Minas Gerais: Conceição do Mato Dentro, Serra da Serpentina, 19°02'13"S, 43°25'30"W , 18- 28.iii.2009, WGS 84, Carga 1 and 2, R.R. Silva, [1♂, 1♀, MZUSP]. Pará: Conceição do Araguaia, 08°15'28"S, 49°15'54"W, 23-25.v.1983, Malaise trap, J.A. Rafael, [4♂, INPA]. Idem, but 19-31.i.1983 [1♂, INPA]. São Paulo: Gália, Estação Ecológica dos Caetetus, Trilha Z1, 22°20ʹS, 49°40ʹW, 26.iv.-0.3.v.2003, Malaise, J.F. Nunes, [1♂, INPA].

Kapala genistriata Schoeninger n. sp. (Fig. 19 A‒E)

Diagnosis. Recognize by a combination of the following characters: a pattern of oblique wide striae from the inferior margin of torulus to the lateral ocelli on the frons and lower face with transversal striae (Fig. 19B); gena with strong tranvesal striae (Fig. 19D); length of rami of Fl2 long, 1.6x head height (Fig. 19A); frenal processes long, 2.3‒2.4x as long as length of axillae and mesoscutellar disc with base black and apical half reddish orange; Gt1 orange with some brown spots.

Description. Male. Body length 5.3 mm; length of mesosoma excluding the frenal processes 2.2‒2.3 mm. Head, mesosoma, coxae and petiole black; frenal processes with base black and apical half reddish orange; scapes yellow, pedicels, flagellomeres and rami brown; legs yellow; wings slightly brown infuscate, venation brown; Gt1 orange with some brown spots. Head. 1.4‒1.5x as broad as high, with sparse setae, more prominent on upper frons and vertex (Fig. 19B). Scrobal depression with irregular striations and punctations. Frons with oblique wide striae from the inferior margin of torulus to the lateral ocelli; lower face with transversal striae (Fig. 19B). Eyes separated by 2.3‒2.6x their height. Malar space 1.1‒1.4x eye height. Gena with strong tranvesal striae. Supraclypeal area and clypeus smooth to even faint striated; anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 1.5‒2.1x longer than broad, 0.2‒0.3x head height; flagellum 10 segmented, transversal, each flagellomere with a single long rami; length of rami of Fl2 1.6x head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, 1.5‒1.7x as high as broad. Midlobe elevated with transversal carina and slightly bilobed in frontal view; lateral lobes elevated to approximately the height of midlobe and with faint longitudinal striae. Axilla and mesoscutellar disc longitudinal striated. Mesoscutellar disc 1.0‒1.1x as long as axilla with medial apex raised 0.8‒1.0x height of frenal processes in lateral view; medial apex carinate with a shallow depression on dorsal margin. Axillula longitudinal striated. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with 88

longitudinal striae (Fig. 19D); lower mesepimeron slightly reticulate; femoral depression crenulate. Callus carinate and pilose. Metapleuron carinate and slightly reticulate. Propodeal disc flat and broad, with shallow depression on dorsal margin. Frenal processes cylindrical and long, 3.1‒3.6x longer than the basal width of mesoscutellum, 2.3‒2.4x as long as length of axillae and mesoscutellar disc, weakly curved in lateral and dorsal view (Figs. 19A, C), frenal processes with longitudinal striae; apex excavated, with ventral region like a flange-shape more elongated (Fig. 19C). Forewing 2.2‒2.6x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.2‒1.3x longer than marginal vein; stigmal vein rectangular, 1.4x longer than broad and postmarginal vein long. Hindwing fringed. Metacoxa semiglobose with a medium line of setae, 1.6‒1.7x longer than broad. Metafemur 6.4‒7.2x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical and finely striated, 6.7‒8.0x longer than broad, 2.4‒2.5x as long as metacoxa. Tergal scar on Gt1 absent.

Female. Unknown.

Etymology. The specific name comes from Latim, genis = cheeks, striata = striae, referring to the strong sculpture on genae.

Biological notes. Host unknown.

Distribution. Brazil (Fig. 19E).

Material examined. Holotype. BRAZIL, Bahia: Andarai, Mata Carrasco, 12°48'25"S 41°19'51"W, 13-14.xii.1990, S.T.P. Amarante, [1♂, MZUSP, n 07570]. Paratype (1♂). Mato Grosso: Chapada dos Guimarães, Colégio Agr. Buriti, 15°20'43"S 55°53'50"W, 8-13.ii.1986, Armadilha Malaise, I.S. Gorayeb, [1♂, MPEG].

Kapala gracilispina Schoeninger n. sp. (Fig. 20 A‒E)

Diagnose. Recognized by a combination of the following characters: fine longitudinal striae on the frons and lower face smooth to even transversal striated (Fig. 20B); length of rami of Fl2 0.9‒1.1x head height; mesoscutellar disc and axilla with a medial longitudinal depression (Fig. 20C); frenal processes very thin with apex rounded (Figs. 20A, C).

Description. Male. Body length 2.7‒3.2 mm; length of mesosoma excluding the frenal processes 1.5‒1.6 mm (Fig. 20A). Head, mesosoma, coxae, petiole and frenal processes black; scapes brown, pedicels, flagellomeres and rami light brown; legs yellow; wings hyaline, venation light brown; Gt1 black. Head. 1.5x as broad as high, with sparse setae, more prominent on frons and vertex (Fig. 20B). Scrobal depression punctate. Frons with fine longitudinal striae; lower face smooth to even transversal striated (Fig. 20B). Eyes separated by 2.0‒2.5x their height. Malar space 1.0‒1.2x eye height. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2 or 3/3. Antenna with 12 segments; scape 2.0‒3.0x longer than broad, 0.2‒0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 0.9‒1.1x head height and 0.9‒1.0x as long as following branch. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view (Fig. 20D), 1.6‒1.7x as high as broad. Midlobe with transversal carina in frontal view; lateral lobes with faint oblique striae and elevated to approximately the height of midlobe. Axilla and mesoscutellar disc longitudinal striated and with a depression crossing them longitudinally in the middle (Fig. 20C). Mesoscutellar disc 1.0‒1.3x as long as axilla with medial apex raised 1.0‒1.5x height of frenal processes in lateral view. Axillula smooth to even faint longitudinal striated. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth except for some longitudinal striae (Fig. 20D); lower mesepimeron with shallow punctate; femoral depression crenulate. Callus carinate and covered by semi-erect setae. Metapleuron with shallow punctate. Propodeum flat, broad and smooth, except by the middle punctate. Frenal processes cylindrical and thin, 2.5‒2.8x longer than the basal width of 90

mesoscutellum, 1.3‒1.6x as long as length of axillae and mesoscutellar disc, curved in lateral and dorsal view, frenal processes with faint irregular longitudinal striae, giving a serrated appearance; apex rounded. Forewing 2.4‒2.5x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.2‒1.3x longer than marginal vein; stigmal vein subrectangular 1.0‒1.7x longer than broad; postmarginal vein absent. Hindwing fringed. Metacoxa 1.5‒1.6x longer than broad, semiglobose with a medium line of setae. Metafemur 5.1‒5.7x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical and reticulate, 5.0‒6.4x longer than broad and 2.1‒2.4x as long as metacoxa. Gt1 smooth and tergal scar present or absent.

Female. Unknown.

Comments. Specimens from Piauí (Brazil) have the escapes black and pedicels, flagellomeres and rami dark-brown. In addition, they also have the lower face with well-marked transverse striae, which extend over the supraclipeal and clypeus area.

Etymology. The specific name comes from Latin, “gracilis” = thin and “spinae” = spine, referring to frenal process thin.

Biological notes. Host unknown.

Distribution. Brazil (Fig. 20E).

Type material. Holotype. BRAZIL, Rio Grande do Norte: Mossoró - Fazenda Santa Júlia, 05°01'10''S, 37°22'56''W, 06.iii.2007, Arm. Malaise, D.R.R. Fernandes e eq., [1♂, INPA]. Paratypes (8♂). Idem, [1♂, INPA]. Maranhão: Carolina, P.N. Chapada Mesas, Cachoeira da Prata, 7°19'S, 47°20'06''W 25-30.vi.2009, Armadilha Malaise, A.L. Costa, M.B. Aguiar-Neto, P.A. Moraes & cols., [1♂, CZMA]. Piauí: Caracol, PARNA Serra das Confusões, Cores da Caatinga, 09°12'48.9''S, 43°27'59.9''W, 1-10.vii.2014, 705 m, Armadilha Suspensa, J.A. Rafael, F. Limeira-de-Oliveira, T.L. Rocha & G.A. Reis, [1♂, CZMA]. Idem, but 10-20.v.2014, [1♂, 91

CZMA]. Idem, but Riacho dos Bois, 09°13'11.9''S, 43°29'26.2''W, 15-31.vii.2013, 575 m, Armadilha de Malaise, J.A. Rafael, F. Limeira-de-Oliveira & T.T.A. Silva, [1♂, CZMA]. Guaribas, Parque Nacional Serra das Confusões, Andorinha, 09°08'27.8''S, 43°33'42.1''W, 20- 31.viii.2013, 515 m, Armadilha de Malaise, J.A. Rafael, F. Limeira-de-Oliveira & T.T.A. Silva, [1♂, CZMA]. 10 Km N Corrente, Fazenda Maracujá, 23-27.xi.1991, Armadilha de Malaise, S.T.P. Amarante & C.F. Martins, [1♂, MZUSP].

Kapala haplospinosa Schoeninger n. sp. (Fig. 21 A‒E)

Diagnose. Recognized by a combination of the following characters: frenal processes soft, light- brown to even reddish-brown (Figs. 21A, C); its apex excavated with ventral region like a flange- shape more elongated (Fig. 21C); pattern of bifurcate striae, giving a rugose appearance on frons (Fig. 21B); medial apex of mesoscutellar disc raised 1.7‒2.0x height of frenal processes and carinate (Fig. 21A). Females with flagellum 8 or 9 segmented and 0.9‒1.0x head height (Fig. 21D).

Description. Male. Body length 2.9‒3.4 mm; length of mesosoma excluding the frenal processes 1.6‒1.8 mm. Head, mesosoma, coxae and petiole black; frenal processes light-brown to even reddish-brown; scapes yellow to even brown, pedicels, flagellomeres and rami brown; legs yellow, except the base of the femora brown; wings hyaline, venation brown; Gt1 light brown with a dark-brown central spot. Head. 1.4‒1.5x as broad as high, with sparse long setae, more prominent on frons and vertex. Scrobal depression with irregular striae. Frons with wide longitudinal striae, some of them bifurcate giving a rugose appearance (anastomosed) (Fig. 21B); lower face transversally striated. Eyes separated by 2.0‒2.3x their height. Malar space 0.8‒1.0x eye height. Supraclypeal area striated. Clypeus smooth. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape 2.3‒2.6x longer than broad, 0.2x head height; flagellum 10 segmented, transversal, each flagellomere with a single long rami; length of rami of Fl2 0.9‒1.2x head height and 0.9‒1.0x as long as following rami. 92

Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.3‒1.4x as high as broad. Midlobe with transversal and longitudinal carina in frontal view; lateral lobes with longitudinal striae. Axilla and mesoscutellar disc longitudinal striated. Mesocutellar disc swollen in lateral view, 1.4‒1.7x as long as axilla with medial apex raised 1.7‒2.0x height of frenal processes in lateral view; medial apex carinate. Axillula longitudinal striated. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with longitudinal striae; lower mesepimeron rugose; femoral depression crenulate. Callus carinate and covered by semi-erect setae. Metapleuron punctate. Propodeum flat, broad and reticulate. Frenal processes cylindrical, 2.1‒2.4x longer than the basal width of mesoscutellum, 1.5‒1.7x as long as length of axilla and mesoscutellar disc, straight in lateral view and, slightly curved in dorsal view; frenal processes with longitudinal irregular striae, giving a serrated appearance (Fig. 21A); apex of frenal processes excavated, with ventral region like a flange-shape more elongated (Fig. 21C). Forewing 2.5‒2.6x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.2‒1.3x longer than marginal vein; stigmal vein rectangular 1.7‒2.0x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa 1.6‒2.1x longer than broad, semiglobose with a medium line of long setae. Metafemur 5.7‒6.6x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical, smooth to even reticulate, 5.7‒7.5x longer than broad and

2.1‒2.7x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 3.1‒3.6 mm; length of mesosoma excluding the frenal processes 1.6‒2.3 mm. Similar to male except for the following: Supraclypeal area smooth to striated. Maxillary/labial palpi 3/3. Antenna with 10 and 11 segments (Fig. 21D); scape 3.0‒3.3x longer than broad and 0.3‒0.4x head height; flagellum 8‒9 segmented, cylindrical and slightly serrated, length of flagellum 0.9‒1.0x head height; Fl2 1.6‒1.7x as long as apical width and 1.0‒1.2x as long as following flagellomere. Medial apex of mesoscutellar disc raised 1.3x height of frenal process in lateral view. Metafemur 6.6‒7.4x longer than broad. Petiole 2.7‒3.4x longer than broad and 1.1‒1.3x as long as metacoxa.

Comments. Some specimens have femora entirely brown. 93

Etymology. The specific name comes from Geek, haplos = simple, referring to frenal process unique within the genus.

Biological notes. Host unknown.

Distribution. Cuba, Dominican Republic, Haiti, Saint Kitts, United States of America (Florida) (Fig. 21E).

Type material. Holotype. USA: Florida: Monroe Co., Big Pine Key, SW 1/4, S4, 0m, 24°41'26"N, 81°22'13"W, 1-30.ix.1985, S&J Peck, mangrove-hardwood transition, Malaise, [1♂, CNC: UCRCENT00300634]. Paratypes (4♀, 1♂). DOMINICAN REPUBLIC: Hato Mayor: Los Haitises P.N., 18°45'51"N, 69°15'25"W, M. Ivie, [1♀, MAIC: UCRCENT00172268]. Prov. Pedernales: Sierra Bahoruco, Alcoa Rd., Km 23-26, 18°29′N 71°25′W, 530-750 m, 14.vii.1990, L. Masner, [1♀, UCRC]. Idem, but 23 Km N Cabo Rojo, 540 m, vii.1990, L. Masner, [1♀, UCRC]. San Juan: Par. Nac. Armando Bermudez, La Compartacion, 2380m, 19°02'00"N, 70°58'00"W, 7-9.vii.1992, M. A. & R. O. Ivie, [1♀, MAIC: UCRCENT00242270]. St. KITTS: Otleys Plantation, 28.x-5.xi.1991. Earl Thomas, malaise trap, [1♂, FSCA].

Additional material examined: CUBA: Upper Yara Valley, 20°16'30"N, 76°57'15"W, 18-28.x, L.C. Scaramuzza, [1♀, MCZC: UCRCENT00172270]. DOMINICAN REPUBLIC: Carbonera Monte, 19°51'02"N, 71°38'43"W, 24.vi.1978, R.O. Schuster & R.S. Rominger, [1♀, UCDC: UCRCENT00172271]. D.N., St. Domingo 7-8 WI, 18°28'44"N, 69°53'26"W, Aug. Busck [1♀, USNM: UCRCENT00172265]. San Cristobal, 4 km NW Villa Altagracia, 300m, 18°41'46"N, 70°11'38"W, 12.iv-6.vii.1992, FIT, M. A. Ivie & D. S. Sikes, [1♀, MAIC: UCRCENT00242269]. W.I.: Duarte, 20 km NE San Francisco de Macoris; Loma Quite Espuela, 300m, 19°20'31"N, 70°12'15"W, vii.1991, malaise trap, [1♀, CNC: UCRCENT00172251]. GRAND CAYMAN ISLANDS: Midland Acres, 0-10m, 19°18'00"N, 81°13'00"W, 16-19.viii.1985, Malaise, R. R. Askew, [UCRC: UCRCENT00302314]. HAITI: -, 18°58'07"N, 72°17'16"W, [1♀, AMNH: UCRCENT00172263]. Camp Perrin nr 1000 ft., 18°19'36"N, 73°51'52"W, 8-27.x.1934, [1♀, MCZC: UCRCENT00172269]. Port-au Prince & vic Haiti, 18°32'20"N, 72°19'43"W, 2.x.1934, 94

Darlington, [1♀, MCZ: UCRCENT00172264]. St. KITTS: Basseterre, 17°17'48"N, 62°42'49"W, 13.iv.1985, [1♀, UCRC: UCRCENT00172266]. USA: Florida: Dade Co., Miami 13603Old Culter Rd., 25°38'41"N, 80°18'08"W, 8.i.1979, yellow pan trap, L.A. Stange, [1♀, FSCA: UCRCENT00172262]. Monroe Co., 29°04'46"N, 81°24'17"W, [1♀, FSCA: UCRCENT00172261]. Big Pine Key, 24°41'32"N, 81°22'22"W, 13.iv.1971, roadside weeds, sweep, W.H. Pierce, [1♀, FSCA: UCRCENT00172260]. Idem, but SW 1/4 S4, 0m, 24°40'12"N, 81°21'14"W, 1-31.viii.1986, mangrove/hardwood transition, malaise trap, S. & J. Peck, [1♀, UCRC: UCRCENT00172257]. Big torch key, 24°43'19"N, 81°26'29"W, 4.ix.1972, Hepner, Conocarpus erecta [1♀, FSCA: UCRCENT00172258]. Cudjoe Key, 1/4 mi S S20, 24°39'53"N, 81°30'21"W, 4.iii-29.iv.1985, hammock forest, malaise & FIT, S & J Peck, [1♀, CNC: UCRCENT00172259]. Middle Torch Key, 0m, 24°41'06"N, 81°24'29"W, 1-30.iii.1986, hammock forest edge, Malaise, S & J Peck, [1♀, CNC: UCRCENT00300636]. N. Big Pine Key, 0m, 24°41'32"N, 81°22'22"W, 1.vii-17.xi.1985, palm woodland, malaise trap, S. & J. Peck, [1♂, UCRC: UCRCENT00092028].

Kapala jalisca Schoeninger n. sp. (Fig. 22 A‒E)

Diagnose. Recognized by a combination of the following characters: mesoscutellar disc swollen and rounded (Figs. 22A, D); medial apex of mesoscutellar disc not elevated above base of frenal processes (Fig. 22D, indicated); frenal process short and thin, which has a U-shape in dorsal view (Fig. 22C); frenal process oblique striated with the apex rounded.

Description. Male. Body length 2.2‒3.2 mm; length of mesosoma excluding the frenal processes 1.3‒1.9 mm. Head, mesosoma, coxae, petiole and frenal processes black with metallic- green reflections; antennae and legs yellow to light-brown; wings hyaline, venation brown; Gt1 dark-brown with apex light-brown, subsequent tergites light-brown. Head. 1.4‒1.5x as broad as high, with sparse setae, more prominent on frons (Fig. 22B). Scrobal depression faint striated. Frons with wide longitudinal striae, which curved toward the lower margin of the eye (Fig. 22B); lower face transversal striated. Eyes separated by 2.0‒2.2x 95

their height. Malar space 0.9‒1.0x eye height. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape 2.2‒2.5x longer than broad, 0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 1.2‒1.4x head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.3‒1.4x as high as broad. Midlobe with transversal and longitudinal carina in frontal view, giving a rough appearance (Figs. 22B, C); lateral lobes with faint oblique striae and elevated to approximately the height of midlobe. Axilla and mesoscutellar disc longitudinal striated. Mesocutellar disc swollen and rounded in lateral view (Figs. 22A, D), 1.6‒1.8x as long as axilla with medial apex not elevated above base of frenal processes (Fig. 22D). Axillula longitudinal striate. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth except for some longitudinal striae; lower mesepimeron smooth; femoral depression crenulate. Callus carinate and covered by semi-erect setae. Metapleuron with shallow punctations. Propodeum flat, broad and with a central depression in the middle, deeper in the upper region, and delimited by carina. Frenal processes short and thin, 2.1‒2.3x longer than the basal width of mesoscutellum, 1.4‒1.5x as long as length of axilla and mesoscutellar disc; slightly curved in lateral view and U-shape in dorsal view; frenal processes oblique striated (Fig. 22C); apex of frenal processes rounded. Forewing 2.1‒2.3x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.1‒1.4x longer than marginal vein; stigmal vein rectangular 2.0‒2.3x longer than broad; postmarginal vein absent. Hindwing fringed. Metacoxa 1.5‒1.9x longer than broad, semiglobose with a medium line of setae. Metafemur 5.8‒6.0x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical, weakly rugose with two faint parallel carinae, 4.1‒4.8x longer than broad and 1.8‒2.0x as long as metacoxa. Gt1 smooth and tergal scar present or absent. Female. Unknown.

Etymology. The specific name refers to type locality of this species, Jalisco state, Mexico.

Biological notes. Host unknown.

Distribution. Mexico (Fig. 22E).

Type material. Holotype. MEXICO, Jalisco: Est. Biologica Chamela, 19º30'N, 105º03'W, 4.vii.2012, REO, EJE Central, Cham 197 [1♂, UNAM]. Paratypes (10♂). Idem, [5♂, UNAM]. Idem, but 3.vii.2016, Cham 194, [3♂, UNAM]. Chamela, Biostation, Camino Calanaria, 19º30'N, 105º03'W, YPT (90), Cham 34, [1♂, UCRC]. Idem [without locallity and date of collection], 19°26′N 99°08′W Gomez Farias, G. Farias Tamps, 112, [♂, UCRC: UCRCENT 320098].

Kapala spinaepplanata Schoeninger n. sp. (Fig. 23 A‒G)

Diagnose. Recognize by a combination of the following characters: frenal processes flattened laterally with longitudinal striae and apex straight (Fig. 23C); frons with a pattern of wide longitudinal striae, curving to inferior margin of torulus and smooth to even transversal striated on the lower face (Fig. 23B).

Description. Male. Body length 3.3 mm; length of mesosoma excluding the frenal processes 1.8 mm. Head, mesosoma, coxae, petiole and frenal processes black; scapes, pedicels, flagellomeres and rami brown; legs light brown; wings hyaline, venation brown; Gt1 black with apex dark-brown. Head. 1.5x as broad as high, densely pilose (Fig. 23B). Scrobal depression striated. Frons with some wide longitudinal striae, which curve to inferior margin of torulus (Fig. 23B); lower face smooth to even transversal striated. Eyes separated by 2.1x their height. Malar space 1.0x eye height. Supraclypeal area and clypeus smooth. Maxillary/labial palpi 3/2. Antenna with 12 segments; scape 2.0x longer than broad, 0.2x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami of Fl2 1.2x head height and 1.0x as long as following rami. Mesosoma. Mesoscutum dorsal half abruptly curved anteriorly, in lateral view, and 1.4x as high as broad. Midlobe with transversal and longitudinal carina in frontal view; lateral lobes smooth to even faint oblique striae and elevated to approximately the height of midlobe. Axilla 97

and mesoscutellar disc longitudinal striated. Mesoscutellar disc 1.1x as long as axilla with medial apex raised 0.7x height of frenal processes in lateral view. Axillula longitudinally striated. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth except for some longitudinal striae; lower mesepimeron smooth; femoral depression weakly crenulate. Callus carinate and covered by semi-erect setae. Metapleuron smooth. Propodeum flat, broad and reticulate. Frenal processes flattened laterally (Fig. 23C, A), 2.4x longer than the basal width of mesoscutellum, 1.6x as long as length of axilla and mesoscutellar disc, slightly curved in lateral view and straight in dorsal view; frenal processes with longitudinal striae (Fig. 23E); apex of frenal processes straight. Forewing 2.2x longer than broad covered by small bristles, except for the base bare; submarginal vein 1.0x longer than marginal vein; stigmal vein rectangular 1.5x longer than broad; postmarginal vein short. Hindwing fringed. Metacoxa 1.7x longer than broad, semiglobose with a medium line of long setae. Metafemur 6.7x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical, smooth with two parallel carinae at base, 5.7‒7.5x longer than broad and 2.1‒2.7x as long as metacoxa. Gt1 smooth and tergal scar absent.

Female. Body length 3.2 mm; length of mesosoma excluding the frenal processes 1.8 mm. Similar to male except for the following: scape yellow; antenna with 10 segments; scape 2.7‒3.7x longer than broad and 0.3x head height; flagellum 8 segmented (Fig. 21D), cylindrical and slightly serrated, length of flagellum 0.9‒1.0x head height; Fl2 2.0‒2.3x as long as apical width and 1.2‒1.4x as long as following flagellomere. Petiole 3.0‒3.8x longer than broad and 1.3‒1.4x as long as metacoxa.

Etymology. The specific name comes from Latim, spinae = spine, applanata = flat, referring to frenal process flat laterally.

Biological notes. Host unknown.

Distribution. Colombia (Fig. 23G).

Holotype: COLOMBIA: Dept. of Valle: Anchicaya 80km EI Cauchal, 18.viii.1979, malaise, J. Monsalve, [1♂, EMEC: UCRCENT00236090]. Paratypes (2♀): Valle del Cauca: Central de Anchicaya 30 km. E. Buenaventura, 560m, 3°52'56"N, 76°48'02"W, 14-16.vii.1975, tropical very wet forest, MT, R.C. Wilkerson, [1♀, FSCA: UCRCENT00318597]. Valle Rio Anchicaya, 400m, 10.ii.1977, Breed & C.D. Michener [1♀, UCRC: UCRCENT00446413].

Species with uncertain position

Kapala dicerodera (Spinola, 1853)

Eucharis dicerodera Spinola, 1853: 43. Type data: Brazil: Pará. Holotype ♂, by monotypy. Type depository: Museo di Zoologia dell'Università di Torino (MRSN). Description of male. Additional citation: Dalla Torre, 1898: 360 (catalog). Kapala dicerodera; Ashmead, 1904a: 473. On page 467, Ashmead refers to the original citation of Eucharis dicerodera as questionably belonging to Kapala, but differently on page 473 refers to K. dicerodera.

Distribution. Neotropical: Brazil [Pará]

Comments. The description by Spinola (1853) is not clear about the characters that, in fact, could separate K. dicerodera from the other species of Kapala and establish the correct position within the genus. The description is comprehensive and, in which could be insert several species of the genus, being impossible its identification. In addition, many collections were carried out in the state of Pará (type locality of the species), yet nothing that could fit the species described by Spinola. Another problem is that the type species is lost, completely impossible to determine the identity of K. dicerodera.

Kapala confusa Schoeninger n. sp. (Figs. 24A‒E)

Diagnosis. Recognized by a combination of the following characters: frons not swollen and with faint longitudinal striae (Fig. 24B), supraclypeal area with transversal striae (Fig. 24B), mesosoma with fine longitudinal striae (Fig. 24C), medial apex of mesoscutellar disc elevated like a keel-shape (Fig. 24D, indicated) and wings brown infuscate.

Description. Male. Body length 2.3‒3.5 mm; length of mesosoma excluding the frenal processes 1.1‒1.5 mm. Head, mesosoma, coxae, petiole and frenal processes black; scapes, pedicels and flagellomeres yellow, rami brown; legs yellow; wings brown infuscate, venation brown; Gt1 dark-brown. Head. 1.4‒1.5x as broad as high, smooth surface with sparse setae, more prominent on upper frons and vertex. Frons flat and with fine longitudinal striae (Fig. 24B). Scrobal depression striated. Eyes separated by 2.0‒2.2x their height. Malar space 1.0x eye height, with oblique fine striae. Supraclypeal area with transversal striae, defined by weakly impressed sulci and slightly swollen (Fig. 24B). Clypeus smooth. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 2.7‒3.0x longer than broad, 0.3x head height; flagellum 10 segmented, transversal, each flagellomere with a single long rami; length of rami of Fl2 1.1‒1.2 head height and 0.9‒1.0x as long as following rami. Mesosoma. Mesoscutum anteriorly smoothly curved, in lateral view, and 1.4x as high as broad. Midlobe with transverse striae and lateral lobes with fine transversal striae (Fig. 24C). Axilla and mesoscutellar disc with fine longitudinal striae. Mesoscutellar disc 1.3‒1.5x as long as axilla with medial apex raised 1.0‒2.0x height of frenal processes in lateral view, medial apex elevated like a keel-shape (Fig. 24D). Axillula with fine longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with weakly striae and some shallow and inconspicuous punctures in upper region, anterior region of mesepisternum with a shallow sulcus; femoral depression crenulate. Callus carinate, colliculate and pilose. Metapleuron carinate and colliculate. Propodeum flat, broad and delimited by carina, surface colliculate with some longitudinal carina in the upper region, superficial depression in the 100

upper middle region. Frenal processes cylindrical and very thin (Fig. 24C) 3.4‒3.7x longer than the basal width of mesoscutellum, 1.5‒1.8x as long as length of axillae and mesoscutellar disc, arched in lateral view and convex in dorsal view; frenal processes with longitudinal striae and with apex acuminate. Forewing 2.4‒2.6x longer broad; covered by small bristles, except for the base bare; submarginal vein 1.0‒1.2x longer than marginal vein; stigmal vein rectangular, 1.2‒1.3x longer than broad and postmarginal vein indistinct. Hindwing fringed, 4 hamulli curved hook-shaped. Metacoxa semiglobose with a medium line of setae, 2.0‒2.2x longer than broad. Metafemur 6.0‒7.2x longer than broad, weakly striated with sparse setae. Metasoma. Petiole cylindrical and colliculate, 6.3‒7.5x longer than broad, 2.1‒2.7x as long as metacoxa. Tergal scar on Gt1 present or absent.

Female. Unknown.

Comments. The specimen from Peru has some variations compared to the specimens from Ecuador, flagellomeres and branches yellow (vs. brown), supraclypeal smooth (vs. striated) and medial apex of the scutellar disc not so high. These specimens contain characteristics that cover the three species complexes proposed so far for the genus Kapala and therefore, we opted in the present work not to insert it in any complex until morphological and molecular phylogenetic studies provided more information.

Etymology. The specific name refers to specimen that has characteristics of the three species complexes.

Biological notes. Host unknown.

Distribution. Ecuador and Peru (Fig. 24E). Material examined. Holotype: ECUADOR: Napo: Cosanga, Yanayacu Biological Station, 02- 08.iv.2015, 0°36’4.09’’S/77°53’19.15’’W, 2.208m, Arm. Varredura, A. Plant & J. Camara, [1♂, INPA]. Paratypes (2♂): ECUADOR: Napo: Idem, PERU: Valle Chanchamayo, 1600m 1.v.1939, C. Weyrauch, [1♂, UCRC_ENT 00433059, IFML]. 101

Acknowledgment

We thank of granting the scholarship from CNPQ (National Counsel of Technological and Scientific Development) and CAPES (Brazilian Federal Agency for Support and Evaluation of Graduate Education) for the PDSE scholarship / visiting student at University California - Riverside (process PDSE - 88881.133943/2016-01). MLO is scholarship of CNPq, Brazil (306100/2016–9).

References

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Heraty, J.M. & Woolley, J.B. (1993) Separate species or polimorphism; a recurring problem in Kapala (Hymenoptera: Eucharitidae). Annals of the Entomological Society of America, 85: 517-531. Howard, R.W., Pérez-Lachaud, G., & LACHAUD J.P. (2001) Cuticular Hydrocarbons of Kapala sulcifacies (Hymenoptera: Eucharitidae) and its host, the Ponerine ant Ectatomma ruidum (Hymenoptera: Formicidae). Annals of the Entomological Society of America, 94(5): 707- 716. Kirby, W.F. (1886) A synopsis of the genera of the Chalcididae, subfamily Eucharinae, with descriptions of several new genera and species of Chalcididae and Tenthredinidae. Journal of the Linnean Society, 20: 28–37. Lachaud, J.P. & Pérez-Lachaud, G. (2009) Impact of natural parasitism by two eucharitid wasps on a potential biocontrol agent ant in southeastern Mexico. Biol. Control, 48: 92–99. Lachaud, J.P. & Pérez-Lachaud, G. (2012) Diversity of species and behavior of hymenopteran parasitoids of ants: A review. Psyche: A Journal of Entomology, 1-24. Latreille, P.A. (1825) Families naturelles de règne animla, exposées succintement et dans um order analytiques avec l’indication de leurs genres. Paris: J.B. Bailliere, 570 pp. Myers, J.C. (1931) Descriptions and records of parasitic Hymenoptera from British Guiana and the West Indies. Bulletin of Entomological Research, 22: 267–277. Murray, E.A. (2014) Systematics and evolution of Eucharitidae (Hymenoptera: Chalcidoidea), with a focus on the New World Kapala. University of California, Riverside, CA. Murray, E.A., Carmichael, A.E., & Heraty, J.M. (2013) Ancient host shifts followed by host conservatism in a group of ant parasitoids. Proceedings of the Royal Society of London B: Biological Sciences, 280 (1759), 20130495. Perez-Lachaud, G., Heraty, J., Carmichael, A. & Lachaud, J. (2006). Biology and Behavior of Kapala (Hymenoptera: Eucharitidae) attaking Ectatomma, Gnamptogenys, and Pachycondyla (Formicidae: Ectatomminae and Ponerinae) in Chiapas, Mexico. Annals of the Entomological Society of America, 99(3): 567-576. Risbec, J. (1954) Chalcidoïdes et Proctotrupoïdes de l'Afrique occid l'Institut Français d'Afrique Noire (A), 16, 1035-1092. Shulz, W.A. (1905) Strandgut. Spolia Hymenopterologica, 1905: 77–269. 104

Shorthouse, D.P. (2010) SimpleMappr, an Online Tool to Produce Publication-Quality Point Maps [WWW document]. URL http://www.simplemappr.net [accessed on 26 December 2015]. Spinola, M. (1853) Compte Rendu des Hyménoptères inédits provenants du voyage entomologique de Mr. Ghiliani dans le para en 1846. Memorie della Reale Accademia delle Scienze de Torino, serie secunda 13: 19–94. Torréns, J. (2013) A review of the biology of Eucharitidae (Hymenoptera: Chalcidoidea) from Argentina. Psyche: A Journal of Entomology, 2013: 1-14. Walker, F. (1839) Monographia Chalciditum. Vol. 2. London. 100 pp. Walker, F. (1862) Notes on chalcidites and characters of undescribed species. Transactions of the Entomological Society of London, 1: 345–397. Walker, F. (1871) Notes on Chalcidiae. Part IV. Chalcididae, Leucospididae, Agaonidae, Eucharidae, Perilampidae, Ormyridae, Encyrtidae. London, 55–70. Walker, F. (1872) Economy of Chalcididiae (continued). The Entomologist, 6: 65–70. Wheeler, W.M. (1907) The polymorphism of ants with an account of some singular abnormalities due to parasitism. Bulletin of the American Museum of Natural History, 23: 1–100. Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. http://dx.doi.org/10.1371/journal.pone.0015991

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Figure 1. Kapala spp. A. head ♀ (frontal view); antenna ♀ (lateral view); C. antenna ♂ (lateral view).

106

Figure 2. Kapala flabellata ♂. D. mesosoma (lateal view); E. mesosoma (dorsal view); F. habitus; G. forewing.

107

Figure 3. Specimen measurements. A. Head (frontal view); B. antenna (lateral, ♂); C. antenna (lateral, ♀); D. habitus (lateral, ♀; E. habitus (dorsal, ♀); F. Forewing. 108

Figure 4. K. cuprea: A. Habitus (lateral, ♂); B. head (front, ♂); C. antenna (lateral, ♀); D. Habitus (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution. 109

Figure 5. K. furcata: A. habitus (lateral, ♀); B. front (head, ♀); C, antenna (lateral, ♀); D. front (head, ♂); E. habitus (lateral, ♂); G. habitus (dorsal, ♂); H. distribution.

110

Figure 6. K. splendens: A. habitus (lateral, ♀); B. antenna (lateral, ♀); C. head (frontal, ♀); D. habitus (dorsal, ♀); E. habitus (lateral, ♂); F. distribution. 111

Figure 7. K. iridicolor: A. habitus (lateral, ♀); B. head (front, ♀); C. antenna (lateral, ♀); D. habitus (lateral, ♂); E. habitus (dorsal, ♀); F. habitus (dorsal, ♂). G. distribution. Red setae = indicating the mesonotum smoothly curved. 112

Figure 8. K. argentina: A. habitus (lateral, ♀); B. head (front, ♀); C. mesosoma (lateral, ♀); D. head (front, ♂); E. habitus (lateral, ♂); F. habitus (dorsal, ♀); G. habitus (dorsal, ♂); H. distribution.

113

Figure 9. K. chacoensis: A. habitus (lateral, ♂); B. head (front, ♂); C. head (front, ♀); D. mesoscutum (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution.

114

Figure 10. K. cynipsea: A. habitus (lateral, ♀); B. head (front, ♀); C. distribution.

115

Figure 11. K. flabellata: A. habitus (lateral, ♂); B. head (front, ♂); C. head (front, ♀); D. mesosoma (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution. 116

Figure 12. K. floridana: A. habitus (lateral, ♂); head (front, ♂); C. mesosoma (lateral, ♂); D. antenna and mesosoma (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution. Red setae = Mesoscutellar disc not elevated above base of frenal process.

117

Figure 13. K. inexagens: A. habitus (lateral, ♀); B. head (front, ♀); C. habitus (dorsal, ♀); D. antenna (lateral, ♀); E. distribution. 118

Figure 14. K. ivorensis: A, habitus (lateral, ♀); B. head (front, ♀); C. antenna and mesosoma (lateral, ♀); D. habitus (lateral, ♂); E. habitus (dorsal, ♀); F. habitus (dorsal, ♂); G. distribution. 119

Figure 15. K. izapa: A. habitus (lateral, ♂); B. head (frontal, ♂); C. mesosoma (lateral, ♂); D. habitus (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution.

120

Figure 16. K. romandii: A. habitus (lateral, ♂); B. head (front, ♂); C. antenna (lateral, ♀); D. head (frontal, ♀); E. habitus (dorsal, ♀); F. mesosoma (lateral, ♂); G. habitus (dorsal, ♂); H. distribution.

121

Figure 17. K. terminalis: A. habitus (lateral, ♂); B. head (front, ♂); C. mesosoma (lateral, ♂); D. antenna (lateral, ♀); E. habitus (lateral, ♀); F. habitus (dorsal, ♂); G. habitus (dorsal, ♀); H. distribution. Red setae = mesoscutellar disc not elevated above base off renal process.

122

Figure 18. K. corcovata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsolateral, ♂); D. mesosoma (front, ♀); E. habitus (lateral, ♀); F. distribution. 123

Figure 19. K. genistriata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma (lateral, ♂); E. distribution. 124

Figure 20. K. gracilispina n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma, ♂); E. distribution. 125

Figure 21. K. haplospinosa n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. head and mesosoma (laterofrontal, ♀); E. distribution. 126

Figure 22. K. jalisca n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesosoma (lateral, ♂); E. distribution. 127

Figure 23. K. spinaepplanata n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. frenal processes (dorsal, ♂); D. mesosoma (lateral, ♀); E. habitus (dorsal, ♂); F. habitus (dorsal, ♀); G. distribution. 128

Figure 24. K. confusa n. sp.: A. habitus (lateral, ♂); B. head (front, ♂); C. habitus (dorsal, ♂); D. mesososma (lateral, ♂); E. distribution. Red setae = mesoscutellar disc elevated like a keel-shape. 129

Capítulo II

Schoeninger, K., Torréns, J., Heraty, J.M.,

Oliveira, M.L. (2018) Filogenia de Kapala

Cameron, 1884 (Hymenoptera: Eucharitidae)

com base em dados morfológicos e moleculares.

130

Filogenia de Kapala Cameron, 1884 (Hymenoptera: Eucharitidae) com base em dados morfológicos e moleculares

KARINE SCHOENINGER1,4, JAVIER TORRÉNS2, JOHN M. HERATY3 & MARCIO L. OLIVEIRA1

Resumo. Kapala Cameron é um dos gêneros mais diversos e especiosos, com espécies distribuídas na região Neotropical e apenas uma espécie disjunta com ocorrência na região Afrotropical. Um estudo filogenético com base em dados morfológicos e moleculares foi realizado com o objetivo de testar a monofilia de Kapala e as relações entre as suas espécies. Para este propósito, 61 caracteres relacionados à morfologia externa foram analisados. Para as análises moleculares, partes de cinco regiões de genes foram amplificadas: 18S, 28S-D2, 28S- D3-D5, COI-nj e COII. Como resultado, foi verificado em todas as análises, morfológica e molecular, que Kapala é parafilético, dividido em três complexos de espécies, que por sua vez, estão divididos por agrupamentos de gêneros distintos. A morfologia de Kapala é altamente conservada e pobre em caracteres informativos, resultando em várias espécies crípticas. Não há informações suficientes acerca de Kapala que permitam dividir o grupo, sendo necessária uma topologia estável e adição de novas informações.

Palavras-chave. Chalcidoidea, clado Kapala, Eucharitinae, Região Neotropical, parasitoides de formigas. 131

Abstract. Kapala Cameron is one of the most diverse and specious genera, with species distributed in the Neotropical region and only a disjunct species occurring in the Afrotropical region. A phylogenetic study based on morphological and molecular data was performed to clear the relationships between species and to test de monophyly of Kapala. For this purpose, 61 characters related to external morphology were analyzed. For the molecular analyzes parts of five gene regions were amplified: 18S, 28S-D2, 28S-D3-D5, COI-nj and COII. As a result, it was verified in all morphological and molecular analyzes that Kapala is paraphyletic, divided into three species complexes, which in turn are divided by groups of different genera. The Kapala morphology is highly conserved and poor in informative characters resulting in several cryptic species. There is not enough information about Kapala that allows dividing the group, requiring a stable topology and adding new information.

Keywords: ant parasitoids, Chalcidoidea, Eucharitinae, Kapala clade, Neotropical region.

Introdução

Todos os membros de Eucharitidae (Hymenoptera: Chalcidoidea) são parasitoides especializados de formigas (Clausen, 1940a, b; Heraty, 2002; Lachaud & Pérez-Lachaud, 2012). Os adultos depositam seus ovos internamente ou sobre o tecido vegetal, e a larva ativa do primeiro instar, denominada planídea, é responsável por ter acesso as formigas hospedeiras (Clausen, 1940; Heraty, 2002). Murray et al. (2013) propuseram que Eucharitidae se originou no Velho Mundo e colonizou o Novo Mundo através de múltiplos eventos de dispersão ocorrendo aproximadamente 20-43 mya. Dentro de Eucharitini (Eucharitinae), um grupo monofilético ataca formigas das subfamílias Ponerinae, Ectatommine e Myrmeciinae (clado PEM). O clado PEM é suportado por análises moleculares e algumas morfológicas (Heraty, 2002; Murray et al., 2013) e é composto por três grupos: clados Chalcura e Schizaspidia do Velho Mundo e o clado Kapala do Novo Mundo. O clado Kapala é um grupo de 13 gêneros, os quais possuem múltiplas modificações morfológicas bizarras. O formato da cabeça, morfologia da antena, padrões de esculturação do tegumento e morfologia dos processos frenais são particularmente variáveis dentro do clado. Os fenótipos extremos são melhor exemplificados pelos processos frenais emparelhados; esses 132

variam desde ligeiramente curvados e linearmente carenados (espécies de Kapala Cameron, 1884) até com carenas circulares a lisas (Lasiokapala Ashmead, 1899 e algumas espécies de Lirata Cameron, 1884); os processos frenais também podem ser dorsoventralmente achatados e estreitamente separados (Dicoelothorax Ashmead, 1899 e Dilocantha Shipp, 1894) até amplamente arqueados formando uma carapaça sobre o metassoma (Galearia Brullé, 1846 e Thoracantha Latreille, 1825). O dimorfismo sexual é principalmente confinado a diferenças na morfologia da antena, metassoma e diferenças na morfologia dos processos frenais, nos quais os processos dos machos são muito mais reduzidos e delgados em relação aos processos das fêmeas (Dicoelothorax, Galearia e Dilocantha). Uma outra modificação mais bizarra ocorre em Isomerala Shipp, 1894, em que os olhos são em forma de “mamilo” e a fronte possui protuberâncias inchadas. Enquanto a maioria dos gêneros do clado Kapala são definidos por características dos processos frenais, o gênero Kapala é reconhecido por sua morfologia relativamente simplificada, porém consistente (Murray, 2014). O gênero Kapala é extremamente comum com um total de 18 espécies válidas, as quais se estendem desde o sul do Estados Unidos ao norte da Argentina, com exceção do Chile (Heraty, 2002; Murray, 2014). Apenas uma espécie, Kapala ivorensis Risbec, possui ocorrência no Velho Mundo e está difundida na região Afrotropical e Madagascar (Heraty, 2002, Murray & Heraty, 2016). Segundo Heraty (2002), o gênero é definido por um conjunto de simplesiomorfias que incluem processos frenais lirifomes com carenas longitudinais ou em espiral (liso em Kapala chacoensis Gemignani, 1947), olhos nus, ramos antenais do macho maiores que a altura da cabeça, flagelômero basal da fêmea raramente mais que duas vezes mais longo que largo e margem posterior do disco mesoescutelar elevada a uma crista proeminente entre as bases dos processos frenais. De acordo com Heraty (2002), em uma análise filogenética que utilizou 100 caracteres morfológicos para os gêneros de Eucharitidae, foi sugerido que Kapala é monofilético, não existindo, porém, sinapomorfias que sustentem sua monofilia. Contudo, através da utilização de cinco marcadores moleculares (18S, 28S-D2, 28S-D3-D5, COI-nj e COII), foi proposto que Kapala é parafilético (Murray et al., 2013; Murray, 2014), sendo dividido em três complexos de espécies, os quais são separados filogeneticamente por dois agrupamentos de gêneros morfologicamente distintos e pelo gênero Isomerala (Murray, 2014.). 133

O desacordo entre os dois tipos de dados (morfológico e molecular) propostos até o momento para o gênero Kapala nos motivou a testar a sua monofilia, bem como verificar as relações filogenéticas entre as suas espécies com base em dados morfológicos e moleculares.

Material e métodos

Dados morfológicos

Escolha dos terminais

Quarenta e três espécies foram utilizadas na análise, sendo 25 de Kapala e 18 pertencentes a 15 gêneros de Eucharitini (Tabela S1). A escolha do grupo externo foi feita com base na filogenia proposta por Heraty (2002), na qual utilizou-se dados morfológicos e, também, mais recentemente, na filogenia de Murray et al., (2013), com dados moleculares. Foi dada preferência para a inclusão de espécies-tipo. As espécies foram representadas por machos e fêmeas e sempre que possível foi utilizado mais de um exemplar para as codificações dos caracteres.

Morfologia e análises dos caracteres

A morfologia de Kapala foi estudada a partir de exemplares alfinetados e secos, examinados em microscópio estereoscópio Nikon modelo SMZ645. A terminologia utilizada para a morfologia em termos gerais segue a proposta por Heraty (2002) e detalhes da esculturação corporal segue a proposta por Harris (1979) e Gibson et al. (1998). O flagelômero basal é contado como Fl2 de acordo com Heraty (2002); os segmentos funiculares incluem Fl2 e os segmentos seguintes, incluindo a clava. As espinhas emparelhadas do mesosoma surgem a partir do frenum e são referidas aqui como processos frenais (carácter 50; Heraty, 2002). Os padrões do tegumento facial compreendem um caráter importante para a definição de complexos de espécies dentro do gênero Kapala. A fronte e a face inferior são consideradas como áreas separadas, definidas como acima e abaixo do torulus. As estrias oblíquas da fronte podem se 134

estender do torulus para os ocelus laterais, seguindo a depressão escrobal antenal ou, inversamente, as estrias podem se estender do ocelo mediano para a gena, seguindo a margem do olho composto. Abaixo do torulus as estrias podem ser oblíquas ou podem ser transversais, seguindo horizontalmente a face inferior adjacente ao clípeo. As estrias dos processos frenais são consideradas longitudinais quando se prolongam além do comprimento do processo e oblíqua quando seguem uma direção espiral externa na metade apical. Para a construção da matriz de caracteres, foram utilizados alguns caracteres já presentes na literatura, como os propostos por Heraty (2002), Heraty & Woolley (1993) e Murray (2014); modificação de caracteres previamente propostos, bem como novos caracteres incorporados a partir de análise extensiva da morfologia dos espécimes. Sessenta e um caracteres foram codificados numa matriz no programa Mesquite® versão 3.2 (Maddison & Maddison, 2017) (SI 2). Durante a codificação, foi utilizado um ponto de interrogação (?) quando o estado era inaplicável e não podia ser codificado para um terminal, por ele não possuir a estrutura na qual o caráter ou o estado fosse definido. Já o símbolo de traço (-), foi usado para estados onde os dados são faltantes e não é possível ter acesso à informação filogenética. Apesar dos programas computacionais não distinguirem entre dados inaplicáveis e faltantes (Strong & Lipscomb, 1999) foi utilizada a distinção não somente para facilitar as futuras análises ou a adição de dados na matriz como proposto por Schuh & Brower (2009), mas também para ficar claro o motivo pelo qual não foi possível codificar determinado estado de caráter. Os caracteres polimórficos foram separados pelo “&” comercial (e.g. 0&1). Cabe ressaltar que, quando não foi possível determinar estados dos caracteres nas estruturas de alguns exemplares, eles foram codificados a partir da informação disponível na literatura. Nas análises, todos os caracteres foram tratados como não ordenados ou não aditivos, de modo que a mudança de um estado de caráter para outro tem o mesmo custo ou peso na análise e constitui a hipótese mais geral sobre a transformação de um estado de caráter (Schuh & Brower 2009). Já os caracteres não informativos foram mantidos e documentados, pois em uma análise futura eles podem ser potenciais sinapomorfias. A polarização dos caracteres foi a posteriori, determinada com enraizamento pelo método de comparação com grupos externos (Farris 1982; Nixon & Carpenter, 1993). 135

As análises filogenéticas por parcimônia foram conduzidas no programa TNT (Goloboff et. al., 2003; Goloboff et al., 2008). Para as análises tradicionais, foi utilizado o algoritmo de Wagner + TBR, com os seguintes parâmetros: randon seed = 10, réplicas = 100 e swapping algorithm = 1000. Para as análises, foi utilizada a pesagem implícita, onde o valor de concavidade (k) de cada caráter é obtido simultaneamente a cada reconstrução e a escolha da melhor árvore é obtida pelo maior valor de ajuste (fit) total, que é a soma do ajuste individual de cada caráter (Schuh & Brower, 2009). Portanto, na pesagem implícita o cladograma ideal não é árvore mais curta como na análise sem pesagem, mas sim a árvore que tem o maior valor de ajuste. Nas análises com pesagem implícita, a escolha do valor de k (7.1875) foi determinada pelo algoritmo setk.run (Arias et al., 2011). Valores de suporte de ramo foram baseados em 1000 réplicas de bootstrap no TNT. A visualização dos cladogramas e a otimização dos caracteres foram realizados no software Winclada 1.00.08 (Nixon, 2002). Os cladogramas foram editados em softwares de vetorização. Para avaliar os resultados das análises filogenéticas, foram usadas medidas como o número de passos (L), índice de consistência (CI) e retenção (RI).

Dados moleculares

A matriz molecular possui um total de 89 espécimes (Tabela S2), com 67 pertencentes à Kapala, representando 18 espécies (10 válidas e 8 novas); espécimes de 14 gêneros como grupo externo, sendo: 10 pertencentes ao clado Kapala mais dois gêneros novos, Torquata n. gen e Ecarinata n. gen. (descritos no Cap. V) e Chalcura nr. ramosa e Schizaspidia aenea, do Velho Mundo. Não foram obtidos dados moleculares para representantes de Parakapala e Liratella (clado Kapala), bem como para as seguintes espécies de Kapala: K. chacoensis, K. cynipsea, K. inexagens, K. splendens, K. confusa n. sp., K. corcovata n. sp., K. gracilispinae n. sp., K. genistriata n. sp., K. jalisca n. sp., K. longicornis n. sp. e K. spinaepplanata n. sp. Todos as sequências moleculares foram obtidas a partir de um número único “voucher” de DNA (D #), os quais estão depositados em um banco de dados interno em arquivos armazenados no FileMakerPro® mantido na UCR (Universidade da Califórnia – Riverside). Para as extrações 136

de DNA, foram utilizados espécimes frescos ou secos. Cabe ressaltar que todo processo de extração, amplificação e sequenciamento de DNA foi realizada pela esquipe do Laboratório de Sistemática de Hymenoptera da Universidade da Califórnia – Riverside (UCR). O DNA dos espécimes foi extraído de forma não destrutiva, utilizando o protocolo chelex-proteinase-K (Walsh et al., 1991) a fim de preservar a integridade das vespas parasitoides e sua retenção como espécimes “voucher”. Partes de cinco regiões de genes foram amplificadas: 18S, 28S-D2, 28S- D3-D5, COI-nj e COII (Tabela S3). Os reagentes e protocolos Qiagen (Valencia, CA) foram utilizados para a realização da PCR. O software Sequencer 5.4.6 (Gene Codes Corp, Ann Arbor, MI) foi utilizado para editar os cromatogramas em sequências finais; cabe ressaltar que os primers não foram incluídos na contagem final da sequência (Tabela S4).

Análises moleculares

Os genes individuais foram alinhados através do MAFFT (Katoh et al., 2005) pela interface do software Mesquite versão 3.2, utilizando as configurações-padrão. Para os genes nucleares ribossomais, foi aplicada a estratégia algorítmica E-INS-I e para os genes mitocondriais, a estratégia algorítmica G-INS-I. Para COI-nj e COII, todos os códons foram traduzidos para aminoácidos funcionais usando o código mitocondrial dos invertebrados. O SequenceMatrix 1.7.7 (Vaidya et al., 2011) foi utilizado para concatenar os genes para uma matriz final. Os genes ribossômicos e mitocondriais foram então concatenados para um comprimento de alinhamento final de 3034 pb. Para efeito de comparação, foi utilizado neste trabalho as mesmas análises e algoritmos utilizados por Murray (2014).

Parcimônia

Análises de parcimônia foram realizadas através do programa TNT (Goloboff et al. 1993, 2003; protocolo de utilização – Goloboff et al. 2008). Os gaps foram codificados como missing data. Foram realizadas buscas heurísticas através do New Technology Search, utilizando 137

simultaneamente os algoritmos Sectorial Search, Ratchet, Drift e Tree fusing (10 rounds). Para o suporte de ramos foi utilizado o suporte de bootstrap com 1000 repetições.

Máxima Verossimilhança (ML)

Máxima verossimilhança (ML) usando RaxML v8.0.24 (Stamatakis et al., 2008) foi implementada através do CIPRES Science Gateway (Miller et al., 2010). Foram realizadas mil repetições rápidas de bootstrap, sendo que os demais parâmetros foram mantidos de forma padrão. Cabe ressaltar, que para as análises de ML os dados foram particionados por gene, sendo que os genes mitocondriais (COI-nj e COII) foram separados nas posições (1+2) e (3).

Resultados

Análise de caracteres morfológicos

A análise dos caracteres realizada através da comparação dos 43 táxons terminais estabeleceu 61 caracteres, sendo: 28 binários e 33 multiestados. Os caracteres são apresentados de acordo com a posição do corpo (cabeça, antena, mesossoma e seus apêndices e metasoma). Segue a abaixo a listagem dos caracteres; os valores apresentados após a descrição do caráter são resultantes da árvore mais parcimoniosa obtida na análise com pesagem implícita dos caracteres.

Lista de caracteres

CABEÇA

1. Cerdas nos olhos compostos: (0) ausentes ou no máximo visíveis somente em alta ampliação (Fig. 1A); (1) presentes (Fig. 1B). [L = 3; CI = 33; RI = 66]. Comentário. As cerdas dos olhos são altamente variáveis e variam desde pequenas até robustas e longas. De acordo com Heraty (2002), embora o tipo de cerda seja importante para a identificação, qualquer particionamento adicional do estado resulta em excesso de polimorfismos dentro dos gêneros. 138

Figura 1. Cerdas nos olhos compostos. (A) cerdas ausentes (Kapala flabellata); B. cerdas presentes (Lirata spinaelevis).

2. Formato do olho composto: (0) globular e convexo; (1) arredondado e pronunciado; (2) com protuberância em forma de “mamilo”. [L = 2; CI = 100; RI = 100]. Comentário. O formato do olho composto globular e convexo entre os gêneros próximos a Kapala é predominante. Já o formato arredondado e pronunciado é encontrado nas espécies dos gêneros Lirata e Neolirata. A presença de uma protuberância em forma de mamilo é uma característica única do gênero Isomerala.

Figura 2. Formato do olho composto. A. globular e convexo (Kapala flabellata); B. arredondado e pronunciado (Lirata spinaelevis n. sp.); C. com protuberância em forma de “mamilo” (Isomerala coronata).

3. Depressão escrobal: (0) rasa com margens laterais arredondadas gradualmente (Fig. 3A); (1) profunda com as margens laterais anguladas, carenadas ou abruptamente arredondadas (Fig. 3B). 139

[L = 2; CI = 50; RI = 0]. Comentário. Esse caráter se refere aos canais paralelos utilizados por Heraty (2002) em análises prévias, e que considerou o presente caráter como uma autapomorfia de Psilocharis (gênero não utilizado na presente análise). A grande maioria dos táxons possuem as margens laterais mais arredondadas gradualmente.

Figura 3. Depressão escrobal. A. depressão escrobal rasa (Colocharis hungi); B. profunda com as margens laterais abruptamente arredondadas (Isomerala coronata).

4. Carena contínua entre o ocelo mediano e ocelos laterais: (0) ausente (Fig. 4A); (1) presente (Fig. 4B). [L = 1; CI = 0; RI = 0]. Comentário. Torréns e Heraty (2013) propuseram um novo gênero, Neolirata, com base na presença de uma carena entre o ocelo médio e os ocelos laterais, sendo uma autapomorfia do gênero. Os demais gêneros não possuem uma carena entre os ocelos médio e laterais.

5. Área supraclipeal: (0) sulco lateral indistinto (ausente) ou fracamente impresso < 1/2 da distância do torulus; (1) sulcos laterais fortemente impressas ou > 1/2 da distância do torulus. [L = 9; CI = 11; RI = 46]. Comentário. Este é um caráter muito difícil de definir devido a uma considerável variação na estrutura da região supraclipeal. Segundo Heraty (2002) e Murray (2014), a definição dessa área pode ser feita pela presença de uma impressão linear clara ou sulco crenulado lateralmente. A área supraclipeal pode estar claramente ausente e não ser distinta de forma alguma, ou pode ser inchada e, muitas vezes, com apenas uma mudança no padrão do tegumento que a distingue como uma região distinta. 140

Figura 4. Carena entre o ocelo mediano e ocelos laterais. A. carena ausente (Kapala iridicolor); B. carena presente (Neolirata daguerrei).

Figura 5. Área supraclipeal (indicada). A. sulco lateral indistinto (Liratella sp.); B. sulco lateral crenulado (Colocharis hungi).

6. Área supraclipeal, padrão do tegumento: (0) liso (Fig. 6A); (1) estriado (Fig. 6B); (2) rugoso (Fig. 6C); (3) pontuado (Fig. 6D). [L = 7; CI = 42; RI = 33]. Comentário. O padrão do tegumento é extremamente variável entre os gêneros. Entre as espécies de Kapala, esse caráter pode variar de liso a estriado, como também ser polimórfico (fêmeas, liso; machos, estriado). 141

Figura 6. Área supraclipeal (indicada), padrão do tegumento. A. liso (Torquata n.gen.); B. estriado (Kapala haplospinosa n. sp.); C. rugoso (Lirata pustula); D. pontuado (Dilocantha sp.).

7. Padrão do tegumento da fronte: (0) liso ou com poucas estrias fracamente impressas (Fig. 6A); (1) estrias grossas, as quais são separadas uma das outras (Fig. 7A); (2) estrias finas, próximas umas das outras (Fig. 7B); (3) rugoso (Fig. 6C); (4) reticulado/pontuado (Fig. 6D). [L = 10; CI = 40; RI = 73]. Comentário. Como já mencionando anteriormente, os padrões do tegumento da fronte compreendem um caráter importante para a definição de gêneros. Para Kapala esse caráter auxilía na definição dos complexos de espécies furcata (7:2), iridicolor (7:0) e romandii (7:1).

8. Padrão do tegumento na face inferior: (0) liso ou com poucas estrias fracamente impressas; (1) estrias grossas, separadas umas das outras; (2) estrias finas, próximas umas das outras; (3) rugoso; (4) reticulado. [L = 9; CI = 44; RI = 75]. Comentário. A fronte e a face inferior são 142

consideradas como áreas separadas, definidas como acima e abaixo do torulus. Assim como o caráter anterior, os padrões do tegumento da face inferior contribuem para o diagnóstico de gêneros, como também são de fundamental importância para a definição dos complexos de espécies dentro de Kapala.

Figura 7. Padrão do tegumento da fronte. A. estrias grossas, as quais são separadas uma das outras (indicado); B. estrias finas, próximas umas das outras (indicado).

9. Padrão das estrias da fronte: (0) longitudinal (Fig. 8A); (1) oblíquo (do triângulo ocelar em direção à margem inferior do olho) (Fig. 8B); (2) oblíquo (do ocelo lateral para a margem inferior do torulus) (Fig. 8C). [L = 2; CI = 100; RI = 100]. Comentário. O padrão de estrias no tegumento da fronte pode ser encontrado somente nos seguintes gêneros: Galearia, Kapala, Lasiokapala, Latina, Lirata e Thoracantha. Em Kapala, os estados 1 e 2 podem ser encontrados dentro do complexo de espécies furcata, já o estado 0 no complexo romandii.

10. Padrão das estrias na face inferior: (0) transversal (Fig. 8A); (1) oblíquo (do torulus em direção a gena) (Fig. 8B). [L = 1; CI = 100; RI = 100]. Comentário. Dentre os gêneros, o estado 0 é comum, já o estado 1 pode ser encontrado somente dentro do complexo de espécies furcata (gênero Kapala).

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Figura 8. Padrão das estrias da fronte. A. longitudinal. B. oblíquo (do triângulo ocelar em direção à margem inferior do olho); C. oblíquo (do ocelo lateral para a margem inferior do torulus).

11. Fronte: (0) achatada (Fig. 9B); (1) inchada/pronunciada (Fig. 9A). [L = 1; CI = 100; RI = 100]. Comentário. A fronte inchada é encontrada apenas em espécies pertencentes ao complexo de espécies iridicolor (gênero Kapala), sendo uma das principais características que definem esse complexo.

Figura 9. Fronte. A. inchada (Kapala longicornis n. sp.); B. achatada (Kapala flabellata).

12. Margem posterolateral da gena: (0) estreita com a margem posterolateral uniformemente arredondada ou reta (Fig. 10A); (1) larga e angulada medialmente, com ou sem uma carena na margem posterior (Fig. 10B, indicado). [L = 3; CI = 33; RI = 71]. 144

13. Número de dígitos do labro (usado como um intervalo; tipicamente variável dentro das espécies): (0) 4-6 dígitos; (1) 7-9 dígitos; (2) 10-13 dígitos. [L = 4; CI = 25; RI = 78]. Comentário. O agrupamento de dígitos, muitas vezes com dois dígitos isolados lateralmente e diversos medialmente, é altamente variável, mas pode ser muito importante para as análises de uma menor quantidade de gêneros mais estreitamente relacionados.

14. Palpo maxilar: (0) ausente; (1) 1-segmentado; (2) 2-segmentado; (3) 3-segmentado. [L = 6; CI = 50; RI = 70].

15. Palpo labial: (0) Ausente; (1) 1-segmentado; (2) 2- segmentado; (3) 3- segmentado. [L = 14; CI = 21; RI = 45]. Comentário. Segundo Heraty (2002), três palpômeros é um estado plesiomórfico.

ANTENA

16. Comprimento do escapo: (0) não atingindo o ocelo médio (Fig. 11A); (1) atingindo o ocelo médio ou excedendo sua margem inferior. [L = 2; CI = 50; RI = 66]. Comentário. É difícil atribuir arbitrariamente um valor para descrever um escapo "longo" versus "curto", mas, em geral, eles são facilmente divididos, com os escapos curtos sendo tão longos quanto largos. Em 145

alguns casos, o ápice do escapo é expandido, resultando na proporção mais baixa (cerca de 3X mais longo que largo), mas um escapo longo geralmente é cinco ou mais vezes mais longo do que largo. Dentro do clado Kapala (Eucharitinae), somente Neolirata e Lirata possuem um escapo que atinge o ocelo médio.

Figura 11. Comprimento do escapo. A. não atingindo o ocelo médio (Parakapala decarloi); B. atingindo o ocelo médio (Lirata spinaelevis n. sp.)

17. Número de flagelômeros, fêmea. Registrados na matriz de acordo com a quantidade que é de 6 a10); ‘0’ indica 10, que é o número máximo. [L = 9; CI = 33; RI = 60]. Comentário. Com base no grupo externo, um funículo com 7 segmentos é plesiomórfico, com o maior número de segmentos resultantes da incorporação de segmentos da clava e, em alguns casos, da adição de novos segmentos. A clava é definida como o segmento terminal ou segmentos que estão intimamente associados (não articulados) e são frequentemente expandidos e distintos dos segmentos funiculares. No entanto, em Eucharitinae os segmentos da clava são muitas vezes perdidos à medida que os segmentos apicais se tornam indiferenciados dos segmentos funiculares e, em alguns casos, apenas o segmento terminal pode ser considerado como clava (Heraty, 2002).

18. Flagelo: relação comprimento/altura da cabeça, fêmea: (0) < 1x; (1) 1.1‒1.4x; (2) 1.41‒1.99x; (3) > 2x. [L = 14; CI = 21; RI = 35].

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19. Flagelômero basal (Fl2) comprimento vs. largura, fêmea: (0) tão longo quanto largo, aproximadamente transversal; (1) 1.5‒3x mais longo que largo; (2) > 3.1x mais longo que largo. [L = 13; CI = 15; RI = 31].

20. Formato de Fl2, fêmea: (0) simples, cilíndrica; (1) serreada; (2) serreada com o ápice terminando em uma ponta afiada; (3) ramificada, alongada. [L = 9; CI = 33; RI = 62]. Comentário. Esse caráter foi inicialmente excluído devido ao viés inerente na codificação para o que parece ser uma característica muito polimórfica. Contudo, a retirada dele causou algumas perturbações nas análises e, por fim, optou-se por deixá-lo.

Figura 12. Formato de Fl2 da fêmea. A. flagelômeros simples/cilíndricos (Dilocantha sp.).; B. serreado (Ecarinata n. gen.); C. serreado com o ápice terminando em uma ponta afiada (Lirata diabla); D. ramificado, alongado (Lasiokapala serrata) 147

21. Comprimento de Fl3 em relação ao comprimento de Fl2, fêmea: (0) menor que Fl2; (1) mesmo comprimento; (2) maior que Fl2. [L = 4; CI = 25; RI = 62]. Comentário. Idem ao caráter anterior.

22. Formato de Fl3, fêmea: (0) simples ou cilíndrica; (1) serreada; (2) serreada oblíqua com o ápice terminando em uma ponta afiada; (3) ramificada, alongada. [L = 10; CI = 30; RI = 56].

23. Número de flagelômeros, macho: (0) 8; (1) 9; (2) 10. [L = 1; CI = 100; RI = 100]. Comentário. A grande maioria dos gêneros possuem 10 flagelômeros ramificados. Apenas Colocharis possui 8.

24. Comprimento do ramo de Fl2 em relação à altura da cabeça, macho: (0) < 1.0x; (1) 1.0‒1.4x; (2) 1.41‒1.99x; (3) > 2x. [L = 7; CI = 28; RI = 54]. Comentário. Esse é um caráter que pode ser considerado extremamente variável nos demais gêneros, contudo auxilía na definição das espécies macho de Kapala.

25. Comprimento de Fl2 vs. largura, não incluindo o ramo flagelar, macho: (0) tão longo quanto largo, aproximadamente transverso; (1) 1.5‒3x mais longo que largo; (2) >3x mais longo que largo. [L = 3; CI = 66; RI = 66]. Comentário. O comprimento de Fl2 é variável e potencialmente mais significativo quando utilizados gêneros de Oraseminae. Contudo, em Eucharitinae, é quase impossível dividir os vários tamanhos, além de um segmento muito curto, de forma significativa, pois um Fl2 transversal ou quadrado está sempre associado a uma antena pectinada, a qualé encontrada nos gêneros em questão na presente análise.

MESOSSOMA

26. Mesoescutelo, em vista lateral: (0) baixo, não projetado sobre a cabeça e com a margem anterior arredondada (Fig, 13A); (1) vertical, projetado sobre a cabeça e com margem anterior suavemente curvada (Fig. 13B); (2) vertical, projetado sobre a cabeça e com margem anterior abruptamente curvada na metade dorsal (Fig. 13C). [L = 3; CI = 66; RI = 90]. Comentário. Em 148

Eucharitinae, mais especificamente dentro do clado Kapala, ocorre uma modificação bizarra do mesoscuto, o qual é elevado sobre o pronoto. Contudo, a definição dessa elevação pode ser um tanto complicada de se estimar. Na presente análise, além da elevação, consideramos a curvatura do mesoscuto, considerando como mais elevado o que está sempre abruptamente curvado a partir da metade dorsal, em vista lateral.

Figura 13. Mesoescutelo em vista lateral. A. baixo, não projetado sobre a cabeça (Chalcura sp.); B. vertical, projetado sobre a cabeça e com a margem anterior suavemente curvada (Kapala ceciliae n. sp.); C. vertical, projetado sobre a cabeça e com margem anterior abruptamente curvada na metade dorsal (Kapala argentina)

27. Lóbulos laterais, em vista lateral: (0) elevado a aproximadamente a altura do lóbulo médio (Fig. 14A); (1) não elevado à altura do lóbulo médio (Fig. 14B). [L = 1; CI = 0, RI = 0].

Figura 14. Lóbulos laterais do mesoescuto, em vista lateral. A. elevado a aproximadamente a altura do lóbulo médio; B. não elevado à altura do lóbulo médio.

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28. Lóbulos laterais, em vista frontal: (0) não projetado (Fig. 15A); (1) projetado (Fig. 15B). [L = 1; CI = 100; RI = 100].

Figura 15. Lóbulos laterais do mesoescuto em vista frontal. A. não projetado (Kapala parafurcata n. sp.); B. projetado (Kapala deltalis n. sp.)

29. Flange do mesoescuto sobre a tégula: (0) flange muito fina ao longo da margem da tégula (Fig, 16A); (1) flange triangular que se projeta posteriormente do canto posterolateral do mesoescuto sobre a tégula (Fig. 16B). [L = 3; CI = 33; RI = 33]. Comentário. A flange é uma estrutura incomum encontrada apenas em Dilocantha, em um grupo de Lirata e em uma nova espécie de Liratella. É considerada como um desenvolvimento separado de outras modificações da margem mesoescutal.

30. Formato do prepecto, dorsolateral: (0) triangular a sub-triangular; (1) estreito e alongado. [L = 3; CI = 33; RI = 0]. Comentário. Esse é um caráter com base no comprimento relativo ventral ou dorsal do complexo pronoto-prepecto.

31. Mesoescutelo: (0) sem um canal mediano; (1) com um canal mediado dando uma aparência de elevação dupla. [L = 1; CI = 100; RI = 100]. Comentário. A presença de um canal mediano no mesoescutelo ocorre somente nas espécies do gênero Lirata. 150

Figura 16. Flange do mesoescutelo. A. flange muito fina ao longo da margem da tégula (indicado) (Parakapala decarloi); B. flange triangular que se projeta posteriormente do canto posterolateral do mesoescuto sobre a tégula (indicado) (Lirata pustula).

32. Formato do mesoescutelo em vista lateral: (0) achatado (Fig. 17A); (1) inchada/convexa (Fig. 17B). [L = 2; CI = 50; RI = 66]. Comentário. O estado 1 é incomum, sendo encontrado somente em duas espécies: Kapala terminalis e Kapala jalisca n. sp.

Figura 17. Formato do mesoescutelo em vista lateral. A. achatado (Kapala inexagens); B. inchado/convexo (Kapala terminalis).

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33. Mesoescutelo, padrão de esculturação do tegumento: (0) carenas longitudinais; (1) carenas oblíquas; (2) liso; (3) rugoso. [L = 10; CI = 30; RI = 0]

34. Margem medial posterior do disco mesoescutelar em vista lateral: (0) arredondada (Fig. 18A); (1) terminando em um ápice elevado, ápice < 1.5x a altura do processo frenal; (2) terminando em um ápice elevado, ápice > 1.5x a altura do processo frenal (Fig. 18B); (3) medialmente deprimida (Fig. 18C). [L = 9; CI = 33; RI = 45]. Comentário. Os estados de caráter 1 e 2 são restritos de Kapala, Isomerala e Neolirata. Contudo, a altura do ápice medial do escutelo pode variar dentro das espécies e entre as espécies de Kapala. Além disso, Kapala terminalis, K. floridana e K. jalisca n. sp. possuem a margem medial posterior do disco mesoescutelar medialmente deprimida.

Figura 18. Margem medial posterior do disco mesoescutelar em vista lateral. A. arredondada (Chalcura sp.); B. terminando em um ápice elevado (Kapala haplospinosa n. sp.); C. medialmente deprimida (Kapala floridana)

35. Projeção da margem posterior do escutelo: (0) curta (Fig. 19A); (1) longa (Fig. 19B). [L = 2; CI = 100; RI = 100]. Comentário. Colocharis é o único membro dentro do clado Kapala com os processos frenais curtos, sendo cada processo não mais que 4x mais longo que largo e são amplamente separados na região basal.

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Figura 19. Projeção da margem posterior do mesoescutelo.A. curto (Colocharis hungi); B. longo (Kapala argentina).

36. Comprimento dos processos frenais em relação ao disco mesoescutelar: (0) menor que o comprimento do mesoescutelo; (1) 1‒2x o comprimento a partir da articulação transescutal (TSA) até o ápice do disco mesoescutelar; (2) > 2x o comprimento a partir da TSA até o ápice do disco mesoescutelar; (3) mais longo que o disco mesoescutelar, mas menor do que 1x o comprimento a partir da TSA até o ápice do disco mesoescutelar. [L = 6; CI = 50; RI = 62].

37. Ápice dos processos frenais: (0) arredondado ou acuminado (Fig. 20A); (1) emarginado (Fig. 20B); (2) emarginado, com uma flange côncava na margem inferior dos processos frenais (Fig. 20C). [L = 11; CI = 18; RI = 57]. Comentário. O ápice dos processos frenais é um caráter que auxilía na definição de espécies do gênero Kapala. Outra questão importante que deve ser mencionada, é que a presença do estado 2 é extremamente comum em espécimes encontrados na América Central, já as espécies as da América do Sul possuem um ápice emarginado sem uma flange côncava.

38. Forma dos processos frenais, fêmea: (0) processo cilíndrico e paralelo; (1) amplo e achatado dorsoventralmente; (2) formando uma carapaça arqueada sobre o metassoma. [L = 2; CI = 100; RI = 100].

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Figura 20. Ápice dos processos frenais. A. arredondado ou acuminado (Kapala iridicolor); B. emarginado (Kapala cavicornis n. sp.); C. emarginado com uma flange côncava na margem inferior dos processos frenais (Kapala haplospinosa n. sp.).

Figura 21. Formato dos processos frenais das fêmeas (imagens superiores em vista lateral e inferiores em vista dorsal). A. cilíndricos e paralelos (Kapala flabellata); B. amplos e achatados dorsoventralmente (Dicoelothorax sp.); C. formando uma carapaça arqueada sobre o metassoma (Galearia latreillei).

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39. Cerdas nos processos frenais: (0) ausente; (1) presente ao longo do comprimento dos processos frenais. [L = 2; CI = 50; RI = 83].

40. Processos frenais, padrão de esculturação do tegumento (aspecto dorsal), fêmea. (0) liso ou aparentemente quase liso (Fig. 22A); (1) com carenas longitudinais ao longo de todo o comprimento ou tornando-se oblíquas no ápice (Fig. 22B); (2) carenas transversais ou circulares (Fig. 22C); (3) rugoso em mais de um quarto do comprimento (Fig. 22D). [L = 4; CI = 75; RI = 0].

41. Processos frenais, padrão de esculturação do tegumento (aspecto dorsal), macho: (0) liso ou aparentemente quase liso; (1) com carenas longitudinais ao longo de todo o comprimento ou tornando-se oblíquas no ápice; (2) carenas transversais; (3) rugoso em mais de um quarto do comprimento; (4) carenas irregulares longitudinais, dando uma aparência serreada. [L = 5; CI = 80; RI = 87].

42. Mesepisterno: (0) liso; (1) com estrias em mais de um quarto da superfície; (2) pontuado. [L = 8; CI = 25; RI = 70]. Comentário. Esse caráter se concentra inteiramente na interligação da margem posterior do prepecto e margem mesepimeral.

43. Mesepimero: (0) liso; (1) com estrias em mais de um quarto da superfície; (2) pontuado. [L = 9; CI = 22; RI = 65].

44. Callus, padrão do integumento: (0) liso ou com pontuações suaves; (1) com carenas; (2) estriado. [L = 5; CI = 40; RI = 66]. Comentário. A área do callus é localizada lateralmente ao sulco pós-espiráculo e acima da metapleura. Essa estrutura possui os mais variados padrões de integumento, sendo que aqui distinguimos apenas as formas mais extremas e marcantes.

45. Callus, pilosidade: (0) nu ou com algumas pequenas cerdas; (1) densamente piloso. [L = 2; CI = 50; RI = 92]. Comentário. A estrutura e localização das cerdas são altamente variáveis e podem fazer com que não seja um caráter muito informativo, a princípio. 155

Figura 22. Processos frenais, padrão de esculturação do tegumento (aspecto dorsal). fêmea. A. liso ou aparentemente quase liso (Lasiokapala serrata). B. com carenas longitudinais ao longo de todo o comprimento ou tornando-se oblíquas no ápice (Kapala cuprea); C. carenas transversais ou circulares (Lirata luteogaster); D. rugoso (Lirata maranhensis).

46. Disco propodeal: (0) convexo (Fig. 23A); (1) achatado (Fig. 23B). [L = 2; CI = 50; RI = 50].

Figura 23. Disco propodeal, em vista lateral. A. convexo (Lirata striatissima); B. achatado (Liratella nigra).

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47. Carena delimitando o disco propodeal: (0) ausente (Fig. 24A); (1) presente (Fig. 24B, indicado). [L = 2; CI = 50; RI = 75]. Comentário. A carena forma uma borda distinta em torno do disco propodeal, o qual é relativamente plano e liso. Em Thoracantha e Galearia, a carena é pronunciada e se encaixa com a face ântero-dorsal do primeiro tergito do metasoma.

Figura 24. Carena delimitando o disco propodeal. A. ausente (Ecarinata n. gen.); B. presente (Kapala iridicolor, indicado).

48. Espiráculo propodeal: (0) circular (Fig. 25A); (1) circular com uma incisão ventral (“formato de fechadura”) (Fig. 25B). [L = 1; CI = 100; RI = 100].

Figura 25. Espiráculo propodeal. A. circular (indicado); B. circular com uma incisão ventral (indicado). Fonte: adaptado de Heraty, 2002. 157

METASSOMA

49. Primeiro tergo metassomal: (0) pouco esclerozado e flexível; (1) endurecido e em forma de placa (Thoracantha e Gallearia). [L = 2; CI = 50; RI = 0]. 50. Cerdas eretas em mais da metade do primeiro tergito metassomal: (0) ausentes; (1) presentes, embora possam ser muito esparsas. [L = 1; CI = 100; RI = 100].

51. Cicatriz tergal: (0) ausente; (1) presente. [L = 9; CI = 11; RI = 42]. Comentário. Uma cicatriz linear ou circular, delimitada por uma ligeira rugosidade da cutícula está presente dorsolateralmente e subapicalmente no Gt1. A cicatriz marca uma conexão esclerosada interna entre Gt1 e Gt2. Quase todos os Eucharitini possuem uma cicatriz tergal linear, enquanto que em Colocharis e alguns Kapala, a cicatriz é circular.

52. Hipopígio (0) com poucas ou nenhuma cerda apical envolvendo o mucro; (1) com um único anel linear de cerdas longas e curvadas apicalmente; (2) com um tufo de cerdas apicais em torno do mucro. [L = 6; CI = 33; RI = 55]. Comentário. Segundo Heraty (2002), a homologia dos estados 0 e 1 é incerta devido às várias colocações e formas das cerdas do hipopígeo.

PERNAS E ASAS

53. Calcar: (0) curvado e bífido; (1) reto e acuminado. [L = 2; CI = 50; RI = 83].

54. Número de esporões das tíbias posteriores: (0) esporão ausente; (1) um esporão; (2) dois esporões. [L = 7; CI = 28; RI = 44]. Comentário. Dois esporões tibiais são amplamente distribuídos em Chalcidoidea e Heraty (2002) presume ser um caráter plesiomórfico.

55. Asas anteriores: (0) uniformemente transparentes ou ligeiramente infuscadas; (1) infuscadas, escuras. [L = 5; CI = 20; RI = 20].

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56. Venação das asas anteriores: (0) distinta; (1) fraca, venação após a veia submarginal geralmente ausente. [L = 3; CI = 33; RI = 71].

57. Veia estigmal: (0) escura; (1) semi-transparente. [L = 4; CI = 25; RI = 62]

58. Comprimento da veia pós-estigmal das asas anteriores: (0) ausente; (1) 3‒4x mais longo que largo; (2) > 5x mais longo que largo. [L = 11; CI = 18; RI = 35].

59. Pilosidade asas anteriores: (0) densamente pilosa, com longas cerdas; (1) moderadamente pilosa, geralmente com cerdas pequenas; (2) cerdas ausentes. [L = 6; CI = 16; RI = 28]. Comentário. A presença ou ausência de cerdas na área basal e área do espéculo da asa é muito variável para ser útil para a análise de grupos superiores, contudo, ela é muito útil quando se trata de gêneros mais relacionados. Além disso, a diferença entre uma asa densamente pilosa e moderadamente pilosa é relativamente fácil de discernir.

60. Cerdas marginais nas asas anteriores: (0) ausente; (1) presente. [L = 2; CI = 50; RI = 66].

61. Cerdas marginais nas asas posteriores: (0) ausente; (1) presente. [L = 2; CI = 50; RI = 50].

Análise cladística morfológica

A análise de parcimônia com pesagem implícita (k =7.1875) resultou em uma árvore mais parcimoniosa com um comprimento de 300 passos (IC = 35; IR = 60; Fig. 26). De acordo com os resultados encontrados, Kapala não foi recuperado como um grupo natural, sendo considerado parafilético. O gênero foi dividido em três complexos de espécies bem definidos, sendo eles: complexo K. iridicolor (clado em verde), complexo K. romandii (clado em azul) e complexo K. furcata (clado em vermelho) (Fig. 26). O complexo de espécies K. iridicolor foi recuperado como monofilético, sendo sustentado por uma sinapormofia [11:1, fronte inchada, Fig. 9A] e duas homoplasias [15:3, palpo labial 3- segmentado; 17:9, número de flagelômeros 9-segmentado]; já o suporte de bootstrap foi baixo 159

(14) (Fig. 26). Esse complexo tem como grupo irmão os clado 2 + clado 3. Internamente podemos verificar a formação de dois pequenos clados, sendo o primeiro composto por K. ipa n. sp.+ (K. ceciliae n. sp.+ K. longicornis n. sp.) e o segundo por K. parairidicolor n. sp. (K. iridicolor + K. cavicornis n. sp.), os quais são sustentados apenas por homoplasias. Além disso, podemos verificar que K. ipa sp. n. é suportada por uma autapormofia [17:0, flagelo 10- segmentado], sendo a única espécie dentro do complexo a possuir essa quantidade de flagelômeros, as demais espécies, possuem somente 9. O complexo de espécies K. roamndii é monofilético, o qual é sustentado apenas por duas homoplasias [8:1, padrão do tegumento da fronte com estrias grossas, as quais são separadas umas das outras; 19:2, flagelômero basal > 3x mais longo que largo] (Fig. 26). Não houve suporte de bootstrap. Na presente análise, o complexo K. romandii é composto por 12 espécies, das quais três são novas (Kapala corcovata n. sp., Kapala jalisca n. sp. e Kapala haplospinosa n. sp.). Internamente podemos observar a formação de pequenos agrupamentos como K. terminalis e K. jalisca n. sp. sustentados por duas homoplasias [32:1; 34:0]; K. floridana + (K. corcovata n. sp. + K. haplospinosa n. sp.) sustentado por uma homoplasia [5:1, área supraclipeal fortemente impressa]; K. izapa + K. romandii, sendo sustentado por duas homoplasias [42:0, mesepisterno liso; 43:0, mesepimero liso] e K. flabellata + K. ivorensis, apesar de serem espécies muito semelhantes morfologicamente, não existem sinapomorfias que sustentem ambas espécies. Espécies do complexo K. romandii também podem ser reconhecidas pela presença de 8 flagelômeros nas fêmeas, mesoescuto elevado, mesoescutelo curto e processos frenais dos machos com carenas irregulares com um aspecto serreado. Apesar de, na presente análise, Kapala inexagens ser recuperada fora do complexo K. romandii (Fig. 26), optou-se por mantê-la dentro desse complexo, uma vez que, suas características morfológicas se assemelham em muito com as das demais espécies do complexo. Além disso, esse resultado se deve principalmente à grande quantidade de missing data para essa espécie em função da baixa quantidade de exemplares analisados (e desses muitos estavam com estruturas quebradas e perdidas), bem como ausência de exemplares machos.

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Figura 26. Cladograma obtido com pesagem implícita (k = 7.1875) dos caracteres através de busca tradional pelo TNT (L = 300); IC = 35; IR = 60). Os círculos pretos representam mudanças únicas, os brancos representam paralelismos ou convergências (homoplasia). As linhas coloridas representam as espécies de Kapala. Valores de bootstrap são mostrados nos retângulos lilases. À direta, a filogenia proposta por Heraty (2002) para o clado Kapala. 161

O complexo de espécies K. furcata também é sustentado por duas homoplasias [15:3, palpo labial com 3 segmentos;17:9, fêmeas com 9 segmentos flagelares] (Fig. 26). Não houve suporte de bootstrap. O complexo de espécies K. furcata tem como grupo irmão o clado formado pelos seguintes gêneros: Dilocantha, Latina, Dicoelothorax, Parakapala, Isomerala, Galearia, Liratella, Lasiokapala, Thoracantha, Neolirata e Lirata. Na presente análise, o complexo K. furcata é composto por 6 espécies, sendo três novas (K. deltalis n. sp., K. parafurcata n. sp. e K. quasimodo n. sp.). O complexo é dividido internamente em dois clados menores, sendo eles: K. furcata + K. splendens, sustentado por duas homoplasias [5:1; 58:0] e um suporte de bootstrap baixo (10) e K. cuprea + K. parafurcata n. sp. + (K. deltalis n. sp. + K. quasimodo n. sp.) sustentado por duas sinapomorfias [9:1, fronte com estrias oblíquas do triângulo ocelar em direção a margem inferior do olhos; 10:1, face com estrias oblíquas do torulus em direção a gena], além de uma suporte baixo de bootstrap (16). De acordo com as análises filogenéticas morfológicas, dois novos gêneros são recuperados (Fig. 26) e propostos: Ecarinata n. gen. e Torquata n. gen. (descrição no Cap. V). Ecarinata n. gen. tem como grupo irmão o clado formado por Colocahris hungi + Colocharis napoana; já Torquata n. gen. tem como grupo irmão os clados 1 + 2 + 3.

Análise molecular: parcimônia

A análise molecular de parcimônia resultou em 12 árvores mais parcimoniosas, as quais estão sumarizadas em uma árvore de consenso estrito (Fig. 27). Assim como na análise morfológica, Kapala não foi recuperado como um grupo natural (Fig. 27). O gênero foi dividido em dois complexos bem definidos (K. furcata e K. iridicolor) os quais estão separados das demais espécies de Kapala por três agrupamentos de gêneros morfologicamente distintos. As espécies que formaram o complexo K. romandii na análise morfológica não correspondem a um grupo natural na análise molecular de parcimônia, sendo que estas formaram pequenos grupamentos de espécies os quais, por sua vez, estão divididos por agrupamentos de gêneros morfologicamente distintos.

O complexo K. furcata foi recuperado como monofilético, com suporte de booststrap de 48 e, tem como grupo irmão o clado formado pelos gêneros Neolirata, Latina, Thoracantha, 162

Dicoelothorax e Lasiokapala e o clado que compõe o complexo de espécies K. iridicolor (Fig. 27). Além disso, podemos observar que as relações entre as espécies K. deltalis n. sp., K. quasimodo n. sp. e K. parafurcata n. sp. não foram recuperadas. O complexo K. iridicolor foi recuperado como monofilético com um suporte de bootstrap de 74. Contudo, as relações internas entre as espécies não foram recuperadas. Torquata n. gen. resultou como monofilético com um suporte de bootstrap de 85. Ecarinata n. gen. também resultou como monofilético, porém sem suporte. Na presente análise, Ecarinata n. gen. corresponde ao grupo irmão de Gallearia, podendo essa relação ser colocada em cheque, uma vez que ambos os gêneros são muito distintos e, nas demais análises Ecaritana corresponde a um gênero mais basal juntamente com Colocharis e Torquata n. gen. (Figs 26, 27) em relação aos demais gêneros.

Análise molecular: máxima verossimilhança

A análise de máxima verossimilhança corrobora com os resultados encontrados nas análises anteriores (morfológica e molecular – parcimônia), nas quais Kapala não corresponde a um grupo natural (Fig. 28). De forma similar, o complexo de espécies K. rmandii é parafilético, sendo que suas espécies formaram pequenos agrupamentos (ramos em azul), que por sua vez, estão divididos pelo clado formado pelos gêneros Dilocantha + Lirata, pela espécie Isomerala bouceky e por Gallearia latreillei (Fig. 28). Kapala romandii foi recuperada como grupo irmão do clado composto pelos gêneros Neolirata, Latina, Thoracantha, Dicoelothorax, Lasiokapala, do complexo furcata e do complexo iridicolor. Os complexos de espécies K. furcata (clado em vermelho) e K. iridicolor (clado em verde) perfazem grupos monofiléticos com suporte de bootstrap de 77 e 98, respectivamente (Fig. 28). Torquata n. gen. resultou como monofilético com um suporte de bootstrap de 100. Ecarinata n. gen. também, porém sem suporte.

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Figura 27. Análise de parcimônia com buscas heurísticas através do New Technology Search, com algoritmos Sectorial Search, Ratchet, Drift e Tree fusing (10 rounds) pelo TNT. Valores de bootstrap são mostrados abaixo dos ramos. Ramos em azul representam o complexo de espécies K. romandii; em vermelho o complexo K. furcata; em verde o complexo K. iridicolor. 164

Figura 28. Análise de máxima verossimilhança de 89 táxons terminais pelo RaxML. Suporte de bootstrap mostrado acima dos ramos. 165

Discussão

Todas as análises filogenéticas (morfológica + molecular: parcimônia e MV) suportam Kapala como parafilético. A ampla variação morfológica existente dentro de Kapala e dentro do clado Kapala resulta em uma mistura de gêneros morfologicamente variáveis, o que provavelmente torna o gênero parafilético. Essa situação, segundo Horandl & Stueessy (2010), é denominada de “budding”, no qual a origem de um novo táxon não afeta a existência ou os caracteres do ancestral original; supõe-se que isso aconteça após eventos de colonização por alguns indivíduos, seguido por uma especiação (peripátrica) rápida e adaptação aos novos ambientes. Um padrão semelhante ao de Kapala foi observado em Orasema (Eucharitidae: Oraseminae), com 57 espécies reconhecidas, e Perilampus (Perilampidae), com 84. Esses dois gêneros também parecem ser grupos morfologicamente coesos, contudo, análises moleculares mostram que ambos são intercalados por vários outros gêneros. Esse padrão de explosões fenotípicas de evolução dentro de um grupo morfologicamente conservado ainda não foi correlacionado com biologia ou história de vida do clado Kapala ou até mesmo para o gênero Kapala. Esse padrão também é enfatizado em Brady et al. (2014), que observaram a heterogeneidade de taxas evolutivas na morfologia das formigas da subfamília Dorylinae (Formicidae), onde, por exemplo, o gênero Cerapachys agora é compreendido com base na retenção coletiva de uma morfologia generalizada, com outros grupos aninhados dentro (por exemplo, as formigas-de-correição e outros gêneros) passando por uma evolução morfológica divergente. No presente estudo, verificou-se uma situação similar, provavelmente devido à incapacidade de se identificar a priori os caracteres filogeneticamente importantes dentre todas as variações fenotípicas e de traduzir efetivamente mudanças morfológicas em estados de caracteres. Em estudo realizado por Murray (2014), com o intuito de traçar a evolução do clado Kapala, utilizando dados morfológicos e moleculares, foi verificada a parafilia de Kapala. A autora recuperou dois clados monofiléticos (complexo K. furcata e K. iridicolor) bem suportados, e as demais espécies de Kapala formando pequenos clados divididos por demais gêneros, tendo uma resolução muito menor. Para tanto, o presente estudo forneceu uma topologia similar à 166

encontrada por Murray (2014), com dois complexos de espécies bem definidos e outro conjunto de espécies formando clados distintos, porém com baixo suporte. O complexo K. iridicolor obteve bons suportes de bootstrap nas análises moleculares (Figs 27, 28), sendo que suas espécies são morfologicamente definidas por uma combinação de caracteres que inclui uma face lisa, fronte inchada, palpo labial com 3-2 segmentos, perfil do mesoescuto baixo, mesossoma piloso e fêmeas com 9-10 flagelômeros. Cinco espécies novas foram estabelecidas para esse complexo, sendo que uma revisão e filogenia molecular mais apuradas são fornecidas no Capítulo III. O complexo K. furcata, que inclui a espécie-tipo do gênero, Kapala furcata (Fabricius), obteve um bom suporte na análise de máxima verossimilhança (Fig. 28). As espécies que compõem esse complexo são morfologicamente definidas por uma combinação de caracteres que inclui fêmeas com 9 flagelômeros (raramente 10), estrias faciais finas e oblíquas, palpo labial com 3-2 segmentos e corpos robustos em comparação aos demais indivíduos que fazem parte do gênero Kapala. Três novas espécies foram incorporadas para esse complexo, sendo que uma revisão e filogenia molecular mais apuradas são fornecidas no Capítulo IV. Na análise morfológica o complexo K. romandii foi recuperado como monofilético, porém de acordo com as análises moleculares, não corresponde a um agrupamento natural, sendo dividido por Galearia latreillei e pelo clado formado por Lirata e Dilocantha. Além disso, pode- se verificar que a variabilidade entre as análises deixa ambiguidade na posição do clado Dilocantha + Lirata e também das relações de Galearia letreillei (Figs 26, 27,28), o que acarreta uma certa preocupação na definição dos limites de Kapala (complexo K. romandii), um vez que, esses clados também possuem baixo suporte, podendo ser facilmente definidas novas relações através de análises mais robustas. Apesar dessa problemática, no presente trabalho optamos por manter o complexo K. romandii, devido ao compartilhamento de características plesiomórficas pelas espécies que integram esse complexo (Fig. 26). Através das presentes análises, fica evidente que o gênero Kapala precisará ser dividido, mas, devido a ausência ou baixo suporte de ramo na espinha dorsal da árvore (a qual envolve as espécies de Kapala), a circunscrição do gênero ainda deve ser sujeita a análises adicionais antes de se implementar grandes mudanças taxonômicas. Outra questão importante é a presença de mais de 20 morfótipos depositados na Universidade da Califórnia – Riverside (UCRC), e que são 167

provenientes de diversas regiões da América Central, que apresentam um claro processo de hibridização, cujos espécimes possuem características dos três complexos de espécies (K. iridicolor, K. furcata e K. romandii). Resumindo, antes de Kapala ser dividido, as seguintes questões devem ser analisadas e levadas em consideração: 1) baixo suporte de ramos em análises iniciais, sendo necessárias novas análises mais robustas; 2) presença de morfótipos com grau de hibridização que devem ser analisados e incluídos em análises futuras; 3) capacidade de codificação de caracteres e procura de novos caracteres morfológicos internos, uma vez que caracteres morfológicos externos não estão sendo muito eficazes e; 4) adição de informações referentes a comportamentos e possíveis hospedeiros.

Conclusão

Com base nas análises filogenéticas, Kapala é parafilético, dividido em três complexos de espécies, que por sua vez, estão divididos por agrupamentos de gêneros distintos. Neste momento, não há informações suficientes acerca de Kapala que permitam dividir o grupo, sendo necessária uma topologia estável. Para isso, a adição de mais informações sobre caracteres internos, comportamento, hospedeiros, além de espécimes adicionais, são necessários para a obtenção de uma topologia robusta. Além disso, estudos futuros incorporando informações genômicas adicionais ajudarão potencialmente a estabelecer limites para o gênero Kapala.

Agradecimentos

Agradecemos a concessão da bolsa do CNPQ (Conselho Nacional de Desenvolvimento Científico e Tecnológico) e da CAPES (Agência Federal de Apoio e Avaliação do Ensino de Pós- Graduação) à bolsista PDSE / visitante da University California - Riverside (processo PDSE - 88881.133943 / 2016 -01).

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Informação suplementar

Gênero Espécie Região País Museu Obs. Chalcura Kirby, 1886 Chalcura sp. OCE Tonga FSCA GE Colocharis hungi Torréns, 2012 NEO Brasil INPA GE Colocharis Heaaty, 2002 Colocharis napoana Heraty,2002 NEO Equador UCRC GE Dicoelothorax Ashmead, 1899 Dicoelothorax platycerus Ashmead, 1904 NEO Brasil INPA GE Dilocantha Shipp, 1894 Dilocantha bennetti Heraty, 1998 NEO Trinidade FSCA GE Galearia Brullé, 1846 Galearia latreillei (Guérin-Menéville, 1838) NEO Brasil MPEG GE Isomerala Shipp, 1894 Isomerala coronata (Westwood, 1874) NEO Brasil MZSP GE Lasiokapala Ashmead, 1899 Lasiokapala spiralicornis Torréns et al., 2016 NEO Brasil INPA GE Latina Koçak & Kemal,2008 Latina rugosa (Torréns, Heraty & Fidalgo, 2007) NEO Argentina FSCA GE Lirata luteogaster Cameron, 1884 NEO Brasil INPA GE Lirata Cameron, 1884 Lirata striatissima (Walker, 1862) NEO Brasil INPA GE Liratella nigra Girault, 1913 NEO Brasil INPA, UCRC GE Liratella Girault, 1913 Liratella n. sp. NEO Brasil DEFS, MPEG GE Neolirata Torréns & Herarty, 2013 Neolirata daguerrei (Gemignani, 1937) NEO Brasil INPA GE Parakapala Gemignani, 1937 Parakapala decarloi Gemignani, 1937 NEO Brasil DEFS, FSCA GE Thoracantha Latreille, 1825 Thoracantha striata Perty, 1833 NEO Brasil INPA GE Torquata n. gen. Torquata n. sp NEO Equador, Peru UCRC, INPA GE Ecarinata n. gen. Ecarinata n. sp. NEO Argentina UCRC GE Kapala Cameron, 1884 Kapala argentina Gemignani, 1933 NEO Argentina, MACN, INPA GI; CR Kapala cavicornis n. sp. NEO Trinidade UCRC GI; CI Kapala ceciliae n. sp. NEO Equador USNM, UCRC GI; CI Kapala chacoensis Gemignani, 1947 NEO Argentina MACN, UCRC GI; CR Kapala corcovata n. sp NEO Brasil INPA GI; CR Kapala cuprea Cameron, 1913 NEO Guiana BMNH GI; CF 172

Kapala cynipsea (Walker, 1862) NEO Brasil BMNH GI; CR Kapala deltalis n. sp. NEO Costa Rica UCRC GI; CF Kapala flabellata (Fabricius, 1804) NEO Brasil ZMUC; UCRC GI; CR Kapala floridana (Ashmead, 1885) NEO USA (Florida) USNM GI; CR Kapala furcata (Fabricius, 1804) NEO Brasil ZMUC; UCRC GI; CF Kapala haplospinosa n. sp. NEO USA (Florida) CNC GI; CR Kapala inexagens (Walker, 1862) NEO Brasil BMNH GI; CR Kapala ipa n. sp NEO Panamá UCRC GI; CI Kapala iridicolor (Cameron, 1904) NEO Nicarágua BMNH GI; CI Kapala ivorensis Risbec, 1954 AFRO Ivory Coast MNHN GI; CR Kapala izapa Carmichael, 2006 NEO México UCRC GI; CR Kapala jalisca n. sp. NEO México UNAM GI; CR Kapala longicornis n. sp. NEO Brasil UFES GI; CI Kapala parafurcata n. sp. NEO Argentina UCRC GI; CF Kapala parairidicolor n. sp. NEO México UCRC GI; CI Kapala quasimodo n. sp. NEO Venezuela UCRC GI; CF Kapala romandii (Guérin-Meneville, 1845) NEO Nicarágua BMNH GI; CR Kapala splendens Ashmead, 1904 NEO Brasil USNM GI; CF Kapala terminalis NEO Cuba USNM GI; CR

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Tabela S2. Lista dos gêneros e espécies do grupo externo e interno utilizados na análise filogenética molecular, sexo, número de identificação de depósito (Numb. ID), DNA voucher, museu de deposito e localidade de coleta.

Se DNA Espécies Num. ID Museu Localidade xo voucher Chalcura sp. UCRCENT_91769 D#0646a UCRC Australia: Northern Territory, 15.iii.2002, J. Heraty Schizaspidia aenea UCRCENT_91398 D#0168 UCRC Australia: NEQ, 11 km N, Ellis Beach, 30.i.1999, J. Heraty Colocharis napoana ♂ UCRCENT_92220 D#1104 USNM Equador: Orellana, Tiputini Biodiversity Sta., 26.x.1998, T.L. Erwin et al. Colocharis napoana ♀ UCRCENT_91450 D#1102 USNM Equador: Napo, Reserva Etinica Waorani, 27.x.1998, T.L. Erwin et al. Torquata n. gen. ♂ UCRCENT_10304 D#1141 UCRC Equador: Napo, Reserva Etinica Woarani, 6.ii.1999, T.L. Erwin et al. Torquata n. gen. ♂ UCRCENT_92029 D#1173 UCRC Equador: Orellana, Tiputini Biodiversity Station, 7.ii.1999, T.L. Erwin et al. Torquata n. gen. ♂ UCRCENT_397251 D#3440 UCRC Peru: Madre de Dios, Los Amigos Bio. St., 22.xii.2010, J. Heraty Ecarinta n. gen. ♀ UCRCENT_00483995 D#4898 IFML Argentina: Tucuman Prov., R 310, route to El Cajon, 25.xi.2005, J. Torréns Dicoelothorax platycerus ♂ UCRCENT_161497 D#2512 UCRC Argentina: Salta, Cabeza de Buey, RN34, 15.iii.2007, J. & J. Heraty, J. Torréns Dilocantha benneti ♂ UCRCENT_412125 D#3807 UCRC Trinidade: Mt. St. Benedict Trail, 19..vii.2013, Heraty & Baker Dilocantha benneti ♀ UCRCENT_00412138 D#3820 UCRC Trinidade: Mt. St. Benedict Trail, 19..vii.2013, Heraty & Baker Galeria latrellei ♀ UCRCENT_161520 D#2251 UCRC Venezuela: Bolivar, Guyana mer. or., Kavanayen vill. env., 26-28.ii.2005 Isomerala bouceki ♀ UCRCENT_412139 D#3821 UCRC Trinidade: Tucuche Tr., Caura Valley, 24.vii.2013, Heraty & Baker Latina rugosa ♀ UCRCENT_000323 D#2509 IMLA Argentina: Salta Latina rugosa ♂ UCRCENT_00091466 D#1073b UCRC Argentina: Salta, Rosario de la Frontera, Hotel Termal , 21.iii.2003, P. Fidalgo Lasiokapala spiralicornis ♀ UCRCENT_333656 D#3600 IMLA Argentina: Santiago del Estero, La Unión, 20.iii.2012, J. Torréns & J. Fidalgo Lasiokapala spiralicornis ♀ UCRCENT_333654 D#3598 IMLA Argentina: Santiago del Estero, La Unión, 20.iii.2012, J. Torréns & J. Fidalgo Lirata luteogaster ♀ UCRCENT_92211 D#1106 UCRC Equador: Napo, Reserva Etinica Waorani, 7.ii.1999, T.L. Erwin et al. Lirata striatissima ♂ UCRCENT_92235 D#0188 CNC Trinidade e Tobago: Maracas Valley, Loango Village, 22.vi-6.vii.1993, S. & J. Peck Lirata striatissima ♂ UCRCENT_161521 D#2264 UCRC Guiana Francesa: Regina Road Roura-Kaw, vii.2004, O. Morvan Neolirata daguerrei ♂ UCRCENT_91838 D#1067b UCRC Argentina: Formosa, RN 11; south of Formosa, 26.iii.2003, J. Heraty Thoracantha striata ♀ UCRCENT_0092195 D#1254 UCRC Brazil: Rhodonia Prov., Rancho Grande Kapala ipa n. sp. ♀ UCRCENT_91879 D# 0381 UCRC Panamá: Nusagandi Lodge, 21.i.2001, L. Masner Kapala parairidicolor n. sp. ♂ UCRCENT_92059 D#1267 UCRC Honduras: Feo. Morazan, Zamorano Campus, 1.vii.2002, D. Yanega Kapala parairidicolor n. sp. ♀ UCRCENT_2411 D#1895 UCRC Mexico: Chiapas, Rosario Izapa, 11.viii.1997, P. Lachaud Kapala parairidicolor n. sp. ♂ UCRCENT_92075 D#0933c UCRC Honduras: Olancho, El Boquerón Nat. Mon., 2.vii.2002, D. Yanega Kapala iridicolor ♂ UCRCENT_92077 D#0933b UCRC Honduras: Olancho, El Boquerón Nat. Mon., 2.vii.2002, D. Yanega Kapala iridicolor ♂ UCRCENT_91816 D#0928a UCRC Colombia: Magdalena, PNN Tayrona Zaino, 28.iv-13.v.2000, R. Henriquez 174

Kapala iridicolor ♂ UCRCENT_92085 D#938a UCRC Ecuador: Esmeraldas, Bilsa Biol Sta., 10.v-4.vi.1996, P. Hibbs Kapala iridicolor ♂ UCRCENT_92125 D#935a UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs Kapala iridicolor ♀ UCRCENT_282474 D#2917 UCRC Costa Rica: Heredia, La Selva Biol., 14.viii.2010, J. Heraty Kapala iridicolor ♂ UCRCENT_91809 D#0940 UCRC Ecuador: Pichincha, Rio Palenque Science Ctr., 25.iv-6.vi.1996, P. Hibbs Kapala ceciliae n. sp. ♀ UCRCENT_92246 D#1138 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 1.x.1996, T.L. Erwin et al. Kapala ceciliae n. sp. ♀ UCRCENT_247782 D#3831 USNM Ecuador: Orellana, Reserva Etnica Waorani, 20.x.2005, T.L. Erwin, M.C. Pimienta Kapala ceciliae n. sp. ♂ UCRCENT_92008 D#1121 UCRC Ecuador: Orellana, Tiputini Biodiversity Sta., 29.vi.1998, T.L. Erwin et al. Kapala ceciliae n. sp. ♀ UCRCENT_247783 D#3841 USNM Ecuador: Orellana, Rio Piraña Bridge, 20.x.2005, T.L. Erwin, M.C. Pimienta et al Kapala cavicornis n. sp. ♂ UCRCENT_92073 936a UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs Kapala cavicornis n. sp. ♂ UCRCENT_175171 D#0937 UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs Kapala cavicornis n. sp. ♀ UCRCENT_92092 D#0947 UCRC Colombia: Cauca, PNN Gorgona Alto el Mirador, 4-24.iii.2000, R. Duque Kapala cavicornis n. sp. ♂ UCRCENT_92081 D#939a UCRC Ecuador: Pichincha, Rio Palenque Science Ctr., 25.iv-6.vi.1996, P. Hibbs Kapala cavicornis n. sp. ♂ UCRCENT_91869 D#0382a UCRC Panamá: Panama, P.N. Soberania Plantation, 21.i.2001, L. Masner Kapala cavicornis n. sp. ♂ UCRCENT_92074 D#0917 UCRC Colombia: Magdalena, PNN Tayrona Zaino, 28.vi-17.vii.2000, R. Henriquez Kapala cavicornis n. sp. ♂ UCRCENT_92055 D#0382b UCRC Panamá: Panama, P.N. Soberania Plantation, 21.i.2001, L. Masner Kapala cavicornis n. sp. ♂ UCRCENT_235921 D#2781 UCRC Colombia: Chocó, PNN UTRIA C. Visitantes, 5-19.vii.2000, J. Perez Kapala cavicornis n. sp. ♂ UCRCENT_92120 D#0929 UCRC Colombia: Magdalena, PNN Tayrona Zaino, 28.iv.-13.v.2000, R. Henriquez Kapala cavicornis n. sp. ♂ UCRCENT_91804 D#0950 UCRC Colombia: Magdalena, PNN Tayrona Pueblito, 12-28.v.2000 Kapala cavicornis n. sp. ♂ UCRCENT_92072 D#0944a UCRC Colombia: Magdalena,, PNN Tayrona Pueblito, 29.vi-14.vii.2000, R. Henriquez Kapala cavicornis n. sp. ♀ UCRCENT_412137 D#3819 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker Kapala cavicornis n. sp. ♂ UCRCENT_412134 D#3816 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker Kapala cavicornis n. sp. ♂ UCRCENT_412135 D#3817 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker Kapala cavicornis n. sp. ♂ UCRCENT_412136 D#3818 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker Kapala furcata ♀ UCRENT_92093 D#1078 UCRC Argentina: Misiones, Santa Ana, near Loreto, 27.iii.2003, J. Heraty Kapala deltalis n. sp. ♂ UCRCENT_92109 D#1388 UCRC Costa Rica: Guanacaste, Guanacaste N.P., Biol. Sta. Cacao, 13.ii.1995, L. Masner Kapala quasimodo n. sp. ♂ UCRCENT_92002 D#0942 UCRC Ecuador: Esmeraldas, Bilsa Biol. Station, 7-19.vi.1996, P. Hibbs Kapala parafurcata n. sp. ♂ UCRCENT_91921 D#1069 UCRC Argentina: Misiones, Santa Ana, near Loreto, 27.iii.2003, J. Heraty Kapala parafurcata n. sp. ♂ UCRCENT_161498 D#2519 UCRC Argentina: Misiones, Rt17, E of 9 de Julio, 26.iii.2007, J. Heraty Kapala parafurcata n. sp. ♂ UCRCENT_92073 D#1068a UCRC Argentina: Salta, Oran, Rd to San Andres along Rio Blanca, 22.iii.2003, J. Heraty Kapala parafurcata n. sp. ♂ UCRCENT_397224 D#3436 UCRC Paraguay: Caazapá, San Rafael Reserve, 8-10.xii.2000, Z.H. Falin Kapala parafurcata n. sp. ♂ UCRCENT_000324 D#2520 UCRC Argentina: Misiones, Rt17, E of 9 de Julio, 25.iii.2007, J. Heraty Kapala parafurcata n. sp. ♀ UCRCENT_91817 D#0711 UCRC Argentina: Misiones, Loreto, Ruinas Jesuiticas, 4.xi.2001, S.O. Martinez, P. Fidalgo 175

Kapala parafurcata n. sp. ♀ UCRCENT_000325 D#2518 UCRC Argentina: Misiones, RN 12, N of Puerto Bosseti, 25.iii.2007, J. Heraty & J. Torréns Kapala parafurcata n. sp. ♀ UCRCENT_91803 D#1086 UCRC Argentina: Misiones, Santa Ana, near Loreto, 27.iii.2003, J. Heraty Kapala cuprea ♂ UCRCENT_412124 D#3806 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker Kapala cuprea ♂ UCRCENT_412123 D#3805 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker Kapala cuprea ♀ UCRCENT_412122 D#3804 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker Kapala cuprea ♂ UCRCENT_412133 D#3815 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker Kapala cuprea ♀ UCRCENT_91807 D#1003 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 2.vii.1995, T.L. Erwin et al. Kapala cuprea ♂ UCRCENT_320768 D#3837 CNC Ecuador: Sucumbíos, Napo River, Sacha Lodge, 3-13.vi.1994, P. Hibbs Kapala cuprea ♀ UCRCENT_247775 D#3838 USNM Ecuador: Orellana, Rio Piraña Bridge, 17.x.2005, T.L. Erwin, M.C. Pimienta et al Kapala cuprea ♂ UCRCENT_320853 D#3852 CNC Ecuador: Sucumbíos, Napo River, Sacha Lodge, 4-14.v.1994, P. Hibbs Kapala cuprea ♂ UCRCENT_92227 D#1004 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 9.vi.1995, T.L. Erwin et al. Kapala cuprea ♂ UCRCENT_247778 D#3833 USNM Ecuador: Orellana, Reserva Etnica Waorani, 17.x.2005, T.L. Erwin, M.C. Pimienta Kapala argentina ♀ UCRCENT_91827 D#1076 UCRC Argentina: Salta, Rosario de la Frontera, Hwy 34, 21.iii.2003, J. Munro Kapala argentina ♀ UCRCENT_10184 D#1064 UCRC Argentina: Salta, Rosario de la Frontera, 21.iii.2003, J. Heraty Kapala argentina ♂ UCRCENT_282476 D#2920 UCRC Argentina, Salta, RN9 Rosario de la Frontera, 13-14.iii.2007, J. Heraty & J. Torréns Kapala flabellata ♂ UCRCENT_320837 D#3872 UCRC Guiana Francesa: PK35, x.2007, J. Cerda Kapala floridana ♂ UCRCENT_92111 D# 0432b UCRC USA: Florida, Marion Co., Juniper Spr Rd., 19.ix.2001, J. Heraty Kapala haplospinosa ♀ UCRCENT_172257 D# 2784 UCRC USA: Florida, Marion Co., Big Pine Key, SW 1/4, S4, 1-31.viii.1986, S. & J. Peck Kapala haplospinosa ♀ UCRCENT_161512 D#2261 UCRC Republica Dominicana: Punta Cana, Reserva, 11-14.xi.2005, L. Masner Kapala ivorensis ♀ UCRCENT_302068 D#2746 UCRC Kenya: Coast, Arabuko-Sokoke Forest, 8-9.i.2010, R. Copeland Kapala ivorensis ♀ UCRCENT2_41581 D#2922 UCRC Republic of Congo: Pool Dept., Abio, Lesio-Louna Pk., 11-18.ix.2008, Sharkey & Braet Kapala ivorensis ♂ UCRCENT_241604 D#2924 UCRC Republic of Congo: Pool Dept., Abio, Lesio-Louna Pk., 30.ix-7.x.2008, Sharkey & Braet Kapala izapa ♂ D#2786 UCRC Mexico: Quintana Roo, Reserva Ecologia El Eden, 19.viii.1998, R. Rodriguez Kapala izapa ♂ UCRCENT_397279 D#2926 UCRC Mexico: Chiapas, Playón de la Gloria, 24.vi.2008 Kapala izapa ♂ UCRCENT_356033 D#2925 UCRC Mexico: Chiapas, Playón de la Gloria, 24.vi.2008 Kapala romandii ♂ UCRCENT_92221 D#0384C CNC Panama: 2 km S torti, Serrania de Maje, 18.i.2001, L. Masner Kapala terminalis ♂ UCRCENT_161511 D# 2260 UCRC Dominican Republic: Punta Cana, Reserva, 11-14.xi.2005 Kapala terminalis ♂ UCRCENT_00091880 D# 1270 UCRC Dominican Republic: Puerto Plata, 32.i.1989, L. Masner Kapala terminalis ♂ UCRCENT_397249 D# 3435 UCRC Puerto Rico: Cerezos, circle 38, 28.II.2004, L Yunes

176

Tabela S3. Lista dos gêneros e espécies do grupo externo e interno utilizados na análise filogenética molecular, sexo, número de identificação de depósito (Numb. ID), DNA voucher, museu de deposito e localidade de coleta.

Genes Espécies Sexo Num. ID DNA voucher 18S 28S-D2 28S-D3-5 COI-nj COII Chalcura sp. UCRCENT_91769 D#0646a x x x x Schizaspidia aenea UCRCENT_91398 D#0168 x x x x x Colocharis napoana ♂ UCRCENT_92220 D#1104 x x x x x Colocharis napoana ♀ UCRCENT_91450 D#1102 x x x x x Torquata n. gen. ♂ UCRCENT_10304 D#1141 x x x x x Torquata n. gen. ♂ UCRCENT_92029 D#1173 x x x x x Torquata n. gen. ♂ UCRCENT_397251 D#3440 x x x x x Ecarinta n. gen. ♀ UCRCENT_00483995 D#4898 x x x x x Dicoelothorax platycerus ♂ UCRCENT_161497 D#2512 x x x Dilocantha benneti ♂ UCRCENT_412125 D#3807 x x x x x Dilocantha benneti ♀ UCRCENT_00412138 D#3820 x x x x x Galeria latrellei ♀ UCRCENT_161520 D#2251 x x x x x Isomerala bouceki ♀ UCRCENT_412139 D#3821 x x x x x Latina rugosa ♂ UCRCENT_00091466 D#1073b x x x x x Latina rugosa ♀ UCRCENT_000323 D#2509 x x x Lasiokapala spiralicornis ♀ UCRCENT_333656 D#3600 x x x x x Lasiokapala spiralicornis ♀ UCRCENT_333654 D#3598 x x x x x Lirata luteogaster ♀ UCRCENT_92211 D#1106 x x x x x Lirata striatissima ♂ UCRCENT_92235 D#0188 x x x x x Lirata striatissima ♂ UCRCENT_161521 D#2264 x x x x x Neolirata daguerrei ♂ UCRCENT_91838 D#1067b x x x x Thoracantha striata ♀ UCRCENT_0092195 D#1254 x x x Kapala ipa n. sp. ♀ UCRCENT_91879 D# 0381 x x x x x Kapala parairidicolor n. sp. ♂ UCRCENT_92059 D#1267 x x x x x Kapala parairidicolor n. sp. ♀ UCRCENT_2411 D#1895 x x x Kapala parairidicolor n. sp. ♂ UCRCENT_92075 D#0933c x x x x x Kapala iridicolor ♂ UCRCENT_92077 D#0933b x x x x x 177

Kapala iridicolor ♂ UCRCENT_91816 D#0928a x x x x x Kapala iridicolor ♂ UCRCENT_92085 D#0938a x x x x x Kapala iridicolor ♂ UCRCENT_92125 D#0935a x x x x x Kapala iridicolor ♀ UCRCENT_282474 D#2917 x x x x x Kapala iridicolor ♂ UCRCENT_91809 D#0940 x x x x Kapala ceciliae n. sp. ♀ UCRCENT_92246 D#1138 x x x x x Kapala ceciliae n. sp. ♀ UCRCENT_247782 D#3831 x x x Kapala ceciliae n. sp. ♂ UCRCENT_92008 D#1121 x x x x Kapala ceciliae n. sp. ♀ UCRCENT_247783 D#3841 x x x Kapala cavicornis n. sp. ♂ UCRCENT_92073 D#0936a x x x x x Kapala cavicornis n. sp. ♂ UCRCENT_175171 D#0937 x x x x x Kapala cavicornis n. sp. ♀ UCRCENT_92092 D#0947 x x x x x Kapala cavicornis n. sp. ♂ UCRCENT_92081 D#0939a x x x x Kapala cavicornis n. sp. ♂ UCRCENT9_1969 D#0920 x x x x x Kapala cavicornis n. sp. ♂ UCRCENT_91869 D#0382a x x x x x Kapala cavicornis n. sp. ♂ UCRCENT_92074 D#0917 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_92055 D#0382b x x x x Kapala cavicornis n. sp. ♂ UCRCENT_92120 D#0929 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_91804 D#0950 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_92072 D#0944a x x x x Kapala cavicornis n. sp. ♀ UCRCENT_412137 D#3819 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_412134 D#3816 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_412135 D#3817 x x x x Kapala cavicornis n. sp. ♂ UCRCENT_412136 D#3818 x x x x Kapala furcata ♀ UCRENT_92093 D#1078 x x x x x Kapala deltalis n. sp. ♂ UCRCENT_92109 D#1388 x x x x x Kapala quasimodo n. sp. ♂ UCRCENT_92002 D#0942 x Kapala parafurcata n. sp. ♂ UCRCENT_91921 D#1069 x x x x x Kapala parafurcata n. sp. ♂ UCRCENT_161498 D#2519 x x x Kapala parafurcata n. sp. ♂ UCRCENT_92073 D#1068a x x x x x Kapala parafurcata n. sp. ♂ UCRCENT_397224 D#3436 x x x x Kapala parafurcata n. sp. ♂ UCRCENT_000324 D#2520 x x x 178

Kapala parafurcata n. sp. ♀ UCRCENT_91817 D#0711 x x x x Kapala parafurcata n. sp. ♀ UCRCENT_000325 D#2518 x x Kapala parafurcata n. sp. ♀ UCRCENT_91803 D#1086 x x x x Kapala cuprea ♂ UCRCENT_412124 D#3806 x x x x Kapala cuprea ♂ UCRCENT_412123 D#3805 x x x x Kapala cuprea ♀ UCRCENT_412122 D#3804 x x x x Kapala cuprea ♂ UCRCENT_412133 D#3815 x x x x Kapala cuprea ♀ UCRCENT_91807 D#1003 x x x x x Kapala cuprea ♂ UCRCENT_320768 D#3837 x x x Kapala cuprea ♀ UCRCENT_247775 D#3838 x x x Kapala cuprea ♂ UCRCENT_320853 D#3852 x x x Kapala cuprea ♂ UCRCENT_92227 D#1004 x x x x x Kapala uúprea ♂ UCRCENT_247778 D#3833 x x x Kapala argentina ♀ UCRCENT_91827 D#1076 x x x x x Kapala argentina ♀ UCRCENT_10184 D#1064 x x x x Kapala argentina ♂ UCRCENT_282476 D#2920 x x x x Kapala flabellata ♂ UCRCENT_320837 D#3872 x x x x Kapala floridana ♂ UCRCENT_92111 D# 0432b x x x x x Kapala haplospinosa ♀ UCRCENT_172257 D# 2784 x x Kapala haplospinosa ♀ UCRCENT_161512 D#2261 x x x x x Kapala ivorensis ♀ UCRCENT_302068 D#2746 x x x x x Kapala ivorensis ♀ UCRCENT_241581 D#2922 x x x x Kapala ivorensis ♂ UCRCENT_241604 D#2924 x x x x Kapala izapa ♂ D#2786 x x x Kapala izapa ♂ UCRCENT_397279 D#2926 x x x Kapala izapa ♂ UCRCENT_356033 D#2925 x x x x x Kapala romandii ♂ UCRCENT_92221 D#0384C x x x x Kapala terminalis ♂ UCRCENT_161511 D# 2260 x x x x x Kapala terminalis ♂ UCRCENT_91880 D# 1270 x x x x x Kapala terminalis ♂ UCRCENT_397249 D# 3435 x x x

179

Tabela SI 4. Primer oligonucleotídeos utilizados neste estudo. Sequencias marcadas com “*” foram modificadas a partir das referências da publicação original.

Gene Primer Sequence References 18S 18S F (mid) 5’-AAA TTA CCC ACT CCC GGC A-3’ Munro et al., 2011 18S R (mid) 5’-TGG TGA GGT TTC CCG TGT T-3’ Munro et al., 2011 18Si F (inside mid) 5’-ATC GCT CGC GAT GTT TAA CT-3’ Heraty et al., 2004 18Si R (inside mid) 5’-AGA ACC GAG GTC CTA TTC CA-3’ Heraty et al., 2004 18S1 F (5’end) 5’-TAC CTG GTT GAT CCT GCC AGT-3’ Ouvrard et al., 2000* 18S4 R (5’end) 5’-GAA TTA CCG CGG CTG CTG G-3’ Schulmeister et al., 2003 18Sa F (3’end) 5’-ATG GTT GCA AAG CTG AAA C-3’ Schulmeister et al., 2003 18S9 R (3’end) 5’-GAT CCT TCC GCA GGT TCA CCT-3’ Ouvrard et al., 2000* 28S D2 D2-3551F 5’-CGG GTT GCT TGA GAG TGC AGC-3’ Campbell et al., 2000* D2Ra R 5’-CTC CTT GGT CCG TGT TTC-3’ Campbell et al., 2000* 28S D3-D5 D3-4046 F 5’-TTG AAA CAC GGA CCA AGG AG-3’ Nunn et al., 1996* D3-4413 R 5’-TCG GAA GGA ACC AGC TAC TA-3’ Nunn et al., 1996* D5-4625 R 5’-CGC CAG TTC TGC TTA CCA-3’ Schulmeister et al., 2003 COI COI-nj F 5’-TAT ATT TTA ATT YTW CCW GGA TTTGG-3’ Simon et al., 1994* COI-MD R 5’-ATT GCA AAT ACT GGA CCT AT-3’ Dowton & Austion, 1997* COII COII-MTD16 F 5’-ATT GGA CAT CAA TGA TAT TGA-3’ Simon et al., 1994* COII-MTD18 R 5’-CCA CAA ATT TCT GAA CAT TGA CCA-3’ Dowton & Austion, 1997*

180

Capítulo III

Schoeninger, K., Torréns, J., Heraty, J.M., Oliveira, M.L. (2018) Revision of Kapala iridicolor species complex (Hymenoptera:

Chalcidoidea: Eucharitidae) with a first phylogenetic approach.

181

Revision of Kapala iridicolor species complex (Hymenoptera: Chalcidoidea: Eucharitidae) with a first phylogenetic approach

KARINE SCHOENINGER1,4, JAVIER TORRÉNS2, JOHN M. HERATY3 & MARCIO L. OLIVEIRA1

1Coordenação de Biodiversidade, Programa de Pós-Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia, CEP 69067-375, Manaus, AM, Brazil. 2CRILAR-CONICET, Entre Ríos y Mendoza, 5301 Anillaco, La Rioja, Argentina. 3Department of Entomology, University of California, Riverside, CA 92521, United States. 4Corresponding author. [email protected]

Short title. Revision of Kapala iridicolor species complex

Abstract. The genus Kapala Cameron (Hymenoptera: Eucharitidae) is numerically abundant and diverse in the Neotropical region. However, its taxonomy is confusing due to the large intraspecific morphological variation, as well as the absence of limits for the diagnosis of the species. Due the importance of establishment the new boundaries of the paraphyletic Kapala, we deem it a primary task to revisit the type species and establish diagnostic characters for identification of species of the K. iridicolor clade. As results, were recognize six species within the iridicolor complex, including K. iridicolor described by Cameron (1904). New species described here include: K. cavicornis Schoeninger & Heraty, n. sp., K. ceciliae Schoeninger & Torréns n. sp., K. ipa Schoeninger & Heraty n. sp., K. longicornis Schoeninger & Oliveira n. sp. and K. parairidicolor Schoeninger & Torréns n. sp. A diagnosis for the K. iridicolor complex is provided, as well as a molecular phylogeny for the species based on 18S, 28S-D2, 28S-D3-D5, COI and COII. Also, an identification key for the species of the iridicolor complex is provided.

Keywords. Ant parasitoids, identification key, Neotropical region, phylogeny, taxonomy.

182

Introduction

Kapala Cameron, 1884 is widespread throughout the Neotropical region, except for Chile and one disjunct species is found in Afrotropical region and Madagascar (Herary, 1985, 2002; Heraty & Woolley, 1993; Murray & Heraty, 2016). This genus exhibits a large amount of external morphological variations and the boundaries need to be better defined. Kapala is phenotypically distinctive from related genera of Kapala clade (13 genera in total) and until now there are no morphological or molecular synapomorphies to unite the genus (Heraty, 2002; Murray, 2014). Kapala is numerically abundant and diverse with 18 valid species (Heraty, 2002; Murray, 2014). According to Heraty (2002) this genus currently is defined by a suite of symplesiomorphies including frenal spines lyre-shaped with longitudinal or spiral carina (except in Kapala chacoencis Gemignani, 1947 whose process is soft without defined carina), eyes bare, male antennal branches 0.8‒1.8X longer than height of head, female basal flagellomeres rarely more than twice as long as broad and posterior margin of the mesoscutellar disc raised to a prominent ridge between the bases of the frenal processes (however, this is considered to have been lost in a few species, for example, in K. floridana (Ashmead, 1885) and K. terminalis Ashmead, 1892). Species of Kapala can be divided into three complexes based on both morphological and molecular data: K. furcata, K. iridicolor and K. romandii (Murray, 2014; Schoeninger et al., in prep.). Based on molecular data only, these clades are separated phylogenetically by clusters of morphologically distinct genera (Murray, 2014; Schoeninger et al., in prep.) The K. iridicolor complex is considered monophyletic with high support in molecular and morphological analysis (Murray, 2014; Schoeninger et al., in prep.) Specimens of this complex have the face smooth, frons slightly swollen, mesoscutum with a low profile, lateral lobes of mesoscutum densely setose and the mesoscutellum elongated (Heraty & Woolley, 1993). However, the extreme variation in morphological characters and the great similarity between species make this species group an especially difficult problem for taxonomy of Kapala. Some of these variations were previously observed by Heraty and Wolley (1993) in specimens from Ecuador and Trinidad. These variations contemplate the apex of the frenal process, which are broad and strongly emarginate in specimens from Trinidad and gradually narrowed to a fine point in specimens from Ecuador. 183

Due the importance of establishment the new boundaries of the paraphyletic Kapala, we deem it a primary task to revisit the type species and establish diagnostic characters for identification of species of K. iridicolor complex. In this study, we defined the K. iridicolor complex, described five new species, as well as a molecular phylogeny for the species based on 18S, 28S-D2, 28S-D3-D5, COI and COII. Also, an identification key for the species of the K. iridicolor complex is provided.

Materials and methods

Molecular phylogenetic analysis

The molecular matrix has a total of 35 specimens (Tables S1, S2), with 30 from K. iridicolor complex, representing five species. Outgroup taxa are from the K. furcata complex with three species (K. furcata, K. cuprea and Kapala. sp. n. sp.). The species are from 9 countries, with only Kapala longicornis n. sp. with no molecular data. The matrix includes three nuclear ribosomal (partial 18S, 28S-D2 and 28S-D3-D5) and two mitochondrial (COI-nj and COII) gene fragments. For the DNA extractions, both fresh and dried specimens were used. Specimens were non-destructively extracted using a chelex-proteinase-K-protocol or DNeasy (Qiagen). Chelex-proteinase-K was performed (Walsh et al., 1991) using non-destructive sampling to preserve the integrity of the parasitoid wasps for retainment as voucher specimens. Portions of five gene regions were amplified: 18S, 28S-D2, 28S-D3-D5, COI and COII; primers are reporter in Table (S3). Qiagen (Valencia, CA) reagents were used for PCR. Sequencer 5.4.6 (Gene Codes Corp, Ann Arbor, MI) was used to edit chromatograms to final sequences; primers were not included in the final sequence counting. All molecular data were obtained from a unique DNA voucher number (D#), which are deposited in the internal laboratory FileMaker v.11® database maintained at UCR. Individual genes were aligned using the MAFFT online server (Katoh et al., 2005) under default settings. For nuclear ribosomal genes, the E-INS-I algorithmic strategy was applied, and for mitochondrial genes, the G-INS-I strategy was applied. SequenceMatrix 1.7.7 (Vaidya et al., 2011) was used to concatenated genes for a final matrix. The dataset was portioned by gene, and 184

COI and COII were split into 1+2 and 3. Ribosomal and mitochondrial genes were then concatenated for a final alignment length of 2588 bp. Maximum likelihood (ML) using RaxML v8.0.24 (Stamatakis et al., 2008) was implemented through the CIPRES Science Gateway (Miller et al., 2010). One thousand rapid bootstrap replicates were performed, with other parameters keep at default.

Taxonomy

Morphological descriptions

The funiculars include Fl2 and the following unfused flagellomeres before the clava, which is defined by 1–3 fused flagellomeres. The paired spines of the mesoscutellum arise from the frenum and are refered here as frenal process (character 50; Heraty 2002). Morphological abbreviations (see Figs 1 and 2, Chap. I): acl, anteclypeus; acr, acropleuron; ao, anterior ocellus; ax, axilla; axg, axillular groove; axl, axillula; cal, callus; clv, clava; cly, clypeus; cx, coxa; es2, mesepisternum; Fl2, flagellomere basal; flg, flagellum; fmd, femoral depression; frl, frenal line, fp, frenal processes; fub, funicle; gen, gena; Gt1, first gastral tergite; iod, interocular distance; lep2, lower mesepimeron; llm, lateral lobe of mesoscutum; mlm, midlobe of mesoscutum; msp, malar space; mv, marginal vein; not, notaulus; pdl, pedicel; pet, petiole; pl3, metapleuron; pmv, postmarginal vein; po, posterior ocellus; ppd, propodeum; pre, prepectus; sca, supraclypeal area; scd, scrobal depression, scp; scape, sct, mesoscutellar disc; smv, submarginal vein; sss, scutoscutellar suclcus; stv, stigmal vein; tgl, tegula; tor, torulus; tsa, transscutal articulation; upe2, upper mesipimeron.

Measurements

brackets], followed by the range. All measurements are given in millimeters and follows (Schoeninger et al., in prep Cap. I).

Specimen imaging

Specimens repository

The following institutions served as source of material and type depositories for specimens examined for this study: The Natural History Museum, London, England (BMNH); Canadian National Collection of Insects, Arachnids and Nematods, Ottawa, Ontario, Canada (CNC); Cornell University Insect Collection, Ithaca, New York, USA (CUIC); University of Guelph Insect Collection, Guelph, Ontario, Canada (DEBU); Escuela Agricola Panamericana, Tegucigalpa, Honduras (EAZP); Essig Museum of Entomology, University of California, Berkeley, California, USA (EMEC); Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, Florida, USA (FSCA); Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (INPA); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ); Mississippi Museum of Natural Science, Mississippi, USA (MNCN); Museo Nacional da Costa Rica (MNCR); Lund University, Lund, Sweden (MZLU); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZSP); National Museum of Natural History, Washington D.C., USA (NMNH); Royal Ontario Museum, Toronto, Ontario, Canada (ROME); Iziko Museum of Capetown, Cape Town, South Africa (SAMC); Universidad Autónoma Nuevo León, Facultad de Ciencias Forestales, Linares, Mexico (UANL); University of California, Davis, California, USA (UCDC); University of California, Riverside, California, USA (UCRC); Coleção Entomológica, Departamento de Biologia, Universidade Federal do Espírito Santo, Brazil (UFES); University of Michigan, Museum of Zoology, Ann Arbor, Michigan, USA (UMMZ); Zoologisches Institute und Zoologisches 186

Museum, Universität von Hamburg, Hamburg, Germany (ZMUH); Zoologische Staatssammlung, Munchen [= Munich], Germany (ZSM); W.F. Barr Entomological Collection, University of Idaho, Moscow, Idaho, USA (WFBM). Most specimens are identified with unique UCRC specimen identifiers that indicate the museum of deposition. All specimens were databased in a FileMaker v.11® database maintained in the Heraty lab. Localities were georeferenced using Google Earth (coordinates in italics) or from the original label data. The distribution map was constructed in Simple Mappr (Shorthouse, 2010).

Results and discussion

Phylogeny

As observed in previous analysis (Schoeninger et al., in prep., Chapter II) the K. iridicolor species complex is confirmed as monophyletic with 100% bootstrap support. All described species appear as monophyletic in the ML analysis. Much of the diversity of the iridicolor species complex is found in South America and, mainly in Central America (Fig. 7). According to the ML analysis we can verify that Kapala ipa n. sp. + Kapala parairidicolor n. sp. correspond as the sister group of the rest of the K. iridicolor complex with 60% bootstrap support. Both species have peculiar characteristics that separate them from the rest of the clade. K. ipa n. sp. is the only species whose frons are not swollen, in addition the females have the long and 10‒segmented flagellum; since the males have the long branch F2, different from the other species of the clade, that have the branch F2 shorter. However, males of K. ipa n. sp. may have the frons with faint longitudinal striae. This latter characteristic is also observed in females and males of K. parairidicolor n. sp. (100% bootstrap support, Fig. 1), however the striations in this species are well marked and may extend to the genae in some specimens from Mexico. Kapala iridicolor is distinguished from the rest by the frons swollen and frenal process very thin with apex acuminate (Fig 4A). In a study by Heraty and Woolley (1993), which addressed the problem of the polymorphism between the species of K. iridicolor and K. romandii Cameron and intermediates, specimens verified a geographic variation occurring within the population that is more apparent 187

in the frenal process. The frenal processes are thick with apex broad and strongly emarginate in specimens from Trinidad and gradually narrowed to a fine point in specimens from Ecuador. According to the ML analysis and an extensive morphological observation of these species, the specimens from Ecuador correspond to K. iridicolor, since this is the only one within the clade that has the fine frenal processes with the apex ending in a fine point (acuminate) and the specimens from Trinidad correspond to a new species K. cavicornis n. sp., described here. Thus, the present study corroborates with the observations made by Heraty and Woolley (1993). In addition, some populations of K. cavicornis n. sp., mainly from Colombia and Panama, comprise some small internal clades that could indicate new species, but morphologically they do not present any external phenotypic variation, reason why it was decided to maintain them as an only species. Kapala ceciliae n. sp. is recognized by a set of characteristics such as long antenna, frenal process narrows with apex emarginated and process with longitudinal striae, which become slightly oblique in apical half (Fig. 4C). This species has a bootstrap support of 100% and occurs only in Ecuador. Although, K. longicornis n. sp. has no molecular data, it fits perfectly into the K. iridicolor complex because it has the swollen and smooth frons (Fig. 2A), the mesoscutum low and smoothly curved anteriorly, in lateral view as well as an elongated scutellum. But, it differs from the others species by the frenal process long exceeding the metasoma and females with flagellum long and F2 much longer than F3.

Taxonomy

Kapala Cameron, 1884

Kapala; Kirby, 1886: 32. [Kirby questions the identity of Brullè’s drawing and suggests that it looks closer to Thoracantha atrata Walker than to either of the Fabricius species. Indeed, the drawing appears closer to Schizaspidia than to what is now recognized as Kapala. The figure legend in Brullè (1846: pl. 38) refers to Chirocerus furcatus Westwood, not Fabricius, and may not, unfortunately, refers to the Fabricius species. Kapala; Ashmead, 1888[45]: 187 (key). Kapala; Ashmead, 1904[238]: 269,270 (key), 472. Capala Schulz, 1906: 150. Unjustified emendation. Holcokapala Cameron, 1913: 117. Synonymy by Heraty, 2002: 167. Kapala; Gemignani, 1933: 485. Subsequent description. Kapala; Heraty, 1985: 84. Subsequent description. Kapala; Heraty, 2002: 167-170. Subsequent description.

Kapala iridicolor species complex

Diagnosis. The iridicolor species complex can be defined by a combination of the following characters: frons swollen (except for Kapala ipa n. sp., which is flat), usually smooth (wide longitudinal striated, only in K. parairidicolor n. sp.) (Figs. 2A, B) and with sparse setae, more prominent on upper frons and vertex; lower face smooth; females with flagellum 9‒10 segmented. Mesoscutum low and smoothly curved anteriorly in lateral view (Fig. 5A, indicated); lateral lobe of mesoscutum smooth and pilose. Scutellum elongated. The dorsolateral projection of prepectus is an elongate triangle, broadly rounded at the apex.

Key to the species of the K. iridicolor complex

1 Frons not swollen (Fig. 3D); antenna of female with 10 flagellomeres (Fig. 3D), flagellum 1.2– 1.3x head high; male with length of branch of F2 > 1.2x head height. Both sexes with axilla and mesoscutellar disc with fine longitudinal carinae close to each other (Fig. 4D) … Kapala ipa n. sp. - Frons swollen (Figs. 3A; 4A, B dorsal view); antenna of female with 9 flagellomeres (Fig. 3B), flagellum 0.9–1.0x head high; male with length of branch F2 < 1.2x head height. Both sexes with axilla and mesoscutellar disc with wide longitudinal carinae widely spaced (Figs. 4B, E) … 2 189

2(1) Both sexes with frons longitudinal striated (Fig. 2B), which extends to the genae, but more pronounced in males … Kapala parairidicolor n. sp. - Both sexes with frons completely smooth (Fig. 2A) … 3

3(2) Frenal processes long, exceeding the length of the metasoma (Figs. 4E; 6C, D); female with

Fl2 long and swollen at apex and 1.7x as long as following flagellomere (Fig. 3E); eyes separated by 2.3-2.4x their height … Kapala longicornis n. sp. - Frenal process short, not exceeding the length of the metasoma (Figs. 4A, B; 5A, C); female with F2 1.1–1.5x as long as following flagellomere; eyes separated by 1.9–2.3x their height … 4

4(3) Female with flagellum 1.1–1.2x head height (Fig. 3C); male with branch of F2 1.5x head height (Fig. 5F). Both sexes with frenal process longitudinally striated become slightly oblique in apical half (Fig. 4C) … Kapala ceciliae n.sp. - Female with flagellum 0.8–1.0x head height (Figs. 3A, B); male with branch of F2 1.0‒1.2x head height (Figs. 5B). Both sexes with frenal process longitudinally striated (Figs. 4A, B) … 5

5(4) Frenal process cylindrical and thin, with apex of frenal process acuminate (Fig. 4A) … Kapala iridicolor (Cameron, 1904) - Frenal process cylindrical and thick, with apex emarginated with an elongate flange like spoon- shaped (Fig. 4B, indicated) … Kapala cavicornis n. sp.

Kapala iridicolor (Cameron, 1904) (Figs. 3A, 4A, 5A‒B)

Lirata iridicolor Cameron, 1904: 60. Type data: Nicaragua: Chinandega. Holotype ♀, by monotype. Type depository: Natural History Museum (BMNH), London, England. Type nº 5.385 (examined). [Holotype relatively complete: right medial leg broken after tibia; left hind leg broken after the coxa and glued to the side of the body; metasoma broken and lost]. Kapala iridicolor; Heraty & Woolley, 1993: 522. [description of both sexes, illustrated.].

190

Diagnosis. Recognized by a combination of the following characters: frons swollen and smooth; mesoscutum smoothly curved anteriorly (Fig. 5A, indicated); frenal process cylindrical and very thin with apex acuminate (Fig. 4A); female with mesoscutellar disc with striae widely spaced and a short flagellum 0.8–0.9x head height (Fig. 3A).

Biological notes. Known to oviposit into closed flower buds of Cordia macrostachya, Cordia cana (Boraginaceae) and Gossypium hirsutum (Malvaceae) (Heraty & Woolley, 1993). Also collected on Gossypium hirsutum L., Hamelia patens Jacq., Melampodium divaricatum L., Onoseris onoseroides (Kunth.) B.L. Rob., Panicum sp., Phoradendron sp.; fructification of Carica papaya L., Passiflora edulis L., Terminalia catappa L.; flowering cactus-like Asclepiad

Distribution. Brazil, Colombia, Costa Rica, Ecuador, El Salvador, Honduras, Mexico, Nicaragua, Panama, Suriname, Peru and Venezuela.

Holotype. Nicaragua: ♀, Chinandega, Baker coll., [BMNH type n°5.385, UCRCENT 310008: BMNH].

Additional material examined. Brazil: 1♂, Roraima: Ilha de Maracá, 3°24'N 61°42'W, 02- 13.v.1987, Malaise, J.A Rafael & equipe, [INPA]. Colombia: 1♂, Magdalena, PNN Tayrona Zaino, 50m, 11°20'01"N, 74°02'01"W, 14-29.vi.2000, Malaise trap, R. Henriques, [UCRCENT00172493: UCRC]. 1♂, Boyacá, Muzo, 5°32'00"N, 74°06'00"W, Westwood, [UCRCENT00245267: ZSMC]. 1♀, Cauca Valley, near Palmira, 1000m, 3°35'02"N, 76°15'01"W, v.1986, A. Braun, [UCRCENT00243536: UCDC]. 1♀, 11 km W. Santa Marta, Pozo Colorado, 11°10'00"N, 74°14'00"W, 25-30.iv.1968, Borys Malkin, [UCRCENT00236312: EMEC]. 2♂, PNN Tayrona Zaino, 50m, 11°20'00"N, 74°02'00"W, 13-30.v.2000, Malaise trap, R. Henriquez, [ -, UCRCENT00092217: UCRC]. 1♀, Valle del Cauca, Cadelaira, Finca San Luis, 1010m, 3°24'00"N, 76°23'00"W, 4.vii.1975, tropical dry forest, Malaise trap, R.C. Wilkerson, [UCRCENT00306477: FSCA]. 1♂, Central de Anchicaya, 3°52'56"N, 76°48'02"W, 28.i.1975, Malaise trap, R. Wilkerson, [UCRCENT00306443: FSCA]. 1♀, Valle Rio Anchicaya, -, 400m, 10.ii.1977, Breed & C.D. Michener, [UCRCENT00446413: UCRC]. Costa Rica: 1♀, Curubande, 10°43'00"N, 85°25'00"W, xi.1981, forest edge, sweep, S.G. Compton, [SAMC: 191

UCRCENT00246690]. 1♀, Guardia, 40m, 10°33'54"N, 85°35'18"W, 21.vii.1967, A.D. Banegas, [UCRCENT00236307: EMEC]. 1♂, Alajuela, Chiles de Aguas Zarcas, café, 300m, 10°23'56"N, 84°20'20"W, i.1990, R. Cespedes, [UCRCENT00274871: UCRC]. 1♂ 1♀, idem but xii.1989, [UCRCENT00302321‒22: UCRC]. 2♂, San Pedro de la Tigra, Cacao, 200m, 10°22'07"N, 84°34'55"W, i-ii.1990, R. Cespedes, [UCRCENT00274879, UCRCENT00302325: UCRC]. 1♀ 5♂, idem but iii-iv.1990, [UCRCENT00243537, UCRC: UCRCENT00274876‒78, UCRCENT00302324, UCRCENT00302326: UCDC]. 4♀, Guanacaste, ACG, Santa Rosa, 300m, 10°50'22"N, 85°37'06"W, 8-18.vi.1995, S. Dadelahi, L. Price & N. Zitani, [UCRCENT00318838‒40, UCRCENT00320089: UCRC]. 5♂, Cerro el Hacha, NW Volcan Orosi, 300m, 10°59'57"N, 85°33'03"W, 1988, [UCRCENT0027487‒75, UCRCENT00302323: UCRC]. 1♀, La Pacifica, 4 km. N.W. Canas, 10°27'16"N, 85°07'54"W, 8.ix.1971, Cordia cana, [UCRCENT00236290: EMEC]. 1♂, Santa Rosa NP, 300m, 10°53'33"N, 85°45'59"W, 16.xi- 7.xii.1985, D.H. Janzen & I.D. Gauld [UCRCENT00320091: UCRC]. 1♂, Santa Rosa NP Bosq. Hum. (11O), 10°53'33"S, 85°45'59"W, [UCRCENT00274882: UCRC]. 1♂, Santa Rosa NP Sn. Emilio (6C), 10°52'15"N, 85°43'38"W, [UCRCENT00274880: UCRC]. 1♂, F. La Selva 3km S Pto. Viejo, 80m, 10°26'00"N, 84°01'00"W, 29-31.vii.1976, E.M. Fisher, [UCRCENT00320316: UCRC]. 1♀, La Selva Biol. Sta., 64m, 10°25'49"N, 84°00'26"W, 14.viii.2010, Hamelia patens and nearby bushes, sweep, J. Heraty, [UCRCENT00282474: UCRC]. 1♂, Puntarenas, Manuel Antonio National Park, 9°23'45"N, 84°08'16"W, 23-28.viii.1986, costal rainforest, screen sweep, L. Masner, [UCRCENT00315911: CNC]. 1♂, Parque Nacional Corcovado, Est. Sirena, 8°30'00"N, 83°36'00"W, 22.iii.1981, H.A. Hespenheide, [UCRCENT00305602: UCRC]. Ecuador: 2♀, Bucay, 304.8m, 2°12'17"S, 79°10'40"W, 4.x.1922, F.X. Williams, [UCRCENT00236291‒92: EMEC]. 1♂, Microwave tower, 1°49'52"S, 78°11'00"W, 6-7.iii.1976, Malaise trap, [UCRCENT00315880, CNC]. 1♂, Esmeraldas, Bilsa Biol. Sta., 500m, 0°20'24"N, 79°42'36"W, 10.v-4.vi.1996, malaise trap, P. Hibbs, [UCRCENT00092085: UCRC]. 1♀, Borbon, 1°05'21"N, 78°59'24"W, 1.ii.1988, Mike Huybensz, [UCRCENT00242352: MCZ]. 1♂, Zapallo Grande, 0°47'00"N, 78°59'00"W, 25-30.x.1987, Mike Huybensz, [UCRCENT00242385: MCZ]. 1♂, Guayas, Isla Puna, 2°46'35"S, 80°08'34"W, 22.iii.1988, Mike Huybensz, [UCRCENT00246691: SAMC]. 1♂, Orellana, 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 216m, 0°39'26"S, 76°27'11"W, 4.ii.1996, terre firme forest, fogging, T.L. Erwin et al., [UCRCENT00320096: UCRC]. 1♀, Tiputini Biodiversity Sta. nr. Yasuni National Park, Trans. 192

Ent., 220-250m, 0°37'55"S, 76°08'39"W, 2.ii.1999, terra firme forest, Fogging, T.L. Erwin et al., [UCRCENT00320093: UCRC]. 2♂, Pichincha, 16km SE Santo Domingo, Tinalandia, 680m, 0°18'17"S, 78°59'00"W, 15-30.vi.1975, S. & J. Peck, [UCRCENT00241041‒42: CNC]. 3♂, 45 km S. Santo Domingo, Rio Palenque, 0°39'05"S, 79°19'33"W, 5.v-25.vii.1985, rain forest, FIT, Peck, [UCRCENT00316114‒16: CNC]. 11♂, idem but 250m, 0°39'05"S, 79°19'33"W, 17- 25.ii.1979, S.A. Marshall, [UCRCENT00316001, UCRCENT00316024, UCRCENT00316052‒54: CNC; UCRCENT00251183‒86, UCRCENT00251194‒95: DEBU]. 3♂ 1♀, idem but 29.v.1975, [UCRCENT00241038‒40, UCRCENT00313652: CNC]. 1♂, 47km S. Santo Domingo, Rio Palenque Station, 200m, 0°39'05"S, 79°19'33"W, [UCRCENT00315875: CNC]. 2♂, Rio Palenque, 200m, 0°36'00"S, 79°21'00"W, 6.iii-1.iv.1996, malaise trap, P. Hibbs, [UCRCENT00172491‒92: UCRC]. 1♂, Rio Palenque, 0°39'05"S, 79°19'33"W, 20.ii.1979, S.A. Marshall, [UCRCENT00251196: DEBU]. 4♂, idem but ii.1976, G. Shewel, [UCRCENT00316048‒51: CNC]. 1♂, idem but Res. Sta., 0°39'05"S, 79°19'33"W, vi-viii.1985, S & J Peck, [UCRCENT00316118: CNC]. 100♂ 4♀ 11?, idem but ii.1983, MT, Sharkey & Masner, [UCRCENT00240971‒73, UCRCENT00240975‒79, UCRCENT00240981‒82, UCRCENT00313612‒16, UCRCENT00313618‒29, UCRCENT00313631‒37, UCRCENT00313639‒40, UCRCENT00313642, UCRCENT00313644‒48, UCRCENT00313651, UCRCENT00313653‒54, UCRCENT00313656‒66, UCRCENT00315812‒13, UCRCENT00315815, UCRCENT00315818‒29, UCRCENT00315831, UCRCENT00315833‒35, UCRCENT00315838‒39, UCRCENT00315841‒44, UCRCENT00315847‒50, UCRCENT00315854, UCRCENT00315861‒64, UCRCENT00316055‒58, UCRCENT00316060‒67, UCRCENT00316069‒70: CNC; UCRCENT00305605, UCRCENT00446428, UCRCENT00446430‒38: UCRC]. 4♂, Rio Palenque Stn., 0°39'05"S, 79°19'33"W, Malaise trap, [UCRCENT00240962‒65: CNC]. 1♀, Tinalandia, 800m, 0°16'47"S, 78°57'48"W, 8.ii.1983, Masner & Sharkey, [UCRCENT00315866: CNC]. 1♂, Zamora Chinchipe, Rio Bombuscaro, 1100m, 4°07'12"S, 78°58'48"W, 26.iv-4.vii.1996, P. Hibbs, malaise trap/pan trap, [UCRC: UCRCENT00091806]. El Salvador: 1♂ 1♀, Quezaltepeque, 13°50'00"N, 89°16'00"W, 1.vii.1963, D. Cavagnaro & M.E. Irwin, [UCRCENT00236305: EMEC, UCRCENT00279932: UCDC]. Honduras: 1♂, C.A., EAP, 14°01'02"N, 87°01'39"W, 13.vi.1982, R.W. Jones, [UCRCENT00241044: CNC]. 2♀, Comayagua, 3 km S Comayagua, 600m, 14°24'47"N, 193

87°37'25"W, 23.vii.1989, R. Cave, [UCRCENT00282629, UCRCENT00282635: EAPZ]. 1♂, Comayagua, 14°27'15"N, 87°38'35"W, 10.vii.1964, G.A. Axtell [UCRCENT00279927: UCDC]. 1♀, El Parais, Moroceli, El Mesias, 14°07'12"N, 86°46'48"W, 20.iii.1991, Carica papaya fructification, M. Avila, [UCRCENT00316462: EAPZ]. 1♀, Rapaco, 630m, 14°04'56"N, 86°54'26"W, 17.v.1993, Pimenta vegetative, B. Castro, [UCRCENT00316463: EAPZ]. 1♂, Francisco Morazan Dept.: 32 Km E. Tegucigalpa, El Zamorano, 14°01'02"N, 87°01'39"W, 24.ix.1985, R. Caballero, [UCRCENT00282632: EAPZ]. 1♂, San Antonio de Oriente, El Zamorano, 14°01'02"N, 87°01'39"W, 19.x.1988, P1assiflora edulis fructificacion, R. Cave, [UCRCENT00282636: EAPZ]. 1♂, El Zamorano, 14°01'02"N, 87°01'39"W, 22-29.iii.1990, coffee plantation ‘baja Inga’, malaise, R. Cave, [UCRCENT00316472: EAPZ]. 1♀, Est. Agr. Pan., Zamorano, 14°00'29"N, 87°00'35"W, 19.vii.1948, in monte, T.H. Hubbell, [UCRCENT00306467: FSCA]. 2♂, El Boquerón Nat. Mon., 14°47'06"N, 86°00'42"W, 2.vii.2002, D. Yanega, [UCRCENT00092077, UCRCENT00172479: UCRC]. 1♂, Tulin, 14°43'00"N, 86°11'00"W, 7.v.1979, C. Henriquez, [UCRCENT00305998: UCRC]. 1♀, Catacamas, Catacamas, 1000m, 14°50'51"N, 85°53'32"W, 28.iv.1993, Phoradendron sp., R. Cave, [UCRCENT00316471: EAPZ]. 1♂, San Francisco de la Paz, El Ocote, 15°26'11"N, 86°35'53"W, 16.vii.1989, Onoseris onoseroides, G. Marquez, [UCRCENT00282634: EAPZ]. 1♂ 1♀, Silca, El Carbonal, 14°47'36"N, 86°26'28"W, 15.x.1988, R. Cave, [UCRCENT00282630‒31: EAPZ]. Mexico: 3♂ 3♀, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 16.viii.2004, E. ruidum nest, G. Perez, [UCRCENT00235924, UCRCENT00318828‒31, ????: UCRCENT00002417: UCRC]. 1♀, idem but 5.vii.2004, on Panicum sp., [UCRCENT00318837: UCRC]. 1♂ 1♀, Tapachula, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 16.vii.2004, Melampodium divaricatum, G. Perez, [UCRCENT00318835‒36: UCRC]. 3♂, Nuevo Leon, Salinas Victoria, 20 km N. Carr. Colombia, 450m, 26°02'03"N, 100°18'18"W, 13.vii.1983, A. Gonzalez, [UCRCENT00320609‒11: UANL]. 1♂, Quintana Roo, Lazaro Cardena, 25 km NNE Leona Vicario Reserva Ecologia El Eden, 21°13'00"N, 87°11'00"W, 10.viii.1998, secondary growth near greenhouse, sweep, R. Rodriguez, [UCRCENT00320099: UCRC]. 1♂, 25 km NNE Leona Vicario Reserva Ecologica El Eden, 21°13'00"N, 87°11'00"W, 19.viii.1998, savannah de cabana, sweep, R. Rodriguez, [UCRCENT00320100: UCRC]. 1♂, Tamualipas, Gomez Farias, 23°01'49"N, 99°08'53"W, [UCRC: UCRCENT00320098]. Nicaragua: 1♀, near Comalapa, 12°16'55"N, 85°30'44"W, 1.xii.1976, from dooryard Gossypium hirsutum, swept, W.H. Cross, 194

[UCRCENT00242467, MMNS]. 1♀, idem but 1.xii.1976, on flowering cactus-like Asclepiad, [UCRCENT00242468: MMNS]. 2♂ 1♀, Chinandega, Chinandega, 12°37'28"N, 87°07'47"W, Baker, [UCRCENT00241291, UCRCENT00241293, UCRCENT00300485: CUIC]. 9♂, Managua, Tipitapa, 50m, 12°11'47"N, 86°05'44"W, 24.ix.1991, Terminalia catapa fructification, R. Caballero, [UCRCENT00316464‒70, UCRCENT00316474‒75: EAPZ]. Panama: 1♂, Cerro Galera, 8°56'00"N, 79°37'00"W, 9.ix.1983, Neutusij, [UCRCENT00245182: ZMUH]. 1♀, David, 40m, 8°25'40"N, 82°25'50"W, 20.vi.1978, G.J. Umphrey, [UCRCENT00306355: UCRC]. 3♂, Canal Zone, 1mi. W Frijoles, 20-50m, 9°10'34"N, 79°48'21"W, 18.vii.1977, R.B. & L.S. Kimsey, [UCRCENT00243502‒04: UCDC]. 1♂, Barro Colorado Island, 20-180m, 9°09'20"N, 79°50'46"W, 1.vii.1976, R.B. & L.S. Kimsey, [UCRCENT00243517: UCDC]. 1♂, idem but 10.viii.1981, [UCRCENT00243514: UCDC]. 1♂, idem but 13.vii.1976, [UCRCENT00243512: UCDC]. 2♂, idem but 15.ix.1976, [UCRCENT00243508, UCRCENT00243510: UCDC]. 2♂, idem but 15.vii.1976, [UCRCENT00243493‒94: UCDC]. 1♂, idem but 17.vi.1981, [UCRCENT00243525: UCDC]. 1♂, idem 19.vii.1976, [UCRCENT00243518: UCDC]. 1♂, idem but 2.vii.1976, [UCRCENT00243506, UCDC]. 1♂, idem but 20.ix.1978, [UCRCENT00243534: UCDC]. 1♂, idem but 20.vi.1981, [UCRCENT00243524: UCDC]. 7♂ 1♀, idem but 20.vii.1976, [UCRCENT00243476‒79, UCRCENT00243481‒84: UCDC]. 2♂, idem but 22.vii.1976, [UCRCENT00243474‒75: UCDC]. 1♂, idem, but 26.vii.1976, [UCRCENT00243520: UCDC]. 1♂, idem but 28.vi.1982, [UCRCENT00243523: UCDC]. 1♂, idem but 3.viii.1976, R.B. & L.S. Kimsey, [UCRCENT00243505: UCDC]. 1♂, idem but 31.vii.1977, [UCRCENT00243535: UCDC]. 1♂, idem but 6.vii.1976, [UCRCENT00243486: UCDC]. 1♂, idem but 9.vii.1976, [UCRCENT00243492: UCDC]. 1♂, idem but 13.ix.1978, R.B. Rimsey, [UCRCENT00243533: UCDC]. 1♂, idem but 3-13.vi.1983, malaise trap, B. Gill, [UCRCENT00315886: CNC]. 1♂, idem, but 9°10'00"N, 79°50'00"W, 23.vii.1977, H.A. Hespenheide, [UCRCENT00305599: UCRC]. 1♂, Fort Clayton, 40m, 9°00'19"N, 79°34'50"W, 6.viii.1978, H.J. Harlan, [UCRCENT00306002: UCRC]. 2♂, Puma Island, 20-70m, 9°14'10"N, 79°54'12"W, 24.vii.1982, R.B. Kimsey, [UCRCENT00243529‒30: UCDC]. 1♂, Tigre Island, 20-60m, 9°13'52"N, 79°54'39"W, 10.vi.1982, R.B. Kimsey, [UCRCENT00243515: UCDC]. 4♂, idem but 21.vi.1982, [UCRCENT00243465‒66, UCRCENT00243470, UCRCENT00243472: UCDC]. 2♂, idem but 24.vii.1982, [UCRCENT00243526‒27: UCDC]. 5♂, idem, but 28.vi.1982, [UCRCENT00243496‒97, UCRCENT00243499‒501: UCDC]. 1♂, idem but 31.v.1982, 195

[UCRCENT00243532: UCDC]. Peru: 1♂, Lima, Callanga, 1200m, 12°34'00"S, 76°19'00"W, Staudinger K., [UCRCENT00320348]. Suriname: 1♀, Paramaribo, 5°49'24"N, 55°10'04"W, 25.viii.1911 [EMEC: UCRCENT00236313]. Venezuela: 1♂, Coje des El Baul, 8°57'16"N, 68°18'04"W, 19.v.1967, J&B Bechyne, [UCRCENT00239429: BMNH]. 1♂ 1♀, idem but 20.vii.1967, [UCRCENT00239422, UCRCENT00239428: BMNH]. 1♂, El Pao, 8°02'26"N, 62°18'04"W, 19.v.1967, J&B Bechyne, [UCRCENT00239430: BMNH]. 3?, Aragua, Rancho Grande N. P., 1100m, 10°22'53"N, 67°37'08"W, 18.viii-3.ix.1992, cloud forest, maxinet, L. Masner, [UCRCENT00418062: ROME; UCRCENT00418062, UCRCENT00446455: UCRC]. 6♂ 1♀, Lara, 1 km E. Barquisineto, 500m, 10°04'38"N, 69°15'18"W, 27.xii.1985, savannah, R. Jones & P. Kovarik, [UCRCENT00241002‒08: CNC].

Kapala cavicornis Schoeninger & Heraty, n. sp. (Figs. 3B, 4B, 5C‒D)

Diagnosis. Recognized by a combination of the following characters: frons swollen and smooth; females with flagellum 9 segmented and flagellum 0.9–1.0x head height (Fig. 3B); frenal process thick with apex emarginated with an elongate flange like spoon-shaped (Fig. 4B, indicated).

Description. Female. Body length 2.8–4.4 mm; length of mesosoma excluding the frenal process 1.4–2.1 mm. Head, mesosoma, coxae, petiole and basis of the frenal process black, remaining of frenal process reddish-brown; scape and pedicel yellow, flagellum brown; legs yellow; wings hyaline to slightly infuscate, venation light brown; Gt1 brown to yellow (Fig. 5C). Head. 1.3–1.4x as broad as high, smooth surface with sparse setae, more prominent on upper frons and vertex. Scrobal depression with smooth surface. Eyes separated by 2.0-2.3× their height. Malar space 0.8–1.2x eye height, smooth except for some faint striae. Supraclypeal area defined by weakly impressed sulci, smooth and slightly swollen. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 segments; scape 3.5– 4.5x longer than broad, 0.3x head height; flagellum 9 segmented, cylindrical and slightly serrated, clava rounded (Fig. 3B); length of flagellum 0.9–1.0x head height; Fl2 2.0x as long as apical width and 1.3x as long as following flagellomere. Mesosoma. Mesoscutum 1.3–1.4x as high as broad (Fig. 5C). Midlobe with transverse carinae and lateral lobes smooth. Axilla and mesoscutellar disc with longitudinal carinae (Fig. 196

4B). Mesoscutellar disc 1.4–1.8x as long as axilla with medial apex raised 0.7–1.3x height of frenal process in lateral view. Axillula with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth, except for some shallow and inconspicuous punctures; lower mesepimeron reticulate; femoral depression crenulate. Callus carinate and densely pilose. Metapleuron carinate and slightly reticulate. Propodeum flat, broad and delimited by carina, surface reticulate to smooth with some longitudinal carinae in the upper region. Frenal process cylindrical and thick, 2.2–2.7x longer than basal width of mesoscutellum, 1.4–1.9x as long as length of axillae and mesoscutellar disc, arched in lateral view and convex in dorsal view; frenal process with longitudinal striae and apex emarginated with an elongate flange like spoon-shaped (Fig. 4B). Forewing 2.4–2.6x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.1–1.4x longer than marginal vein; stigmal vein rectangular, 1.3–2.0x longer than broad and postmarginal vein indistinct. Hindwing fringed. Metacoxa semiglobose with medial row of setae, 1.8x longer than broad. Metafemur 5.2–7.3x longer than broad, smooth with sparse setae. Metasoma. Petiole weakly flattened dorsally and with parallel longitudinal carinae, 2.5–

3.0x longer than broad, 1.3–1.4x as long as metacoxa. Tergal scar on Gt1 present or absent.

Male. Body length 2.4–3.1 mm, length of mesosoma excluding the frenal process 1.2–1.7 mm (Fig. 5D). Similar to female except for following: fore wings infuscated at the apex; frenal process with longitudinal carinae to even slightly oblique in the apical half. Head 1.4–1.5 mm as broad as high; eyes separated by 1.8–2.1x their height; maxillary/labial palpi 3/3 and 3/2 segmented; antenna with 12 segments; scape 2.0–2.3x longer than broad; length of branch of Fl2 0.8–1.2x head height and 0.8–1.0x as long as following branch. Frenal process 2.0–3.0x longer than the basal width of mesoscutellum, 1.1–1.8x as long as length of axillae and mesoscutellar disc. Metacoxa 1.5–2.0x longer than broad. Petiole 4.2–6.3x longer than broad, 1.8–2.5x as long as metacoxa.

Remarks. Variations in coloration between specimens of different populations can be found, this ranges from completely black (populations of Colombia, Panama and Venezuela) to completely reddish brown (Trinidad) or the latter coloration restricting the coxae and frenal process (Colombia, Ecuador and Trinidad). 197

Etymology. The specific name comes from Latin “cavus” = excavated, hollow; “cornis” = horn, referring to frenal process apex excavated.

Biological notes. Host unknown.

Distribution. Brazil, Bolivia, Colombia, Ecuador, Panama, Trinidad and Venezuela.

Holotype. Trinidad: ♀, Simla Res.St., 250m, 10°41'34"N, 61°17'23"W, 22.vii.2013, Heraty & Baker, station, Malaise [UCRCENT00412137: UCRC].

Paratypes. Trinidad: 3♂, Curepe, 10°38'00"N, 61°24'00"W, 17.xii.1978 [UCRCENT00105798, UCRCENT00105811, UCRCENT00105826: UCRC]. 2♂, idem but, 26-28.iii.1979, malaise trap, [UCRCENT00092286, UCRCENT00092287: UCRC]. 1♂, idem but, 10°38'47"N, 61°24'56"W, malaise trap, [UCRCENT00305640: UCRC]. 1♂, idem but, 21.vii.1978, [UCRCENT00446511: UCRC]. 1♀, idem but, 22-25.xi.1977, W. & E. Mason [UCRCENT00320185: UCRC]. 2♂, idem but, 23-27.xi.1977, MT [CNC: UCRCENT00316252: CNC, UCRCENT00306554: UCRC]. 5♂, idem but, 5-8.xii.1977, [UCRCENT00305619, UCRCENT00305632, UCRCENT00305634, UCRCENT00305637, UCRCENT00316357: UCRC]. 2♂, 2.xii.1977, Mason, [UCRCENT00446468, UCRCENT00446473: UCRC]. 2♂, Simla Res. St., 250m, 10°41'34"N, 61°17'23"W, 22.vii.2013, Heraty & Baker, station, malaise [UCRCENT00412135, UCRCENT00412136: UCRC].

Kapala ceciliae Schoeninger & Torréns, n. sp. (Figs. 3C, 4C, 5E‒F)

Diagnosis. Recognized by a combination of the following characters: female with flagellum long,

1.1‒1.2x head height and Fl2 cylindrical (Fig. 3C), 2.0‒2.6X as long as apical width; both sexes with frenal process narrow with apex emarginated (Fig. 4C), process longitudinally striate, which become slightly oblique in apical half.

198

Description. Female. Body length 2.5–3.0 mm; length of mesosoma excluding the frenal process 1.2–1.5 mm. Head, mesosoma, coxae, petiole and frenal process black; scape and pedicel yellow, flagellomeres brown; legs pale yellow; wings hyaline to slightly infuscate near the stigmal region, venation light brown; Gt1 reddish brown (Fig. 5E). Head. 1.3–1.4x as broad as high, smooth surface with sparse setae. Scrobal depression with faint striae. Eyes separated by 1.9–2.1x their height. Malar space 1.0x eye height, smooth with some faint striae. Supraclypeal area and clypeus smooth. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 3.0– 4.0x longer than broad, 0.2–0.3x head height, smooth with sparse setae; flagellum 9 segmented (Fig. 3C), cylindrical and slightly serrated, clava rounded; length of flagellum 1.1–1.2x head height; Fl2 cylindrical, 2.0–2.6x as long as apical width and 1.3–1.5x as long as following flagellomere. Mesosoma. Mesoscutum 1.1–1.4x as high as broad. Midlobe with transverse striae and lateral lobes smooth. Axilla and scutellar disc with wide longitudinal striae (Fig. 4C); scutellar disc 1.5–1.7x as long as axilla with medial apex raised 1.0–1.3x height of frenal process in lateral view. Axillula with longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth with some finelly puctations; lower mesepimeron reticulate; femoral depression deep and crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeum flat, broad and reticulate with some longitudinal carinae in upper region. Frenal process cylindrical, 2.3–2.8x longer than basal width of scutellum, 1.7‒1.9x as long as length of axillae and scutellar disc, arched in lateral view and convex in dorsal view; frenal process with longitudinal striae, which become slightly oblique in apical half; apex of frenal process emarginated (Fig. 4C). Forewing 2.3–2.5x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.0–1.2x longer than marginal vein; stigmal vein rectangular, 1.7–2.2x longer than broad and postmarginal vein long. Hindwing fringed. Metacoxa semiglobose with median row setae, 1.6– 2.2x longer than broad. Metafemur 5.5–7.3x longer than broad, smooth with sparse setae.

Metasoma. Petiole cylindrical, weakly reticulate with some striae, 3.5–4.0x longer than broad and 1.6–1.7x as long as metacoxa. Tergal scar on Gt1 present.

199

Male. Body length 2.3 mm; length of mesosoma excluding the frenal process 1.1 mm. (Fig. 5F) Similar to female except for the following: malar space 0.8x eye height. Antenna with 12 segments; scape 2.5x longer than broad; length of branch of F2 1.5x head height and 0.9x as long as following branch. Scutellar disc 1.1x as long as axilla. Frenal process 3.4x longer than basal width of scutellum. Petiole 6.0x longer than broad and 2.0x as long as metacoxa.

Remarks. Some females have the last segment of the clava partially divided (only in the dorsal region of the antenna) giving a 3-segmented clava appearance and the impression of flagellum with 10 segments. However, the division of the last segment of the clava is not complete.

Etymology. The specific name is in honor to Cecilia Dorfey, colleague, friend and the person who showed me the fantastic world of the Hymenoptera parasitoids.

Biological notes. Host unknown.

Distribution. Ecuador (Fig. 7C).

Holotype. Ecuador: ♀, Orellana, Rio Piraña Bridge, Reserva Etnica Waorani, Onkone Gare Camp, 216.3m, 0°39'25.7"S, 76°27'10.8"W, 20.x.2005, T.L. Erwin, M.C. Pimienta et al, terra firme forest, Fogging, Lot 3056, [UCRCENT00247782: USNM].

Paratypes. Ecuador: 1♂, Napo, Reserva Etinica Waorani, 1km S Onkone Gare Camp, 220m, 0°39'10"S, 76°26'00"W, 23.x.1998, T. L. Erwin et al., terre firme forest, fogging, [UCRCENT00092333: UCRC]. 1♀, idem but, Trans. Ent., 216m, 0°39'26"S, 76°27'11"W, 1.x.1996, Lot 1690 [UCRCENT00247738: USNM]. 1♀, Orellana, 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 216m, 0°39'26"S, 76°27'11"W, 4.ii.1996, T.L. Erwin et al., terra firme forest, fogging, Lot 1409, [UCRCENT00320095: UCRC]. 1♀, idem but, 5.x.1995, Lot 1192 [UCRCENT00320094: UCRC]. 1♀, idem but, 6.x.1995, Lot 1223, [UCRCENT00320092: UCRC]. 1♀, Rio Piraña Bridge, Reserva Etnica Waorani, Onkone Gare Camp, 216.3m, 0°39'25.7"S, 76°27'10.8"W, 20.x.2005, T.L. Erwin, M.C. Pimienta et al, terra firme forest, Fogging, Lot 3056, [UCRCENT00247783: USNM]. 1♂, Tiputini Biodiversity Sta. nr. Yasuni 200

National Park, 220-250m, 0°37'55"S, 76°08'39"W, 29.vi.1998, T.L. Erwin et al., Fogging, Lot 1802 [UCRCENT00092008: UCRC]. 1♀, Transect 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 220m, 0°39'25.7"S, 76°27'10.8"W, 1.x.1996, T.L. Erwin et al., terra firme forest, fogging, Lot 1684, [UCRCENT00092246: USNM].

Kapala ipa Schoeninger & Heraty, n. sp. (Fig. 3D, 4D, 6A‒B)

Diagnosis. Recognized by a combination of the following characters: female with antenna long, 1.2–1.3x head height and flagellum 10 segmented (Fig. 3D); male with length of branch of F2 1.2–1.4x head height; both sexes with fine longitudinal striae in axilla, scutellar disc and frenal process (Fig. 4D).

Description. Female. Body length 2.3–3.8 mm; length of mesosoma excluding the frenal process 1.2–2.0 mm. Head, mesosoma, coxae and frenal process black; scape, pedicel and legs pale-yellow; flagellomeres brown; wings hyaline, venation brown; Gt1 brown to dark brown (Fig. 6A). Head. 1.4–1.5x as broad as high, smooth surface with sparse setae. Scrobal depression with faint striae. Eyes separated by 1.9–2.2x their height. Malar space 1.0x eye height, smooth with some faint striae and punctuations. Supraclypeal area and clypeus smooth. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 3.0–3.5x longer than broad, 0.3x head height, smooth with sparse setae; flagellum 10 segmented (Fig. 3D), cylindrical and slightly serrated, clava rounded; length of flagellum 1.2– 1.3x head height; F2 1.5–2.0x as long as apical width and 1.0–1.4x as long as following flagellomere. Mesosoma. (Fig. 6A), mesoscutum 1.2–1.4x as high as broad. Midlobe and lateral lobes with transverse striae. Axilla and scutellar disc with fine longitudinal striae; scutellar disc 1.5– 1.8x as long as axilla with medial apex raised 0.5–0.8x height of frenal process in lateral view. Axillula with fine longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth with some finelly punctuations; lower mesepimeron reticulate; femoral depression weakly crenulate. Callus carinate and 201

reticulate, covered by semi-erect setae. Metapleura reticulate and delimited by carina. Propodeum flat, broad, and weakly reticulate. Frenal process cylindrical, 2.1–3.1x longer than basal width of scutellum, 1.3‒1.8x as long as length of axillae and scutellar disc, slightly arched in lateral view and slightly convex in dorsal view; frenal process with fine longitudinal striae, apex of frenal process slightly emarginated (Fig.4D). Forewing 2.6–2.7x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.1–1.4x longer than marginal vein; stigmal vein rectangular, 1.2–2.5x longer than broad and postmarginal vein indistinct. Hindwing fringed. Metacoxa semiglobose with row setae median, 1.8–2.5x longer than broad. Metafemur 6.2–7.0x longer than broad, smooth with sparse setae.

Metasoma. Petiole flattened dorsally and weakly reticulate with some fine striae, 3.7–

4.7x longer than broad and 1.5–1.8x as long as metacoxa. Tergal scar on Gt1 absent.

Male. Body length 2.7–3.2 mm, length of mesosoma excluding the frenal projections 1.4– 1.7 mm (Fig. 6B). Similar to female except for the following: frons smooth to even faint longitudinal striated; eyes separated by 1.8–2.0x their height; scape 2.0–3.5x longer than broad; flagellum 10 segmented, transversal and each flagellomere with a single long branch; length of branch of F2 1.2–1.4x head height and 0.9–1.0x as long as following branch. Forewing 2.2–2.5x longer than broad. Metafemur 5.7–7.0x longer than broad. Petiole cylindrical, 5.2–7.0x longer than broad and 2.0–2.1x as long as metacoxa.

Remarks. This species is very similar to Kapala ceciliae n. sp. but differ by F2 1.5–2.0x as long as apical width (vs. F2 long, 2.0–2.6x as long as apical width); lateral lobe striated (vs. lateral lobe smooth); scutellar disc with medial apex raised 0.5–0.8x height of frenal process (vs. scutellar disc with apex raised 1.0–1.3x height of frenal process); frenal process with fine longitudinal striae (vs. frenal process with wide longitudinal striae, which become slightly oblique in basal half).

Etymology. The specific name is in honor of the best style of beer ever made the “Indian Pale Ale”, popularly known as IPA.

202

Biological notes. Host unknown.

Distribution. Costa Rica, Panama, Tobago (Fig. D)

Holotype. Panama: ♀, Nusagandi Lodge, 390m, 9°20'31"N, 78°59'38"W, 21.i.2001, L. Masner, open road, sweep, [UCRCENT00091879: UCRC].

Paratypes. Costa Rica: 3♂, Limon, 4km NE Bribri, 50m, 9°39'03"N, 82°48'16"W, ix-xi.1989, 89-11-01-1, Paul Hanson, [UCRCENT00274869, UCRCENT00302318‒19: UCRC]. 1♀, Puntarenas, P.N. Manuel Antonio, Quepos, 80m, x.1991, G. Varela, [INBIOCRI00501853: INBIO]. 1♀, R.B. Carara, Estac. Bijogoal, 500m, 10°21'51"N, 84°12'25"W, x.1989, P. Hanson, [UCRCENT00302320: UCRC]. Panama: 3♀, Com de San Blas, 2 km N of Nusagadi, 320- 400m, 9°21'29"N, 78°58'42"W, 14.i.2001, screen sweep, 2001/044, M. Yoder & J. B. Woolley, [UCRC: UCRCENT00172512‒14: UCRC]. 1♀, Canal Zone, Fort Randolph, 5m, 9°23'05"N, 79°52'59"W, 23.i.1929, C.H. Curran, [UCRCENT00236304: EMEC]. Tobago: 1♀, 1 1/8mi ESE Adelphi, 150m, 11°12'12"N, 60°42'24"W, 15.ii.1977, secondary scrub along pasture edge, sweep, P. Feinsinger, [UCRCENT00306444: FSCA].

Kapala longicornis Schoeninger & Oliveira, n. sp. (Fig. 2A, 3E, 4E, 6C‒D)

Diagnosis. Recognized by a combination of the following characters: frenal process long, exceeding the length of the metasoma (Figs. 4E; 6C‒D), process with longitudinal striae, which become strongly oblique in the apical half; F2 of female long and swollen at apex and 1.7x as long as following flagellomere (Fig. 3E) and eyes separated by 2.3–2.4x their height.

Description. Female. Body length 3.4–4.1 mm; length of mesosoma excluding the frenal process 1.7–2.0 mm. Head, mesosoma, coxae and petiole black; frenal process reddish brown; scape, pedicel and legs yellow, flagellomeres brown; wings hyaline, venation brown; Gt1 dark brown (Fig. 6C). Head. 1.3x as broad as high, smooth surface with sparse setae more prominent on frons and vertex. Scrobal depression smooth. Eyes separated by 2.3–2.4x their height. Malar space 1.0x 203

eye height, with some faint striae and punctuations. Supraclypeal area and clypeus smooth. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 segments; scape 3.5x longer than broad, 0.3x head height, smooth with sparse setae in dorsal region; flagellum 9 segmented (Fig. 3E), cylindrical and slightly serrated; length of flagellum 1.0x head height, F2 long and swollen at apex, 1.7–2.0x as long as apical width and 1.7x as long as following flagellomere. Mesosoma. (Fig. 6C); mesoscutum 1.2–1.4x as high as broad. Midlobe with transverse striae and lateral lobes smooth. Axilla and scutellar disc with longitudinal striae (Fig. 4E); scutellar disc 1.5x as long as axilla with medial apex raised 0.7–0.8x height of frenal process in lateral view. Axillula with some weakly longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth; lower mesepimeron weakly reticulate; femoral depression crenulate. Callus carinate and reticulate, covered by semi-erect setae. Metapleura reticulate and delimited by carina. Propodeum flat, broad, and weakly reticulate. Frenal process cylindrical and long exceeding the length of metasoma, 2.4–2.8x longer than basal width of scutellum, 1.7-1.9x as long as length of axillae and scutellar disc, slightly arched in lateral view and slightly convex in dorsal view; frenal process with longitudinal striae, which become strongly oblique in the apical half; apex emarginated (Fig. 4E, 6C). Forewing 2.6–2.75x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.1–1.3x longer than marginal vein; stigmal vein rectangular oblique, 1.3–1.7x longer than broad and postmarginal vein short. Hindwing fringed. Metacoxa semiglobose with row setae median, 1.7–2.0x longer than broad. Metafemur 6.5x longer than broad, smooth with sparse setae. Metasoma. Petiole flattened dorsally and reticulate with some fine striae, 3.2–3.4x longer than broad and 1.6–1.7x as long as metacoxa. Tergal scar on Gt1 present.

Male. Body length 4.3 mm, length of mesosoma excluding the frenal projections 2.0 mm (Fig. 6D). Similar to female except for the following: head 1.5x as broad as high; scape 2.3x longer than broad; flagellum 10 segmented, transversal and each flagellomere with a single long branch; length of branch of F2 1.3x head height and 0.9x as long as following branch. Frenal process 3.0x longer than the basal width of scutellum. Metafemur 5.4x longer than broad. Petiole 6.2x longer than broad and 2.3x as long as metacoxa. 204

Remarks. A couple collected in Rio de Janeiro (Sta. Maria Magdalena) has the flagellum 8 segmented and a short body length (1.8‒2.1 mm), reddish-brown coloration and completely yellow antennae. However, they have the frenal process long, with longitudinal striae, that become oblique in the apical half; the female has F2 long and swollen at apex.

Etymology. The specific name comes from Latin “longus” = long, “cornis” = spine, referring to long frenal process, that exceeding the length of the metasoma.

Biological notes. Host unknown.

Distribution. Brazil (Fig. E).

Holotype. Brazil: ♀, Espírito Santo, Sta.[Santa] M.[Maria] de Jetibá, Faz.[enda] Clarindo Krüger, 20°04'27.9''S, 40°44'51.3''W, 29.xi–06.xii.2002, Malaise T7, Tavares, Azevedo & eq., [UFES].

Paratypes. Brazil: 1♂, Espírito Santo, Dom.[ingos] Martins, 20°21'38.87"S, 40°39'35.32"W, 05- 12.vii.2003, Malaise, R. Kawada, [UFES]. 1♂ 1♀, Rio de Janeiro, Sta.[Santa] Maria Madalena, Parque Estadual do Desengano, 21°59'03''S, 41°57'0''W, 16-19.iv.2002, 560m, Arm. Malaise, Trilha pt. 7, A.M. Penteado-Dias & eq., [MZUSP].

Kapala parairidicolor Schoeninger & Torréns n. sp. (Fig. 2B, 3F, 4F, 6E‒F)

Diagnosis. Recognized by a combination of the following characters: frons with wide longitudinal striae (Fig. 2B), which extends to genae; frenal process with apical half slightly oblique striate with apex emarginated (Fig. 4F); lateral lobe with transversal striae (Figs. 6E, F).

Description. Female. Body length 1.9 mm; length of mesosoma excluding the frenal process 1.5 mm. Head, mesosoma, coxae and petiole black; frenal process black in basal half and 205

brown in apical half; scape, pedicel and flagellum light brown; legs pale yellow; wings hyaline, venation yellow.; Gt1 light brown (Fig. 6E). Head. 1.4x as broad as high, smooth surface with sparse setae, more prominent on upper frons and vertex. Frons with longitudinal striae (Fig. 2B); lower face smooth. Eyes separated by 2.0x their height. Scrobal depression with longitudinal striae. Malar space 0.8x eye height, with weakly transversal striae. Supraclypeal area smooth, defined by weakly impressed sulci and slightly swollen. Clypeus smooth. Anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 segments; scape 3.0x longer than broad, 0.3x head height; flagellum 9 segmented (Fig. 3F), slightly serrated, clava rounded; length of flagellum 0.9x head height; Fl2 2.0x as long as apical width and 1.3x as long as following flagellomere. Mesosoma. (Fig. 6E); mesoscutum 1.4–1.5x as high as broad. Midlobe with transverse striae and lateral lobes smooth. Axilla and scutellar disc with longitudinal striae (Fig. 4F). Scutellar disc 1.3x as long as axilla with medial apex raised 1.5x height of frenal process in lateral view. Axillula smooth. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth; lower mesepimeron reticulate; femoral depression crenulate. Callus reticulate and covered by semi-erect setae. Metapleuron carinate and reticulate. Propodeum flat, broad and delimited by carina, surface weakly reticulate. Frenal process cylindrical and thin 2.6x longer than the basal width of scutellum, 1.3x as long as length of axillae and scutellar disc, arched in lateral view and convex in dorsal view; frenal process with longitudinal striae and apex rounded (Fig. 4F). Forewing 2.6x longer than broad; covered by small bristles, except for the base bare; submarginal vein 1.0x longer than marginal vein; stigmal vein rectangular, 1.5x longer than broad and postmarginal vein absent. Hindwing fringed. Metacoxa with a medium line of setae, 2.0x longer than broad. Metafemur 7.0x longer than broad, smooth with sparse setae. Metasoma. Petiole cylindrical and reticulate with lateral carina, 3.3x longer than broad,

1.2x as long as metacoxa. Tergal scar on Gt1 absent.

Male. Body length 2.5–3.1 mm; length of mesosoma excluding the frenal process 1.3–1.6 mm (Fig. 6F). Similar to female except for the following: Gt1 light brown with a dark brown spot. Antenna with 12 segments; scape 2.3–3.0x longer than broad, 0.2–0.3x head height; flagellum 10 206

segmented, transversal and each flagellomere with a single long branch; length of branch of F2 0.9–1.2 head height and 0.9–1.0x as long as following branch. Axillula with longitudinal striae. Mesepisternum and upper mesepimeron with weakly striae and some shallow and inconspicuous punctures in upper region. Propodeum with a superficial depression in the middle region. Forewing 2.2–2.4x longer than broad; submarginal vein 1.1–1.2x longer than marginal vein; stigmal vein 2.0x longer than broad. Petiole reticulate with some faint longitudinal striae, 5.5–

7.0x longer than broad, 2.1–2.6x as long as metacoxa. Tergal scar on Gt1 present or absent.

Remarks. Some specimens from Mexico (Chiapas) have frons with weak longitudinal striae, in addition to an individual that has the frenal processes with longitudinal striae. The scutellar disc, in the population of Mexico (Chiapas), is more swollen and has a higher medial apex, compared to individuals from Honduras.

Etymology. From Greek “para” = near of, due to the similarity with Kapala iridicolor species.

Biological notes. Female collected in nest of Gnamptogenys regularis? Or hartmanni. Also, associated with: Melampodium divaricatum (Rich.) and Panicum sp.

Distribution. Costa Rica, Honduras and Mexico (Fig. 7F).

Holotype. Mexico: ♀, Chiapas, Rosario Izapa, 14°58'00"N, 92°09'00"W, 11.viii.1997, P. Lachaud, host: Gnamptogenys regulars? or hartmanni [UCRCENT00002411: UCRC].

Paratypes. Costa Rica: 1♂, Guanacaste, Santa Rosa NP, 300m, 10°53'33"N, 85°46'00"W, 28.xii.1985-18.i.1986, D.H. Janzen & I.D. Gauld, [UCRCENT00446454: UCRC]. 1♂, Santa Rosa NP Hacienda (4C), 10°53'33"N, 85°45'59"W [UCRCENT00274881: UCRC]. Honduras: 1♂, Fco. Morazan, San Antonio de Oriente, El Zamorano, 14°01'02"N, 87°01'39"W, 30.vii.1999, C. Lemus & H. Cuestas, [UCRCENT00316476: EAZP]. 1♂, Feo. Morazan, Zamorano Campus, 840m, 14°00'43"N, 87°00'50"W, 1.vii.2002, D. Yanega, [UCRCENT00092059: UCRC]. 1♂, Olancho, El Boquerón Nat. Mon., 14°47'06"N, 86°00'42"W, 2.vii.2002, D. Yanega, [UCRCENT00092075: UCRC]. Mexico: 2♂, Chiapas: 11 km. No. Tapachula, 15°00'23"N, 207

92°15'43"W, 20.v.1980 [UCRCENT00306536, UCRCENT00407559: WFBM]. 2♂, Tapachula, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 16.viii.2004, G. Perez, E. ruidum nest, [UCRCENT00318828, UCRCENT00318832: UCRC]. 1♂, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 19.vii.2002, G. Perez, Melampodium divaricatum [UCRCENT00318834: UCRC]. 1♀, Canton Leoncillos, 15m, 14°46'24"N, 92°24'24"W, 5.vii.2004, G. Perez, on Panicum sp. [UCRCENT00318837: UCRC].

Acknowledgment

References

Ashmead, W.H. (1888) Descriptions of three new eucharids from Florida, with a generic table of the Eucharidae. Entomologica Americana, 3: 186–188. Ashmead, W.H. (1904) Classification of the chalcid flies, or the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected in South America by Herbert H. Smith. Memoirs of the Carnegie Museum, 1: i–ix, 225–551, pls. xxxi–xxxix. Bruleè, M.A. (1846) A. LePeletier de Saint-Fargeau, Histoire Naturelle des Insectes par M. LeConte. Hyménopterès. IV. Prais, 680 pp. Cameron, P. (1884) Hymenoptera (Families Tenthredinidae-Chrysididae). In Goddman, F.D., and Salvin, D., Biologia centrali-americana. Insecta Hymenoptera. London: Taylor and Francis. 1:1‒487 + 20pls. Cameron, P. (1904) New Hymenoptera mostly from Nicaragua. Invertebrata pacifica, 1: 46 Cameron, P. (1913) The Hymenoptera of the Georgetown Museum, Part V. Journal of Royal Agricultural Society, Demerara 3: 105–137. 208

Desmarest, E. (1860) Chenu, Jean Charles, Encylopédie d'histoires naturelles. Paris: E. Girard and A. Boitte. Gemignani, E.V. (1933) La familia Eucharidae (Hym. Chalcidoidea) en la Republica Argentina. Anales del Museo Nacional de Historia Natural de Buenos Aires, 37: 477–493. Gibson, G.A.P., Read, J.D. & Fairchild, R. (1998) Chalcid wasps (Chalcidoidea): illustrated glossary of positional and morphological terms. Available from: http://www.canacoll.org/Hym/Staff/Gibson/apss/chglintr.htm (accessed 22 March 2016) Harris, R.A. (1979) A glossary of surface sculpturing. Occasional papers in Entomology, 28:1 31. Heraty, J.M. (1985) A revision of the Nearctic Eucharitinae (Hymenoptera: Chalcidoidea: Eucharitidae). Proceedings of the Entomological Society of Ontario, 85: 61-103. Heraty, J.M. (2002) A Revision of the Genera of Eucharitidae (Hymenoptera: Chalcidoidea) of the World. Memoirs of the American Entomological Institute, 68: 368 pp. Heraty, J.M. & Woolley, J.B. (1993) Separate species or polimorphism; a recurring problem in Kapala (Hymenoptera: Eucharitidae). Annals of the Entomological Society of America, 85: 517-531. Katoh, K., Kuma, K., Toh, H. & Miyata, T. (2005) MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic. Acids Res., 33, 511-8. Kirby, W.F. (1886) A synopsis of the genera of the Chalcididae, subfamily Eucharinae, with descriptions of several new genera and species of Chalcididae and Tenthredinidae. Journal of the Linnean Society, 20: 28–37. Latreille, P.A. (1825) Families naturelles de règne animla, exposées succintement et dans um order analytiques avec l’indication de leurs genres. Paris: J.B. Bailliere, 570 pp. Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 1-8. Murray, E.A. (2014) Systematics and evolution of Eucharitidae (Hymenoptera: Chalcidoidea), with a focus on the New World Kapala. University of California, Riverside, CA. 209

Murray, E.A., Carmichael, A.E., & Heraty, J.M. (2013) Ancient host shifts followed by host conservatism in a group of ant parasitoids. Proceedings of the Royal Society of London B: Biological Sciences, 280 (1759): 20130495. Murray, E. A. & Heraty, J. (2016) Invading Africa: a novel transoceanic dispersal by a New World ant parasitoid. Journal of Biogeographic, 43: 1750‒1761. Sequencher® version 5.4.6 DNA sequence analysis software, Gene Codes Corporation, Ann Arbor, MI USA. Access in: http://www.genecodes.com Schoeninger, K., Torréns, J., Heraty, J. & Oliveira, M. (2018) Filogenia de Kapala Cameron, 1884 (Hymenoptera: Eucharitidae) com base em dados morfológicos e moleculares. In prep., Cap. I. Schoeninger, K., Torréns, J., Heraty, J. & Oliveira, M. (2018) Taxonomy of Kapala (Hymenoptera: Eucharitidae): New species, redescriptions and identification key. In prep., Cap. II. Shulz, W.A. (1905) Strandgut. Spolia Hymenopterologica, 1905: 77–269. Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57: 758-71. Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics, 27, 171-180. Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. http://dx.doi.org/10.1371/journal.pone.0015991 Walsh, P.A., Metzger, D.A. & Higuchi, R. (1991) Chelex 100 as a medium for simple extraction of DNA for PCR-based typing from forensic material. Biotechniques, 10: 506-513.

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Figure 2. Front head of Kapala iridicolor complex. A. Kapala longicornis n. sp. (front smooth); B. Kapala parairidicolor n. sp. (presence of striae on the frons).

Figure 3. Antenna of females of Kapala iridicolor complex, in lateral view. A. Kapala iridicolor; B. Kapala cavicornis n. sp.; C. Kapala ceciliae n. sp.; D. Kapala ipa n. sp.; E. Kapala longicornis n. sp.; F. Kapala parairidicolor n. sp.

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Figure 4. Dorsal view of Kapala iridicolor complex (females). A. K. iridicolor; B. K. cavicornis n. sp.; C. K. ceciliae n. sp.; D. K. ipa n. sp.; E. K. longicornis n. sp.; F. K. parairidicolor n. sp.; Red seta indicating the apex of frenal processes emarginated with an elongate flange like spoon-shaped.

213

Figure 5. Lateral view of Kapala iridicolor complex A‒B. K. iridicolor (♀ and ♂, respectively); C‒D. K. cavicornis n. sp. (♀ and ♂); E‒F. K. ceciliae n. sp. (♀ and ♂). Red seta indicating the mesoscutum low and smoothly curved anteriorly in lateral view.

214

Figure 6. Lateral view of Kapala iridicolor complex. A‒B. K. ipa n. sp. (♀ and ♂, respectively); C‒D. K. longicornis n. sp. (♀ and ♂); E‒F. K. parairidicolor n. sp. (♀ and ♂).

215

Figure 7. Distribution of the species of K. iridicolor complex. A. K. iridicolor; B. K. cavicornis n. sp.; C. K. ceciliae n. sp.; D. K. ipa n. sp.; E. K. longicornis n. sp.; F. K. parairidicolor n. sp. 216

Supplementary information

Table S1. Locality, museum, identification number (ID) and DNA voucher number information for the 35 taxa of iridicolor complex and outgroup (furcata complex) used in the molecular analysis.

DNA voucher Species Sex ID number Museum Locality number K. ipa n.sp. ♀ UCRCENT91879 D# 0381 UCRC Panamá: Nusagandi Lodge, 21.i.2001, L. Masner K. parairidicolor n.sp. ♂ UCRCENT92059 D#1267 UCRC Honduras: Feo. Morazan, Zamorano Campus, 1.vii.2002, D. Yanega K. parairidicolor n.sp. ♀ UCRCENT2411 D#1895 UCRC Mexico: Chiapas, Rosario Izapa, 11.viii.1997, P. Lachaud K. parairidicolor n.sp. ♂ UCRCENT92075 D#0933c UCRC Honduras: Olancho, El Boquerón Nat. Mon., 2.vii.2002, D. Yanega K. iridicolor ♂ UCRCENT92077 D#0933b UCRC Honduras: Olancho, El Boquerón Nat. Mon., 2.vii.2002, D. Yanega K. iridicolor ♂ UCRCENT91816 D#0928a UCRC Colombia: Magdalena, PNN Tayrona Zaino, 28.iv-13.v.2000, R. Henriquez K. iridicolor ♂ UCRCENT92085 D#938a UCRC Ecuador: Esmeraldas, Bilsa Biol Sta., 10.v-4.vi.1996, P. Hibbs K. iridicolor ♂ UCRCENT92125 D#935a UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs K. iridicolor ♀ UCRCENT282474 D#2917 UCRC Costa Rica: Heredia, La Selva Biol., 14.viii.2010, J. Heraty K. iridicolor ♂ UCRCENT91809 D#0940 UCRC Ecuador: Pichincha, Rio Palenque Science Ctr., 25.iv-6.vi.1996, P. Hibbs K. ceciliae n.sp. ♀ UCRCENT92246 D#1138 UCRC Ecuador: Orellana, Res. Etnica Waorani, 1.x.1996, T.L. Erwin et al. K. ceciliae n.sp. ♀ UCRCENT247782 D#3831 USNM Ecuador: Orellana, Res. Etnica Waorani, 20.x.2005, T.L. Erwin, et al K. ceciliae n.sp. ♂ UCRCENT92008 D#1121 UCRC Ecuador: Orellana, Tiputini Biodiversity Sta. , 29.vi.1998, T.L. Erwin et al. K. ceciliae n.sp. ♀ UCRCENT247783 D#3841 USNM Ecuador: Orellana, Res. Etnica Waorani, 20.x.2005, T.L. Erwin, M.C. et al K. cavicornis n.sp. UCRCENT92073 D#0936a UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs K. cavicornis n.sp. ♂ UCRCENT175171 D#0937 UCRC Ecuador: Pichincha, Rio Palenque, 6.iii-1.iv.1996, P. Hibbs K. cavicornis n.sp. ♀ UCRCENT92092 D#0947 UCRC Colombia: Cauca, PNN Gorgona Alto el Mirador, 4-24.iii.2000, R. Duque K. cavicornis n.sp. ♂ UCRC92081 D#0939a UCRC Ecuador: Pichincha, Rio Palenque Science Ctr., 25.iv-6.vi.1996, P. Hibbs K. cavicornis n.sp. ♂ UCRCENT91969 D#0920 UCRC Colombia: Magdalena, Tayrona Cañaveral, 28.vi-17.vii.2000, R. Henriquez K. cavicornis n.sp. ♂ UCRCENT91869 D#0382a UCRC Panamá: Panama, P.N. Soberania Plantation, 21.i.2001, L. Masner K. cavicornis n.sp. ♂ UCRCENT92074 D#0917 UCRC Colombia: Magdalena, Tayrona Zaino, 28.vi-17.vii.2000, R. Henriquez K. cavicornis n.sp. ♂ UCRCENT92055 D#0382b UCRC Panamá: Panama, P.N. Soberania Plantation, 21.i.2001, L. Masner K. cavicornis n.sp. ♂ UCRCENT235921 D#2781 UCRC Colombia: Chocó, PNN UTRIA C. Visitantes, 5-19.vii.2000, J. Perez K. cavicornis n.sp. ♂ UCRCENT92120 D#0929 UCRC Colombia: Magdalena, PNN Tayrona Zaino, 28.iv.-13.v.2000, R. Henriquez 217

K. cavicornis n.sp. ♂ UCRCENT91804 D#0950 UCRC Colombia: Magdalena, PNN Tayrona Pueblito, 12-28.v.2000 K. cavicornis n.sp. ♂ UCRCENT92072 D#0944a UCRC Colombia: Magdalena, Tayrona Pueblito, 29.vi-14.vii.2000, R. Henriquez K. cavicornis n.sp. ♀ UCRCENT412137 D#3819 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker K. cavicornis n.sp. ♂ UCRCENT412134 D#3816 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker K. cavicornis n.sp. ♂ UCRCENT412135 D#3817 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker K. cavicornis n.sp. ♂ UCRCENT412136 D#3818 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker K. longicornis n.sp. UFES Espírito Santo: Sta. M. de Jetibá, 29.xi–06.xii.2002, Tavares e eq. K. furcata ♀ UCRENT92093 D#1078 UCRC Argentina: Misiones, Santa Ana, near Loreto, 27.iii.2003, J. Heraty K. parafurcata n.sp. ♂ UCRCENT161498 D#2519 UCRC Argentina: Misiones, Rt17, E of 9 de Julio, 26.iii.2007, J. Heraty K. parafurcata n.sp. ♂ UCRCENT397224 D#3436 UCRC Paraguay: Caazapá, San Rafael Reserve, 8-10.xii.2000, Z.H. Falin K.a cuprea ♀ UCRCENT91807 D#1003 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 2.vii.1995, T.L. Erwin et al. K. cuprea ♂ UCRCENT92227 D#1004 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 9.vi.1995, T.L. Erwin et al.

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Table S2. Taxa, identification number, DNA voucher number information and the respective genes used for molecular analysis. Genes Species Sex ID number DNA voucher number 18S 28S-D2 28S-D3-5 COI-nj COII Kapala ipa n.sp. ♀ UCRCENT91879 D# 0381 x x x x Kapala parairidicolor n.sp. ♂ UCRCENT92059 D#1267 x x x x Kapala parairidicolor n.sp. ♀ UCRCENT2411 D#1895 x x x Kapala parairidicolor n.sp. ♂ UCRCENT92075 D#0933c x x x x Kapala iridicolor ♂ UCRCENT92077 D#0933b x x x x Kapala iridicolor ♂ UCRCENT91816 D#0928a x x x x Kapala iridicolor ♂ UCRCENT92085 D#0938a x x x x Kapala iridicolor ♂ UCRCENT92125 D#0935a x x x x Kapala iridicolor ♀ UCRCENT282474 D#2917 x x x x x Kapala iridicolor ♂ UCRCENT91809 D#0940 x x x Kapala ceciliae n.sp. ♀ UCRCENT92246 D#1138 x x x x Kapala ceciliae n.sp. ♀ UCRCENT247782 D#3831 x x x Kapala ceciliae n.sp. ♂ UCRCENT92008 D#1121 x x x Kapala ceciliae n.sp. ♀ UCRCENT247783 D#3841 x x x Kapala cavicornis n.sp. UCRCENT92073 D#0936a x x x x Kapala cavicornis n.sp. ♂ UCRCENT175171 D#0937 x x x x Kapala cavicornis n.sp. ♀ UCRCENT92092 D#0947 x x x x Kapala cavicornis n.sp. ♂ UCRC92081 D#0939a x x x Kapala cavicornis n.sp. ♂ UCRCENT91969 D#0920 x x x x Kapala cavicornis n.sp. ♂ UCRCENT91869 D#0382a x x x x Kapala cavicornis n. sp. ♂ UCRCENT92074 D#0917 x x x Kapala cavicornis n.sp. ♂ UCRCENT92055 D#0382b x x x Kapala cavicornis n.sp. ♂ UCRCENT92120 D#0929 x x x Kapala cavicornis n.sp. ♂ UCRCENT91804 D#0950 x x x Kapala cavicornis n.sp. ♂ UCRCENT92072 D#0944a x x x Kapala cavicornis n.sp. ♀ UCRCENT412137 D#3819 x x x x Kapala cavicornis n.sp. ♂ UCRCENT412134 D#3816 x x x x Kapala cavicornis n.sp. ♂ UCRCENT412135 D#3817 x x x x Kapala cavicornis n.sp. ♂ UCRCENT412136 D#3818 x x x x 219

Kapala furcata ♀ UCRENT92093 D#1078 x x x x Kapala parafurcata n.sp. ♂ UCRCENT161498 D#2519 x x x Kapala parafurcata n.sp. ♂ UCRCENT397224 D#3436 x x x x Kapala cuprea ♀ UCRCENT91807 D#1003 x x x x Kapala cuprea ♂ UCRCENT92227 D#1004 x x x x

220

Capítulo IV

Schoeninger, K., Murray, E., Torréns, J., Heraty,

J.M, Oliveira, M.L. (2018) Taxonomy of Kapala furcata species complex (Hymenoptera:

Chalcidoidea: Eucharitidae): New species and identification key

221

Taxonomy of Kapala furcata species complex (Hymenoptera: Chalcidoidea: Eucharitidae): New species and identification key

KARINE SCHOENINGER1,2, ELIZABETH MURRAY2, JOHN M. HERATY3, JAVIER TORRÉNS4 and MARCIO L. OLIVEIRA1

1Coordenação de Biodiversidade, Programa de Pós-Graduação em Entomologia, Instituto Nacional de Pesquisas da Amazônia, CEP 69067-375, Manaus, AM, Brazil. 2Department of Entomology, Cornell University, Ithaca, NY 14853, United States. 3CRILAR-CONICET, Entre Ríos y Mendoza, 5301 Anillaco, La Rioja, Argentina. 4Department of Entomology, University of California, Riverside, CA 92521, United States. 5Corresponding author. [email protected]

Short title. Taxonomy of Kapala furcata species complex

Abstract. Kapala Cameron (Chalcidoidea, Eucharitidae) has a wide distribution in the Neotropical region and exhibits a large amount of external morphological variation. In this study we improved the knowledge about Kapala by reviewing the furcata species complex. As results, were recognize six species within the furcata complex, including K. furcata described by Fabricius (1804), K. cuprea Cameron, 1903 and K. splendens Ashmead, 1904 providing a new diagnosis for these. Three new species are described here: K. deltalis Murray & Schoeninger, n. sp., K. parafurcata Murray & Schoeninger, n. sp. and K. quasimodo, Murray & Schoeninger n. sp.. A diagnosis for the K. furcata complex is provided, as well as a molecular phylogeny for the species based on 18S, 28S-D2, 28S-D3-D5, COI and COII. Also, an identification key for the species of the furcata complex is provided.

Keywords. Ant parasitoids, identification key, mitochondrial genes, Neotropical region, taxonomy.

222

Introduction

Kapala Cameron (Chalcidoidea, Eucharitidae) is the one of the most commonly encountered genera of eucharitids wasps in the Neotropical region (Heraty & Woolley, 1993; Heraty, 2002; Murray, 2014; Murray & Heraty, 2016). Is numerically abundant and diverse with 18 species described and valid until now. Kapala exhibits a large amount of intraspecific external morphological variation and because of this the boundaries of most species have yet not been established. It has long been recognized that there is high diversity and few diagnostic characters across the genus (Heraty & Wolley, 1993; Murray, 2014). This genus is phenotypically distinctive from related genera, there are no morphological or molecular synapomorphies to unite the genus (Heraty, 2002; Murray, 2014; Schoeninger et al., in prep.). and in molecular, morphological or combined (morphological and molecular) analyses Kapala is not monophyletic, despite its phenotypic similarity. According to the analyses Kapala was divided into three complexes of species, which are furcata species complex, iridicolor species complex and romandii species complex (Murray, 2014; Schoeninger et al., in prep). These complexes are separated by clusters of morphologically distinct genera (Murray, 2014; Schoeninger et al., in prep.). The type species, Kapala furcata (Fabricius, 1804) was described from a single female collected in South America (Fabricius, 1804). The name has been consistently misapplied to many museum specimens and has led to problems association both biological data and geographic limits (Heraty, 2002). The furcata species complex is monophyletic with high support in molecular analyses. In this study we improved the knowledge about Kapala by reviewing the furcata species complex, setting new boundaries for this complex, describing three new species, providing an idetification key, and increasing the knowledge about its geographical distribution.

Material and methods

Molecular phylogenetic analysis

The molecular matrix has a total of 23 specimens (Table S1), with 21 from furcata complex, representing five species. Outgroup taxa are from the iridicolor complex with two 223

species of K. iridicolor Cameron, 1904. The species of furcate complex are from 5 countries, with only Kapala splendens with no molecular data. The matrix includes three nuclear ribosomal (partial 18S, 28S-D2 and 28S-D3-D5) and two mitochondrial (COI-nj and COII) gene fragments (Table S2). For the DNA extractions, both fresh and dried specimens were used. Specimens were non-destructively extracted using a chelex- proteinase-K-protocol or DNeasy (Qiagen). Chelex-proteinase-K was performed (Walsh et al., 1991) using non-destructive sampling to preserve the integrity of the parasitoid wasps for retainment as voucher specimens. Portions of five gene regions were amplified: 18S, 28S-D2, 28S-D3-D5, COI and COII; primers are reporter in Table. Qiagen (Valencia, CA) reagents were used for PCR. Sequencer 5.4.6 (Gene Codes Corp, Ann Arbor, MI) was used to edit chromatograms to final sequences; primers were not included in the final sequence counting. All molecular data were obtained from a unique DNA voucher number (D#), which are deposited in the internal laboratory FileMaker v.11® database maintained at UCR. Individual genes were aligned using the MAFFT online server (Katoh et al., 2005) under default settings. For nuclear ribosomal genes, the E-INS-I algorithmic strategy was applied, and for mitochondrial genes, the G-INS-I strategy was applied. SequenceMatrix 1.7.7 (Vaidya et al., 2011) was used to concatenated genes for a final matrix. The dataset was portioned by gene, and COI and COII were split into 1+2 and 3. Ribosomal and mitochondrial genes were then concatenated for a final alignment length of 2341bp. Maximum likelihood (ML) using RaxML v8.0.24 (Stamatakis et al., 2008) was implemented through the CIPRES Science Gateway (Miller et al., 2010). One thousand rapid bootstrap replicates were performed, with other parameters keep at default.

Taxonomy

Morphological descriptions

The basal flagellomere is counted as Fl2 following Heraty (2002). The funiculars include Fl2 and 224

the following unfused flagellomeres before the clava, which is defined by 1–3 fused flagellomeres. The paired spines of the mesoscutellum arise from the frenum and are refered here as frenal processes (character 50; Heraty 2002). Morphological abbreviations (see Figs 1 and 2, Chap. I): acl, anteclypeus; acr, acropleuron; ao, anterior ocellus; ax, axilla; axg, axillular groove; axl, axillula; cal, callus; clv, clava; cly, clypeus; cx, coxa; es2, mesepisternum; Fl2, flagellomere basal; flg, flagellum; fmd, femoral depression; frl, frenal line, fp, frenal processes; fub, funicle; gen, gena; Gt1, first gastral tergite; iod, interocular distance; lep2, lower mesepimeron; llm, lateral lobe of mesoscutum; mlm, midlobe of mesoscutum; msp, malar space; mv, marginal vein; not, notaulus; pdl, pedicel; pet, petiole; pl3, metapleuron; pmv, postmarginal vein; po, posterior ocellus; ppd, propodeum; pre, prepectus; sca, supraclypeal area; scd, scrobal depression, scp; scape, sct, mesoscutellar disc; smv, submarginal vein; sss, scutoscutellar suclcus; stv, stigmal vein; tgl, tegula; tor, torulus; tsa, transscutal articulation; upe2, upper mesipimeron.

Measurements

Specimen imaging

225

Specimens repository

The following institutions served as source of material and type depositories for specimens examined for this study: American Museum of Natural History, New York, USA (AMNH); American Museum of Natural History, New York, USA (BMNH); California Academy of Sciences, San Francisco, California, USA (CAS); Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA (CMNH); Canadian National Collection of Insects, Arachnids and Nematods, Ottawa, Ontario, Canada (CNC); Florida State Collection of Arthropods, Division of Plant Industry, Gainesville, Florida, USA (FSCA); Instituto Fundación Miguel Lillo, Tucuman, Argentina (IFML); Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Brasil (INPA); Los Angeles County Museum of Natural History, Los Angeles, California, USA (LACM); Coleção Entomológica do “Instituto Biológico de Ribeirão Preto” – APTA, São Paulo, Brasil (LPRP); Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (MACN); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (MCZ); Museo de La Plata, Universidad Nacional de La Plata, Division Entomologia, La Plata, Argentina (MLPA); Museu Nacional da Costa Rica, San José, Costa Rica (INBIO collection) (MNCR); Museu Paraense Emílio Goeldi, Belém, Pará, Brazil (MPEG); Lund University, Lund, Sweden (MZLU); Museum and Institute of Zoology, Polish Academy of Science, Warsaw, Poland (MZPW); Museu de Zoologia de São Paulo, São Paulo, Brazil (MZUSP); Naturhistorisches Museum Wien, Wien, Austria (NHMW);Royal Ontario Museum, Toronto, Ontario, Canada (ROME); Staatliches Museum für Naturkunde, Stuttgart, Germany (SMNS); University of California, Riverside, California, USA (UCRC); University of Connecticut, Storrs, Connecticut, USA (UCMS); Coleção Entomológica, Departamento de Biologia, Universidade Federal do Espírito Santo, Brazil (UFES); United States National Museum, Washington D.C., USA (USNM); Zoologisk Museum, Copenhagen, Denmark (ZMUC). Specimens are identified with unique UCRC specimen identifiers that indicate the museum of deposition. All specimens were databased in a FileMaker v.11® database maintained in the Heraty lab. Localities were georeferenced using Google Earth (coordinates in italics) or from the original label data. The distribution map was constructed in Simple Mappr (Shorthouse, 2010). 226

Results and discussion

Phylogeny

We are able to recognize four discrete clades within the K. furcata complex in ML analysis (Fig. 1). As in our previous analyses with a much larger taxon sampling (Schoeninger et al., in prep., Chapter II) the furcata complex is monophyletic with high bootstrap support. The furcata complex is distributed from southern Mexico to central Argentina (Fig. 6). Kapala furcata is distinguished from the other species of the complex by the pattern of striations on the face (Fig. 2A), which are oblique from the lower margin of the torulus towards the lateral ocelli. The same pattern can also be observed in K. splendens (Fig. 2B), however, it was not possible to obtain molecular data for this species. K. furcata overlaps with K. parafurcata, but K. parafurcata is easily-recognized due to the weaker oblique striations and longer frenal process. Kapala deltalis and K. quasimodo form a phylogenetic grade, but share similar facial striation with weak striae weakly punctate. They also show a gradient in the morphology of the lateral lobes, with K. deltalis having extremely large and conical lobes (Fig. 3C) and K. quasimodo having less pronounced lobes (Fig. 3A). K. parafurcata and K. cuprea comprise distinct species, which form two different clades. Both species have a pattern of striae identical on the face, however exhibit differences in the frenal processes. K. cuprea has the short and thick frenal processes with longitudinal striae in the basal half and oblique striae in the apical half (Fig. 5C), whereas K. parafurcata has the longest and finest frenal processes and longitudinal striae (Fig. 5D). Differences in the geographical distribution of both species can also be observed. The geographic distribution of K. cuprea is northern South America and K. parafurcata is distributed centrally to southerly (Fig. 6).

Taxonomy

Chirocera Desmarest, 1860: 161, fig. 141. Unjustified emendation of Chirocerus Brullè, with reference to Chirocera furcatus. Kapala Cameron, 1884: 102-103. Replacement name. Kapala; Kirby, 1886: 32. [Kirby questions the identity of Brullè’s drawing and suggests that it looks closer to Thoracantha atrata Walker than to either of the Fabricius species. Indeed, the drawing appears closer to Schizaspidia than to what is now recognized as Kapala. The figure legend in Brullè (1846: pl. 38) refers to Chirocerus furcatus Westwood, not Fabricius, and may not, unfortunately, refers to the Fabricius species. Kapala; Ashmead, 1888[45]: 187 (key). Kapala; Ashmead, 1904[238]: 269,270 (key), 472. Capala Schulz, 1906: 150. Unjustified emendation. Holcokapala Cameron, 1913: 117. Synonymy by Heraty, 2002: 167. Kapala; Gemignani, 1933: 485. Subsequent description. Kapala; Heraty, 1985: 84. Subsequent description. Kapala; Heraty, 2002: 167-170. Subsequent description.

Kapala furcata species complex

Diagnosis. The furcata species complex can be defined by a combination of the following characters: face with fine striae; facial setae spars, more prominent on upper frons, with a row of short transverse setae on anteclypeus; labial palpi 2-3 segmented and females with 9-10 flagellomeres. When there are 10 flagellomeres present, F8-F10are generally broader than long. There is a variation in facial striae (Fig. 2); most species have the frons adjacent to scrobes with oblique striae, traversing between middle ocellar triangle medially to the lower eye margin and lower face with oblique striae from inferior margin of torulus to the posterior genal margin. Two species, K. furcata and K. splendens, however, has striae oblique from the inferior margin torulus to the lateral ocelli and the lower face with transverse striae adjacent to the clypeus (Figs 2A, B). Many of the specimens in this complex have thick frenal processes and robust bodies (Fig. 5).

228

Key of identification to Kapala furcata species complex

1 Lateral lobes of mesoscutum subconical and projecting dorsolaterally to the approximate height of the midlobe (Fig. 3C) …………………………………………………………………………………… Kapala deltalis n. sp. - Lateral lobes lower than height of midlobe and not conical (Fig. 3D) ….…………………………………. 2

2(1) Striation of frons oblique from inferior margin of torulus to the lateral ocelli (Figs 2A, B); lower face with transverse striae adjacent to the clypeus. Facial striation may be widely spaced. Maxillary and labial palpomere formula 2/2 or 3/3 …..………………………………………………………………. 3 - Facial striation oblique from medial ocelli to gena and continuing as oblique on face below torulus (Figs 2C, D). Facial striation closely spaced. Maxillary and labial palpomere formula 3/3…………………………………………………………………………………………………………………………..………………… 4

3(2) Gena with margin broad and angulate. Flagellum of females 8‒9 segmented (Fig. 2B); length of flagellum 1.0–1.1x head height; males with length of rami of F2 1.1X head height. Frenal processes 1.0–1.8x longer than the basal width of scutellum. Body color metallic green to blue ………………………………………………………………………………………………………. K. splendens Ashmead - Gena with margin not broad and angulate. Flagellum of females 9–10 segmented; length of flagellum 0.8–1.0x head height; males with length of rami of F2 1.2–1.3x head height. Frenal processes 1.9–2.3x longer than the basal width of scutellum. Body color black ………………………….. ………………………………………………………………………………………………………………… K. furcata (Fabricius)

4 Face with fine shallow striae, in combination with weak punctuations below torulus (Fig. 2E). Lateral lobes enlarged and midlobes raised with two medial swellings (Fig. 3A). Frenal processes often parallel or U-shape in dorsal view (Fig. 5F). Only males known … Kapala quasimodo n. sp. - Facial striation more deeply impressed. Lateral lobes not enlarged. Females nearly always with 9 flagellomeres, but rarely appearing as 10 in ventral view ………………………………………………………. 5

5 Frenal processes longitudinal striate, extending from base to tips; process 1.3‒1.8x the length of mesoscutellum. Females with 9 or very rarely 10 flagellomeres ………… Kapala parafurcata n. sp. 229

- Frenal processes longitudinal in basal half, and in apical half oblique or spiraling outwards around spines (Fig 5C); processes 1.0‒1.3x the length of mesoscutellum ……………………………………. ………………………………………………………………………………………………….. Kapala cuprea Cameron, 1913

Kapala cuprea Cameron, 1913 (Figs 2C, 3D, 4C, 5C)

Kapala cuprea Cameron, 1913: 116‒117. Type data: Guiana. Holotype ♂, [by original designation]. Type depository: Natural History Museum, London, England (BMNH); type n° 5.390, examined). [Description of the male]. [Holotype relatively complete: exemplar pinned on a quadrangular carboard; region of the axilla and apex of the midlobe dirty with a green gum (like); digits broken and lost; right hind leg broken after coxa, right medial leg broken after femur]. Thoracantha cynipsea Walker, 1862. Type data: Brazil: Villa Nova. Syntype male. Type depository: Natural History Museum, London, England (BMNH); type n° 5.636b. (Withdraw K. cynipsea syntypes in part., only one male). Kapala cynipsea; Ashmead, 1904a: 403 (taxonomic position).

Diagnosis. Recognized by a combination of the following characters: frons and face with oblique striations (Fig. 2C); frenal processes short and stout, 1.0‒1.3x as long as length of mesoscutellum; frenal processes longitudinal striate in basal half and with strong oblique striae in apical half (Fig. 5C).

Biological notes. Associate with Ponerinae (Pachycondyla crassinoda Latreille) (Meyers, 1931).

Distribution. Bolivia, Brazil, Ecuador, French Guiana, Guyana, Peru, West Indies (Fig. 6).

Holotype. GUIANA [probably British Guiana]: P. Cameron Coll., 1914‒110, [1♂, Type Hym. 5.390, BMNH: UCRCENT00310005]. Examined. Additional material examined. Bolivia: Santa Cruz Dept.: 5km SSE Buena Vista, Hotel Fauna y Flora, 440m, 17°29'56"S, 63°39'08"W, 6-15.x.2003, forest, flight intercept trap, S. & J. Peck, 230

[2♂, CNC: UCRCENT00300676, UCRCENT00300677]. Buena Vista, 17°27'32"S, 63°39'33"W, 8.vii.1973, C. Potter, L. Styange & E. Demares, [2♂, IMLA: UCRCENT00242080, UCRCENT00242081]. Estac. Experimental General Saavedra, 430m, 17°47'38"S, 63°11'00"W, 9.vii.1972, C. Porter & L. Stange, [1♂, IMLA: UCRCENT00313142]. Brazil: Amapá: [Pedra Branca do] Amapari – Tucano 2, 0°46'10''N, 51°57'13'', 6-7.xi.1993, Malaise, W. França, [3♂, MPEG]. Idem, 11-15.xi.1993, J. Pena, [5♂, MPEG]. Idem, but serraria, 10-14.xi.1995, equipe, [3♂, MPEG]. Bahia: Porto Seguro, PARNA Monte Pascoal, 16°53'33.65"S, 39°13'0.96"W, 12.viii-2.ix.2013, Malaise, [1♂, UFES]. Mato Grosso: [Campo Novo do Parecis], [Salto] Utiariti, Rio Papagaio, 13°40'30''S,57°53'31''W, x.1966, Lenko & Pereira, [1♂, n° 07549, MZUSP]. Mato Grosso do Sul: [Dois Irmãos do] Buriti, 20°41'23''S, 55°16'43''W, 8.ii.1961, J. & B. Bechyné, [1♂, MPEG]. Pará: Aldeia Coraci 12 km W Caninde, Rio Gurupi, 1°48'38"S, 46°24'03"W, 16- 26.iv.1963, B. Malkin, [1♂, AMNH: UCRCENT00237855]. Benevides, PA 408 - Km 06, 01°21'30''S, 48°14'42''W, 23.iv.1981, Fligth trap, I.S. Gorayeb, [1♂, MPEG]. Idem, 24.iv.1981, [1♂, MPEG]. Idem, 25.iv.1981, [1♂, MPEG]. Rio Araguaia, 19-31.1983, Malaise, J.A. Rafael, [1♂, INPA]. Santarém, 0°57'27"S, 46°59'38"W, [1♀, CMNH: UCRCENT00172445]. Turcurui, 3°42'00"S, 49°42'00"W, i.1978, M. Alvarenga, [1♀, UCRC: UCRCENT00305549]. Pernambuco: Caruaru, Serra dos Cavalos, 800 m, 8°22'23''S, 36°01'04''W, 7.viii.2015, Varredura, J.A. Rafael & P.C. Grossi, [1♂, INPA]. Roraima: Ilha de Maracá, Rio Uraricoera, 3°24'N, 61°42'W, 19- 24.vii.1987, Malaise, J.A. Rafael & L.S. Aquino, [1♂, INPA]. Ecuador: Napo: Sacha, 1°04'59.3"S, 77°37'05"W, 9.iii.1983, L. Huggert, [1♂, MZLU: UCRCENT00242568]. Orellana: 1 km S. Onkone Gare Camp, Reserva Etnica Waorani, 216m, 0°39'25.7"S, 76°27'10.8"W, 2.vii.1995, terre firme forest, fogging, T.L. Erwin et al., [1♀, UCRC: UCRCENT00091807]. Idem, but 9.vii.1995 [1♀, ????: UCRCENT00092227]. Rio Piraña Bridge, Reserva Etnica Waorani, Onkone Gare Camp, 216.3m, 0°39'25.7"S, 76°27'10.8"W, 17.x.2005, terre firme forest, Fogging, T.L. Erwin, M.C. Pimienta et al., [1♂ 1♀, USNM: UCRCENT00247778, UCRCENT00247775]. Sucumbíos: Napo River, Sacha Lodge, 230m, 0°30'00"S, 76°30'00"W, 13-23.iv.1994, malaise trap, P. Hibbs, [1♂, CNC: UCRCENT00320767]. Idem, but 290m, 3- 13.vi.1994, [1♂, CNC: UCRCENT00320768]. Idem, but 4-14.v.1994, [1♂, CNC: UCRCENT00320853]. Guyana: Bartica, Kartabo, 0-30m, 5°45'15"N, 57°42'16"W, 24.iii.1924 [1♀, AMNH: UCRCENT00238159]. Peru: Madre de Dios: Los Amigos Bio. St., 300m, 12°33'44.4"S, 70°05'47.1"W, 25.xii.2010, trail 10, swp, J. Heraty, [1♂, UCRC: 231

UCRCENT00301932]. Rio Tambopata Res., 30 km air SW Pto. Maldonado, 290m, 12°50'00"S, 69°20'00"W, 6.xii.1982, J.J. Anderson, [1♂, FSCA: UCRCENT00411912]. Trinidad: 10°41'26"N, 61°13'16"W, v.2004, [1♂, UCRC: UCRCENT00172511]. Brasso Seco, Rd to Paria Bay, 148m, 10°44'57"N, 61°15'53"W, 25 Jul 2013, forest, swp, Heraty & Baker, [2♂ 1♀, UCRC: UCRCENT00412122‒24]. Curepe, 10°38'48"N, 61°24'56"W, 21.vii.1978, malaise trap, [1♂, UCRC: UCRCENT00305559]. nr. Chaguaramas, 400m, 10°43'01"N, 61°11'52"W, 24.xi.1977, W. & E. Mason, [1♂, CNC: UCRCENT00425741]. Simla Field Station, Arima Valley, 10°41'49"N, 61°17'06"W, 8-9.iii.1977, tropical rain forest, malaise trap, P. Feinsinger, [1♀, FSCA: UCRCENT00306377]. Simla Res. St., 250m, 10°41'34"N, 61°17'23"W, 22.vii.2013, station, Malaise, Heraty & Baker, [1♂, UCRC: UCRCENT00412133]. Tunapuna, Mt. St. Benedict, 500m, 10°40'20"N, 61°23'51"W, 21-8.vii.1993, Mt. Tabor rainforest, Malaise, S&J Peck, [15♂, UCRC: UCRCENT00320102‒03, UCRCENT00320105‒17]. U.S. Virgin Islands: St. Thomas, 1500m, 18°20'17"N, 64°53'39"W, 1859, Thoreg, [1♂, NMW: UCRCENT00317112].

Kapala furcata (Fabricius, 1804) (Figs 2A, 4A, 5A)

Eucharis furcata Fabricius, 1804: 158. Type locality: Am.: mer. [=America Meridionali, South America]. Holotype ♀, by monotypy. Lectotype designated by Heraty, 2002:171 [invalid Lectotype designation, ICZN Art. 74.2]. Type depository: Zoologisk Museum, Copenhagen, Denmark (ZMUC); Type n° ZMUC: 00241187 (image examined). [Holotype not complete: specimen pinned with body dirty; right wings broken and lost; apical half of left frenal processes broken and lost; anterior legs broken and lost]. Thoracantha furcata; Walker, 1839: 22. Chirocerus furcatus; Brullè, 1846: 571. [Kirby (1886) questions the identity of Brullé’s drawing and suggests that it looks closer to Thoracantha atrata Walker than to either the Frabricius species. According Heraty, the drawing appears closer to Kapala sulcifacies (Cameron). The figure legend in Brullè (1846: pl. 38) refers to Chirocerus furcatus Westwood, not Fabricius, and probably refers to a species identified by Westwood. Chirocera furcata; Chenu [Desmarest], 1860: 161, illustrated. 232

Schizaspidia furcata; Walker, 1871: 66. Schisaspidia furcata; Walker, 1872: 65, [misspelling of generic epithet]. Kapala furcata Cameron, 1884: 103, pl. 5, fig. 17. [Change of combination and designation by monotypy as type of Kapala].

Diagnosis. Recognized by a combination of the following characters: frons with oblique wide striae from the torulus to the lateral ocelli (Fig. 2A), and with transverse striae adjacent to clypeus, often deeply impressed across ventral half of face. Female with 9-10 flagellomeres. The medial apex of mesoscutellar disc is lower than other species of the furcata clade, and in lateral view is < 1x height of the base of the frenal processes.

Distribution. Argentina, Brazil and Uruguay.

Holotype. Am.: mer. [=America Meridionali, South America]: Schmidt, [1♀, ZMUC 00241187, ZMUC]. Image examined.

Additional material examined. Argentina: Misiones: Santa Ana, near Loreto, 84m, 27°20'11"S, 55°31'51"W, 27.iii.2003, humid forest, J. Heraty, [1♀, UCRC: UCRCENT00092093]. Salta: Finca El Rey, 1000m, 24°42'00"S, 64°38'00"W, ii.1953, N. Kusnezov, [1♀, IMLA: UCRCENT00313148]. Tucuman: Siambon, 26°42'00"S, 65°27'00"W, i.1945, D. Olea, [1♂, IMLA: UCRCENT00242082]. Tucuman, 26°48'30"S, 65°13'03"W, i.1947, Cordoba, [1♀, IMLA: UCRCENT00313138]. Brazil: Rio Grande do Sul: -, 29°32'04"S, 53°23'26"W, Stieglmayr, [4♂ 6♀, NMW: UCRCENT00242559‒62, UCRCENT00242564, UCRCENT00242566, UCRCENT00317114‒15, UCRCENT00317118‒19]. CPCN Pro-Mata, 325m, 29°30'00"S, 50°10'00"W, 4.iv.1997, J. Ketterl, [1♂, SMNS: UCRCENT00318556]. Idem, but 30.iv.1997, [1♂, SMNS: UCRCENT00318557]. Santa Catarina: Corupa (Hansa Humbolt), 26°25'34"S, 233

49°14'36"W, xii.1940, A. Maller, [1♀, AMNH: UCRCENT00238157]. Pinhal, 27°14'00"S, 51°55'00"W, iv.1947, A. Maller, [1♂, AMNH: UCRCENT00238160]. Rio Natal, 26°21'00"S, 49°18'00"W, i.1945, A. Maller, [3♀, AMNH: UCRCENT00238151, UCRCENT00238153‒54]. Rio Vermelho, 27°29'28"S, 48°24'32"W, ii.1945, A. Maller, [2♀, AMNH: UCRCENT00238150, AMNH: UCRCENT00238152]. Idem, but vi.1958, Dirings, [1♀, MZUSP n° 07555, MZUSP]. São Paulo: Ribeirão Grande, Parque Estadual Intervales, 24°16’23.6S, 48°25’21.08’’W, 22.i.2010, Malaise - Ponto 1, N.W. Perioto e eq., [1♂ 1♀, LPRP]. Idem, but Malaise - Ponto 5, [1♂, LPRP]. Idem, but 21.i.2011, Malaise - Ponto 5, [1♂, LPRP]. Idem, 21.iii.2011, [2♂, LPRP]. Idem, 22.iv.2011, [1♂, LPRP]. Uruguay: Tacuarembo: 40 km NW Tacuarembo, 200-300m, 31°29'45"S, 56°18'08"W, 2-9.ii.1963, J.K. Bouseman, [2♂, AMNH: UCRCENT00237807, AMNH: UCRCENT00238155].

Kapala splendens Ashmead, 1904 (Figs 2B, 4B, 5B)

Kapala splendens Ashmead, 1904a: 473. Type data: Brazil: Chapada. Holotype ♀, by original designation. Type depository: National Entomological Collection, National Museum of Natural History (USNM), Washington, DC, USA; type nº 8084, (examined). [Description of female, illustrated. Holotype relatively complete: apex of antennae broken and lost; left middle leg broken and lost; coxa and hind legs broken and lost; apex of right and left forewings broken; right and left hindwing with apical half broken; abdomen and petiole broken and glued erroneously below the frenal processes].

Diagnose. Recognized by a combination of the following characters: body very robust than the others Kapala species; margin of gena broad and angulate (Fig. 2B); frons with fine oblique striae from lateral ocelli curving towards the supraclypeal area and face with fine transversal striae (Fig. 2B); flagellum 8 or 9 segmented; head and mesosoma brilliant metallic green and metasoma blue with green reflections. Biological notes. Unknown.

Distribution. Argentina and Brazil (Fig. 6). 234

Holotype. BRAZIL: Chapada, in April and November, H.H. Smith coll., [1♀, USNMENT 00809391, type n° 8084].

Additional material examined. Argentina: 38°30'42.01"S, 63°38'23.89"W, [1♀, MLPA]. Misiones: San José, 27°46'12.18"S, 55°46'49.33"W, xii.1942, Bridarolli, S.J., [1♂, MACN]. Brazil: Goiás: Alto Paraíso de Goiás, BR 010, 18°56'48''S, 47°25'53''W, 1400m, 28.xii.2011, V. Kanamura & K. Ramos, [1♀, MZUSP]. Goiás, 15°56'02''S, 50°08'24''W, 26.iii.1978, A. Raw, [1♀, UCRC_ENT 297983, BMNH]. Mato Grosso: Chapada, 15°27'29.90"S, 55°45'8.85"W, H.H. Smith, [2♀, Acc.2966, UCRC_ENT 00240757, CMNH; Acc. 2966, UCRC_ENT 00240756, CMNH]. Rio de Janeiro: 22°54'54.90"S, 43°10'7.27"W, [1♀, UCRC_ENT 00416661, USNM].

New species descriptions

Kapala deltalis Murray & Schoeninger, n. sp. (Figs 2F, 3C, 4D, 5E)

Diagnose. Recognized by a combination of the following characters: distinctive subconical dorsolateral swelling of the mesoscutal sidelobes (Fig. 3C) and by apex of the mesoscutellar disc with a medially acute, sharp apical crest (Fig. 4D). Another species in the furcata complex, K. quasimodo n. sp., also has enlarge lateral lobes, but they do not reach the height of the midlobe.

Description. Female. Body length 4.9‒5.8 mm; length of mesosoma excluding the frenal processes 2.7‒3.1 mm. Head, mesosoma, coxae, petiole and frenal processes black; scapes and pedicels yellow; flagellum light brown; femora pale yellowish to light brown; wings hyaline to an even light infuscation, venation brown; Gt1 black, subsequent tergites orange. Head. 1.4–1.5x as broad as high, with few sparse setae. Scrobal depression with faint irregular striae. Face with a pattern of fine uniform striations and punctations (Fig. 2F); frons adjacent to scrobes with oblique striae, traversing between middle ocellar triangle to lower eye margin; lower face with striae broadly curved from torulus to posterior genal margin. Eyes separated by 2.2–2.3x their height. Malar space 0.9–1.1x eye height. Supraclypeal area and clypeus smooth; anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 or 12 segments; scape 3.0–4.0x longer than broad, 0.3x head height, 235

smooth surface; flagellum 9–10 segmented, basal flagellomeres cylindrical or slightly serrate, clava rounded and sometimes subdivided resulting in 10 flagellomeres; length of flagellum 1.0–

1.2x head height; Fl2 2.0–2.7x as long as apical width and 1.2–1.6x as long as following flagellomere. Mesosoma. Mesoscutum anteriorly abruptly curved in dorsal half, in lateral view, and 1.3–1.5x as high as broad. Midlobe with fine, regular transverse striae and with flattened anterior face; lateral lobes subconical and projecting dorsolaterally to the height of the midlobe (Fig. 3C). Axilla and mesoscutellar disc with shallow, regular longitudinal striae. Mesoscutellar disc 1.1– 1.2x as long as axilla with medial apex raised 1.1–1.6x height of frenal processes in lateral view and bordered by a strong carina posteriorly (Fig. 4D). Axillula smooth and eventually with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum smooth and shining except for weak rugose crenulate ventrally; upper mesepimeron smooth or with weak punctations, femoral depression crenulate. Callus slightly swollen, carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeal disc flat and broad, with shallow areolate crenulations on dorsal margin. Frenal processes cylindrical, 2.1–2.6x longer than the basal width of mesoscutellum, 1.4–1.5x as long as length of axilla and mesoscutellar disc; frenal processes, arched in lateral view and convex in dorsal view; frenal processes with longitudinal striae and with apex emarginate (Fig. 5E). Forewing 2.5–2.8x longer than broad covered by small bristles, with the exception of the base that is bare; submarginal vein 1.0–1.3x longer than marginal vein; stigmal vein 1.5–2.0x longer than broad; posmarginal vein absent. Metacoxa semiglobose 1.7–2.1x longer than broad, with medial row of setae. Metafemur 6.8–7.8x longer than broad, smooth with sparse appressed setae. Metasoma. Petiole cylindrical and with fine parallel longitudinal carinae, slightly flattened dorsally, petiole 3.1–4.1x longer than broad and 1.3–1.7x as long as metacoxa. Gt1 bare except for sparse minute setae; tergal scar present.

Male. Body length 4.6–5.2 mm; length of mesosoma excluding the frenal processes 2.4– 2.8 mm. Similar to female except for the following: Eyes separated by 1.9–2.1x their height. Antenna with 12 segments; scape 2.5–3.0x longer than broad; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami Fl2 1.2–1.4x head height 236

and 0.9x as long as following rami. Petiole 4.7–5.3x longer than broad and 2.0–2.7x as long as metacoxa.

Etymology. From the Greek letter delta, Δ. The shape of the mesosoma resembles an inverted triangle from the anterior aspect.

Biological notes. Unknown.

Distribution. Costa Rica and México (Fig. 6).

Holotype. Costa Rica: Guanacaste: Macizo Miravalles, Estación Cabro Muco, 1100m, 10°43'00"N, 85°08'34"W, 23-28.ix.2003, J. Azofeifa, Tp de Luz, L N 299769 411243 #75484 [1♀, INBIO: INBIOCRI03778914, deposited in UCRC].

Paratypes (7♀, 46♂): Costa Rica: Alajuela: P.N. Rincon de la Vieja, 2 km N Colonia Blanca, 800m, 18-28.vi.1992, III curso Parataoxon, L-N 308800, 397800, [1♂ 1♀, INBIO: INBIOCRI00703908, INBIOCRI00704253]. Guanacaste: Est. Cacao, 2 km SW del Cerro Cacao, 800-1400m, 10°55'59"N, 85°28'03"W, 7-18.ii.1995, L N 323100 375800 #5320, M. Iobo, [2♂, INBIO: INBIOCRI00235220‒21]. Idem, but 1500m, iii.1996, #8400, C. Moraga, Sombrereta, [1♂, INBIO: INBIOCRI00473635]. Est. Cacao, SW side Volcan Cacao, 1000-1400m, 10°55'42"N, 85°28'06"W, xi-xii.1989, URCG R. Blanco & C. Chaves, [6♂, INBIO: INBIOCRI00143717, INBIOCRI00143797, INBIOCRI00144236, INBIOCRI00144249, INBIOCRI00144251, INBIOCRI00146357]. Idem, but x.1989, 323300, 375700, [9♂ 2♀, INBIO: INBIOCRI00097313, INBIOCRI00097315, INBIOCRI00097317, INBIOCRI00097320, INBIOCRI00097322‒23, INBIOCRI00097325, INBIOCRI00097331, INBIOCRI00097333‒34, INBIOCRI00097341]. Estac. Mengo, SW Volcan Cacao, 1100m, 10°56'03"N, 85°27'22"W, 1988-1989, [2♂, INBIO: UCRCENT00305583, UCRCENT00305593]. Idem, but vii.1988, P. Hanson, [1♂, INBIO: UCRCENT00305586]. Guanacaste N.P., Biol. Sta. Cacao, 900m, 10°35'03"N, 85°22'46"W, 13.ii.1995, screen sweep, L. Masner, [11♂ 1♀, CNC: UCRCENT00316290‒95, UCRC: UCRCENT00305703‒08]. Macizo Miravalles, Estación Cabro Muco, 1100m, 10°43'00"N, 85°08'34"W, 22.ix-5.x.2003, MT #2, L_N 299769_411243 #75755, 237

J.D. Gutierrez, [2♂, INBIO: INBIOCRI03983169‒70]. Idem, but 24.ix-5.x.2003, MT #1, L_N 299769_411243 #75731, B. Hernandez, [4♂ 1♀, INBIO: INBIOCRI03730848, INBIOCRI03983084, INBIOCRI03983089, INBIOCRI03983129, INBIOCRI03983131]. Rio San Lorenzo, Tierras Morenas, 1050m, x.1995, L_N 287800_427600 #6405, G. Rodriguez, [1♂, INBIO: INBIOCRI00453881]. Puntarenas: San Luis, Monteverde, 1000-1350m, 10°16'31"N, 84°47'40"W, ii.1995, L_N_250850_449250 #4393, Z. Fuentes, [1♂, INBIO: INBIOCRI00165650]. San Luis, Monteverde, 1040m, 10°16'25"N, 84°49'27"W, iv.1994, L-N 449250_250850 #2845, Z. Fuentes, [1♂, INBIO: INBIOCRI00796351]. Idem, but 24.viii- 15.ix.1992, L-N 250850, 449250, F.A. Quesada, [1♀, INBIO: INBIOCRI00856129]. Idem, but vii.1992, L-N 250850, 449250, Z. Fuentes, [1♂, INBIO: INBIOCRI00722852]. Idem, but, viii.1992, [1♂, INBIO: INBIOCRI00754455]. Idem, but xii.1992, [1♂, INBIO: INBIOCRI00958023]. Mexico: Chiapas: 32 km N of Ocozocoautla, on road to Malpaso, 762m, 16°59'01"N, 93°30'17"W, 6.x.1974, D.E. & J.A. Breedlove, [1♂ 1♀, CASC: CASENT02174570, UCRCENT00292310].

Kapala parafurcata Murray & Schoeninger, n. sp. (Figs 2D, 3B, 4F, 5D)

Diagnose. Recognized by a combination of the following characters: face with regularly-spaced, oblique striations on the frons and lower face (Figs 2D, 3B) in combination with frenal processes tapering to apex and 1.3‒1.8x as long as length of mesoscutellum (Fig 5D). Distinguished from K. cuprea which has frenal processes 1.0‒1.3x as long as length of mesoscutellum.

Description. Female. Body length 2.6‒5.2 mm; length of mesosoma excluding the frenal processes 2.4‒3.1 mm. Head, mesosoma, coxae, petiole and frenal processes black; scapes and pedicels yellow, flagellum light brown; legs yellowish to light brown; wings hyaline, venation brown; Gt1 dark-brown, subsequent tergites light brown. Head. 1.4–1.5x as broad as high, with few sparse setae, more prominent in frons. Scrobal depression reticulate with some faint longitudinal striae. Frons adjacent to scrobes with oblique striate, traversing between middle ocellar triangle to lower eye margin (Fig. 2D); lower face with striae broadly curved from torulus to posterior genal margin. Eyes separated by 2.0–2.2x their 238

height. Malar space 0.8–1.0x eye height. Supraclypeal area and clypeus smooth; anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 11 or 12 segments; scape 3.0–3.5x longer than broad, 0.3x head height; flagellum 9–10 segmented, basal flagellomeres cylindrical or slightly serrate, clava rounded and sometimes subdivided resulting in

10 flagellomeres; length of flagellum 0.9–1.0x head height; Fl2 1.5–1.7x as long as apical width and 1.2–1.5x as long as following flagellomere. Mesosoma. Mesoscutum anteriorly abruptly curved in dorsal half (Fig. 4F), in lateral view, and 1.3–1.6x as high as broad. Midlobe with regular transverse striae; lateral lobes with regular longitudinal striae and elevated to approximately the height of medial midlobe. Axilla and mesoscutellar disc with longitudinal striae (Fig. 5D). Mesoscutellar disc 1.4–1.5x as long as axilla with medial apex raised 1.0x height of frenal processes in lateral view. Axillula longitudinal striate. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron with weak to strong transverse striae; femoral depression crenulate. Callus slightly swollen, carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeal disc flat and broad, with shallow areolate crenulations on dorsal margin. Frenal processes cylindrical, 2.0–2.4x longer than the basal width of mesoscutellum, 1.0–1.5x as long as length of axilla and mesoscutellar disc; frenal processes arched in lateral view and parallel convexly rounded in dorsal view, longitudinally carinate and frenal processes tapering to apexes which are emarginate (Fig. 5D). Forewing 2.4–2.6x longer than broad covered by small bristles, with the exception of the base that is bare; submarginal vein 1.0–1.2x longer than marginal vein; stigmal vein 1.5–2.0x longer than broad; postmarginal vein very short. Metacoxa semiglobose 1.8–2.0x longer than broad, with medial row of setae. Metafemur 5.7–6.2x longer than broad, smooth with sparse appressed setae. Metasoma. Petiole cylindrical and with fine parallel longitudinal carinae, slightly flattened dorsally, petiole 3.0–4.3x longer than broad and 1.1–1.4x as long as metacoxa. Gt1 bare except for sparse minute setae; tergal scar present.

Male. Body length 3.9‒4.3 mm; length of mesosoma excluding the frenal processes 2.1‒2.3 mm. Similar to female except for the following: scape 1.8–2.8x longer than broad; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami

Fl2 1.2–1.3x head height and 0.9‒1.0x as long as following rami. Frenal processes cylindrical, 239

2.5–2.8x longer than the basal width of mesoscutellum, 1.5–1.7x as long as length of axilla and mesoscutellar disc; petiole 4.0‒5.0x longer than broad and 1.8–2.0x as long as metacoxa

Etymology. The name derives from the close similarity to K. furcata in body shape, and due to many museum specimens of this species previously having been identified as “near K. furcata”.

Biological notes. Unknown.

Distribution. Argentina, Brazil, Paraguay, Peru (Fig. 6).

Holotype: Argentina: Misiones: Loreto, Ruinas Jesuiticas, 27°29'59"S, 55°31'59"W, 4.xi.2001, malaise trap, S.O. Martinez, P. Fidalgo, [1♀, UCRC: UCRCENT00091817, deposited in UCR].

Paratypes (90♂, 15♀): Argentina: Buenos Aires, 34°36'30"S, 58°22'23"W, Antigua, [1♂, MACN: UCRCENT00242244]. Corrientes: Las Marias, Ca. Virasoro, 28°05'33"S, 56°02'42"W, 7.xi.1971, C. Porter, [2♂, IMLA: UCRCENT00242086, UCRCENT00313140]. Misiones: Aristobulo del Valle, 27°05'51"S, 54°53'47"W, 28.xi.1951, Montes & Willink, [1♂, IMLA: UCRCENT00313136]. Cataratas del Iguazu, 25°40'00"S, 54°27'00"W, 5-9.xi.1970, C. Potter & L. Stange, [7♂, IMLA: UCRCENT00242084‒85, UCRCENT00274414‒15, UCRCENT00274417, UCRCENT00313134‒35]. Dos de Mayo, 27°02'00"S, 54°39'00"W, xii.1973, Fritz, [2♂, AMNH: UCRCENT00237787‒88]. Iguazu, 25°55'25"S, 54°21'51"W, 30.i- 13.iii.1945, Hayward, Willink & Golbach, [2♀, IMLA: UCRCENT00313139, UCRCENT00313141]. Loreto, Ruinas Jesuiticas, 285m, 27°30'00"S, 55°32'00"W, 26.i- 20.ii.2001, S.O. Martinez, P. Fidalgo, MT, [1♂, UCRC: UCRCENT00302400]. Los Helechos, 350m, 27°32'05"S, 55°05'07"W, v.1949, Duret, [1♂, AMNH: UCRCENT00237791]. Parque Nacional Iguazu, 200m, 25°40'48"S, 54°27'00"W, 2-7.xii.2003, B.V., malaise trap, Brown & G. Kung, [1♂, UCRC: UCRCENT00316349]. Puerto Bemberg, 25°55'10"S, 54°35'08"W, [1♂, MACN: UCRCENT00242245]. Reserva de Vide Silvestre Urugua-I, 400m, 25°58'28"S, 54°06'59"W, 7-9.xii.2003, malaise trap, B. Brown & G. Kung, [3♂, UCRC: UCRCENT00172510, UCRCENT00172515‒16]. RN 12, N of Puerto Bosseti, 221m, 25°48'20"S, 54°32'19"W, 25.iii.2007, Aurac.for., H07-025, J.&J. Heraty & J. Torréns, [1♀, 240

UCRC: UCRCENT00000325]. Rt17, E of 9 de Julio, 212m, 26°24'02"S, 54°27'54"W, 26.iii.2007, roadside, H07-031, J.&J. Heraty & J. Torrens, [2♂, UCRC: UCRCENT00000324, UCRCENT00161498]. Santa Ana, near Loreto, 84m, 27°20'11"S, 55°31'51"W, 27.iii.2003, J. Heraty, humid forest, H03-038, [2♂ 1♀, UCRC: UCRCENT00091803, UCRCENT00091921, UCRCENT00172312]. Salta: Oran, Rd to San Andres along Rio Blanca, 399m, 23°05'30"S, 64°21'57"W, 22.iii.2003, sclerophyll scrub, H03-015, J. Heraty, [1♂, UCRC: UCRCENT00092070]. Brazil: Mato Grosso: Maracaju, 21°37'08"S, 55°10'02"W, iv-v.1937, G. Fairchild, [1♂ 1♀, MCZ: UCRCENT00242335, UCRCENT00242337]. Paraná: Prudentópolis, 25°12'56"S, 50°58'08"W, 23-25.ii.1969, C. Porter & A. Garcia, [1♂, MCZ: UCRCENT00242336]. Pernambuco: Caruaru, 900m, 8°16'51"S, 35°58'32"W, vi.1972, J. Lima [1♂, ROME: UCRCENT00242713]. Rio de Janeiro: Rio de Janeiro, 22°54'13"S, 43°12'35"W, Westwood, [1♂, ZMUC: UCRCENT00245090]. Santa Catarina: -, 26°25'34"S, 49°14'36"W, Luderwaldt, [1♂ 1♀, MZPW: UCRCENT00242646, UCRCENT00242680]. Nova Teutonia, 300-500m, 27°11'00"S, 52°23'00"W, 1.iv.1937, Fritz Plaumann, [1♂, LACM: UCRCENT00242214]. Idem, but 30.iii.1966, [1♂, CNC: UCRCENT00300619]. Idem, but 9.ii.1967, [1♀, CNC: UCRCENT00300603]. Idem, but i.1969, [2♂, CNC: UCRCENT00300627, UCRCENT00300629]. Idem, but ii.1968, [3♂, CNC: UCRCENT00300605, UCRCENT00300612 ‒13]. Idem, but iii.1968, [3♂, CNC: UCRCENT00300606, UCRCENT00300618, UCRCENT00300623]. Idem, but iii.1969, [1♂ 2♀, CNC: UCRCENT00300601‒02, UCRCENT00300611]. Idem, but iii.1971, [2♂, CNC: UCRCENT00300614, UCRCENT00300624]. Idem, but iv.1968, [5♂ 1♀, CNC: UCRCENT00172444, UCRCENT00300604, UCRCENT00300607, UCRCENT00300609, UCRCENT00300616, UCRCENT00300628]. Idem, but xii.1968, [7♂, CNC: UCRCENT00300608, UCRCENT00300610, UCRCENT00300620‒22, UCRCENT00300625‒26]. São Paulo: Am. Brasiliense Cerradão Clube Náutico, 21°42'12.1"S, 48°01'24.7"W, 5.v.2000, Varredura Vegetação (5 min.) horas 12:15, M.T. Tavares et al., [1♂, UFES: UFES00002697]. Idem, but 10:44, [2♂, UFES: UFES00002695, UFES00002696]. Idem, but 11:55, [1♂, UFES: UFES00002694]. Idem, but 14:15, [1♂, UFES: UFES00002698]. Idem, but 10:38, [1♂, UFES: UFES00002701]. Idem, but Arm. Corredor, 21°42'21.1"S, 48°01'24.7"W, 5-10.v.2000, Malaise-ponto 2, M.T. Tavares et al., [1♂, UFES: UFES00002708]. Paraguay: Salto del Guaira (sw), 24°03'24"S, 54°18'30"W, 8.xii.1971, L. Pena, [1♂, CNC: 241

UCRCENT00300617]. Idem, but 3.xii.1971, [1♂, CNC: UCRCENT00247548]. San Bernardino, 25°16'12"S, 57°19'12"W, ii, Fiebrig, [1♂ 1♀, NMW: UCRCENT00317116, ZSMC: UCRCENT00245260]. San Pedro: Carumbe, 125m, 25°33'00"S, 56°40'00"W, 1.ii-8.iii.1966, R. Golbach, [1♂, IMLA: UCRCENT00274416]. Rio Ypane, Cororo, 23°26'22"S, 56°30'57"W, xii.1983, M.A Fritz, [1♂, AMNH: UCRCENT00237786]. Colonia Independencia, 25°41'43"S, 56°15'34"W, 27.iii.1951, [6♂, MACN: UCRCENT00242227‒32]. Paso Yobai, 200-230m, 25°43'31"S, 55°59'50"W, 15.ii.1951, [1♂, MACN: UCRCENT00242226]. Pirapo, 26°51'19"S, 55°32'29"W, 28.xii.1971, L. Pena, [4♂, CNC: UCRCENT00300615, UCRCENT00300630‒32]. Puerto Stroessner, 25°30'31"S, 54°40'31"W, 7.xii.1971 [1♂, CNC: UCRCENT00247547]. Alto Paraná: Central Forest, 12 km N of Ciudad del Este, 25°25'29"S, 54°37'01"W, 29.iv-5.v.1986, R.E. Woodruff, [1♂ 1♀, FSCA: UCRCENT00411949‒50]. Caazapá: Estancia Condesa/Toro Blanco, San Rafael Reserve, 110m, 26°19'11"S, 55°39'57"W, 8-10.xii.2000, flight intercept trap, Z.H. Falin, [1♂, ????: UCRCENT00397274]. Guaira: -, 25°52'52"S, 56°17'35"W, xii.1950, Carl Pfannl & Foerster, [2♂, AMNH: UCRCENT00237798‒99]. Colonia Independencia, 25°41'43"S, 56°15'34"W, iv.1951, Foerster, [3♂, AMNH: UCRCENT00237792‒93, UCRCENT00237898]. Idem, but xii.1950, [1♂, AMNH: UCRCENT00237797]. Villarrica, 25°47'04"S, 56°27'03"W, vi, F. Schade, [1♂, MCZ: UCRCENT00242334]. Itapua: 17 km N. Hohenau, 200m, 26°57'14"S, 55°32'32"W, 2-4.ii.1983, E.G. Riley, [1♀, AMNH: UCRCENT00237790].

Kapala quasimodo Murray & Schoeninger, n. sp. (Figs 2E, 3A, 4E, 5F)

Diagnose. Recognized by a combination of the following characters: finely striate and punctate face (Fig. 2E), lateral lobes enlarged and two dorsal swellings of the midlobe of the mesoscutum (Fig. 3A); distance between the frenal processes typically as wide at apex as at base, forming a “U-shape”; frenal processes weakly carinate (Fig. 5F). Another species in furcata complex, K. deltalis n. sp., has much larger lateral lobes, reaching the height of the mid lobe. Description. Male (female unknown). Body length 4.2‒5.1 mm; length of mesosoma excluding the frenal processes 2.4‒2.7 mm. Head, mesososma, coxae, petiole and frenal processes black; scapes yellow, flagellum brown; legs pale yellow; wings hyaline to an even light infuscation, venation light brown to brown; Gt1 black, subsequent tergites dark orange. 242

Head. 1.4–1.5x as broad as high, with few sparse setae more prominent in frons. Scrobal depression transversal striate. Face with a pattern of fine uniform striations and punctations (Fig. 2E); frons adjacent to scrobes with oblique striae, traversing between middle ocellar triangle medially to lower eye margin; lower face with striae broadly curved from torulus to posterior genal margin. Eyes separated by 1.9–2.1x their height. Malar space 0.8–0.9x eye height. Supraclypeal area and clypeus smooth; anteclypeus with a row of short transverse setae. Maxillary/labial palpi 3/3 segmented. Antenna with 12 segments; scape 3.0–3.7x longer than broad and 0.3x head height; flagellum 10 segmented, transversal and each flagellomere with a single long rami; length of rami Fl2 1.2–1.3x head height and 0.9x as long as following rami. Mesosoma. Mesoscutum anteriorly abruptly curved in dorsal half (Fig. 4E), in lateral view, and 1.3–1.4x as high as broad. Midlobe with fine, regular transverse striae and raised with two medial swellings; lateral lobes enlarged and with regular longitudinal striae (Fig. 3A). Axilla and mesoscutellar disc with longitudinal striae. Mesoscutellar disc 1.4–1.8x as long as axilla with medial apex raised 1.1–1.4x height of frenal processes in lateral view. Axillula longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum with weak longitudinal striae and crenulate ventrally; upper mesepimeron with weak punctations and longitudinal striae; lower mesepimeron reticulate; femoral depression crenulate. Callus slightly swollen, carinate and reticulate, covered by semi-erect setae. Metapleuron reticulate. Propodeal disc flat and broad, with shallow areolate crenulations on dorsal margin. Frenal processes cylindrical, 2.4–2.8x longer than the basal width of mesoscutellum, 1.5–1.6x as long as length of axilla and mesoscutellar disc; frenal processes often parallel or U-shape in dorsal view and longitudinally carinate (Fig. 5F), apex rounded. Forewing 2.4–2.6x longer than broad covered by small bristles, with the exception of the base that is bare; submarginal vein 1.2–1.3x longer than marginal vein; stigmal vein 1.5x longer than broad; postmarginal vein absent. Metacoxa semiglobose 1.5–1.7x longer than broad, with medial row of setae. Metafemur 6.2– 7.0x longer than broad, smooth with sparse appressed setae. Metasoma. Petiole cylindrical and with fine parallel longitudinal carinae, slightly flattened dorsally, petiole 5.8–6.6x longer than broad and 2.1–2.3x as long as metacoxa. Gt1 bare except for sparse minute setae; tergal scar present.

243

Etymology. Specific name “in apposition”, named for Quasimodo, Victor Hugo’s protagonist of the Hunchback of Notre Dame. This species has projections of the mesosoma mid and lateral lobes, giving it a hunched appearance.

Biological notes. Unknown.

Distribution. Ecuador and Venezuela (Fig. 6).

Holotype: Venezuela: Aragua: Rancho Grande N.P., 1100m, 10°24'13"N, 67°34'26"W, 18.viii- 3.ix.1992, cloud forest, maxinet, L. Masner, [1♂, UCRC: UCRCENT00397263, deposited in UCR].

Paratypes (22♂): Ecuador: Esmeraldas: Bilsa Biol. Sta., 500m, 0°20'24"N, 79°42'36"W, 10.v- 4.vi.1996, malaise trap, P. Hibbs, [1♂, UCRC: UCRCENT00092084]. Idem, but 7-19.vii.1996, malaise trap/flight intercept trap, [1♂, UCRC: UCRCENT00092002]. Venezuela: Aragua: Rancho Grande N.P., 1100m, 10°24'13"N, 67°34'26"W, 18.viii-3.ix.1992, cloud forest, maxinet, L. Masner, [17♂, UCMS: UCRCENT00397267, UCRC: UCRCENT00172446‒60, UCRCENT00397264]. Rancho Grande, 1150m, 10°04'00"N, 67°32'36"W, 4.vii-9.viii.1986, B. Gill [3♂, CNC: UCRCENT00425738‒40].

Acknowledgment

244

References

Ashmead, W.H. (1904) Classification of the chalcid flies, or the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected in South America by Herbert H. Smith. Memoirs of the Carnegie Museum, 1: i–ix, 225–551, pls. xxxi–xxxix. Bruleè, M.A. (1846) A. LePeletier de Saint-Fargeau, Histoire Naturelle des Insectes par M. LeConte. Hyménopterès. IV. Prais, 680 pp. Cameron, P. (1884) Hymenoptera (Families Tenthredinidae-Chrysididae). In Goddman, F.D., and Salvin, D., Biologia centrali-americana. Insecta Hymenoptera. London: Taylor and Francis. 1:1‒487 + 20pls. Cameron, P. (1904) New Hymenoptera mostly from Nicaragua. Invertebrata pacifica, 1: 46 Cameron, P. (1913) The Hymenoptera of the Georgetown Museum, Part V. Journal of Royal Agricultural Society, Demerara 3: 105–137. Clausen, C.P. (1940) The oviposition habits of the Eucharidae (Hymenoptera). Journal of the Washington Academy of Sciences, 30: 504–516. Clausen, C.P. (1941) The habits of the Eucharidae. Psyche: A Journal of Entomology, 48, 57-69. Fabricius, J.H. (1804) Systema Piezatorum. Brunsvigae: Carolum Reichard. 2: xiv + 30, 440pp. Gibson, G.A.P., Read, J.D. & Fairchild, R. (1998) Chalcid wasps (Chalcidoidea): illustrated glossary of positional and morphological terms. Available from: http://www.canacoll.org/Hym/Staff/Gibson/apss/chglintr.htm (accessed 22 March 2016) Harris, R.A. (1979) A glossary of surface sculpturing. Occasional papers in Entomology, 28:1 31. Heraty, J.M. (2002) A Revision of the Genera of Eucharitidae (Hymenoptera: Chalcidoidea) of the World. Memoirs of the American Entomological Institute, 68: 368 pp. Heraty, J.M. & Woolley, J.B. (1993) Separate species or polimorphism; a recurring problem in Kapala (Hymenoptera: Eucharitidae). Annals of the Entomological Society of America, 85: 517-531. Kirby, W.F. (1886) A synopsis of the genera of the Chalcididae, subfamily Eucharinae, with descriptions of several new genera and species of Chalcididae and Tenthredinidae. Journal of the Linnean Society, 20: 28–37. 245

Katoh, K., Kuma, K., Toh, H. & Miyata, T. (2005) MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic. Acids Res., 33, 511-8. Myers, J.C. (1931) Descriptions and records of parasitic Hymenoptera from British Guiana and the West Indies. Bulletin of Entomological Research, 22: 267–277. Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 1-8. Murray, E.A. (2014) Systematics and evolution of Eucharitidae (Hymenoptera: Chalcidoidea), with a focus on the New World Kapala. University of California, Riverside, CA. Murray, E. A. & Heraty, J. (2016) Invading Africa: a novel transoceanic dispersal by a New World ant parasitoid. Journal of Biogeographic, 43: 1750‒1761. Sequencher® version 5.4.6 DNA sequence analysis software, Gene Codes Corporation, Ann Arbor, MI USA. Access in: http://www.genecodes.com Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57: 758-71. Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics, 27, 171-180. Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. Walker, F. (1839) Monographia Chalciditum. Vol. 2. London. 100 pp. Walker, F. (1862) Notes on chalcidites and characters of undescribed species. Transactions of the Entomological Society of London, 1: 345–397. Walker, F. (1871) Notes on Chalcidiae. Part IV. Chalcididae, Leucospididae, Agaonidae, Eucharidae, Perilampidae, Ormyridae, Encyrtidae. London, 55–70. Walker, F. (1872) Economy of Chalcididiae (continued). The Entomologist, 6: 65–70. Walsh, P.A., Metzger, D.A. & Higuchi, R. (1991) Chelex 100 as a medium for simple extraction of DNA for PCR-based typing from forensic material. Biotechniques, 10: 506-513.

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Figure 2. Kapala furcata complex faces. A. K. furcata (♀); B. K. splendens (♀); C. K. cuprea (♀); D. K. parafurcata n. sp.; E. K. quasimodo n. sp. (♂); F. K. deltalis n. sp. (♀).

Figure 3. Head and mesososma of K. furcata complex. A. K. quasimodo n. sp. (♂); B. K. parafurcata n. sp. (♀); C. K. delatlis n. sp. (♀); K. cuprea (♀).

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Figure 4. Kapala furcata complex lateral habitus of mesosoma. A. K. furcata (♀); B. K. splendens (♀); C. K. cuprea (♀); D. K. deltalis n. sp. (♀); E. K. quasimodo n. sp. (♂); F. K. parafurcata n. sp. (♀).

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250

Figure 6. Species distribution map, from holotype and paratype material. Points mapped in SimpleMappr.

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Supplementary information

Table S1. Locality, museum, identification number (ID) and DNA voucher number information for the 35 taxa of furcata complex and outgroup (iridicolor complex) used in the molecular analysis. DNA voucher Species Sex ID number number Museum Locality K. furcata ♀ UCRENT_92093 D#1078 UCRC Argentina: Misiones, Santa Ana, 27.iii.2003, J. Heraty K. deltalis sp. n. ♂ UCRCENT_92109 D#1388 UCRC Costa Rica: Guanacaste, Biol. Sta. Cacao, 13.ii.1995, L. Masner K. quasimodo sp.n. ♂ UCRCENT_92002 D#0942 UCRC Ecuador: Esmeraldas, Bilsa Biol. Station, 7-19.vi.1996, P. Hibbs K. parafurcata sp.n. ♂ UCRCENT_91921 D#1069 UCRC Argentina: Misiones, Santa Ana, 27.iii.2003, J. Heraty K. parafurcata sp.n. ♂ UCRCENT_161498 D#2519 UCRC Argentina: Misiones, Rt17, 26.iii.2007, J. Heraty K. parafurcata sp.n. ♂ UCRCENT_92073 D#1068a UCRC Argentina: Salta, Oran, 22.iii.2003, J. Heraty K. parafurcata sp.n. ♂ UCRCENT_397224 D#3436 UCRC Paraguay: Caazapá, San Rafael Reserve, 8-10.xii.2000, Z.H. Falin K. parafurcata sp.n. ♂ UCRCENT_000324 D#2520 UCRC Argentina: Misiones, Rt17, 25.iii.2007, J. Heraty Argentina: Misiones, Loreto, Ruinas Jesuiticas, 4.xi.2001, Martinez & K. parafurcata sp.n. ♀ UCRCENT_91817 D#0711 UCRC Fidalgo Argentina: Misiones, RN 12, N of Puerto Bosseti, 25.iii.2007, Heraty & K. parafurcata sp.n. ♀ UCRCENT_000325 D#2518 UCRC Torréns K. parafurcata sp.n. ♀ UCRCENT_91803 D#1086 UCRC Argentina: Misiones, Santa Ana, near Loreto, 27.iii.2003, J. Heraty K. cuprea ♂ UCRCENT_412124 D#3806 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker K. cuprea ♂ UCRCENT_412123 D#3805 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker K. cuprea ♀ UCRCENT_412122 D#3804 UCRC Trinidad: Brasso Seco, Rd to Paria Bay, 25.vii.2013, Heraty & Baker K. cuprea ♂ UCRCENT_412133 D#3815 UCRC Trinidad: Simla Res.St., 22.vii.2013, Heraty & Baker K. cuprea ♀ UCRCENT_91807 D#1003 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 2.vii.1995, T.L. Erwin et al. K. cuprea ♂ UCRCENT_320768 D#3837 CNC Ecuador: Sucumbíos, Napo River, Sacha Lodge, 3-13.vi.1994, P. Hibbs K. cuprea ♀ UCRCENT_247775 D#3838 USNM Ecuador: Orellana, Reserva Etnica Waorani, 17.x.2005, T.L. Erwin et al K. cuprea ♂ UCRCENT_320853 D#3852 CNC Ecuador: Sucumbíos, Napo River, Sacha Lodge, 4-14.v.1994, P. Hibbs K. cuprea ♂ UCRCENT_92227 D#1004 UCRC Ecuador: Orellana, Reserva Etnica Waorani, 9.vi.1995, T.L. Erwin et al. K. cuprea ♂ UCRCENT_247778 D#3833 USNM Ecuador: Orellana, Reserva Etnica Waorani 17.x.2005, T.L. Erwin et al K. iridicolor ♀ UCRCENT_282474 D#2917 UCRC Costa Rica: Heredia, La Selva Biol., 14.viii.2010, J. Heraty K. iridicolor ♂ UCRCENT_92077 D#0933b UCRC Honduras: Olancho, El Boquerón Nat. Mon., 2.vii.2002, D. Yanega 252

Table S2. Taxa, identification number, DNA voucher number information and the respective genes used for molecular analysis.

Genes Species Sex ID number DNA voucher number 18S 28S-D2 28S-D3-5 COI-nj COII Kapala iridicolor ♂ UCRCENT92077 D#0933b x x x x x Kapala iridicolor ♀ UCRCENT282474 D#2917 x x x x x Kapala furcata ♀ UCRENT92093 D#1078 x x x x x Kapala deltalis sp. n. ♂ UCRCENT92109 D#1388 x x x x Kapala quasimodo sp.n. ♂ UCRCENT92084 D#0938b x x x Kapala quasimodo sp.n. ♂ UCRCENT92002 D#0942 x x x x Kapala parafurcata sp.n. ♂ UCRCENT91921 D#1069 x x x x x Kapala parafurcata sp.n. ♂ UCRCENT161498 D#2519 x x x Kapala parafurcata sp.n. ♂ UCRCENT92073 D#1068a x x x x x Kapala parafurcata sp.n. ♂ UCRCENT397224 D#3436 x x x Kapala parafurcata sp.n. ♂ UCRCENT324 D#2520 x x Kapala parafurcata sp.n. ♀ UCRCENT91817 D#0711 x x x Kapala parafurcata sp.n. ♀ UCRCENT325 D#2518 x x Kapala parafurcata sp.n. ♀ UCRCENT91803 D#1086 x x x x Kapala cuprea ♂ UCRCENT412124 D#3806 x x x x Kapala cuprea ♂ UCRCENT412123 D#3805 x x x x Kapala cuprea ♀ UCRCENT412122 D#3804 x x x x Kapala cuprea ♂ UCRCENT412133 D#3815 x x x x Kapala cuprea ♀ UCRCENT91807 D#1003 x x x x x Kapala cuprea ♂ UCRCENT320768 D#3837 x x Kapala cuprea ♀ UCRCENT247775 D#3838 x x Kapala cuprea ♂ UCRCENT320853 D#3852 x x Kapala cuprea ♂ UCRCENT92227 D#1004 x x x x x Kapala cuprea ♂ UCRCENT247778 D#3833 x x

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Capítulo V

Schoeninger, K., Murray, E., Torréns, J., Heraty,

J.M., Oliveira, M.L. (2018) Check the count: Two new genera of Eucharitidae (Hymenoptera,

Chalcidoidea) for the Neotropical Region.

254

Check the count: Two new genera of Eucharitidae (Hymenoptera, Chalcidoidea) for the Neotropical Region

KARINE SCHOENINGER1,5, ELIZABETH MURRAY2, JAVIER TORRÉNS3, JOHN M. HERATY4 & MARCIO L. OLIVEIRA1

Abstract. Two new genera are described based on phylogenetic morphological and molecular analyzes: Ecarinata n. gen. and Torquata n. gen. According to phylogenetic analyzes, both genera belong to the Kapala clade. A diagnosis and a discussion about affinities with other genera are provided. The species Ecarinata sp.1 n. sp. and Torquata sp.1 n. sp. are described for the first time. Also, an updated identification key for Neotropical genera of Eucharitidae is provided.

Keywords. Ant parasitoids, Eucharitinae, identification key, taxonomy.

Introduction

Four subfamilies and 56 genera are recognized for Eucharitidae and they are distributed in almost every zoogeographical region of the World (Heraty, 2002; Torréns et al., 2016). Murray et., (2013) proposed that Eucharitidae originated in the Old World and colonized the New World through multiple dispersal events occurring approximately 20-43 mya (Murray et al., 2013). All members of Eucharitidae (Hymenoptera: Chalcidoidea) are specialized parasitoids of ants (Clausen, 1940; Heraty, 2002; Torréns et al., 2016). Females lay their eggs inside or on 255 plant tissues (leaf buds, ﬂoral buds, underside of leaf or into fruits), either individually or in masses (Clausen, 1940). Deposited remotely from the host, it is the active frst-instar larva (planidium) that is responsible for getting into the ant nest through various associations with foraging adult ants (Torréns, 2013). Once in contact with the larval ant host, the planidium either remains as an external parasite or partially burrows into the host (Heraty & Murray, 2013). After development on the host pupa, the adults must emerge and leave the nest (Clausen, 1923). Given the aggressive behavior of some ants, some specimens of Eucharitidae developed long projections after the mesoscutellum and these projections (frenal processes; Heraty, 2002) may protect the wings during specimen transport out of the nest and might account for their independent development in different Eucharitini (Eucharitinae) (Heraty et al., 2015). Within Eucharitini, a monophyletic group attacks ponerinae, ectatominae and myrmeciinae ants (PEM clade). The PEM clade is supported by molecular and some morphological analyses (Heraty, 2002; Murray et al., 2013) and is composed of three groups: the Old World Chalcura and Schizaspidia clades and the New World Kapala clade. Across the Kapala clade multiple bizarre morphological modifications have lead to the description of numerous genera and currently 13 genera are recognized for this clade (Murray, 2014). Head shape, antennal morphology, sculpture patterns and frenal processes morphology are particularly variable across the clade. The extreme phenotypes are best exemplified by the paired frenal processes; they range from those in Kapala which are slightly curved and linearly carinate, to forms in other genera that range from circularly striate to smooth (Lasiokapala and some Lirata), dorsoventrally flattened and narrowly separated (Dicoelothorax and Dilocantha), to broadly arched and forming a carapace over the metasoma (Galearia and Thoracantha). Here, two new genera are described based on phylogenetic morphological and molecular analyzes (Chap. II). According to their morphological characteristics and phylogenetic analyzes, both genera belong to the Kapala clade. A genus described here, Torquata n. gen., was previously mentioned in Murray's thesis (2014) and was part of her phylogenetic analyzes on the Kapala clade. Also, an updated identification key for Neotropical genera of Eucharitidae is provided.

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Materials and methods

Morphological descriptions

The morphological description follows Heraty (2002), and details on sculpture from Harris (1979) and Gibson et al. (1998). Morphological terms and abbreviations are from Heraty (2002), Gibson et al. (1998) and Hymenoptera Anatomy Ontology (HAO; Yoder et al. 2010). The funiculars include Fl2 and the following unfused flagellomeres before the clava, which is defined by 1–3 fused flagellomeres. The paired spines of the mesoscutellum arise from the frenum and are refered here as frenal process (character 50; Heraty 2002). Morphological abbreviations (see Figs 1 and 2, Chap. I): acl, anteclypeus; acr, acropleuron; ao, anterior ocellus; ax, axilla; axg, axillular groove; axl, axillula; cal, callus; clv, clava; cly, clypeus; cx, coxa; es2, mesepisternum;

Fl2, flagellomere basal; flg, flagellum; fmd, femoral depression; frl, frenal line, fp, frenal processes; fub, funicle; gen, gena; Gt1, first gastral tergite; iod, interocular distance; lep2, lower mesepimeron; llm, lateral lobe of mesoscutum; mlm, midlobe of mesoscutum; msp, malar space; mv, marginal vein; not, notaulus; pdl, pedicel; pet, petiole; pl3, metapleuron; pmv, postmarginal vein; po, posterior ocellus; ppd, propodeum; pre, prepectus; sca, supraclypeal area; scd, scrobal depression, scp; scape, sct, mesoscutellar disc; smv, submarginal vein; sss, scutoscutellar suclcus; stv, stigmal vein; tgl, tegula; tor, torulus; tsa, transscutal articulation; upe2, upper mesipimeron.

Measurements

All measurements and abbreviations follow Heraty & Woolley (1993), Heraty (2002) and Murray (2014). Measurements were made through a Zeiss Stemi SV6® with a 1.5X objective lens. The measures and ratios between structures are given as the value of the holotype [in brackets], followed by the range. All measurements are given in millimeters and follows (Schoeninger et al., in prep Cap. I).

Specimen imaging

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Specimens repository

The following institutions served as source of material and type depositories for specimens examined for this study: INPA, Coleção de Invertebrados do Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil; UCRC, University of California, Riverside, California, USA. Specimens are identified with unique UCRC specimen identifiers that indicate the museum of deposition. All specimens were databased in a FileMaker v.11® database maintained in the Heraty lab. Localities were georeferenced using Google Earth (coordinates in italics) or from the original label data.

Results

Revised key for the Neotropical genera of Eucharitidae (Modified from Heraty 2002 and Heraty 2006)

1 Prepectus separated from pronotum (Fig. 1A). First gastral sternite with transverse furrow delimiting small crescentic anterior region. Dorsal occipital usually rounded, rarely with carina… Orasema Cameron (Oraseminae) - Prepectus fused to pronotum (Figure 1.B), sometimes with crenulate sulcus along line of fusion. First gastral sternum evenly rounded without a transverse sulcus… 2 258

Figure 1. Mesosoma in profile; A. Prepectus separated from pronotum by a suture; B. Prepectus fused to pronotum

Eucharitini (Eucharitinae)

2(1) Scutellum rounded or projecting as a flattened flange (Fig. 2A) … 3 - Scutellum with prominent single or paired process (Fig. 2B) … 7

Figure 2. Mesosoma in dorsal view. A. Scutellum rounded or with a rounded flange; B. Scutellum with prominent process.

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3(2) Scutellum projecting as a flattened flange, slightly upturned flange that is about as long as broad (Fig. 3A) … Carletonia Heraty - Scutellum rounded (Fig. 3B) or slightly projecting as a sharp flange or rounded bidentate lobe; if a process is present, then frenal line distinctly above or below its lateral margins … 4

Figure 3. Mesosoma in dorsal view. A. Scutellum with a flattened flange; B. Scutellum rounded

4(3) Frenum rounded or with broad, emarginated flange (Fig. 4A) … 5 - Frenum with short bifurcating spines (Fig. 4B, frenum in grey) … 6

Figure 4. Mesosoma in profile. A. Frenum broad; B. Frenum with short bifurcating spines

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5(4) Prepectus reaching tegula (Fig. 5A). Mesoscutum smooth. Flagellar segments simple … Pseudometagea Ashmead - Prepectus not reaching tegula (Fig. 5B). Mesososma areolate. Flagellar segments simple, serrate or pectinate in females; always pectinate in males … Pseudochalcura Ashmead

Figure 5. Mesosoma in profile. A. Prepectus reaching tegula; B. Prepectus not reaching tegula

6(4) Flagellomeresof females simple (Fig 6B). Propodeal process blunt; callus and process with strong laminate carina, rarely rugose. Hind tibia with two spurs. Body yellow or orange with brown to black patterns … Obeza Heraty - Flagellomeres of females serrate or lobate (Fig. 6A); segments of male pectinate. Propodeal process forming strong ridge or long tapering spines; callus rugose. Hind tibia with one spur. Body usually black or with metallic luster, rarely patterned black and reddish-orange … Lophyrocera Cameron

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Figure 6. Antenna in profile. A. Funicular segments pectinate; B. Funicular segments simple.

7(2) Propodeal spiracle circular … 8 - Propodeal spiracle strongly incised with margins of incision raised … 10

8(7) Frenal process short and blunt (no more than 2.0‒3.0x as long as broad) (Fig. 7A). Females with flagellum 6 segmented, and its weakly serrate basally. The flagellum of the males has 8 segments … Colocharis Heraty - Frenal process long and usually extending to apex of metasoma (Fig. 7B). Females with flagellum 9 segmented, and its strongly serrate basally (Fig. 17B). The flagellum of the males has 10 segments … Ecarinata Schoeninger & Torréns n. gen.

Figure 7. Mesosoma in dorsal view. A. Frenal processes short and blunt; B. Frenal processes long.

10(7) Scutoescutellar sulcus filled with a dense pacth of long, hook tipped, golden hairs. Frenal processes of female broad and flattened or cylindrical; male processes thin and cylindrical … Dilocantha Ashemad 262

- Scutoescutellar bare or at most with a few scattered hairs or setae … 11

11(10) Frenal processes flattened (Fig. 8A), triangular or dome-shaped; processes forming a carapace over the metasoma and narrowly separated medially … 12 - Frenal processes cylindrical and broadly separated along their entire length (Fig. 8B) … 14

Figure 8. Mesosoma in dorsal view. A. Frenal processes flat; B. Frenal processes cylindrical.

12(11) Frenal processes of female dome-shaped and broadly rounded over metasoma (Fig. 9A), frenal process of male acute apically with a broad nearly contiguous base. Head closely fitted with mesosoma. Eye flat and flush with the margin of head … Gallearia Brullé - Frenal processes of both sexes flat or triangular (Fig. 9B). Head loosely articulating with mesosoma. Eye broadly rounded and slightly protruding … 13

Figure 9. Specimen in profile. A. Galearia; B Dicoelothorax.

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13(12) Eyes bare. Funicle of female with 5‒6 segments. Frenal processes flat (Fig. 10A), subtruncate and striate. Propodeum smooth, callus strongly produced posterodorsally as a sharp flange … Dicoelothorax Ashmead - Eyes with erect setae. Funicle of female with 8 segments. Frenal processes triangular, acute apically and longitudinally ribbed (Fig. 10B). Propodeum of female flat and margined by a strong carina shaped to receive metasoma, callus not apparent … Thoracantha Latreille

Figure 10. Mesosoma in dorsal view. A. Frenal processes dorsoventrally flat; B. Frenal processes triangular.

14(11) Eyes medially tuberculated (Fig. 11A). Frons raised and hemispherical lateral to scrobes and finely reticulate or granulate … Isomerala Shipp - Eyes broadly rounded (Fig. 11B). Frons flat, rarely slightly raised but then the frons is completely smooth … 16

Figure 11. Head in full-face view. A. Isomerala; B. Generalized form, several genera.

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15(14) Frenal processes cylindrical and smooth along most of their length (Fig. 12A). Funicle of female with 6‒7 segments, basal segments pectinate and apical 2‒3segments fused into a distinct clava … Lasiokapala Ashmead - Frenal processes variously sculptured, but not smooth (Fig. 12B). Funicle of females with 5‒6 segments, either simple or serrate, and without a distinct clava … 16

Figure 12. Mesosoma in dorsal view. A. Frenal processes cylindrical and smooth; B. Frenal processes not smooth.

16(15) Basal flagellomere in males more than 5x longer than broad, excluding branch. Scape long and reaching at least to the top of the median ocellus but usually exceeding vertex. Eyes with erect bristles (Fig. 13A) … 17 - Basal flagellomere of male transverse basally, never elongate. Scape not reaching top of median ocellus. Eyes with short setae or bare (Fig. 13B) … 18

Figure 13. Head in full-face view. A. Eyes with erect bristles; B. Eyes with short setae.

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17(16) Mesoscutellum broadly rounded without a distinct median depression. Transverse carina between median and lateral ocelli extending towards the inner margin of the eyes … Neolirata Torréns & Heraty - Mesoscutellum distinctly humped with a median longitudinal depression. Transverse carina between median and lateral ocelli absent … Lirata Cameron

18(16) Face smooth except for fine reticulation around median ocellus. Head, excluding eyes, and body with long, erect bristles. Mesoscutellum smooth and shining dorsally (Fig. 14A). Fore wing with sparse microtrichiae and no marginal fringe … Liratella Girault - Face even more sculptured, rarely smooth. Head and body with short erect or semi-appressed setae. Mesoscutellum longitudinally ribbed dorsally (Fig. 14B). Fore wing moderately to densely pilose, sometimes with minute marginal fringe … 19

Figure 14. Mesosoma in dorsal view. A. Scutellum smooth and shining; B. Scutellum longitudinally ribbed.

19(18) Head surface covered with dense, fine erect or decumbent setae. Females with two medial incisions over the posterior margin of the first gastral tergite (Fig. 15A, di: double medial incision) … Latina Koçak & Kemal - Head surface with sparse setae. Females with one medial incision over the posterior margin of the first gastral tergite (Figure 15B) … 20

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Figure 15. Metasoma in dorsal view. A. Two medial incisions over the posterior margin of the first gastral tergite; B. One medial incision over the posterior margin of the first gastral tergite.

20(19) Hind femur dark brown basally and yellow apically. Axillular groove deep and curved medially (Fig. 16A). Frenal processes thin and closely spaced. Females with 5 short serrate funicular segments. Male with flagellar branches short and thick … Parakapala Gemignani - Hind femur usually yellow, rarely dark brown basally. Axillular groove shallow and linear (Fig. 16B). Frenal processes long, robust and broadly spaced. Females usually with 7‒9 funicular segments. Males with flagellar branches long and thin … 21

Figure 16. Mesosoma in dorsal view. A. Axillular groove deep and curved medially; B. Axillular groove shallow and linear.

21(20) Maxillary and labial palpi 2/2 segmented (Fig. 18C). Frenal processes with helical twisted carinae in the entire length (Fig. 18D) … Torquata Schoeninger & Murray n. gen. - Maxillary and labial palpi 3/2 or 3/3 segmented. Frenal processes usually with longitudinal carina in the entire length … Kapala Cameron 267

Taxonomy

Ecarinata Schoeninger & Torréns, n. gen.

Type species. Ecarinata sp.1 Schoeninger & Torréns n. sp.

Etymology. The name comes from Latin “carina” linked with the letter "e" which signifies absent of carina, referring to the absence of carina bordering the propodeum.

Diagnosis. Recognized by absence of a carina delimiting the propodeum; a circular propodeal spiracle (also found in Colocharis); frons and lower face smooth; labrum with 5‒6 digits; maxillary palpi 2 segmented and long and labial palpi 2 segmented. Females with flagellum 9 segmented; scape narrow and elongate, reaching at most to ventral margin of median ocellus

(also found in Lirata and Neolirata); basal flagellomere (Fl2) long and serrated, Fl2 3.0x as long as apical width. The male has a relatively long flagellum, length of rami Fl2 1.6‒1.9 head height.

Ecarinata sp.1 Schoeninger & Torréns, n. sp. (Figs. 17A‒F)

Description. Female. Body length 2.0–3.1 mm; length of mesosoma excluding the frenal processes 1.1–1.2 mm. Head, mesosoma, coxae, petiole and frenal processes black; antennae and legs yellow; wings hyaline, venation light-brown; Gt1 with base of tergite brown and rest ligth- brown, subsequent tergites light-brown. Head. In frontal view subtriangular, 1.4‒1.6 as long as high. Eyes rounded to protruding and bare, separated by 2.6–2.7x their height. Median ocellus in line with lateral ocelli. Frons and lower face smooth and with densely scattered appressed hairs; scrobal depression shallow and broadly impressed. Occiput weakly striate and dorsal margin with prominent dorsal carina. Interocular area narrow and produced as narrow median ridge. Malar space 1.1–1.2x eye height with faint transversal striae. Supraclypeal area and clypeus smooth. Mandibles falcate with 3/2 formula. Labrum with 5‒6 digits. Maxillary palpi 2 segmented and long, labial palpi 2 segmented. Antenna 11 segmented; scape narrow and elongate, reaching at most to ventral 268 margin of median ocellus, scape 4.0‒4.5 longer than broad and 0.3x head height; flagellum 9 segmented (Fig. 17B), cylindrical and the first flagellomeres serrated, clava long and rounded; length of flagellum 1.4–01.5x head height; Fl2 3.0x as long as apical width and 1.5–1.8x as long as following flagellomere. Mesosoma. Pronotum abutting mesoscutum, no overlap of sclerites. Mesocutum anteriorly smoothly curved (Fig. 17A), in lateral view and 1.2–1.7x as high as broad; dorsum moderately setose; notauli deeply impressed and transversely carinate. Midlobe with transverse striae and dorsally rounded in frontal view; lateral lobes striate. Transcutal articulation (TSA) present. Mesoscutoscutellar sulcus (SSS) transverse, not meeting TSA. Axilla longitudinal stiate; lateral axillar lobe small. Mesoscutellar disc longitudinal striate, mesoscutellar disc 1.5–1.8x as long as axilla with medial apex raised 0.1x height of frenal process in lateral view. Axillula smooth to with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth; lower mesepimeron weakly reticulate; femoral depression crenulate. Callus carinate and weakly reticulate, covered by some semi-erect setae. Metapleuron carinate and slightly reticulate. Propodeal disc flat with a sculpture colliculate and without a bordering carina (Fig. 17C). Frenal processes cylindrical and short, 2.4‒2.7x as long as basally broad of mesoscutellum, 1.5–1.6x as long as length of axillae and mesoscutellar disc, straight in lateral and dorsal view; frenal processes longitudinal striate and apex rounded (Fig. 17E). Forewing 2.3–2.5x longer than broad; covered by small bristles, except for the base that is bare; submarginal vein 1.2–1.3x as long as marginal vein; stigmal vein stout and rectangular, 1.5–1.8x longer than broad, postmarginal vein absent. Hindwing fringed, 4 hamulli curved hook-shaped. Metacoxa semiglobose with a medial line of setae, 1.6‒1.7x longer than broad. Metafemur 6.0–6.4x longer than broad, weakly striate with sparse setae. Calcar robust curved and bifid, hind tibia with 2 spurs. Metassoma. Petiole cylindrical to subcylindrical, colliculate, petiole 4.7–5.0x longer than broad and 1.9–2.0x as long as metacoxa; base of petiole with small dorsal carina. Gastral terga smooth with a few punctuations, tergal scar absent.

Male. Body length 1.8–2.2 mm; length of mesosoma excluding the frenal processes 1.0– 1.1 mm (Fig. 17F). Similar to female except for the following: Frons striate near to scrobal 269 depression; lower face transversal striate (Fig. 17D). Antenna 12 segmented; scape narrow and elongate, reaching at most to ventral margin of median ocellus, scape 3.0‒3.5 longer than broad and 0.4x head height; flagellum 10 segmented, each flagellomere with a single long rami; length of rami Fl2 1.6‒1.9 head height, rami of Fl2 about as long as Fl3. Frenal processes 1.7–1.8x as long as length of axillae and mesoscutellar disc. Metacoxa 1.6‒2.0x longer than broad. Petiole 6.2‒6.5x longer than broad and 2.1–2.6x as long as metacoxal.

Biological notes. Host unknown.

Distribution. Argentina.

Material examined. Holotype: ARGENTINA: Tucumán Pr., Los Chorrillos, 26°18'40.4''S, 64°57'55.1''W, 22.i.2004, Trans. Yunga-chaco, J. Torréns & P. Fidalgo, [♀, deposited in UCRC]. Paratypes (1♀ 2♂): ARGENTINA: Tucumán Pr., ñorco, Fica Reyes, 13.i.2004, P. Fidalgo [1♀ 1♂, INPA]. Los Chorrillos, 26°18'40.4''S, 64°57'55.1''W, 22.i.2004, Trans. Yunga-chaco, J. Torréns & P. Fidalgo, [2♂, INPA].

Torquata Schoeninger & Murray n. gen.

Type species. Torquata sp.1 Schoeninger & Murray

Etymology. The name comes from Latin “torques” that means “twisted neck-chain”, referring to the helical twisted carinae in the frenal processes.

Diagnosis. Recognized by front and lower face smoothly; maxillary palpi long, 2 segmented and labial palpi short, 2 segmented; frenal processes with helical twisted carinae and apex rounded; mesocutum anteriorly smoothly curved, in lateral view; wings brown infuscate. The male has a relatively long flagellum, length of rami of Fl2 1.4‒1.5 head height.

Discussion. Torquata n. gen. is composedof a clade of Peruvian and Ecuadorian specimens previously identified as Kapala. There are additional (non-molecular) specimens from Bolivia, 270

Brazil, Paraguay and Venezuela. This genus was determined to be a new genus in the Kapala clade based on having a smooth face coupled with males with flabellate antennae and 2/2 maxillary and labial palp formula, with the latter two features unique in the Kapala clade. Morphologically, these appear similar to species in the Kapala iridicolor complex which also have a smooth face and low mesoscutum profile, but in analyses molecular and morphological hey are distinct.

Torquata sp.1 Schoeninger & Murray, sp. n. (Figs. 18A‒D)

Description. Male known and female unknown. Body length 2.7–3.0 mm; length of mesosoma excluding the frenal process 1.3–1.6 mm. Head, mesosoma, coxae, petiole and frenal process black; antennae dark-brown; legs yellow; wings brown infuscate; Gt1 dark-brown and subsequent tergites light-brown. Head. In frontal view subtriangular, 1.5‒1.6 as long as high. Eyes rounded to protruding and bare, separated by 1.9–2.1x their height. Median ocellus in line with lateral ocelli. Frons and lower face smooth and shining with a few scattered appressed hairs (Fig. 18B); scrobal depression shallow and broadly impressed. Occiput weakly striate and dorsal margin with prominent dorsal carina. Interocular area narrow and produced as narrow median ridge. Malar space 0.7–1.0x eye height, smooth. Supraclypeal area and clypeus smooth. Maxillary palpi 2 segmented and long, labial palpi 2 segmented (Fig. 18C). Antenna of male 12 segmented; scape narrow and elongate, reaching at most to ventral margin of median ocellus, scape and 2.0‒2.5 longer than broad and 0.3x head height; flagellum 10 segmented, each flagellomere with a single long rami; length of rami Fl2 1.4‒1.5x head height, rami of Fl2 about as long as Fl3. Mesosoma. Pronotum abutting mesoscutum, no overlap of sclerites. Mesocutum anteriorly smoothly curved, in lateral view and 1.2–1.3x as high as broad; dorsum moderately setose; notauli deeply impressed and transversely carinate. Midlobe with transverse striae and dorsally rounded in frontal view; lateral lobes weakly striate. Transcutal articulation (TSA) present. Scutoscutellar sulcus (SSS) transverse, not meeting TSA. Axilla longitudinal stiate; lateral axillar lobe small. Scutellar dis longitudinal striate, scutellar disc 1.3–1.7x as long as axilla 271 with medial apex raised 1.0–1.3x height of frenal process in lateral view. Axillula smooth to with faint longitudinal striae. Prepectus triangular with elongated posterior region and rounded apex. Mesepisternum and upper mesepimeron smooth (Fig. 18C); lower mesepimeron weakly reticulate; femoral depression weakly crenulate. Callus carinate and weakly reticulate, covered by semi-erect setae. Metapleuron carinate and slightly reticulate. Propodeal disc flat with a weakly sculpture colliculate and a single bordering carina, that is most distinct laterally. Frenal process cylindrical, 2.5‒3.1x as long as basally broad of scutellum, 1.5–1.7x as long as length of axillae and scutellar disc, straight in lateral view and convex in dorsal view; frenal process oblique striate and with apex rounded (Fig. 18D). Forewing 2.2–2.4x longer than broad; covered by small bristles, except for the base that is bare; submarginal vein 1.0–1.1x as long as marginal vein; stigmal vein oblique 1.2–1.7x longer than broad, postmarginal vein indistinct. Hindwing fringed, 4 hamulli curved hook-shaped. Metacoxa semiglobose with sparse setae, 1.7‒2.0x longer than broad. Metafemur 6.0–8.0x longer than broad, weakly striate with sparse setae. Calcar robust curved and bifid, hind tibia with 2 spurs. Metassoma. Petiole cylindrical to subcylindrical, dorsal reticulate and lateral with some fine striae, petiole 4.0–5.0x longer than broad and 2.1–3.6x as long as metacoxa; base od petiole with small dorsal carina. Gastral terga smooth with a few punctuations, tergal scar absent.

Biological notes. Host unknown.

Distribution. Ecuador and Peru.

Material examined. Holotype: PERU: Madre de Dios, Los Amigos Bio. St., 255m, 12°33'31.6''S, 70°05'35''W, 26.xii.2010, swp trail 10&28 H10-165, J. Heraty, [1♂, deposited in UCRC: UCRCENT00397284]. Paratypes (3♂): ECUADOR: Napo, Reserva Etinica Woarani, 1Km S, Onkone Gare Camp, 220m, 7.ii.1999, T.L. Erwin, et al., [1♂, UCRC: 92192]. PERU: Madre de Dios¸ Los Amigos Bio. St., 231m, 12°34'17.1''S, 70°05'43.4''W, 22.xii.2010, trail 13, 15, 14, 19, H10-152, J. Heraty, [1♂, UCRC: UCRCENT00397251]. Madre de Dios, Tamboapata Research Center, 13.14S, 69.61W, 16-22.vii.2010, MT, B. Bown, G. Kung, [1♂, UCRC: UCRCENT00235922]. 272

Acknowledgment

References

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Murray, E.A. (2014) Systematics and evolution of Eucharitidae (Hymenoptera: Chalcidoidea), with a focus on the New World Kapala. University of California, Riverside, CA. Murray, E.A., Carmichael, A.E., & Heraty, J.M. (2013) Ancient host shifts followed by host conservatism in a group of ant parasitoids. Proceedings of the Royal Society of London B: Biological Sciences, 280 (1759), 20130495. Torréns, J. (2013) A review of the biology of Eucharitidae (Hymenoptera: Chalcidoidea) from Argentina. Psyche: A Journal of Entomology, 2013: 1-14. Torréns, J., Heraty, J.M., Murray, E. & Fidalgo, P. (2016) Biology and phylogenetic placement of a new species of Lasiokapala Ashmead from Argentina (Hymenoptera: Eucharitidae). Systematic Entomology, 41: 596‒606. Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE 5(12): e15991. http://dx.doi.org/10.1371/journal.pone.0015991

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Figure 17. Ecarinata sp.1 n. sp. A. Habitus (lateral, ♀); B. anetnna (lateral, ♀; Fl2 indicating); C. propodeum (carina absent); D. head (front, ♂); E. habitus (dorsal, ♀); F. habitus (lateral, ♂).

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Figure 18. Torquata sp.1 n. sp. A. habitus (lateral view); B. head (front); C. Maxillary and labial palpi (lateral view); D. Mesososma (dorsal view)

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SÍNTESE

Kapala, de acordo com analises filogenéticas morfológicas e moleculares, é parafilético, o qual é dividido em três complexos de espécies, sendo eles: complexo de espécies K. furcata, complexo K. iridicolor e complexo K. romandii. Os dois primeiros complexos foram recuperados como monofilético com alto suporte de bootstrap. O complexo K. romandii, foi recuperado como monofilético somente na análise morfológica, já nas análises moleculares este não corresponde a um agrupamento natural. Apesar dessa problemática, no presente trabalho optamos por manter o o complexo romandii como um agrupamento artificial devido ao compartilhamento de características plesiomórficas pelas espécies que o integram, sendo de grande valia para o entendimento da diversidade deste gênero. Fica evidente que o gênero Kapala precisará ser dividido, mas, devido a ausência ou baixo suporte de ramo na espinha dorsal da árvore (a qual envolve as espécies de Kapala), a circunscrição do gênero ainda deve ser sujeita a análises adicionais antes de se implementar grandes mudanças taxonômicas. Como por exemplo, amostragem molecular adicional de espécimes provenientes do Brasil; adição de caracteres morfológicos internos (musculares e padrão de carenas internas); adição de informações referentes a comportamentos e possíveis hospedeiros, entre outros. O número de espécies conhecidas de Kapala teve um incremento de 14 para 29 espécies, já incluindo as mudanças taxonômicas. Este número pode estar subestimado uma vez que para a realização deste estudo foram examinados apenas espécimes depositados nas coleções, as quais não contemplam todas as áreas geográficas da região Neotropical. É possível que com a realização de coletas em áreas ainda não amostradas, novas espécies venham a ser encontradas.

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