D E A F G Oxysarcodexia Townsend, 1917 (D...

CARINA MARA DE SOUZA

DIVERSIDADE DE ESPÉCIES E ABORDAGEM FILOGENÉTICA DO GÊNERO

Oxysarcodexia Townsend, 1917 (DIPTERA: SARCOPHAGIDAE)

CAMPINAS

2014

iii

UNIVERSIDADE ESTADUAL DE CAMPINAS

INSTITUTO DE BIOLOGIA

CARINA MARA DE SOUZA

DIVERSIDADE DE ESPÉCIES E ABORDAGEM FILOGENÉTICA DO GÊNERO

Oxysarcodexia Townsend, 1917 (DIPTERA: SARCOPHAGIDAE)

Orientadora: Profa. Dra. Patrícia Jacqueline Thyssen

CAMPINAS

2014

vii

Aos meus pais, razão de tudo.

"O que sabemos é uma gota, o que ignoramos é um oceano"

Isaac Newton

ix

AGRADECIMENTOS

À minha família (especialmente papai Ariozano (in memorian), mamãe Maria José, e irmãos

Cassiano e Carolina) por compartilharem comigo esta etapa, por todo o apoio moral, amoroso e financeiro

durante este período e em todas as minhas decisões. Meu pai, apesar de não estar mais presente neste final,

foi de essencial à execução deste trabalho, além de um “ajudante” muito especial durante parte das coletas.

Sem o apoio e incentivo da minha mãe eu não teria o alicerce e força necessários para concluir essa

caminhada. Obrigada por serem meus exemplos, acreditarem em mim, pela compreensão nos momentos

de ausência (que foram muitos...), pelos conselhos e por todas as palavras motivadoras que me consolaram

e deram força nos momentos mais difíceis.

À inestimável Dra. Patricia J. Thyssen, grande amiga e mestre, pela orientação, oportunidade de

crescimento profissional e pessoal, apoio incondicional em todas as empreitadas dessa jornada, um “anjo-

da-guarda” na minha vida, tanto acadêmica quanto pessoal.

Ao Dr. Arício X. Linhares pelo apoio na execução desse projeto, pela convivência e singular

oportunidade de aprendizado, além das contribuições durante a qualificação, exame prévio e defesa.

Ao Dr. Thomas Pape pela oportunidade de estágio em seu laboratório e aprendizado no mundo

dos sarcofagídeos.

À Dra. Roseli Tuan pelos ensinamentos na área de biologia molecular.

À Dra. Cátia A. Mello-Patiu pela imensurável contribuição durante o estudo das Oxysarcodexia, a

afetuosa receptividade nas várias trocas de e-mails, em minha visita ao Museu Nacional e nos encontros

acadêmicos, além das relevantes contribuições na avaliação da tese.

Aos Drs. Marlene T. Ueta, Silmara Allegretti, Júlio Mendes, Carolina Reigada, Cláudio J. Von

Zuben, Paulo R. Bunde, Sérgio F. dos Reis e José Roberto Pujol-Luz pela disponibilidade em compor as

bancas dos exames de qualificação, prévio e defesa e por suas contribuições nestas etapas.

Aos amigos do Laboratório de Entomologia L2B, André Savino, Carolina Palanch, Cauê Mira,

Daniel Brancoli (Carioca), Danilo Ferraz, Fábio Rezende (Frango), Jandui Amorim, Maicon Grella

(Maicola), Marina Klemm (Nina), Marcela Alonso (Qualy), Mariana Nassu (Dori), Maria Lígia Paseto

(Armaria), Rafael Cedro, Thamiris Smania pelo ótimo ambiente de trabalho, pelas discussões científicas,

momentos de descontração e imensurável ajuda em coletas, criações de moscas, edição de imagens, dicas

na redação, dentre outras atividades, sem os quais parte do trabalho não seria possível. Prezadas amizades.

Aos queridos amigos feitos em Copenhagen, Eliana Buenaventura, Dave Cheung, Daniel

Whitmore, Ken Puliafico e Nesrine Akkari pelas aventuras no exterior e o apoio numa fase difícil.

Aos estimados colegas Karine Vairo, pelo material de Manaus; Matheus Camargo, pelo auxílio na

edição das imagens; Taís Madeira e Thamiris Barbosa, pela colaboração nos experimentos moleculares.

Aos prezados amigos da Pós-Graduação em Parasitologia e do departamento, em especial Luciana

Franceschi, Aline Rimoldi, Laura Gisloti, Rodrigo Labello e Raquel Palasio.

Aos amigos distantes, porém sempre presentes, que conviveram com minha ausência em vários

momentos e nunca se afastaram ou deixaram de me apoiar.

Aos demais professores e técnicos do departamento, em especial Tacilda S. Nalon (Tata), João

Batista e Letícia Duart, pelas contribuições durante as disciplinas e na realização deste trabalho.

À Universidade Estadual de Campinas por prover condições para minha formação acadêmico-

profissional e pelos subsídios fornecidos a esta pesquisa.

À CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) pela concessão da

bolsa de estudos (doutorado regular e programa sanduíche no exterior processo #0906/12-3).

xi

RESUMO

Um conspecto taxonômico com base nos machos de 85 espécies válidas pertencentes ao

gênero Oxysarcodexia Townsend, 1919 (Diptera: Sarcophagidae) é apresentado. Uma diagnose

do gênero e de cada espécie constituinte, distribuição geográfica, dados biológicos – quando

disponíveis –, e um apêndice pictórico são também inclusos nesse conspecto. Além do exame de

espécimes de Oxysarcodexia, uma ampla e minuciosa busca na literatura foi realizada para

abranger toda a informação disponível para esse gênero. Além disso, foram descritas seis

espécies novas de distribuição neotropical. Foi também realizada uma análise filogenética

baseada em caracteres morfológicos provenientes do exame de machos de 54 espécies

pertencentes ao gênero Oxysarcodexia a fim de investigar a assimetria fálica vista em algumas

espécies e a ocorrência de possíveis agrupamentos interespecíficos dentro do gênero. A

assimetria também foi estudada por meio de espécies modelo, com o auxílio da técnica de

microscopia eletrônica de varredura rotacional. A assimetria detectada foi do tipo direcional

sinistral, presente em oito espécies, restringindo-se aos lobos terminais da vésica e sendo

considerada uma homoplasia. Quanto aos agrupamentos de espécies, a topologia encontrada nas

árvores mais parcimoniosas obtidas mostrou certa similaridade com o proposto na literatura,

embora sem completa correspondência. Espera-se, com esse estudo, auxiliar no processo de

identificação dessas espécies, assim como contribuir para o melhor conhecimento desse gênero

de moscas.

Palavras-chave: Sarcofagídeo; Díptero – Morfologia; Inseto – Identificação; Microscopia

eletrônica de varredura.

xiii

ABSTRACT

A taxonomic conspectus based on male specimens of 85 valid species of the genus

Oxysarcodexia Townsend, 1919 (Diptera: Sarcophagidae) is presented. A diagnosis for the genus

and for each species, the geographical distribution, the biological information – when available –,

and a pictorial appendix are included in this conspectus. Besides the species examination, a

thorough and wide search on the literature was performed in order to gather all information

available for this genus. Additionally, six new species of Neotropical distribution were described.

It was also performed a phylogenetic analysis based on males morphological characters of 54

species belonging to Oxysarcodexia genus, in order to study the phallic asymmetry present on

some species and the occurrence of possible interspecific grouping within the genus. The

asymmetry was also investigated through model species, with the help of rotational scanning

electron microscopy technique. The asymmetry detected was of sinistral directed type, present in

eight species, restricted to the terminal lobes of the vesica, and considered a homoplasy. About

the species grouping, the topology found on the most parsimonious trees obtained showed certain

similarity with the propositions of the literature, although without complete correspondence. We

expected with this study contribute with the species identification process and with the best

knowledge of this flesh fly genus.

Key-words: Sarcophagidae; Diptera – Morphology; Insects – Identification; Scanning electron

microscopy.

xiv

SUMÁRIO

1. INTRODUÇÃO ........................................................................................................................... 1

2. REVISÃO BIBLIOGRÁFICA ....................................................................................................... 3

3. OBJETIVOS .............................................................................................................................. 9

4. METODOLOGIA ...................................................................................................................... 11

5. CAPÍTULO I ........................................................................................................................... 13

Abstract ...................................................................................................................................... 13

Resumo ...................................................................................................................................... 14

Introduction ............................................................................................................................... 15

Material and Methods ................................................................................................................ 16

Generic Definition ..................................................................................................................... 18

Remarks ..................................................................................................................................... 19

Genus Diagnosis ........................................................................................................................ 20

Species awaiting status confirmation ........................................................................................ 23

Taxonomy .................................................................................................................................. 23

Final considerations ................................................................................................................. 132

References ............................................................................................................................... 132

Appendices .............................................................................................................................. 147

6. CAPÍTULO II ........................................................................................................................ 179

RESUMO ................................................................................................................................... 179

INTRODUÇÃO ............................................................................................................................ 180

MATERIAL E MÉTODOS ............................................................................................................. 182

RESULTADOS ............................................................................................................................ 183

DISCUSSÃO ............................................................................................................................... 185

REFERÊNCIAS BIBLIOGRÁFICAS ................................................................................................ 190

APÊNDICE ................................................................................................................................. 204

7. CONSIDERAÇÕES FINAIS ..................................................................................................... 217

8. REFERÊNCIAS BIBLIOGRÁFICAS ......................................................................................... 219

1

1. INTRODUÇÃO

O gênero Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae) está distribuído

majoritariamente na região Neotropical, sendo frequentemente relatada a associação destas

moscas a substratos em decomposição, como fezes e carcaças, despertando grande interesse

em diversas áreas como a forense, médica e veterinária. Apesar disso, não é incomum a

identificação das espécies ficar restrita ao nível genérico ou ser errônea, devido à existência de

espécies com caracteres morfológicos muito similares e facilmente confundíveis quando

inexiste familiaridade com as diferentes espécies.

Nesse contexto foi elaborado o primeiro capítulo desta tese, apresentando um conspecto

taxonômico para machos de Oxysarcodexia. Os dados foram formatados de modo a

possibilitar o fácil reconhecimento das diferentes espécies a partir do oferecimento de uma

diagnose do gênero, listagem das espécies válidas, sinonimização de nomes, origem de

materiais tipo e seus depositários, diagnose de cada espécie, discussões sobre caracteres

morfológicos similares para diferentes espécies, distribuição geográfica, dados biológicos e

um banco de dados pictórico a fim de facilitar o reconhecimento das diferentes espécies. Este

capítulo contemplou também a descrição de seis espécies novas a partir do exame de material

depositado em museus (Oxysarcodexia n. sp. 1, coletada na Colômbia e Equador;

Oxysarcodexia n. sp. 2, coletada na Guiana Francesa; Oxysarcodexia n. sp. 3, coletada na

Costa Rica; Oxysarcodexia n. sp. 4, coletada no Brasil; Oxysarcodexia n. sp. 5, e

Oxysarcodexia n. sp. 6, ambas coletadas no Equador).

A partir da análise morfológica dos machos, especialmente da terminália, observou-se a

ocorrência de assimetria na genitália. Por ser uma condição não usualmente observada para os

insetos, que, por via de regra, apresentam simetria bilateral, a investigação desta condição

motivou a estruturação do segundo capítulo. Foi proposta uma análise filogenética baseada em

uma matriz de caracteres morfológicos compilados a partir do estudo de machos de diferentes

espécies, a fim de rastrear a ocorrência da assimetria no gênero Oxysarcodexia. Além disso,

micrografias rotacionais foram produzidas a partir de imagens de microscopia eletrônica de

varredura para Oxysarcodexia fringidea (Curran & Walley, 1934), Oxysarcodexia timida

(Aldrich, 1916) e Oxysarcodexia varia (Walker, 1836), utilizadas como modelos para a

análise das alterações morfológicas consequentes da assimetria.

2

Espera-se que a investigação da diversidade e a abordagem filogenética para as espécies

de Oxysarcodexia contribuam para o melhor entendimento deste grupo e de suas relações

interespecíficas, representando uma motivação para a realização de outros estudos.

3

2. REVISÃO BIBLIOGRÁFICA

A ordem Diptera (Insecta: Arthropoda) compreende cerca de 160.590 espécies,

incluindo aproximadamente 3.800 espécies fósseis, consideradas subestimadas (Zhang, 2013).

Na região Neotropical há a ocorrência de mais de 31.000 espécies (Evenhuis et al., 2007 apud

Amorim, 2009). Estima-se, no entanto, que esse número seja bem maior (Brown et al., 2009),

até cinco vezes mais para algumas famílias (Amorim, 2009), devido a áreas ainda não

amostradas, espécies coletadas que ainda aguardam descrição e ao número restrito de

taxonomistas estudando determinadas espécies (Rafael et al., 2009).

A família Sarcophagidae (Diptera: Muscomorpha) possui aproximadamente 355 gêneros

e em torno de 3.100 espécies descritas mundialmente, sendo que cerca de 870 espécies

ocorrem na região Neotropical e 270 apenas no Brasil (Evenhuis et al. 2007 apud Pape et al.

2009; Carvalho et al. 2012). É reconhecidamente dividida em três subfamílias,

Miltogramminae, Paramacronychiinae e Sarcophaginae, com uma possível relação de grupo-

irmãos entre as duas últimas e monofiletismo reconhecido filogeneticamente para

Sarcophaginae (Pape, 1996).

A subfamília Sarcophaginae engloba moscas com tamanho variável, embora a coloração

externa seja razoavelmente homogênea, consistindo em um tórax acinzentado com três faixas

pretas e abdômem com padrão enxadrezado. A distribuição e diversidade de espécies de

sarcofagídeos são, de maneira geral, diretamente proporcionais a regiões de clima quente,

sendo, consequentemente, bastante reduzidas em regiões subárticas, assim como em ambientes

insulares e, como esperado, inexistentes nas regiões árticas (Shewell, 1987; Pape, 1996). Os

hábitos das espécies inclusas nessa subfamília são amplamente diversificados, variando entre a

coprofagia, a saprofagia, a predação e o parasitismo (Shewell, 1987; Pape, 1996; Pape &

Dahlem, 2010; Carvalho et al., 2012).

Roback (1954), ao estudar a evolução e taxonomia de Sarcophaginae com base na

similaridade morfológica de caracteres externos do corpo, especialmente os da terminália dos

machos, propôs a divisão desta subfamília em duas tribos, Agriini e Sarcophagini, e diversas

subtribos. Partindo deste estudo, Lopes (1969) propôs uma nova divisão que considera a

nomeação de seis tribos: Microcerellini, Notochaetini, Raviniini, Sarcophagini, Sarcophagulini

e Tephromyiini. Ao considerar o desenvolvimento da mandíbula e do arco clipeal das larvas

de primeiro estádio, associado à análise morfológica de machos e fêmeas, especialmente de

4

caracteres da genitália, Lopes (1982) acrescentou as tribos Cuculomyiini, Impariini e

Sarcodexiini (por revisão de status) e Sarothromyiopsini (por nova proposição), resultando em

11 tribos inclusas na subfamília Sarcophaginae. Essa divisão foi corroborada por Verves

(1989), que revisou as subtribos e apresentou uma chave para a identificação das tribos

Cuculomyiini, Emblemasomatini, Impariini, Johnsoniini, Microcerellini, Protodexiini,

Raviniini, Sarcodexiini, Sarcophagini, Sarothromyiini e Sarothromyiopsini.

O gênero Oxysarcodexia Townsend, 1917 é um dos gêneros de Sarcophaginae que

apresenta maior riqueza de espécies na região Neotropical (Lopes, 1969; Verves, 1989; Pape,

1996). Juntamente com Nephochaetopteryx Townsend, 1934, Orosarcophaga Townsend,

1927 [naquele momento ainda considerado um gênero e, posteriormente, reclassificado como

subgênero de Lepdidodexia Brauer & Bergenstamm, 1891 (Pape, 1996)], Oxyvinia Dodge,

1966 e Ravinia Robineau-Desvoidy, 1863, acomodado na tribo Raviniini.

A presença de uma projeção triangular (em forma de “dente”) do tubo fálico, logo acima

da vésica, é considerada a única apomorfia que caracteriza o monofiletismo de Oxysarcodexia

(Giroux et al., 2010). As espécies inclusas nesse gênero são consideradas, de maneira geral,

coprófagas, mas podem também ser encontradas associadas a carcaças (Pape & Dahlem 2010;

Carvalho et al. 2012).

Os dípteros muscóideos, de maneira geral, inclusos os sarcofagídeos, têm acompanhado

a população humana ao longo da história, estabelecendo durante muito tempo associações

harmônicas e desarmônicas. Essas relações eventualmente repercutem em problemas tanto

para a área da saúde quanto para a econômica, por exemplo, quando ocasionam e/ou veiculam

doenças e óbito ao atingirem o homem ou seus animais domésticos (Greenberg, 1973).

As relações que os dípteros possuem com o ambiente humano podem ser classificadas

como sinantropia, um fenômeno ecológico diretamente ligado à biocenose humana, a qual é

influenciada pelas ações antrópicas sobre o ambiente e refletida pelas condições de

antropobiocenoses (habitações humanas e seus animais domésticos) e agrobiocenoses

(pastagens, clareiras florestais, monoculturas e policulturas) (Greenberg, 1971; Ferreira,

1978). Nessas condições, na procura por disponibilidade de recursos alimentares ou sítios para

deposição de seus ovos ou larvas, os dípteros podem encontrar-se associados ao lixo ou às

fezes, tornando-se potenciais carreadores e propagadores de patógenos como vírus, bactérias,

protozoários e helmintos (Greenberg, 1971; 1973; Thyssen et al., 2004).

5

Outras espécies acarretam danos diretos à saúde, quando são agentes causadores de

miíases, popularmente conhecidas como bicheiras (Linhares & Thyssen, 2007). Esta doença é

caracterizada pela infestação, em geral do tecido cutâneo e subcutâneo de vertebrados, por

estágios imaturos de moscas que se alimentam por um curto período de tecidos vivos ou

necrosados de seus hospedeiros (Zumpt, 1965). Apesar das varejeiras estarem mais

comumente envolvidas nesses quadros, dípteros sarcofagídeos também podem ocasionar

miíase humana. Um caso de miíase uretral causada por larvas de Peckia (Sarcodexia) lambens

(Wiedemann, 1830) foi relatado em Belém, Pará (Leão et al. 1996) (Diptera: Sarcophagidae).

Em São João de Meriti, Rio de Janeiro, um caso de infestação auricular, cujas envolvidas

pertenciam à espécie Oxysarcodexia amorosa (Schiner, 1868) (Diptera: Sarcophagidae), foi

reportado por Figueiredo e colaboradores (2002). Casos de miíases humanas (orotraqueal,

auricular e furuncular na região da axila, por exemplo) causados por larvas de Wohlfahrtia

magnifica (Diptera: Sarcophagidae) tem sido relatados com certa frequência em países

europeus (por exemplo, Çiftçioğlu, Altintaș & Haberal 1997; Tuygun et al. 2009; Yazgi et al.

2009). Além disso, as espécies de sarcofagídeos Sarcophaga (Bercaea) haemorrhoidalis,

Sarcophaga (Bercaea) cruentata (Meigen, 1826) e Peckia (Peckia) chrysostoma (Wiedemann,

1830) também já foram associadas a casos de míiases humanas (por exemplo, Guimarães &

Papavero 1999; Turhan et al. 2007; Dutto & Bertero 2010).

Os tecidos de animais mortos podem ser atrativos para uma grande variedade de insetos

sarcossaprófagos, sobretudo dípteros, pelo fato da matéria orgânica constituir uma fonte

alimentar com alto valor proteico para o desenvolvimento de imaturos e nutrição dos adultos

(Nuorteva, 1977; Smith, 1986). Desse modo, a partir dos trabalhos realizados na segunda

metade do século XIX por Mégnin (1894), levantando informações biológicas e ecológicas

para demonstrar o potencial sobre a contribuição dos insetos em investigações de caráter

médico-legal, foi criada a Entomologia Forense. Nessa vertente, os dípteros ganham maior

destaque por serem os primeiros a chegar a uma cena de crime, já que possuem órgãos

sensitivos altamente especializados para a detecção de odores e podem ovipor ou larvipor em,

aproximadamente, até 10 minutos após a morte (Campobasso et al., 2001).

Alguns dos inventários da dipterofauna de determinados ambientes e de trabalhos sobre

os dípteros encontrados visitando ou criando-se em carcaças e cadáveres em decomposição

resultaram em listagens nas quais a identificação taxonômica em nível de espécie dos

6

sarcofagídeos, incluindo aqueles pertencentes ao gênero Oxysarcodexia, não foi alcançada em

sua maioria. Nesses casos, os dados registrados foram referentes apenas à abundância e

frequência de “morfotipos” em ambientes urbanos e silvestres (Carrera 1944; Cornaby 1974;

Dias et al., 1984; Monteiro-Filho & Penereiro, 1987; Souza & Linhares, 1997; Carvalho et al.,

2000; Couri et al., 2000; Pamplona et al., 2000; Carvalho & Linhares, 2001; Carvalho et al.,

2004; Perez et al., 2005; Costamagna et al., 2007; Cruz 2008; Amat 2010; Battán-Horenstein

et al., 2010; Beuter et al., 2012; Sousa et al., 2011; Battán-Horenstein et al., 2012; Ramírez-

Mora et al., 2012).

O produto da morfotipagem, isto é, a criação de categorias abstratas como „morfotipos‟,

as quais não apresentam especificidade necessária para diferenciar um ser de outro, prejudica

o avanço de quaisquer outros trabalhos que investiguem a biologia, ecologia ou dinâmica dos

mais diversos organismos na natureza. Isto explicaria porque, com a grande diversidade de

espécies que há no Brasil, um conhecimento mais aprofundado relativo à bionomia,

identificação e classificação dos dípteros de importância médica, veterinária e forense ainda é

incipiente e a realização de tais estudos é de grande relevância para a formação de bancos de

dados.

Contudo, vale ressaltar que a identificação de diferentes espécies nem sempre é fácil e

corriqueira. A diversidade e as minúsculas diferenças morfológicas observadas entre as várias

espécies, além da ausência de chaves taxonômicas para certos grupos e a insuficiência na

descrição dos caracteres morfológicos em algumas já existentes, aliados ao pequeno número

de especialistas/taxonomistas existente para determinados grupos são alguns dos fatores que

contribuem para a complexidade do processo de identificação (Liu & Greenberg, 1989; Rafael

et al., 2009).

A sistemática filogenética surge, então, numa tentativa de dimensionar a diversidade por

meio da descrição da variedade e da busca por um padrão de ordem nos processos envolvidos

na diversificação dos organismos, visando à ordenação por meio de um sistema hierárquico

(Amorim, 2002). A região Neotropical, por exemplo, abrange uma complexidade florística e

faunística enorme, tornando a reconstrução dos padrões envolvidos no processo contínuo de

mudanças que geram a diversidade das espécies uma tarefa difícil (Amorim, 2012). Isso

mostra, em suma, a relevância do conhecimento mais detalhado das espécies no que diz

respeito aos seus caracteres morfológicos, biológicos ou ecológicos, assim como de variações

7

intraespecíficas e das relações interespecíficas com as demais espécies para várias áreas do

conhecimento, como a entomologia médica, veterinária e forense, a biogeografia, a evolução,

entre outras.

9

3. OBJETIVOS

1. Realizar um conspecto taxonômico sobre o gênero Oxysarcodexia Townsend, 1917

(Diptera: Sarcophagidae) incluindo informações inerentes a cada espécie, como nome válido e

sinonímias, localidade onde o material tipo foi coletado e instituição depositária, diagnose para

identificação dos machos, distribuição geográfica e informações biológicas, quando

disponíveis.

2. Produzir um banco de dados pictórico para machos de espécies de Oxysarcodexia.

3. Estudar a ocorrência de assimetria na genitália de alguns machos pertencentes a este

gênero.

4. Propor uma hipótese de relação filogenética para as espécies de Oxysarcodexia com

base na morfologia externa dos machos.

11

4. METODOLOGIA

Para a elaboração de um conspecto taxonômico e uma matriz de caracteres morfológicos

foram examinados espécimes machos pertencentes às espécies do gênero Oxysarcodexia.

Essas moscas foram provenientes de material de coleção científica depositado nas seguintes

instituições:

CE-TdeA – Centro Tecnológico de Antioquia, Instituição Universitária (Medellín,

Colômbia);

L2B-DBA – Coleção de referência do Laboratório de Entomologia do Departamento de

Biologia Animal, Universidade Estadual de Campinas (Campinas, Brasil)

MNRJ – Museu Nacional/Universidade Federal do Rio de Janeiro (Rio de Janeiro, Brasil);

NRM – Museu de História Natural Sueco (Estocolmo, Suécia);

ZMUC – Museu de Zoologia de Copenhagen (Copenhagen, Dinamarca).

O registro pictórico da terminália dessas espécies, em vista lateral, posterior e anterior

(este último, sempre que possível), assim como do hábito em vista lateral, foi realizado com o

auxílio de uma câmera digital Leica DELUX 3®

(10 megapixels) acoplada a um

estereomicroscópio Leica S8AP0®

e de uma câmera digital Carl Zeiss AXIOCAM MRc®

(5

megapixels) acoplada a um esteromicroscópio Carl Zeiss STEREO DISCOVERY.V12®

. Cada

fotografia foi produzida após a aquisição de imagens com foco estendido, agrupadas com o

auxílio do programa Zerene Stacker®

. Para as espécies sem material disponível para exame, a

melhor ilustração disponível na literatura foi apresentada, com edições sempre que necessário.

Além da informação pictórica acerca de cada espécie, uma minuciosa revisão da

literatura publicada sobre o gênero Oxysarcodexia foi realizada para a compilação de uma lista

taxonômica incluindo as espécies válidas, sinonímias, localidade tipo, depositários do material

tipo, diagnose, distribuição geográfica e notas sobre a biologia de cada espécie, sempre que

disponível. A descrição dos caracteres morfológicos externos seguiu terminologia proposta

por McAlpine (1981), ao passo que para a terminália masculina, seguiu-se as proposições de

Mello-Patiu & Pape (2000) e Giroux et al. (2010).

A matriz de caracteres para machos de Oxysarcodexia foi produzida com o auxílio do

programa Mesquite versão 2.75 (Maddison & Maddison, 2011), incluindo caracteres

morfológicos externos e da terminália, escolhidos após exame comparativo entre as espécies.

12

As análises filogenéticas foram realizadas com auxílio do programa TNT (Goloboff et al.,

2008), por meio de análises por busca tradicional e por novas tecnologias de busca. As árvores

mais parcimoniosas foram geradas considerando pesos iguais e pesos implicado para os

carcteres, incluindo os valores de suporte Bremer para os ramos, tanto para a análise da matriz

completa quanto da matriz considerando apenas caracteres da terminália.

Para analisar as alterações morfológicas consequentes da assimetria, as espécies

Oxysarcodexia fringidea (Curran & Walley, 1934), Oxysarcodexia timida (Aldrich, 1916) e

Oxysarcodexia varia (Walker, 1836), que apresentam vésicas assimétricas, foram submetidas

à microscopia eletrônica de varredura (MEV) para a produção de micrografias rotacionais. A

terminália dos espécimes foi cuidadosamente dissecada, clarificada em ácido lático 80% e

devidamente preparada para a produção de micrografias em microscópio eletrônico de

varredura do tipo “JEOL-JSM-6335F”, localizado no Museu de Zoologia da Universidade de

Copenhagen, Copenhagen, Dinamarca. A confecção de micrografias rotacionais interativas foi

realizada de acordo com a metodologia proposta por Cheung et al. (2013). As imagens foram

então agrupadas em uma animação e as micrografias rotacionais disponibilizadas na web, com

o auxílio de um plug-in específico.

13

5. CAPÍTULO I

ON Oxysarcodexia (DIPTERA: SARCOPHAGIDAE): A TAXONOMIC CONSPECTUS WITH THE

DESCRIPTION OF SIX NEW SPECIES

SOBRE O GÊNERO Oxysarcodexia (DIPTERA: SARCOPHAGIDAE): CONSPECTO TAXONÔMICO

COM A DESCRIÇÃO DE SEIS ESPÉCIES NOVAS

CARINA MARA DE SOUZA1, THOMAS PAPE

2 & PATRICIA JACQUELINE THYSSEN

3

1 Department of Animal Biology, University of Campinas - UNICAMP, POB 6109, PC 13083-

970, Campinas, São Paulo, Brazil. E-mail: [email protected]

2 Natural History Museum of Denmark, Universitetsparken 15, DK - 2100 Copenhagen,

Denmark. E-mail: [email protected]

3 Department of Microbiology and Biology, Federal University of Pelotas - UFPel, POB 354,

PC 96010-900, Pelotas, Rio Grande do Sul, Brazil. E-mail: [email protected]

Abstract

The genus Oxysarcodexia (Diptera: Sarcophagidae) is one of the most species-rich

genera of Neotropical flesh flies, with a few species occurring also in Nearctic, Australasian

and Oceanian Regions. Species within this genus are considered dung-breeding, although a

great part of biological information is still unknown. A taxonomical conspectus based on male

specimens of Oxysarcodexia genus is presented, including diagnosis of each species,

geographical distribution and biological data, when available, besides a pictorial appendix. It

is currently recognized a total of 89 valid species belonging to Oxysarcodexia genus, including

six new here described: Oxysarcodexia n. sp. 1 (Colombia and Ecuador), Oxysarcodexia n.

sp. 2 (French Guiana), Oxysarcodexia n. sp. 3 (Costa Rica), Oxysarcodexia n. sp. 4 (Brazil),

Oxysarcodexia n. sp. 5 (Ecuador) and Oxysarcodexia n. sp. 6 (Ecuador). On the moment,

14

Oxysarcodexia aureiceps (Macquart, 1855), Oxysarcodexia dorisae Dodge, 1965,

Oxysarcodexia flavifrons (Macquart, 1846) and Oxysarcodexia neivae Mattos, 1919 are

considered species awainting status confirmation once original descriptions are based only on

female specimens, some of type material are in very bad conditions, and until a wide and

thorough study of females is conducted.

Key words: Flesh flies, dung-breeding flies, Neotropical Region, new records.

Resumo

O gênero Oxysarcodexia (Diptera: Sarcophagidae) é um dos gêneros de sarcofagídeos

que apresenta grande riqueza de espécies que ocorrem majoritariamente na região Neotropical,

com poucas ocorrendo também nas regiões Neártica, Australásia e Oceânica. As espécies

inclusas nesse gênero são consideradas coprófagas, embora a biologia de muitas espécies

ainda permaneça desconhecida. Um conspecto taxonômico com base nos machos do gênero

Oxysarcodexia é apresentado, incluindo diagnose de cada espécie, distribuição geográfica e

dados biológicos, quando disponíveis, além de um apêndice pictórico. São reconhecidas 89

espécies válidas de Oxysarcodexia, incluindo seis espécies novas que se encontram aqui

descritas: Oxysarcodexia n. sp. 1 (Colômbia e Equador), Oxysarcodexia n. sp. 2 (Guiana

Francesa), Oxysarcodexia n. sp. 3 (Costa Rica), Oxysarcodexia n. sp. 4 (Brasil),

Oxysarcodexia n. sp. 5 (Equador) e Oxysarcodexia n. sp. 6 (Equador). Momentaneamente as

espécies Oxysarcodexia aureiceps (Macquart, 1855), Oxysarcodexia dorisae Dodge, 1965,

Oxysarcodexia flavifrons (Macquart, 1846) e Oxysarcodexia neivae Mattos, 1919 foram

consideradas espécies aguardando confirmação de status devido às descrições originais serem

baseadas exclusivamente em fêmeas, alguns dos materiais tipo estarem comprometidos por

problemas de conservação e até que um amplo e minucioso estudo das fêmeas desse gênero

seja realizado.

Palavras-chave: Sarcofagídeos, moscas coprófagas, região Neotropical, novos registros.

15

Introduction

Sarcophagidae (Diptera: Brachycera) is a family comprising three subfamilies,

Paramacronychiinae, Miltogramminae and Sarcophaginae, with about 3,100 described species.

Approximately 870 have been recorded for the Neotropical Region (Evenhuis et al. 2007 apud

Pape et al. 2009). However, richness of flesh flies might follow the same pattern of Diptera in

general, which is estimated be much higher than the number of described species. The low

number of taxonomists and areas still poorly or not sampled are limiting factors to the

knowledge of actual number of species (Brown 2009; Carvalho et al. 2012). The ecological

relationships of flesh flies vary from parasitism (such as myiasis and parasitoids), predatism

(on other insects, snails, earthworms or spider egg sacs) to feeding in decomposing organic

matter (as vertebrate/invertebrate carcasses and feces) (Pape & Dahlem 2010; Carvalho et al.

2012).

Oxysarcodexia, a genus belonging to the subfamily Sarcophaginae, is one of the most

species-rich genera of Neotropical Sarcophagidae, with about 80 species (Pape 1996; Soares

& Mello-Patiu 2010) and a few occur also in Nearctic, Australasian and Oceanian Regions

(Lopes 1973b, Lopes & Tibana 1987; Pape 1996). Species of this genus play an important

ecological role by being associated with decomposing organic material, as feces of mammals

or birds (dung-breeding) and carcasses (attracted fauna and carrion-breeding) (Pape & Dahlem

2010; Carvalho et al. 2012). Furthermore, it is one of the most frequently recorded and, in

many cases, also most diverse flesh fly genus in about 80 studies recorded in the literature,

dealing with forensic entomology, synanthropy or surveys of dipteran species, especially in

the Neotropical Region (e.g. Dodge & Seago 1954; Linhares 1981; D‟Almeida 1984; Dias et

al. 1984a; Oliveira-Costa et al. 2001; Barros et al. 2008; Rosa et al. 2011). Despite of this

wide occurrence, these flies remain identified in many papers only to genus level (e.g.

Cornaby 1974; Wolff et al. 2001; Pérez, Duque & Wolff 2005; Amat 2010; Horenstein et al.

2010), evidencing the issue of identifying species with a high level of homologies and

similarities. Establishment and maintenance of colonies of different Oxysarcodexia species at

laboratory is also a difficult task, mainly due to the difficulty of having flies to mate under

artificial conditions (Lopes 1973b).

Thus, the aim of this work was review the genus Oxysarcodexia, in order to provide

taxonomic, morphologic, pictorial and biologic information of the species included in this

16

taxon, with several species that have medical, veterinary and forensic importance. This

taxonomic conspectus is focused basically on male adults. Immatures are poorly known and

are scantily documented. Larval descriptions are found only for six species (see Knipling

1936; Lopes 1943; Wharton & Moon 1979; Lopes & Leite 1986; 1987; Leite & Lopes 1987),

although information on larval stages is reported by Lopes (1973b) for another 24 species

mention no details though. Females are known and have been described for some species (see

Walker 1857; Lopes 1933; 1938; 1939; 1946b; 1973a; 1973b; 1975a; 1975b; 1975c; 1975d;

1976; 1978; 1985; Blanchard 1939; 1942; Lopes & Albuquerque 1955; Dodge 1956; 1965;

1966; 1968; Tibana & Mello 1983a; 1985; Lopes & Tibana 1987; 1991; Mulieri et al. 2010),

although they are very difficult to identify due to the high similarity and variability observed

in chaetotaxy and genital morphology of close species. For several other species, females are

still unknown or there is a lack of more detailed studies that relate them to males already

described.

Material and Methods

Specimens here studied are deposited in the following institutions:

CE-TdeA – Tecnológico de Antioquia, Institución Universitaria (Medellín, Colombia);

MNRJ – Museu Nacional/Universidade Federal do Rio de Janeiro (Rio de Janeiro, Brazil);

NRM – Swedish Museum of Natural History (Stockholm, Sweden);

ZMUC – Zoological Museum of Copenhagen (Copenhagen, Denmark).

Digital photographs were taken of the lateral habitus, male terminalia in lateral,

anterior (whenever possible) and posterior views for the already described species with

specimens available for examination and also for the newly described species, using a digital

camera Leica DELUX 3™

(10 megapixels), mounted on a Leica S8AP0™

stereoscope or using

a digital camera Carl Zeiss AXIOCAM MRc™

(5 megapixels), mounted on a Carl Zeiss

STEREO DISCOVERY.V12™

stereoscope. For each view, exposures were taken with

extended deep-focus and were stacked using Zerene Stacker™

software. Species for which no

specimens were available for study were documented using information from the literature and

the best illustration of the male terminalia already published were presented after edition.

17

In addition to the study of specimens, a thorough review of the literature about

Oxysarcodexia until December 2013 was performed for compiling a taxonomic list of valid

species, in alphabetical order. Taxonomic data are given with the following structure: valid

name; synonyms; type locality; depository of type specimens; diagnosis; remarks, whenever

relevant; geographical distribution; and, whenever possible, notes about the species biology.

The distributional data were based on the Catalogue of the Sarcophagidae of the World (Pape

1996), on additional records of the literature and on inclusion of localities where specimens

studied in the present conspectus were collected, with the following structure: biogeographic

region, countries and their states/provinces (alphabetically ordered). New records of

occurrence are underlined. All pictorial information is given in plates for each species, in the

appendix, also in alphabetical order.

Depositories of the species are cited by the following acronyms (plus the three

institutions listed previously):

AMNH – American Museum of Natural History, Department of Entomology (New York,

New York, USA);

BMNH – The Natural History Museum, Department of Entomology (London, England,

United Kingdom);

CAS – California Academy of Sciences, Department of Entomology (San Francisco,

California, USA);

DEI – Deutsches Entomologisches Institut, Deutschen Akademie der

Landwirtswissenschaften zu Berlin (Eberswalde, Brandenburg, Germany);

MZUSP – Museu de Zoologia, Universidade de São Paulo (São Paulo, São Paulo, Brazil);

NMW – Naturhistorisches Museum Wien (Vienna, Austria);

SMN – Staatliches Museum für Naturkunde (Stuttgart, Germany);

UCC – University of Concepción (Concepción, Biobio, Chile);

UKaL – University of Kansas, State Biological Survey of Kansas Invertebrate Collection

(Lawrence, Kansas, USA);

UPRG – Universidad Nacional “Pedro Ruiz Gallo”, Departamento de Fitotecnia, Museo de

Entomología (Lambayeque, Lambayeque, Peru);

18

USNM – United States National Museum of Natural History, United States National

Entomological Collection (Washington, D.C., USA).

Terminology follows McAlpine (1981) for external characters and Mello-Patiu & Pape

(2000) and Giroux et al. (2010) for male terminalia. Measurement of the body length was

obtained by including the head (without considering the antennae), thorax (from the neck to

the posterior margin of the scutellum), and abdomen (from the anterior margin of abdominal

tergite 2 to the posterior margin of epandrium) lengths, in order to offset bias caused by any

eventual curvature of the specimen. Type specimens were deposited in the Diptera collections

of the MNRJ, NRM and ZMUC, as pointed out specifically in each description. Label

information of the new species and photographed specimens were transcribed without any

modification; a forward slash (/) was used to separate individual labels, whereas a double

forward slash (//) was used to separate different specimens of a same species. Any necessary

additional comment was given inside brackets and the depository in parenthesis at the end of

transcription label information.

Generic Definition

In order to group some species which were included previously in Sarcophaga genus

only for the absence of a specific taxon, Townsend proposed in 1917 the genus Oxysarcodexia

(Townsend 1917). Dasyproctia Enderlein, 1928, Hybopygia Enderlein, 1928, Apelophyla Hall,

1938 and Xarcophaga Dodge, 1968 are considered synonymous of Oxysarcodexia (Lopes

1946b; Dodge 1966; Lopes 1975c). The type species of this genus is Oxysarcodexia peltata

(Aldrich, 1916). Specific characters for species identification are located almost exclusively in

male terminalia (Lopes 1946b).

Broader studies of this genus based on male adults, including descriptions of new

species, list of names, geographic distribution, new combination of taxa and dichotomous key

were performed by Lopes (1946b), Dodge (1966) and Lopes & Tibana (1987). A catalogue of

Sarcophagidae species of the world provide a list of Oxysarcodexia species and their known

distribution until 1995 (see Pape 1996). The study of tergites 6+7, nowadays recognized as

tergite 7 (T7), of 38 species is the most comprehensive approach for females, classified them

into three different groups based on syntergite form: undivided (with 21 species), partially

19

divided into two plates (with 12 species) or membranous (with 5 species) (Tibana & Mello

1985). Other studies in this way are scarce, remaining only a few descriptions and revisions

scattered in several papers, generally covering only for one species (e.g. Lopes 1938; 1946b;

1973a; Blanchard 1939; Dodge 1956; Mulieri et al. 2010).

Oxysarcodexia is considered a monophyletic genus by the autapomorphy of the lateral

triangular extension of the phallic tube (“tooth-like”) above the vesica. Characters as post-

cranium concave, tergite 5 (T5) entirely yellow, juxta microvillose, and median stylus curved

towards distal end of the phallus are considered homoplasies (Giroux et al. 2010). Character

states as postalar wall setose; male with ctenidium in mid femur of flattened spines; tegula

darkish and basicosta lighter (generally light brownish); male sternite 5 (ST5) deeply cleft

with almost parallel sides, with a few exceptions; penis unsegmented; phallus with three

conducting styli and with the “tooth-like” extension above the vesica; vesica elongated,

conspicuous and always well constituted are pointed out in the literature as generic structures

that, occurring together, allow the correct recognition of this taxon (Lopes 1946b; Dodge

1966; Pape 1996; Carvalho & Mello-Patiu 2008; Silva & Mello-Patiu 2008).

Remarks

Juxta. According to Roback (1954), the juxta is a ventral (i.e. apical) appendage of the

“corpus” (i.e. phallic tube) that can be immovable (i.e. fused to the phallic tube), partially or

completely movable. From this standpoint, Oxysarcodexia, beside Agria Robineau-Desvoidy,

1830, Cistudinomyia Townsend, 1917, Ravinia Robineau-Desvoidy, 1863, Angiometopa

Brauer & Bergenstamm, 1889 and Wohlfahrtia Brauer & Bergenstamm, 1889, is considered

having no developed juxta (Roback 1954). However, a later phylogeny approach of Giroux et

al. (2010) redefined this structure as an apical extension of the posterior side of the

distiphallus, which, even without a visible landmark as structural divisions or a groove, for

example, is considered arising from the base of the median stylus. Therefore, Sarcophaginae

subfamily, and consequently Oxysarcodexia genus, is recognized as having a juxta. Our

examination of different Oxysarcodexia species comes in agreement with this later

proposition.

20

Genus Diagnosis

Oxysarcodexia Townsend, 1917

Apelophyla Hall, 1938

Dasyproctia Enderlein, 1928

Hybopygia Enderlein, 1928

Xarcophaga Dodge, 1968

Male. Total length ranges, in average, from 5 to 12mm.

Head. Fronto-orbital, parafacial and postocular plates generally with gold

microtomentum, usually intense, but sometimes pale and/or with a few silvery shades due to

natural conditions or conservation issues; parafacial with a few darkish short setae; occiput

blackish with silvery microtomentum, black setae in dorsomedial and lateral areas and a few

golden setae in ventromedial area; postocular plates with golden or silvery microtomentum;

frontal vitta darkish with a row of frontal setae present until half high of pedicel, not strongly

divergent, and varying among 7–8 and 12–13; inner vertical seta well-developed, outer

vertical seta not differentiated from postocular setae; ocellar setae smaller than or equal in size

to uppermost frontals; 1 reclinate fronto-orbital seta, with variable length and proclinate seta

absent (only one exception, Oxysarcodexia orbitalis Dodge, 1966); gena and postgena darkish

with golden microtomentum, generally intense, but sometimes pale and both with black setae;

antenna dark brown, first flagellomere with brownish microtomentum; arista darkish brown

and long plumose on basal half or basal ¾ ; palpus dark brown; strong vibrissa.

Thorax. Grayish with 3 black vittae and lightly pale or strong golden microtomentum

sometimes is more intense laterally, at humeral region. Chaetotaxy: acrostichals: 0+1;

dorsocentrals unequally developed: 2–3 well developed and 1–3 smaller + 2 well

differentiated and 1–3 smaller or 3 well differentiated (a small seta among these 3 can be

present); intra-alars: 2+2 (not unusual one of them weaker than the other); supra-alars: 2+3;

postalars: 2; postpronotals: 3; notopleurals: 4 (2 large primaries and 2 smaller subprimaries);

katepisternals: 3 with middle one weaker and inserted slightly below the others; meropleurals:

6–12; postalar wall setose; apical scutellar seta can be present or not; subapical: 1; lateral: 1;

basal: 1; discal: 1; proepisternum always bare; prosternum setose.

21

Wings. Hyaline, black tegula, R1 bare (only one exception, Oxysarcodexia

chaetopygialis (Williston, 1896)), R4+5 setulose in proximal ½, ⅔ or ¾ of distance to r-m,

costal spine not differentiated and third costal sector without ventral setulae.

Legs. color generally blackish, but sometimes brownish or yellowish; fore femur with a

row of setae on dorsal, posterodorsal, and posteroventral surfaces; fore tibia with 1 strong and

3 weaker anterodorsal, 1 dorsal, 1 posterior and 1 posteroventral setae; mid femur with 5

median anterior and 2 pre-apical posterior setae, 5 anteroventral and 6 posteroventral long

setae, ctenidium of flattened spines apically on posteroventral surface; mid tibia with 2

anterodorsal, 1 posterodorsal, 1 posteroventral and 1 anterodorsal setae; hind femur with 1

strong pre-apical dorsal and 1 posterodorsal seta, 1 anteroventral and 1 anterodorsal row of

long setae, and 1 posteroventral row of decreasing size setae up to the distal ⅓, approximately;

hind tibia with 2 anteroventral, one dorsal, 2 posterodorsal and 3 anterodorsal setae;

tarsomeres of fore, mid and hind legs with a weak ventral golden micromentum.

Abdomen. Silvery or yellow-silveryish microtomentum, sometimes with golden

microtomentum more intense on lateral margin of the tergites; tergite 2 (T2) with silvery (most

common) or yellow-silveryish microtomentum; tergite 3 (T3) with 1–3 lateral marginal setae;

tergite 4 (T4) with 1–4 lateral marginal and 0–2 median marginal setae; tergite 5 (T5)

completely yellow and with about 16–23 strong setae along the posterior margin; sternites 2–4

oblong or square with scattered setulae of variable length, generally stronger along the

posterior edge; sternite 5 (ST5) with a median deep cleft of almost parallel edges (only a few

exceptions – 8 species – which have V-shaped edges) forming arms of variable width (thin,

medium, large), microtomentum variable (completely or partially yellow or dark) and with the

presence of window (space between the arms at ventroposterior edge of T5 level) and setosity,

setae or both along the arms, especially along the posteroapical edge.

Terminalia (Fig. 1). Syntergosternite 7+8 yellow brownish with golden

microtomentum, scattered short black setulae, and 4–10 marginal well developed setae.

Epandrium is yellowish, generally intense, but sometimes pale or blackish, though still

showing golden microtomentum; presence of black setae. Surstylus triangular (enlarged base

and narrow apex) or oblong (elongated dorsoventrally with base not very enlarged than the

apex) and discal setulae present, although a strong seta not always present. Cercus, in lateral

view, straight, sinuous or bent backwards (more rare), with the apex, which can be darker than

22

the rest of the cercus, pointed, expanded (apical expansion straight, oblique or concave) or

normal (i.e. with the same size as the median area), and with the presence of setae dorsally

(length of setae can be variable) and setosity ventrally; in posterior view, conformation

parallel or divergent, especially at apical portion, a remarkable constriction at the middle

portion of the cercus and a lateromedial pads of setae on the apex can be present in some

species. Postgonite is dark brownish, slender, slightly curved, with apex shape variable

(square, pointed), and sometimes with the presence of a seta. Pregonite is dark brownish,

always broader than postgonite, generally curved and with base and apex of same size or

expanded at the base and narrower at the apex (narrowing abruptly or smoothly); apex shape

variable (sharped, pointed, rounded) and mostly darker (blackish) than the base. Phallus well

developed and sclerotized, without division between basi- and distiphallus (i. e. unsegmented),

with the posterodorsal face more sclerotized than the anteroventral; presence of a lateral

triangular extension of the phallic tube (“tooth-like”) above the vesica; distiphallus

ventroapical margin, in lateral view, smooth or serrated; ventroapical concavity at the

distiphallus can be observed in some species; distiphallic apical shape conic, rounded or

square/oblong; specific structures at the distiphallus, as lateroapical furrow, ventroapical

projections, lateral lobes and dorsoapical membranous formation (“swelling-like”) can be

observed in some species. Median stylus coming towards the phallic tube apex and is

connected by dorsal roads (as named by Roback (1954) and pointed out by Silva & Mello-

Patiu (2008)) to the vesica. Juxta bifurcated, rough laterally, smooth ventrally and joined to

the phallic tube. Presence of a conspicuous and highly modified vesica inserted at

approximately the middle of the ventromedial portion of the phallic tube; a basal branch

comes out from the distiphallic tube and splits mediolaterally into two lateral branches that can

end up in filament (at most tapering to the apex), rounded or square terminal lobes, with

normal or reduced sizes. The vesica is articulated and presents a great variety of

ornamentation. In some species, a median projection with an angular or rounded peak can be

present at the basal branch before the mediolateral division of the vesica. Lateral lobes, i.e.

division of the vesica coming from or close to the basal branch of the vesica, placed laterally

to phallic tube is another vesica structural modification seen in some species. The whole

vesica is well sclerotized, although terminal lobes can be more membranous. Spines can be

present along the edges, only on ventral surface, only on dorsal surface or in both surfaces of

23

terminal lobes. Despite of the great shape variation of the vesica, shape as the “leaf-like”

terminal lobes is one of the most common. In only 8 species vesica is asymmetric, having left

side modified somehow (e.g. reduced, displaced in transverse plane, etc.).

Species awaiting status confirmation

O. aureiceps, O. dorisae, O. flavifrons and O. neivae

Descriptions of O. aureiceps (Macquart, 1855) (and incorrect spelling O. aurescens

(Lopes &Tibana 1987)), O. dorisae Dodge, 1965 (and incorrect spelling O. dorissae (Lopes

&Tibana 1987)), O. flavifrons (Macquart, 1846) and O. neivae Mattos, 1919 are based only on

female specimens. Lopes (1946b) considered O. flavifrons probably synonymy with O. varia

(Walker, 1836), although, in a posterior study (Lopes & Tibana 1987), this species was

ascribed as “inquaerendae”, i.e., with doubtful description. Therefore, for the moment, it is

wiser consider these species lacking status confirmation until a wide and deep study of

females is conducted. Oxysarcodexia aureiceps is considered “inquirenda” for being assigned

to Oxysarcodexia genus although presenting doubtful identity (Lopes & Tibana 1987; Pape

1996) and also presenting very bad conditions of preservation according to Aldrich (1930).

Taxonomy

Oxysarcodexia n. sp. 1

(Appendix 1)

Type locality. ECUADOR: Napo Province.

Depository of type material. ZMUC.

Description. Male. Total length = 8.30–8.50 mm (n=2). Head. Fronto-orbital,

parafacial and postocular plates with gold microtomentum; occiput blackish with silvery

microtomentum, black setae in dorsomedial and lateral areas and a few golden setae in

ventromedial area; front about 0.08x head width at level of ocellar triangle; frontal vitta

darkish with row of 9–11 frontal setae; inner vertical seta well-developed, outer vertical seta

not differentiated; ocellar setae equal in size to uppermost frontals; 1 reclinate fronto-orbital

seta, with variable length (⅓ larger than the frontals or equal in size) and proclinate seta

absent; gena and postgena black, but with, respectively, golden and silvery microtomentum,

24

and both with black setae; antenna dark brown, first flagellomere with brownish

microtomentum and about 2.00x longer than pedicel; arista brown with the middle portion

lighter and long plumose on basal ¾; palpus dark brown. Thorax. Grayish with lightly pale or

stronger golden microtomentum, more intense laterally; chaetotaxy: acrostichals 0+1,

dorsocentrals 3+5, intra-alars 2+2, supra-alars 2+3, postalars 2, postpronotals 3, notopleurals 4

(2 large primaries and 2 smaller subprimaries), katepisternals 3 with the middle one weaker

and inserted slightly below the others, meropleurals 8–9, postalar wall setose, scutellum with 1

apical, 1 subapical, 1 lateral, 1 basal and 1 discal seta; prosternum setose on the ⅔ distal.

Wings. Hyaline, black tegula, R1 bare, R4+5 setulose in proximal ⅔ of distance to r-m, costal

spine not differentiated and third costal sector without ventral setulae. Legs. blackish brown;

fore femur with a row of setae on dorsal, posterodorsal, and posteroventral surfaces; fore tibia

with one strong and 3 weaker anterodorsal, 1 dorsal, 1 posterior and 1 posteroventral setae;

mid femur with 5 median anterior and 2 pre-apical posterior setae, 5 anteroventral and 6

posteroventral long setae, ctenidium of flattened spines apically on posteroventral surface; mid

tibia with 2 anterodorsal, 1 posterodorsal, 1 posteroventral and 1 anterodorsal setae; hind

femur with 1 strong pre-apical dorsal and 1 posterodorsal seta, 1 anteroventral and 1

anterodorsal row of long setae, and 1 posteroventral row of decreasing size setae up to the

distal ⅓, approximately; hind tibia with 2 anteroventral, 1 dorsal, 2 posterodorsal and 3

anterodorsal setae; tarsomeres of fore, mid and hind legs with a weak ventral golden

micromentum. Abdomen. Dark brownish with golden microtomentum; tergite 2 with silvery

microtomentum; T3 with 1 marginal lateral seta; T4 with 1 marginal lateral and 1 median

marginal seta; T5 with about 17–23 strong setae along the posterior margin; sternites 2–4

oblong with scattered setulae, stronger along the edges; ST5 with a median deep cleft, with

edges almost parallel. Terminalia. Syntergosternite 7+8 yellow brownish with golden

microtomentum, scattered short black setulae and 8–10 marginal bristles; epandrium yellowish

with golden microtomentum and black setae; surstylus triangular with enlarged base and

narrow apex and sparse marginal and discal setulae; cercus, in lateral view, straight with a

darker apical expansion and, in posterior view, with apex slightly divergent; postgonite

slender, slightly curved, with a seta on the distal ⅓, approximately, and square apex; pregonite

broader, curved and with sharp apex. Phallus well sclerotized, without division between basi-

and distiphallus; oval distiphallic apex; juxta bifurcated, rough laterally and smooth ventrally;

25

vesica well sclerotized and well developed, with a ventrobasal projection, slightly curved

dorsally and, in anterior view, leaf-like branches with spines in dorsal surface (absent

anteriorly).

Female. Unknown.

Etymology. The specific epithet is given based on the grass leaf-like shape of the

vesica; formed by joint the Latin words agros = plant (grass-like) and frons = leaf.

Remarks. Cerci and phallus shape of Oxysarcodexia n. sp. 1 present similarities to

those seen on Oxysarcodexia amorosa (Schiner, 1868), O. berlai, Oxysarcodexia similata

Lopes & Tibana, 1987 and Oxysarcodexia xanthosoma (Aldrich, 1916). The main differences

among them are observed in the vesica, which in Oxysarcodexia n. sp. 1 is more spinous and

doesn‟t have the same basal portion of terminal lobes as the others, and on distiphallus

ventroapical area, without the presence of a ventroapical cleft, seen in lateral view on O.

amorosa, O. similata and O. xanthosoma.

Distribution. NEOTROPICAL. Colombia (San Martin) and Ecuador (Napo Province).

Biology. Unknown.

Material examined. ♂ [holotype]: ECUADOR: Napo Province: Yasuní National Park:

Yasuní Research Station: 76° 36‟W 00° 38‟S: 3–20 XI 1998: T. Pape & B. Viklund / NRM-

DIPT 0014467 (NRM) // ♂ [paratype]: COLOMBIA: Amazonas PNN Amacayanu, Camino a

San Martin; 3°41‟N 70°15‟W; 1–10.iii.2004, sweepnet; T. Pape & D. Arias. Id# 4325 / NRM-

DIPT 0014650 (NRM).


(Appendix 2)

Type locality. FRENCH GUIANA: Montsinery.


Description. Male. Total length = 6.8 mm. Head. Fronto-orbital, parafacial and

postocular plates with gold microtomentum; occiput blackish with silvery microtomentum,

black setae in dorsomedial and lateral areas and a few golden setae in ventromedial area; front

about 0.1x head width at level of ocellar triangle; frontal vitta blackish, with row of 9–11

frontal setae; inner vertical seta well-developed, outer vertical seta 0.2x as long as the inner

one and as long as a postocellar seta; ocellar setae about 0.3x as long as the frontal setae; 1

26

reclinate fronto-orbital seta equal in size to frontals and proclinate seta absent; gena and

postgena blackish with silvery microtomentum and black setae; antenna dark brown, first

flagellomere with pale golden microtomentum and about 3.3x as long as pedicel; arista dark

brown with the middle portion lighter and long plumose on basal ⅔; palpus dark brown.

Thorax. Grayish with pale golden microtomentum, slightly more intense laterally; chaetotaxy:

acrostichals 0+1, dorsocentrals 3+5, intra-alars 2+2, supra-alars 2+2, postalar 1, postpronotals

3, notopleurals 4 (2 large primaries and 2 smaller subprimaries), katepisternals 3 with the

middle one weaker and inserted slightly below the others, meropleurals 7, postalar wall setose,

scutellum with 1 apical, 1 subapical, 1 lateral, 1 basal and 1 discal seta; prosternum bare.

Wings. Hyaline, black tegula, R1 bare, R4+5 setulose in proximal ⅔ of distance to r-m, costal

spine small, but distinct and third costal sector without ventral setulae. Legs. blackish brown,

fore femur with a row of setae on dorsal and anteroventral surfaces; fore tibia with 1 dorsal, 1

anterior, 1 anterodorsal and 1 posteroventral setae; mid femur with 5 median anterior and 2

pre-apical posterior setae, ctenidium of flattened spines apically on posteroventral surface, 1

posteroventral and 1 anteroventral row of long setae; mid tibia with 1 anterior, 2 anterodorsal,

1 posterodorsal, 1 anteroventral and 1 posteroventral setae; hind femur with 1 posterodorsal

seta, 1 anteroventral and 1 anterodorsal row of long setae; hind tibia and hind tarsomeres lost

on both sides. Abdomen. Dark brownish with golden microtomentum more intense laterally;

T3–4 with 2 marginal lateral setae; T5 with about 20 strong setae along the posterior margin;

ST2–4 oblong with a pair of strong setae on posterior margin; ST5 with a median deep cleft,

with edges almost parallel. Terminalia. Syntergosternite 7+8 yellow brownish with sparse

golden microtomentum, scattered short black setulae and 9 marginal bristles; epandrium

yellow brownish with sparse golden microtomentum and short black setulae; surstylus

triangular with enlarged base and narrow apex and sparse marginal and discal setulae; cercus,

in lateral view, straight with a darker apical expansion and, in posterior view, with apex

slightly divergent; postgonite slender, slightly curved and with a square apex; pregonite

broader, longer, curved and with pointed apex. Phallus well sclerotized, without division

between basi- and distiphallus; rounded distiphallic apex; juxta with a rugous aspect; vesica

well sclerotized with, in lateral view, median lobes turned on ventroapical direction, tiny

spines along the edges and on dorsal surface and, in anterior view, chicken wing-like

branches.

27

Female. Unknown.

Etymology. The specific epithet refers to the vesica shape in lateral and anterior views,

which reminds chicken wings. From Latin, alectorius is an adjective relative to peacock.

Remarks. Vesica shape is very similar to that observed in Oxysarcodexia angrensis

(Lopes, 1933), although Oxysarcodexia n. sp. 2 present rounded distiphallus apex and cerci

with enlarged apex contrasting with the conic distiphallus apex and cerci with pointed apex of

O. angrensis.

Distribution. NEOTROPICAL. French Guiana (Montsinery).

Biology. Unknown.

Material examined. ♂ [holotype]: FRENCH GUIANA: Montsinery; “Emerald

Jungle”; Carrefour du Gallion; 01.i.2003, M. Kotrba / NRM-DIPT 0014601 (NRM).


(Appendix 3)

Type locality. COSTA RICA: Puntaneras, Monteverde.


Description. Male. Total length = 9.5–10.0 mm (n=3). Head. Fronto-orbital,

parafacial and postocular plates with gold microtomentum; occiput blackish with silvery

microtomentum, black setae in dorsomedial and lateral areas and golden setae in ventromedial

area; front about 0.1x head width at level of ocellar triangle; frontal vitta dark, slight brownish

anteriorly, with row of 7–9 frontal setae; inner vertical seta well-developed, outer vertical seta

not differentiated; ocellar setae equal in size to uppermost frontals; 2 reclinate fronto-orbital

setae equal in size to frontals and proclinate seta absent; gena and postgena black, but with,

respectively, golden and silvery microtomentum, and both with black setae; antenna dark

brown, first flagellomere with brownish microtomentum and about 2.3x longer than pedicel;

arista dark brown with the middle portion lighter and long plumose on basal ¾; palpus dark

brown. Thorax. Grayish with lightly pale golden microtomentum; chaetotaxy: acrostichals

0+1, dorsocentrals 2+4, intra-alars 2+2, supra-alars 3+3, postalars 2, postpronotals 3,

notopleurals 4 (2 large primaries and 2 smaller subprimaries), katepisternals 3 with the middle

one weaker and inserted slightly below the others, meropleurals 10, postalar wall setose,

scutellum with no apical, 1 subapical, 1 lateral and 1 discal seta; prosternum setose at edges.

28

Wings. Hyaline, black tegula, R1 bare, R4+5 setulose in proximal ⅓ of distance to r-m, costal

spine not differentiated and third costal sector without ventral setulae. Legs. blackish brown,

fore femur with a row of setae on dorsal, posterodorsal, and posteroventral surfaces; fore tibia

with 2 anterodorsal, 1 posterior and 1 posteroventral setae; mid femur with 5 median anterior

and 3 pre-apical posterior setae, ctenidium of flattened spines apically on posteroventral

surface, 1 posteroventral and 1 anteroventral row of long setae; mid tibia with 1 anterior, 2

posteroventral and 1 posterodorsal setae; hind femur with a strong anteroventral and a pre-

apical posterior seta, 1 posteroventral and 1 anterodorsal row of long setae; hind tibia with 1

anterior row of long setae, 2 anteroventral, 1 dorsal, 3 posterodorsal and 2 posterior setae;

tarsomeres of fore, mid and hind legs with a ventral golden micromentum, weaker in mid legs.

Abdomen. Blackish gray with silvery microtomentum more intense laterally; T3 with 2

marginal lateral setae; T4 with 3 marginal lateral setae; T5 with about 18 strong setae along

the posterior margin; ST2–4 square with a pair of strong setae on posterior margin; ST5 with a

median deep V-shaped cleft. Terminalia. Syntergosternite 7+8 large, brownish with golden

microtomentum laterally, scattered short black setulae and 12 marginal bristles; epandrium

yellowish with short black setulae and golden microtomentum; surstylus oblong with narrow

base and apex, enlarged on ⅔ portion, with sparse marginal and discal setulae and a long and

slender colorless setae at apex; cercus, in lateral view, slightly sinuous with blackish preapical

expansion and pointed apex; postgonite slender and slightly curved, especially at the apex;

pregonite broad with narrower rounded apex. Phallus well sclerotized, without division

between basi- and distiphallus; juxta bifurcated with small spines along the inner edges, except

at the apex; 2 lateral styli and 1 median stylus both tubular-shaped and partially hidden by the

juxta; vesica well sclerotized and well developed, with an angular ventroapical projection

about 1.2 mm longer than distiphallic apex and, in anterior view, enlarged base and bifurcated

apex with small spines and horseshoe-like branches.

Female. Unknown (see remarks).

Etymology. The specific epithet is related to the right angle seen on vesica

conformation. From the Latin, angulosus = with corners or angles.

Remarks. Cerci of Oxysarcodexia perneta (Walker, 1861) is similar to that found on

Oxysarcodexia n. sp. 3. However, Oxysarcodexia n. sp. 3 terminalia constitution is very

peculiar, especially for the vesica shape, not found in any other species of the genus. A female,

29

labelled “COSTA RICA, San José; Cerro Muerte, 16km S; Empalme, 2600m; III–VI 1990

Hanson / COSTA RICA INBIO; CRI001; 107347” is mostly likely the female of this species

because of the blackish occiput with silvery microtomentum, black setae in dorsomedial and

lateral areas and, even sparser than in male, golden setae in ventromedial area and the body

color pattern (grayish thorax lightly pale with golden microtomentum, abdomen blackish gray

with silvery microtomentum and genital segments yellowish with golden microtomentum)

very similar to those found in males. Nevertheless, without a more exhaustive study of female

comparative morphology we prefer to exclude it from “formal type status”.

Distribution. NEOTROPICAL. Costa Rica (Puntaneras, San José, and Punta).

Biology. Unknown.

Material examined. ♂♂ [holotype]: COSTA RICA: Punt. Monteverde 1840m; Cerro

Amigos; 14.VI.2000, T. Pape / INB0003086811 (ZMUC) // [paratype]: Est, Cuerici. Send. El

Carbon, 4.6km al E. de Villa Mills, Prov. San Jose, COSTA RICA 2600m. 26–31 OCT / 1995.

B. Gamboa, L_S_389550_ / 500050 #6326 / COSTA RICA INBIO; CRI002; 363463 (ZMUC)

// [paratype]: Estac. Pitier, Prov. Punta, COSTA RICA. 1679m. 5–18 ENE 1995. G. Fonseca,

L_N_330900_577400 #5950 (ZMUC).


(Appendix 4)

Type locality. BRAZIL: Rio de Janeiro.

Depository of type material. MNRJ.


postocular plates with golden microtomentum; occiput blackish with golden microtomentum,

black setae in dorsomedial and lateral areas and a few golden setae in ventromedial area; front

about 0.1x head width at level of ocellar triangle; frontal vitta blackish, with a row of 14

frontal setae; inner vertical seta well-developed, outer vertical seta 0.3x as long as the inner

one; ocellar setae 0.8x as long as the uppermost frontal setae; 1 reclinate fronto-orbital seta

2.2x as long as the uppermost frontal and proclinate seta absent; gena and postgena blackish

with golden microtomentum and black setae; antenna dark brown, first flagellomere with pale

golden microtomentum and about 2.3x as long as pedicel; arista brown and long plumose on

basal ⅔; palpus dark brown. Thorax. Greyish with golden microtomentum; chaetotaxy:

30

acrostichals 3+1, dorsocentrals 3 (first weaker) + 3, intra-alars 2+2, supra-alars 2+3, postalars

2, postpronotals 3, notopleurals 4 (2 large primaries and 2 smaller subprimaries),

katepisternals 3 with the middle one weaker and inserted slightly below the others,

meropleurals 8–9, postalar wall setose, scutellum with 1 apical, 1 subapical, 1 lateral, 1 basal

and 1 discal seta; prosternum with few setae along the edges. Wings. Hyaline, black tegula,

R1 bare, R4+5 setulose in proximal ¾ of distance to r-m, costal spine not differentiated and

third costal sector without ventral setulae. Legs. darkish brown, fore femur with 2 rows of

setae on posterodorsal and 1 on posteroventral surfaces; fore tibia with 3 dorsal, 1

posterodorsal, 2 posteroventral and 1 posterior setae; mid femur with 6 median anterodorsal, 2

pre-apical posterior setae, 1 anteroventral and 1 posteroventral row of setae and ctenidium of

flattened spines apically on posteroventral surface; mid tibia with 1 dorsal, 2 posterodorsal, 1

posterior, 1 posteroventral, 1 ventral, 1 anterior and 2 anterodorsal setae; hind femur with 1

posterodorsal and 1 dorsal seta, 1 anteroventral and 1 anterodorsal row of long setae, 1 apical

posteroventral row of decreasing setae and 1 anterior row of decreasing setae; hind tibia with 2

anteroventral, 3 anterodorsal, 2 dorsal, 1 anterior and 2 posterodorsal setae; tarsomeres of fore

and hind legs with a weak ventral golden micromentum. Abdomen. Grayish with silvery

microtomentum; T2 with 1 marginal lateral seta, T3 with 2 marginal lateral setae and T4 with

4 marginal lateral and 1 median marginal setae; T5 with about 20 strong setae along the

posterior margin; ST2–4 oblong with scattered setulae on posterolateral and posterior edges;

ST5 with a median deep cleft with almost parallel arms. Terminalia. Syntergosternite 7+8

dark brownish with golden microtomentum and scattered black setulae and 8 marginal bristles;

epandrium dark brownish with golden microtomentum, short black setulae and 2 strong setae

dorsoapically; surstylus oblong, elongated dorsoventrally, with sparse marginal and discal

black setulae; cercus, in lateral view, with a basal expansion, square and slightly curved apex;

postgonite slender, slightly curved apically and with square apex; pregonite broader, slightly

curved and rounded apex. Phallus sclerotized, without division between basi- and distiphallus;

distiphallic apex oblique, with lateral, dorsal and ventral projections, smooth edges and a wide

opening ventroapically; vesica well sclerotized, coming out pre-apically from the phallic tube

as a short branch that fold vetroposteriorly culminating in flattened elongated lobes with

microspines on dorsal surface.

Female. Unknown.

31

Etymology. The specific epithet is given in honor of Ariozano de Souza, father of the

first author.

Remarks. Presence of enlarged apex of the phallus as observed in Oxysarcodexia n.

sp. 4, Oxysarcodexia favorabilis (Lopes, 1935), Oxysarcodexia fraterna (Lopes, 1946),

Oxysarcodexia n. sp. 5, Oxysarcodexia nitida Soares & Mello-Patiu, 2010, Oxysarcodexia

notata Soares & Mello-Patiu, 2010, Oxysarcodexia pallisteri Dodge, 1966, Oxysarcodexia

peruviana (Lopes, 1973), Oxysarcodexia vittata (Lopes, 1946) and Oxysarcodexia xon

(Dodge, 1968) (all previously considered as belonging to “Xarcophaga group” and remarked

by Soares & Mello-Patiu (2010), except for Oxysarcodexia n. sp. 4, O. fraterna,

Oxysarcodexia n. sp. 5 and O. peruviana which we included in this group for presenting the

same characteristic). Vesica with “flower-like” shape is another similarity seen on

Oxysarcodexia n. sp. 4, O. favorabilis and Oxysarcodexia n. sp. 5.

Distribution. NEOTROPICAL. Brazil (Rio de Janeiro).

Biology. Unknown.

Material examined. ♂ [holotype]: BRAZIL: RJ Tijuca Forest nr Rio; 7–30.ix.1993,

T.Pape / NRM-DIPT 0014641 (MNRJ).


(Appendix 5)

Type locality. ECUADOR: Napo Province



postocular plates with pale golden microtomentum; occiput blackish with silvery



blackish, with a row of 8–9 frontal setae; inner vertical seta well-developed; outer vertical seta

not differentiated; ocellar setae 1.3x as long as the uppermost frontal setae; 1 reclinate fronto-

orbital seta 2.3x as long as the uppermost frontal and proclinate seta absent; gena and postgena

blackish with pale golden microtomentum and black setae; antenna dark brownish, first

flagellomere with pale golden microtomentum and about 2.2x as long as pedicel; arista brown

and long plumose on basal ¾; palpus dark brown. Thorax. Greyish with pale golden

32

microtomentum; chaetotaxy: acrostichals 0+1, dorsocentrals 3 (first weaker) + 3, intra-alars

2+2, supra-alars 2+3 (middle one stronger), postalars 2, postpronotals 3 (the lateromedial one

weaker), notopleurals 4 (2 large primaries and 2 smaller subprimaries), katepisternals 3 with

the middle one weaker and inserted slightly below the others, meropleurals 9, postalar wall

setose, scutellum without apical, 1 subapical, 1 lateral (weaker), 1 basal and 1 discal seta;

prosternum with few setae along the edges. Wings. Hyaline, black tegula, R1 bare, R4+5

setulose in proximal ⅔ of distance to r-m, costal spine not differentiated and third costal sector

without ventral setulae. Legs. darkish brown, fore femur with a row of setae on posterodorsal

and posteroventral surfaces, and a row of decreasing setae on posterior surface; fore tibia with

1 dorsal, 2 posterodorsal, 1 anterodorsal and 1 anterior setae; mid femur with 5 anterodorsal, 2

pre-apical posterior, 4 anteroventral, 4 posteroventral setae and ctenidium of flattened spines

apically on posteroventral surface; mid tibia with 2 posterodorsal, 1 posteroventral, 1 ventral,

1 anteroventral and 2 anterodorsal setae; hind femur with 1 posterodorsal and 1 dorsal seta, 1

anteroventral and 1 anterodorsal row of long setae, 1 apical posteroventral row of decreasing

setae; hind tibia with 1 anteroventral, 3 anterodorsal, 1 dorsal and 2 posterodorsal setae;

tarsomeres of fore, mid and hind legs with a weak ventral golden micromentum. Abdomen.

Grayish with pale golden microtomentum; T2 with silvery microtomentum laterally; T2–3

with 1 marginal lateral seta; T4 with 3 marginal lateral and 1 median marginal setae; T5 with

about 18 strong setae along the posterior margin; ST2–4 square with scattered setae; ST5 with

a median deep cleft with almost parallel arms. Terminalia. Syntergosternite 7+8 yellow

brownish with golden microtomentum and scattered short black setulae and 8 marginal

bristles; epandrium yellow brownish with golden microtomentum, one strong dorsoapical seta

and short black setulae; surstylus triangular with enlarged base and sparse marginal black

setulae; cercus, in lateral view, sinuous with pointed apex; postgonite slender, curved apically

and with pointed apex; pregonite broader, slightly curved and rounded apex. Phallus

sclerotized, flower-like shaped, without division between basi- and distiphallus; distiphallic

apex expanded, with lateral and ventral projections, smooth edges and wide opening apically;

vesica sclerotized, flattened, oblong, coming out medially from the phallic tube and with

microspines ventroapically.

Female. Unknown.

33

Etymology. The specific epithet is given in honor of Maria José Maia de Souza,

mother of the first author.

Remarks. See “remarks” section on Oxysarcodexia n. sp. 4.

Distribution. NEOTROPICAL. Ecuador (Napo).

Biology. Unknown.

Material examined. ♂ [male holotype]: ECUADOR: Napo Province: / Yasuní

National Park: / Yasuní Research Station: / 76° 36‟W 00° 38‟S: 3–20 / XI 1998: T. Pape & B.

Viklund // NRM-DIPT / 0014490 (NRM).


(Appendix 6)

Type locality. ECUADOR: Napo Province.



postocular plates with pale gold microtomentum; occiput blackish with silvery



blackish, with row of 10–11 frontal setae; inner vertical seta well-developed, outer vertical

seta 0.43x as long as the inner one and as long as a postocellar seta; ocellar setae equal in size

to uppermost frontals; 1 reclinate fronto-orbital seta 1.6x as long as than the frontals and

proclinate seta absent; gena and postgena blackish with golden and silvery microtomentum,

respectively, and black setae; antenna blackish, first flagellomere with pale golden

microtomentum and about 2.00x longer than pedicel; arista dark brown and long plumose on

basal ¾; palpus dark brown. Thorax. Greyish with pale golden microtomentum slightly more

intense on the sides; chaetotaxy: acrostichals 0+1, dorsocentrals 4 (first and third weaker) + 4

(first and second weaker), intra-alars 2+2, supra-alars 2+3, postalars 2, postpronotals 3,

notopleurals 4 (2 large primaries and 2 smaller subprimaries), katepisternals 3 with middle one

weaker and inserted slightly below the others, meropleurals 7–8, postalar wall setose,

scutellum with 1 apical, 1 subapical, 1 lateral, 1 basal and 1 discal seta; prosternum with few

setae along the edges. Wings. Hyaline, black tegula, R1 bare, R4+5 setulose in proximal ⅔ of

distance to r-m, costal spine not differentiated and third costal sector without ventral setulae.

34

Legs. dark brownish, fore femur with a row of setae on dorsal, posterodorsal and anteroventral

surfaces; fore tibia with 1 dorsal, 2 anterodorsal, 2 posterodorsal and 1 posteroventral setae;

mid femur with 4 median anterior and 2 pre-apical posterior setae, 4 anteroventral and 4

posteroventral long setae, ctenidium of flattened spines apically on posteroventral surface; mid

tibia with 2 posterodorsal, 1 posterior, 1 posteroventral, 1 anteroventral and 1 anterodorsal

setae; hind femur with 1 dorsal, 1 posterior and 7 anterodorsal setae, 1 anteroventral and 1

anterodorsal row of long setae, 1 apical posteroventral row of decreasing setae; hind tibia with

1 anterior, 2 anteroventral, 3 anterodorsal, 1 dorsal and 2 posterodorsal setae; tarsomeres of

fore and hind legs with a weak ventral golden micromentum. Abdomen. Brownish with

golden microtomentum; tergite 2 with silvery microtomentum; T3 with one marginal lateral

seta; T4 with 1 marginal lateral and 1 median marginal seta; T5 with about 24 strong setae

along the posterior margin; ST2–4 square with a pair of strong setae on posterior margin and

scattered setulae; ST5 with a median deep cleft, with edges almost parallel. Terminalia.

Syntergosternite 7+8 yellow brownish with golden microtomentum and scattered short black

setulae and 8 marginal bristles; epandrium yellow brownish with sparse golden

microtomentum and short black setulae; surstylus oblong, elongated ventrodorsally with

sparse marginal and discal setulae and a long and slender colorless setae at apex;

cercusstraight with a darker apical expansion, in lateral view, andwith apex slightly divergent,

in posterior view; postgonite slender, slightly curved and with pointed apex; pregonite

broader, curved and pointed apex. Phallus well sclerotized, without division between basi- and

distiphallus; distiphallic apex curved ventrodorsally in an obtuse angle; juxta bifurcated,

rugous laterally, smooth ventrally and knurled apically; vesica well sclerotized, small, knurled

along the edges and, in anterior view, with short and almost flat triangular branches.

Female. Unknown.

Etymology. The specific epithet is given based on conformation of the vesica and

distiphallus, which is clearly separated and, in lateral view, give the impression of a cleft

presence. The word rima, in Latin, means cleft, slit.

Remarks. Oxysarcodexia n. sp. 6 is also very much alike to Oxysarcodexia admixta

(Lopes, 1933); Oxysarcodexia n. sp. 6 differences are noticed on distiphallus median margin

of serrated aspect; on more angulated distiphallus apical shape and on vesica longer

posteroanteriorly, reaching distiphallus ventroapical margin.

35

Biology. Unknown.

Distribution. NEOTROPICAL. Ecuador (Napo).

Material examined. ♂ [male holotype]: ECUADOR: Napo Province: Yasuní National

Park: Yasuní Research Station: 76° 36‟W 00° 38‟S: 3–20 XI 1998: T. Pape & B. Viklund /

NRM-DIPT 0014488 (NRM).

Oxysarcodexia admixta (Lopes, 1933)

Sarcophaga admixta Lopes, 1933

(Appendix 7-A)

Type locality. Brazil, Rio de Janeiro, Angra dos Reis. Lopes, 1933: 156.


Diagnosis. Male. Postocular plate with silvery microtomentum. Ocellar setae well-

developed. Thorax generally with golden microtomentum, sometimes more intense on

humeral region, contrasting with the silvery microtomentum of the abdomen. Two well

differentiated + 2 smaller dorsocentral post-sutural setae; apical scutellar seta present. Legs

blackish. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral marginal

setae. T5 with golden microtomentum only laterally. ST5 with parallel cleft edges and setosity

on the arms. In lateral view, cerci with straight shape, apex expanded with oblique upwards

edge and presence of setae in full ventral extension. Apical shape of the cerci (i.e. last ⅓

portion of the cercus), in posterior view, smaller than the middle part. Cerci conformation

(posterior view) divergent. In lateral view, ventroapical margin of distiphallus smooth; apical

shape of distiphallus conic; dorsal shape straight. Pregonite with same color (brownish) along

the entire extension and with expanded base narrowing smoothly until the apex, same as

postgonite. Vesica with symmetrical branches; terminal lobes reduced, filamentous at most

tapering to apex, with sclerotized texture; presence of spines only along the edges and

presence of a rounded median projection of the main vesica branch.

Remarks. Cerci and general shape of the phallus are very similar to Oxysarcodexia

carvalhoi Lopes, 1946; differences are seen basically on the apex of distiphallus, without a

ventroapical concavity in O. admixta, and on the vesica, more developed in O. carvalhoi,

although still reduced in comparison to other species of the genus. Oxysarcodexia admixta

also presents similarities on terminalia, especially on distiphallus apex (rounded shape) and on

36

vesica (spinous and with apical triangular apex on terminal lobes), in comparison to

Oxysarcodexia avuncula (Lopes, 1933), Oxysarcodexia berlai Lopes, 1975, O. carvalhoi,

Oxysarcodexia diana (Lopes, 1933) and Oxysarcodexia ventricosa (Wulp, 1895) (species

included in “ventricosa group”, according to Lopes (1975c), that considers also morphology of

females for grouping them). See also “remarks” section of Oxysarcodexia n. sp. 6. Female is

classified by Tibana & Mello (1985), according to the shape of the T6+7, as syntergite

undivided.

Distribution. NEOTROPICAL. Argentina (Misiones), Brazil (Distrito Federal, Goiás,

Mato Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina, São

Paulo).

Biology. It is reported in the literature the attractiveness of O. admixta by human feces,

fish, mouse and pig carcasses, chicken viscera (especially rotting liver), marine animals,

banana plus brown sugar and rotten Syagrus comosa (Mart.) Mart. (Arecaceae), a species of a

palmtree (at coastal area) (Lopes 1973b; Dias et al. 1984c; Mendes & Linhares 1993;

D‟Almeida & Lima 1994; Oliveira et al. 2002; Vairo et al. 2011). This fly has already been

found rearing on human feces and mouse carcass (Lopes 1973b; D‟Almeida 1994), and on

agar plus powder milk for 24h then transferred to meat, under laboratory conditions (Lopes

1973b). At our laboratory, it was reared on minced bovine meat, although larviposition was

observed also on rotten fish, with adults emerging after 15–19 days (6 days from larvae to

pupa stage and 9–13 days until adult emergence). This species was also associated to the gum

of Terminalia argentea Mart & Zucc (Combretaceae), a Brazilian pioneer tree, collecting or,

sometimes, ingesting its exudate, probably due to the high concentration of complex

carbohydrates present in this resource (Boff et al. 2008).

Material examined. ♂♂: BRAZIL: São Paulo, Jundiaí, edge of Serra do Japi;

28.X.2011; A. G. Savino (L2B-DBA) // BRAZIL: São Paulo, Jundiaí, Serra do Japi;

24.I.2012; M. D. Grella (L2B-DBA).

Oxysarcodexia adunca Lopes, 1975

(Appendix 7-B)

Type locality. Brazil, Espírito Santo, Conceição da Barra. Lopes 1975c: 475.


37

Diagnosis. Male. Postocular plate with pale golden microtomentum. Ocellar setae

smaller than the superior frontal seta. Thorax gray, sometimes yellowish gray, especially at

humeral region. Two well differentiated + 2 smaller dorsocentral post-sutural setae; apical

scutellar seta absent. Legs blackish. Abdomen with silvery microtomentum, although golden

microtomentum is observed laterally on T4 and on entire extension of T5. T3 with 1 lateral

marginal seta. T4 with 1 median marginal and 1 lateral marginal setae. T5 with golden

microtomentum along the entire extension. ST5 with parallel cleft edges, presence of setosity

and setae at the apex of the arms. Cerci, in lateral view, sinuous with expanded apexes of

oblique upwards edges and darker than the median and basal portions. Setae ventrally on the

cerci (lateral view) absent only in middle portion. Apical shape of the cerci (last ⅓ portion of

the cercus; posterior view) with the same size as the middle part. Conformation of the cerci in

posterior view is divergent. In lateral view, ventroapical margin of distiphallus smooth;

distiphallic apical shape rounded; dorsal shape straight. Remarkable presence of microspines

at distiphallus apex. Pregonite with expanded base and sudden narrowing at apex which is

darker; same as postgonite, except for the color, which is the same in full extension. Vesica

with symmetrical branches; terminal lobes well-developed, rounded, with membranous

texture, presence of spines only on ventral surface and presence of an angular median

projection of the main vesica branch.

Remarks. This species presents similar cerci as Oxysarcodexia cyaniforceps (Hall,

1933), but differs by the apical distiphallus shape, pilose in O. adunca, and by the constitution

of the vesica (Lopes 1975c).

Distribution. NEOTROPICAL. Brazil (Bahia, Espírito Santo, Rio de Janeiro),

Ecuador (Napo Province).

Biology. Unknown.

Material examined. ♂♂: Linhares, Espírito Santo, Brasil / P. C. Elias; VI-72 /

paratypus / Oxysarcodexia adunca; n. sp.; ♂; Det. H. S. Lopes / NRM-DIPT 0014213 (NRM)

// ECUADOR: Napo Province, Yasuní National Park, Yassuní Research Station, 76°36‟W

00°38‟S; 3–20.XI.1998; T. Pape & B. Viklund / NRM-DIPT 0014473 (NRM).

Oxysarcodexia afficta (Wulp, 1895)

Sarcophaga afficta Wulp, 1895

38

(Appendix 7-C)

Type locality. Mexico, Morelos, Cuernavaca; Mexico, Veracruz, Atoyac; México,

Yucatán. Wulp, 1895: 269.

Depository of type material. BMNH.

Diagnosis. Male. Postocular plate with pale golden microtomentum. Ocellar setae not

well-developed. Thorax generally with grayish microtomentum. Dorsocentral post-sutural

setae with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta absent.

Legs blackish. Abdomen grayish with shades of pale golden microtomentum laterally. T3 with

2 lateral marginal setae. T4 with 2 median marginal and 2 lateral marginal setae. T5 with

golden microtomentum only laterally. ST5 with parallel cleft edges and setosity on the arms.

Cerci, in lateral view, sinuous with expanded apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) present in basal half. Apical shape of the cerci (last ⅓ portion of the

cercus; posterior view) with the same size as the middle part. Conformation of the cerci in

posterior view is divergent. In lateral view, ventroapical margin of distiphallus smooth;

distiphallic apical shape conic; dorsal shape of distiphallus straight, and presence of a small

dorsoapical swelling. Pregonite with expanded base narrowing smoothly until the apex, which

is darker; same as postgonite, except for the color, which is the same in full extension. Vesica

with symmetrical branches; terminal lobes well-developed, filamentous at most tapering to

apex, with sclerotized texture and presence of spines only on ventral surface.

Remarks. Oxysarcodexia afficta is considered close to Oxysarcodexia conclausa

(Walker, 1861), Oxysarcodexia thornax (Walker, 1849) and Oxysarcodexia timida (Aldrich,

1916), however easily separated by the unalike vesica shape (Lopes 1946b). Female was


partially divided into two plates.

Distribution. NEARTIC. Mexico (Morelos, San Luis Potosí). NEOTROPICAL.

Colombia, Costa Rica, Ecuador, Mexico (Chiapas, Jalisco, Sinaloa, Veracruz, Yucatán),

Panama.

Biology. Unknown. Information available is only about the collection of the holotype,

using traps for collecting fruit flies (Lopes 1946b).

39

Material examined. ♂♂: N. Perucho; (Otavalo) Ecuador; 200m. I-1971; L. E. Peña

col. / Oxysarcodexia afficta (Wulp); ♂; Det. H. S. Lopes / NRM-DIPT 0014214 (NRM) //

Mexico, VU94; Chiapa de Corzo; COMEXA, 09.11.2010; A. Grzywack, leg (ZMUC).

Oxysarcodexia amorosa (Schiner, 1868)

Sarcophaga amorosa Schiner, 1868

(Appendix 8-A)

Type locality. Brazil. Schiner, 1868: 314.

Depository of type material. NMW.


well-developed. Thorax with golden microtomentum, as well as the abdomen, which is more

intense on T5. Dorsocentral post-sutural setae with 2 well differentiated and 1–3 anterior setae

smaller. Apical scutellar seta present. Legs brownish. T3 with 3 lateral marginal setae. T4 with

1 median marginal and 2 lateral marginal setae. T5 with golden microtomentum in part. ST5

with parallel cleft edges and setosity on the arms. Cerci, in lateral view, straight with expanded

apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view) in full extension.

Apical shape of the cerci (last ⅓ portion of the cercus; posterior view) smaller than the middle

part. Conformation of the cerci in posterior view is divergent. In lateral view, distiphallus

presents ventroapical concavity, with smooth margins; distiphallic apical shape rounded; and

dorsal shape of distiphallus straight. Pregonite with the same color along the entire extension

and with expanded base narrowing smoothly until the apex. Postgonite with expanded base

and sudden narrowing at apex. Vesica with symmetrical branches; terminal lobes well-

developed, filamentous at most tapering to apex, with sclerotized texture, spines on ventral

surface, presence of lateral lobes (i.e. division of the vesica coming from or close to the basal

branch of the vesica, placed laterally to phallic tube) and presence of a rounded median


Remarks. Phenotype variability in different male populations has already been

reported in literature (Lopes 1973b). Oxysarcodexia inflata Lopes, 1975 is considered very

close related to O. amorosa, especially by the color, but the differences are recognized on

distiphallus apex and on vesica conformation (Lopes 1975c). Differences on O. similata

terminalia, in comparison to O. amorosa, are the presence of a slender anteroventral projection

40

directed apically (lateral view) and the large and less oblique internal margins of apical plates

(ventral view) on the distiphallus, and, on vesica constitution, the enlarged and rounded basal

portion, with the presence of spines. Apical plates of O. xanthosoma differs from the

previously mentioned by the serrated edge (Lopes & Tibana 1987). See “remarks” section of

Oxysarcodexia n. sp. 1 for further discussion. Female was classified by Tibana & Mello

(1985), according to the shape of the T6+7, as syntergite partially divided into two plates.

Distribution. NEARTIC. Mexico (San Luis Potosí, Sonora). NEOTROPICAL. Brazil

(Amapá, Amazonas, Bahia, Ceará, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Rio de

Janeiro, Roraima, Santa Catarina, São Paulo), Colombia (Antioquia), Costa Rica, Ecuador,

Guyana, Mexico (Jalisco), Panama (Barro Colorado Island), Peru (Tingo Maria).

Biology. Oxysarcodexia amorosa has been reared on feces of different animals as dog,

felid and human (Lopes 1973b; D‟Almeida 1994), on dead mammal not specified (Dodge

1968) and on fish head and bones (Lopes 1973b). This species has already been reared

successfully at laboratory (Lopes 1973b) and on dead shrimp under non-natural conditions

(D‟Almeida 1989). Using curdled milk as food resource, larvae reared in 3 days and the adults

emerged after 12 days (Lopes 1973b). This fly has shown developmental preference by low

humidity, being found near to dwellings (Lopes 1973b). Attractiveness has been reported to

human and other vertebrates feces, fish, crab, marine animals, rotten Syagrus comosa (Mart.)

Mart. (Arecaceae), a species of a palmtree and rotten banana plus brown sugar (at coastal

areas), chicken viscera, rat carcass, rotting liver, rotting beef lung and squid (Lopes 1973b;

Lopes 1975a; Mendes & Linhares 1993; Pamplona et al. 2000; Oliveira et al. 2002; Sousa et

al. 2011; Ramírez-Mora et al. 2012). Larva of O. amorosa has also been reported involved in

a case of auricular infestation in São João de Meriti, Rio de Janeiro (Figueiredo et al. 2002),

although this species is not recognized as a causing-myiasis agent.

Material examined. ♂♂: Angra dos Reis; E. do Rio; Brasil / Det. H. S. Lopes;

10.10.72 / NRMDIPT 0014218 (NRM) // Angra dos Reis, E. do Rio, Brasil / H. S. Lopes

6.X.71 / Oxysarcodexia amorosa (Schiner) Det. H. S. Lopes ♂.

Oxysarcodexia angrensis (Lopes, 1933)

Sarcophaga angrensis Lopes, 1933

Sarcophaga articulata Hall, 1933

41

Sarcophaga kartabo Curran & Walley, 1934

(Appendix 8-B)



Diagnosis. Male. Postocular plate with golden microtomentum. Ocellar setae well-

developed. Thorax grayish with pale golden microtomentum, as well as abdomen in which

golden microtomentum is more evident laterally. Dorsocentral post-sutural setae with 2 well

differentiated and 1–3 anterior setae smaller. Apical scutellar seta absent. Legs blackish. T3

with 3 lateral marginal setae. T4 with 1 median marginal and 2 lateral marginal setae. Golden

microtomentum at entire extension of T5. ST5 with parallel cleft edges with setosity and setae

on the apex of the arms. Cerci, in lateral view, sinuous with pointed apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) absent only in middle portion.

Apical shape of the cerci (last ⅓ portion of the cercus; posterior view) with the same size as

the middle part. Conformation of the cerci in posterior view is parallel. In lateral view,

presence of ventroapical concavity on the distiphallus with smooth margins; distiphallic apical

shape conic; dorsal shape of distiphallus sinuous, and presence of a small dorsoapical

swelling. Pregonite with expanded base narrowing smoothly until the apex, which is darker;

same as postgonite, except for the color, which is the same in full extension. Vesica with

symmetrical branches; terminal lobes well-developed, filamentous at most tapering to apex,

with sclerotized texture, presence of spines on both ventral and dorsal surfaces and presence of

an angular median projection of the main vesica branch.

Remarks. See “remarks” section of Oxysarcodexia n. sp. 2. Female was classified by

Tibana & Mello (1985), according to the shape of the T6+7, as syntergite partially divided into

two plates. Description of morphological structures of larvae from 1st, 2

nd and 3

rd instars has

been provided by Lopes (1943).

Distribution. NEOTROPICAL. Brazil (Amapá, Amazonas, Goiás, Mato Grosso, Mato

Grosso do Sul, Minas Gerais, Pará, Rio de Janeiro, São Paulo), Colombia (Antioquia), Costa

Rica, Ecuador, Guyana, Panama, Peru, Roraima, Trinidad and Tobago (Trinidad), Venezuela.

Biology. In addition to human feces, attraction of adults of O. angrensis have been

reported by fish, mouse and rat carcasses, marine animals, rotting beef lung, chicken viscera,

and, at costal area, for crabs, rotten Syagrus comosa (Mart.) Mart. (Arecaceae), a species of a

42

palmtree, and rotten banana plus brown sugar (Lopes 1973b; Moretti et al. 2008; Sousa et al.

2011; Ramírez-Mora et al. 2012). No differences in attractiveness of this species by human

feces, chicken viscera and mouse carcasses baits were observed by Linhares (1981). Human

feces have seen being used as a natural rearing substrate (Lopes 1973b; D‟Almeida 1994).

Agar plus powder milk and another medium with a kind of jelly called “gelose” plus horse

blood serum and commercial yeast have been used successfully as artificial rearing sources for

the establishment and maintenance of colonies in laboratory. The developing from 1st instar to

adults was reached in 14–17 days using the first medium, whereas with the last medium

occurred an increase of the larval length in comparison to larvae reared in the same medium

without the yeast (Lopes 1973b). Meat has also been used successfully for rearing O.

angrensis (Lopes 1943). Adults have been collected, besides actively, with malaise and

Shannon traps (Lopes & Tibana 1991) and also associated to cadavers (Oliveira-Costa et al.

2001). Sunlight instead of shaded areas is also referred as a preference of this species

(Linhares 1981). Lopes (1973b) pointed out O. angrensis preference by high humidity based

on self-experience in flesh flies collection done in Brazil, in different periods. However,

Yepes-Gaurisas et al. (2013), collecting flies in a forest area of Antioquia, Colombia, observed

higher abundance of this species in dry season, with strong attraction for fish and also

considering this species asynanthropic.

Material examined. ♂♂: BRAZIL: São Paulo, Mogi Guaçu, Campininha, 18.IX.2011;

C. G. P. Lima, M. D. Grella, N. M. Jimenez / Oxysarcodexia VIII; Mogi Guaçu-SP;

18/11/2011 (L2B-DBA) // BRAZIL: São Paulo, Campinas, UNICAMP, 30.XI.2011; C. M.

Souza / Oxysarcodexia sp.; Campinas-SP; 30/11/2011; 12 (L2B-DBA).

Oxysarcodexia augusta Lopes, 1946

(Appendix 8-C)

Type locality. Brazil, Rio de Janeiro, Guanabara, Méier. Lopes, 1946b: 84.

Depository of type material. Unknown.

Diagnosis. Male. Postocular plate with golden microtomentum. Ocellar setae not well-

developed. Thorax with golden microtomentum, more intense at humeral region. Three well-

differentiated dorsocentral post-sutural setae; one smaller can be present among these others.

Apical scutellar seta present. Legs blackish. Abdomen grayish with silvery microtomentum.

43

T3 with 2 lateral marginal setae. T4 with 1 median marginal and two lateral marginal setae. T5

with silvery microtomentum only. ST5 with parallel cleft edges and with setosity and setae on

the apex of the arms. Cerci, in lateral view, straight with expanded apexes of oblique upwards

edges. Setae ventrally on the cerci (lateral view) present in full extension. Apical shape of the

cerci (last ⅓ portion of the cercus; posterior view) smaller than the middle part. Conformation

of the cerci in posterior view is divergent. In lateral view, presence of ventroapical concavity

on distiphallus with serrated margins; presence of lateroapical furrow; distiphallic apical shape

rounded; sinuous distiphallic dorsal shape. Presence of lateroapical expansions, in dorsal view,

on the distiphallus. Pregonite with expanded base narrowing smoothly until the apex, which

presents same color along the entire extension. Postgonite with expanded base and sudden

narrowing at apex. Vesica with symmetrical branches; terminal lobes well-developed,

filamentous at most tapering to apex, with sclerotized texture; presence of spines only on

ventral surface; and presence of a rounded median projection of the main vesica branch.

Remarks. Three dorsocentral post-sutural setae are seen in O. augusta, Oxysarcodexia

bikini Dodge, 1966, Oxysarcodexia corolla Dodge, 1965, Oxysarcodexia flavipes Lopes &

Tibana, 1987, Oxysarcodexia grandis Lopes, 1946, Oxysarcodexia jamesi Dodge, 1968, O.

nitida, O. notata, O. peruviana, Oxysarcodexia plebeja Lopes, 1946, Oxysarcodexia

terminalis (Hall, 1937), Oxysarcodexia varia (Walker, 1836), and O. vittata. Female was


undivided.

Distribution. NEOTROPICAL. Argentina (Misiones), Brazil (Minas Gerais, Rio de

Janeiro, Roraima, Santa Catarina, São Paulo).

Biology. This species has already been reared under laboratory conditions (Lopes

1973b). However, no further information about its development could be found. Attractiveness

has been reported to human feces, chicken viscera, fish, mouse carcass, marine animals,


palmtree (at coastal area) (Lopes 1969; Lopes 1973b; Mendes & Linhares 1993). Malaise and

Shannon traps have been used to collect this species (Lopes & Tibana 1991), as well as active

collections. At Rio de Janeiro Zoological Garden, this fly had been collected using a wind

oriented trap (W.O.T.), baited with rotting liver (Oliveira et al. 2002).

44

Material examined. ♂: BRAZIL: Minas Gerais, Belo Horizonte, Est. Ecológica,

UFMG Campus; 2–22.VIII.1993; S. D. Gaiman / NRM-DIPT 0014231 (NRM).

Oxysarcodexia aura (Hall, 1937)

Sarcophaga aura Hall, 1937

(Appendix 9-A)

Type locality. Bolivia, northern Chiquitos. Hall, 1937: 372.

Depository of type material. DEI.

Diagnosis. Male. Postocular plate with silveryish microtomentum. Ocellar setae well-

developed. Thorax with golden microtomentum, more intense at humeral region. Dorsocentral

post-sutural setae with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar

seta absent. Legs brownish. Abdomen darkish brown with golden microtomentum. T3 with 3

lateral marginal setae. T4 with 1 (weak) median marginal and 2 lateral marginal setae. T5 with

golden microtomentum laterally. ST5 with V-shaped cleft edges and with setae along the

edges of the arms. Cerci, in lateral view, straight with pointed apexes of concave edges.

Presence of setae ventrally on the cerci (lateral view) only in basal half. Apical shape of the

cerci (last ⅓ portion of the cercus; posterior view) with the same size as the middle part.

Conformation of the cerci in posterior view is divergent. In lateral view, presence of

ventroapical concavity on distiphallus with smooth margins; distiphallic apical shape rounded;

and straight distiphallic dorsal shape. Pregonite with base and apex with same size and color.

Postgonite with expanded base narrowing smoothly until the apex. Vesica with symmetrical

branches; terminal lobes well-developed, with rounded shape and sclerotized texture; without

any spines.

Remarks. Initially it has been identified as Amesothyrsus chilensis, Enderlein, 1928,

revised as Sarcophaga sp. Engel, 1931 and then identified, by the proposition of a new name,

as Sarcophaga aura Hall, 1937. This species is easily recognized by the very peculiar cerci,

straight in lateral view and very slender and spaced in posterior view; by the distiphallus with

a singular conformation; and also by the vesica shape.

Distribution. NEOTROPICAL. Bolivia, Brazil (Distrito Federal, Mato Grosso, Mato

Grosso do Sul, Minas Gerais).

45

Biology. Species attracted to chicken viscera (Lopes 1980) and pig carcasses exposed

in areas of Brazilian Cerrado (“savanna-like” vegetation) (Barros et al. 2008; Rosa et al.

2011).

Material examined. ♂♂: Campinas, Est. De Goyaz, Borgmeier et Lopes 14-XII-35 /

Sarcophaga aura ♂, Hall, 8.5.7., Det. H. S. Lopes / NRM-DIPT 0014232 (NRM) // Brasil:

MG Pirapora 20–29.xii.1978 C. B. Carvalho col. / Hybopygia aura (Hall) Det. H. S. Lopes

(MNRJ) // O. aura Faz. Do Glória/10 [from Uberlândia, Minas Gerais state, Brazil] (L2B-

DBA); O. aura Faz. Do Glória/10 [from Uberlândia, Minas Gerais state, Brazil] (L2B-DBA).

Oxysarcodexia aurata (Macquart, 1851)

Sarcophaga aurata Macquart, 1851

Sarcophaga taitensis Schiner, 1868

Sarcophaga obtusifrons Thomson, 1869

Sarcophaga vesica Hall, 1933

Dasyproctia auricauda Enderlein, 1928

Oxysarcodexia lapitana Lehrer & Barbet, 2008 [not available name]

(Appendix 9-B)

Type locality. Oceania, probably Society Islands, Tahiti [“Océanie”]. Macquart, 1851:

207.

Depository of type material. MNHN.


developed. Thorax and abdomen with silvery microtomentum. Dorsocentral post-sutural setae

with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta absent. Legs

blackish. T3 with 3 lateral marginal setae. T4 with 1 median marginal and 3 lateral marginal

setae. T5 with golden microtomentum in part. ST5 with parallel cleft edges and with setae on

the apex of the arms. Cerci, in lateral view, sinuous with normal apexes (same size as median

area) of concave edges. Setae ventrally on the cerci (lateral view) present at apical ⅓ portion.

Apical shape of the cerci (last ⅓ portion of the cercus; posterior view) with the same size as

the middle part. Conformation of the cerci in posterior view is divergent. In lateral view,

distiphallus ventroapically with smooth margins; distiphallic apical shape rounded and straight

distiphallic dorsal shape. Presence of lateral lobes on distiphallus and of a large distiphallic

46

dorsoapical swelling. Pregonite with base and apex with the same size and apex darker than

the base. Postgonite with expanded base narrowing smoothly until the apex. Vesica with

symmetrical branches; terminal lobes well-developed, filamentous at most tapering to apex,

with partially membranous texture; presence of spines only on ventral surface; and presence of


Remarks. Pape (2008), after revising nominal species of Sarcophagidae described by

Macquart from the Australasian/Oceanian region, proposed O. taitensis as a junior synonym of

O. aurata, although the first name have been widely used in several manuscripts.

Oxysarcodexia lapitana Lehrer & Barbet, 2008, a relatively recent name proposed, is indeed

O. aurata. In spite of material used for the proposition of the name O. lapitana has not seen,

the Lehrer & Barbet‟s illustration (Lehrer & Barbet 2008, fig. 1) provided very clear and

detailed elements enabling comparison between species and the statement of this synonym. It

is also important to stress O. lapitana wasn‟t correctly proposed, once all the International

Code of Zoological Nomenclature requirements weren‟t fulfil and, therefore, it is considered

not available. Oxysarcodexia aurata presents a large membranous dorsoapical swelling on

distiphallus as observed on Oxysarcodexia eberti Lopes & Tibana, 1987, Oxysarcodexia

galeata (Aldrich, 1916), Oxysarcodexia intona (Curran & Walley, 1934), Oxysarcodexia

peltata (Aldrich, 1916) and Oxysarcodexia varia (Walker, 1836). Cerci shape (lateral view)

and the large membranous dorsoapical swelling on the distiphallus are similar characters seen

in O. aurata and O. peltata. This species was included, besides Oxysarcodexia culminata

(Aldrich, 1916), Oxysarcodexia culmiforceps Dodge, 1966, Oxysarcodexia fringidea (Curran

& Walley, 1934), O. intona and O. peltata, in “peltata group”, e.g., possesses the genital

tergite of females reduced, membranous and with 2 spiracles, sternites 6+7 and 8 narrower and

the tergite 5 with lateral edges touching themselves ventrally (Lopes 1975c; Tibana & Mello

1983a). Female was classified by Tibana & Mello (1985), according to the shape of the T6+7,

as syntergite membranous.

Distribution. NEOTROPICAL. Chile (Cauquenes), Colombia (Antioquia), Costa

Rica, Ecuador, Galápagos Islands (Albermale, Charles, Chatham, Duncan, Indefatigable,

Narborough, San Salvador), Panama (Barro Colorado Island), Peru.

AUSTRALASIAN/OCEANIAN. Fiji, French Polynesia (Austral Islands, Marquesas Islands,

Society Islands, Tubuai Islands), Tahiti, Tonga, Western Samoa.

47

Biology. Attraction of this species is reported by chicken viscera and fish (Ramírez-

Mora et al. 2012; Yepes-Gaurisas et al. 2013). In Antioquia, Colombia, Yepes-Gaurisas et al.

(2013) observed a frequency increase of O. aurata with the rain and a close relation with

human settlements due to the high synanthropic index detected.

Material examined. ♂♂: NEW CALEDONIA: Province Nord. Poindimié, near the

coast, 24–28.xi.2001, Johanson, Pape & Viklund / NRM-DIPT 0014339 (NRM) // Society Is.,

Tahiti Punaaina, 4.1.1978, N. H. L. Krauss leg. (ZMUC).

Oxysarcodexia avuncula (Lopes, 1933)

Sarcophaga avuncula Lopes, 1933

Sarcophaga avunculus Blanchard (1935), incorrect subsequent spelling of avuncula

Lopes, 1933

(Appendix 9-C)

Type locality. Brasil, Rio de Janeiro, Rio de Janeiro, Manguinhos. Lopes, 1933: 156.



developed. Thorax with pale golden microtomentum, more evident at humeral portion.

Dorsocentral post-sutural setae with 2 well differentiated and 1–3 anterior setae smaller.

Apical scutellar seta present. Legs blackish. Abdomen grayish with golden microtomentum

more intense at the lateral portion of T3 and full extension of T4. T3 with 3 lateral marginal

setae. T4 with 1 median marginal and 2 lateral marginal setae. T5 with silvery microtomentum

only. ST5 with parallel cleft edges and with setosity and setae on the apex of the arms. Cerci,

in lateral view, sinuous with pointed apexes of concave edges. Setae ventrally on the cerci

(lateral view) present at apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the

cercus; posterior view) with the same size as the middle part. Conformation of the cerci in

posterior view is divergent. In lateral view, presence of ventroapical concavity on distiphallus

with smooth margins; distiphallic apical shape rounded; straight distiphallic dorsal shape.

Pregonite with expanded base narrowing smoothly until the apex as well as postgonite, except

the color which is darker in the apex on the pregonite. Vesica with symmetrical branches;

terminal lobes well-developed, with rounded shape, partially membranous texture; presence of

48

spines only along the edges; and presence of an angular median projection of the main vesica

branch.

Remarks. See “remarks” section of O. admixta. A good and detailed male terminalia

comparison of the sympatric species not unusually misidentified, O. avuncula, O. confusa, O.

diana and O. parva, is given by Silva & Mello-Patiu (2008). Female was classified by Tibana

& Mello (1985), according to the shape of the T6+7, as syntergite undivided. Larvae are

characterized by the presence of sinuous ribbons of festoon on psedocephalon and

conspicuous teeth on maxillae (Leite & Lopes 1987; Lopes & Leite 1986; 1987).

Distribution. NEOTROPICAL. Argentina (Misiones), Bolivia, Brazil (Ceará, Distrito

Federal, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerias, Rio de Janeiro, Santa

Catarina, São Paulo, Pernambuco), Colombia (Antioquia), Ecuador, Mexico (Oxaca),

Paraguay, Peru, Trinidad and Tobago (Trinidad).

Biology. It has been reared on cattle dung (Mendes & Linhares 2002). This species

also presents attraction for human feces, chicken viscera, fish, mouse and pig carcasses,

marine animals and banana plus brown sugar (Lopes 1973b; Lopes 1975a; Dias et al. 1984c;

Mendes & Linhares 1993; Ramírez-Mora et al. 2012; Yepes-Gaurisas et al. 2013). It has been

collected in urban (collection made by the authors) and forest environments (Vasconcelos &

Araújo 2012). This species was also associated to the gum of a Brazilian pioneer tree,

Terminalia argentea Mart & Zucc (Combretaceae), collecting or, sometimes, ingesting its

exudate, probably due to the high concentration of complex carbohydrates present in this

resource (Boff et al. 2008).

Material examined. ♂♂: BRAZIL: São Paulo, Campinas, Sousas, 13.IV.2011; C. M.

Souza, D. L. Brancoli, F. Rezende / O. avuncula (L2B-DBA) // BRAZIL: Minas Gerais,

Extrema, Serra do Lopo, Pedra das Flores, 27.II.2012; A. G. Savino, M. P. Nassu, M. D.

Grella / Oxysarcodexia B, Extrema-MG; 27/02/2012; 3 (L2B-DBA).

Oxysarcodexia bakeri (Aldrich, 1916)

Sarcophaga bakeri Aldrich, 1916

(Appendix 10-A)

Type locality. Cuba, Habana. Aldrich, 1916: 270.

Depository of type material. USNM.

49


developed. Thorax with golden microtomentum. Dorsocentral post-sutural setae with 2 well

differentiated and 1–3 anterior setae smaller. Apical scutellar seta present. Legs blackish.

Abdomen grayish with golden microtomentum. T4 with 2 median marginal setae. T5 with

golden microtomentum. ST5 with parallel cleft edges and with setosity. Cerci, in lateral view,

straight with pointed apexes of oblique upwards edges. Setae ventrally on the cerci (lateral

view) present in full extension. Apical shape of the cerci (last ⅓ portion of the cercus; in

posterior view) with the same size as the middle part. Divergent conformation of the cerci in

posterior view. In lateral view, distiphallus with serrated margins ventroapically; distiphallic

apical shape conic; straight distiphallic dorsal shape and presence of ventroapical projections.

Presence of a remarkable lighter area dorsoapically on the distiphallus. Pregonite with

expanded base and sudden narrowing at apex, which is darker than the base. Postgonite with

expanded base narrowing smoothly until the apex. Vesica with symmetrical branches; terminal

lobes well-developed, filamentous at most tapering to apex, with sclerotized texture; presence

of spines only along the edges; and presence of a rounded median projection of the main

vesica branch.

Remarks. This species is considered similar to Oxysarcodexia modesta Lopes, 1946,

diverging from this by the presence of apical scutellar setae, setosity on syntergosternite 7+8

more abundant and peculiarities on the distiphallus (Lopes 1946b). Female was classified by


two plates. This species has been intercepted in imported container with cargoes from

Americas in Shanghai Port, China (Deng et al. 2011).

Distribution. NEARTIC. Mexico (Baja California Sur, Sonora, Zacatecas), USA

(Texas). NEOTROPICAL. Bahamas (New Providence), Brazil (Bahia, Distrito Federal, Goiás,

Mato Grosso, Pernambuco, Roraima), Chile (Tarapacá), Colombia (Antioquia), Cuba,

Dominica, Ecuador, El Salvador, Guadalupe, Haiti, Jamaica, Mexico (Chiapas, Guerrero,

Sinaloa, Tabasco, Yucatán), Peru, Puerto Rico, Turks & Caicos Island. PALEARTIC. China

(Shangai).

Biology. This species has already been reared at laboratory, using agar plus powder

milk, developing from 1st instar to adults in 15–17 days (Lopes 1973b). Attractiveness has

been reported by human feces, fermented fruits (grapes), fish, chicken viscera, and liver

50

(Flores & Dale 1995; Ramírez-Mora et al. 2012). Oxysarcodexia bakeri was collected on a

residential complex of Valle de Aburrá, surround of Medellín, Colombia (Salazar-Ortega et al.

2012), pointing a possibly relationship of this species with human settlements, which was

corroborated by a high synanthropic index found by Yepes-Gaurisas et al. (2013) in a

collection of this fly at another urban area of Medellín.

Material examined. ♂♂: Oxysarcodexia bakeri sp 24 / 7CP2 [from Colombia] (CE-

TdeA) // Oxysarcodexia bakeri sp 20 / sp20 TdeA 846 [from Colombia] (CE-TdeA) // Cuba,

Ancón Nw de Viñales Pirnasdel Rio J. Holmsn 26.VIII.65 / Oxysarcodexia bakeri Aldr. B.

Rohdendorf det. 1970. VI / bakeri Cuba (MNRJ).

Oxysarcodexia berlai Lopes, 1975

(Appendix 10-B)

Type locality. Brazil, Pará, Belém, Utinga. Lopes 1975c: 473.



well-developed. Thorax with golden microtomentum, more intense at humeral region.


Apical scutellar seta present. Legs brownish. Abdomen grayish with silvery microtomentum.

T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral marginal setae. T5

with silvery microtomentum. ST5 with parallel cleft edges and with setosity and setae on the

apical half of the arms. Cerci, in lateral view, straight with expanded apexes of straight edges.

Setae ventrally on the cerci (lateral view) absent only in middle portion. Apical shape of the

cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.

Conformation of the cerci in posterior view is divergent. In lateral view, presence of

ventroapical concavity on distiphallus with serrated margins; distiphallic apical shape

rounded; sinuous distiphallic dorsal shape. In dorsal view, presence of lateroapical expansions

on the distiphallus. Pregonite with expanded base and sudden narrowing at apex, which

presents same color along the entire extension. Postgonite with expanded base narrowing

smoothly until the apex. Vesica with symmetrical branches; terminal lobes well-developed,

filamentous at most tapering to apex, with sclerotized texture; presence of spines only on


51

Remarks. The main difference between O. berlai and Oxysarcodexia cuernavaca

Dodge, 1966 is the cerci conformation, parallel in first species and divergent in the second one

(Lopes 1975c) and genitalia shape (Dodge 1966). See “remarks” section of Oxysarcodexia n.

sp. 1 for further discussion.

Distribution. NEOTROPICAL. Brazil (Maranhão, Pará, Pernambuco), Colombia.

Biology. Unknown.

Material examined. ♂♂: Avispas, Madre de Dios, PERU, 10–20.IX.1962; L. Pena

400m / Oxysarc. Berlai; ♂; Lopes; Det. H. S. Lopes / NRM-DIPT 0014262 (NRM) // Igarapé

PARAQUEÚ Rosário, MARANHÃO BRASIL / Berla / 20/22-XI-70 / Paratype /

Oxysarcodexia berlai ♂ n.sp. Det. H. S. Lopes (MNRJ).

Oxysarcodexia bicolor Lopes, 1946

(Appendix 10-C)

Type locality. Brazil, Rio de Janeiro, Itatiaia. Lopes, 1946b: 127.



developed. Thorax with golden microtomentum, contrasting with the silveryish

microtomentum of the abdomen. Dorsocentral post-sutural setae with 2 well differentiated and

1–3 anterior setae smaller. Apical scutellar seta absent. Legs brownish. T3 with 1 lateral

marginal seta. T4 with 1 median marginal and 2 lateral marginal setae. T5 without golden

microtomentum. ST5 with parallel cleft edges and with setosity. Cerci, in lateral view, sinuous

with normal (i.e. same size as median area) apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) absent only in middle portion. Apical shape of the cerci (last ⅓

portion of the cercus; posterior view) smaller than the middle part. Conformation of the cerci

in posterior view is parallel. Presence of a remarkable constriction at middle portion of the

cerci, in posterior view. In lateral view, distiphallus with smooth margins ventroapically;

distiphallic apical shape rounded and sinuous distiphallic dorsal shape. Presence of a large

dorsoapical swelling on the distiphallus, in lateral view. Lateroapical expansions on

distiphallus present, in dorsal view. Pregonite with same color along the entire extension and

expanded base narrowing smoothly until the apex. Postgonite with expanded base and sudden

narrowing at apex. Vesica with symmetrical branches; terminal lobes reduced, with rounded

52

shape and membranous texture; presence of spines on both ventral and dorsal surfaces; and

presence of a rounded median projection of the main vesica branch.

Remarks. The unusual cerci shape (in lateral view) draws the attention in this species.

Female was classified by Tibana & Mello (1985), according to the shape of the T6+7, as

syntergite undivided.

Distribution. NEOTROPICAL. Argentina (Buenos Aires), Brazil (Rio de Janeiro, São

Paulo).

Biology. This species has already been reared on laboratory (Lopes 1973b), although

no further information about its development is available on the literature. It is attracted by

human and dog feces and rotten cow liver (Lopes 1973b; Mulieri et al. 2008; Mulieri et al.

2010; Mulieri et al. 2011). This fly has been collected in woodland of Buenos Aires coastline,

Argentina (Mulieri et al. 2008) and in suburban and rural areas of Almirante Brown district,

near to Buenos Aires, pointing a possibly level of asynanthropy. The occurrence of this

species was higher during warmer months (Mulieri et al. 2011). Oxysarcodexia bicolor is also

considered a flower visitor of Scutia buxifolia Reissek, shrubs of Rhamnaceae family (Mulieri

et al. 2010).

Material examined. ♂♂: Alto da Mosela, Petrópolis, 1200m, E. do Rio, Brasil / H. S.

Lopes, 20.XII.70 / NMR-DIPT 0014265 (NRM) // Cult 878 / Petrópolis, Tq. E. do Rio, Brasil

H. S. Lopes 5/♀ / Oxysarcodexia bicolor Lopes Det. H. S. Lopes (MNRJ) // Campos do

Jordão / Est. São Paulo [Brasil] 1600m 3-1945 Wygodzinsky leg. / Oxysarcodexia bicolor

Lop. Det. H. S. Lopes (MNRJ).

Oxysarcodexia bikini Dodge, 1966

(Appendix 11-A)

Type locality. Chile. Dodge, 1966: 685.

Depository of type material. UCC.

Diagnosis. Male. Postocular plate with silvery microtomentum. Ocellar setae not well-

developed. Thorax and abdomen with silvery microtomentum. Three well-differentiated

dorsocentral post-sutural setae; one smaller can be present among them. Apical scutellar seta

present. Legs blackish. T3 with 2 lateral marginal seta. T4 with 1 median marginal and 3

lateral marginal setae. ST5 reddish yellow with short arms. Cerci straight with expanded

53

apexes of oblique upwards edges, in lateral view. Setae ventrally on the cerci (lateral view)

absent only in middle portion. Apical shape of the cerci (last ⅓ portion of the cercus; in

posterior view) smaller than the middle part. Conformation of the cerci is parallel in posterior

view. Presence of a remarkable constriction at middle portion of the cerci, also in posterior

view. In lateral view, distiphallus with ventroapical concavity with serrated margins;

distiphallic apical shape rounded and sinuous distiphallic dorsal shape. Pregonite and

postgonite with expanded base narrowing smoothly until the apex and same color along the

entire extension. Vesica symmetric; terminal lobes reduced, with rounded shape, sclerotized

texture and presence of spines. This diagnosis is based on original description given by Dodge

(1966), on diagnosis provided by Lopes (1973a), and on the examination of one specimen (as

specified below) presenting some conservation issues (“oily” aspect compromising original

microtomentum and color, T5 partially broke and terminalia on glycerin lacking ST5).

Remarks. This species was included by Lopes (1973a) in “paulistanensis group” due

to the similarities, as anterior view of the vesica and distiphallus apex, shared with

Oxysarcodexia injuncta (Walker, 1857), Oxysarcodexia paulistanensis Prado & Fonseca, 1932

and Oxysarcodexia riograndensis Lopes, 1946 species. Features as last genital sternite small

and enclosed in the hind margin of ST8, genital tergite entire and a broad membranous region

between the genital and anal segments are ascribed for O. bikini female (Dodge 1966; Lopes

1973a). See also “remarks” section on O. augusta for further discussion.

Distribution. NEOTROPICAL. Chile (La Araucanía).

Biology. Unknown.

Material examined. ♂: El Camelo, Santiago, CHILE 29-XI-1954 L. E. Peña /

Oxysarc. bikini ♂ Dodge Det. H. S. Lopes (MNRJ).

Oxysarcodexia carvalhoi Lopes, 1946

(Appendix 11-B)

Type locality. Brazil, Minas Gerais, Cordisburgo. Lopes, 1946b: 92.



developed. Thorax with golden microtomentum, more intense laterally at humeral region, as

well as the abdomen, in which is more evident laterally on T4, and on entire extension of T5.

54


Apical scutellar seta present. Legs blackish. T3 with 2 lateral marginal seta. T4 with 1 median

marginal and 2 lateral marginal setae. T5 with golden microtomentum. ST5 with parallel cleft

edges and with setosity. Cerci, in lateral view, sinuous with expanded apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) present on apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.

Conformation of the cerci in posterior view is divergent. In lateral view, distiphallus with

smooth margins ventroapically, presence of a lateroapical furrow and ventroapical projections.

Distiphallic anteroapical shape strongly concave, apical shape conic and straight distiphallic

dorsal shape. Pregonite and postgonite with same color along the entire extension and

expanded base narrowing smoothly until the apex. Vesica with symmetrical branches; terminal

lobes reduced, filamentous at most tapering to apex and sclerotized texture; presence of lateral

lobes (i.e. division of the vesica coming from or close to the basal branch of the vesica, placed

laterally to phallic tube) and spines only ventral surface; and presence of a rounded median


Remarks. See O. admixta “remarks” section. Female was classified by Tibana &

Mello (1985), according to the shape of the T6+7, as syntergite undivided.

Distribution. NEOTROPICAL. Brazil (Amapá, Ceará, Mato Grosso, Minas Gerais,

Pará, Rio de Janeiro, São Paulo), Ecuador, Guyana.

Biology. This fly has been collected using different baits as human feces, chicken

viscera, mouse and rat carcasses, fish, banana and brown sugar (Lopes 1975a; Dias et al.

1984c; Moretti et al. 2008). Human feces, mouse and rat carcasses were the substrates

reported being used for rearing this species under natural conditions (Lopes 1973b; Moretti et

al. 2008). At laboratory, O. carvalhoi has been reared on agar plus powder milk, developing

from 1st instar to adults in 14–17 days (Lopes 1973b).

Material examined. ♂♂: BRAZIL: São Paulo, Campinas, UNICAMP, 05.XII.2011;

C. M. Souza / Oxysarcodexia sp. 1; Campinas-SP; 05/12/2011; 15 (L2B-DBA) // BRAZIL:

São Paulo, Campinas, UNICAMP, 05.XII.2011; C. M. Souza / Oxysarcodexia sp. 1;

Campinas-SP; 05/12/2011; 15 (L2B-DBA).

Oxysarcodexia chaetopygialis (Williston, 1896)

55

Sarcophaga chaetopygialis Williston, 1896

(Appendix 11-C)

Type locality. Saint Vincent. Williston, 1896: 366.

Depository of type material. AMNH.


developed. Thorax with grayish with pale golden microtomentum laterally. Dorsocentral post-

sutural setae with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta

absent. Legs brownish. Abdomen with silvery microtomentum. T3 with 2 lateral marginal

setae. T4 with undifferentiated median marginal and 1 lateral marginal seta. T5 without golden

microtomentum. ST5 with parallel cleft edges and with setosity and setae along the edges of

the arm. Cerci sinuous with pointed apexes of oblique upwards edges, in lateral view. Setae

ventrally on the cerci (lateral view) present on apical ⅓ portion. Apical shape of the cerci (last

⅓ portion of the cercus; in posterior view) same size as the middle part. Conformation of the

cerci in posterior view is parallel. In lateral view, distiphallus short and broad, with smooth

margins ventroapically; distiphallic apical shape square/oblong and sinuous distiphallic dorsal

shape. Pregonite broad, with same color along the entire extension (blackish) and base and

apex with same size, which can, depends on the genitalia exposure, hide the vesica. Postgonite

with expanded base narrowing smoothly until the apex. Vesica with symmetrical branches;

terminal lobes reduced, with rounded shape and membranous texture; without spines; but with

the presence of a rounded median projection of the main vesica branch.

Remarks. This species is recognized by the sinuosity of the cerci and the notable

distiphallus of light yellow color and enlarged ventroapically. Pregonite is dark brownish and

very broad, not unusually “hiding” the vesica. As already stressed by Dodge (1965), O.

chaetopygialis is the unique species of the genus presenting setulae on R1, even though a very

few (about 1–3 setulae).

Distribution. NEOTROPICAL. Jamaica (Saint Vincent).

Biology. Unknown.

Material examined. ♂♂: Majorca 1500‟ St. Vicent, W. I., VII–VIII.1972 / Malaise

trap, A. D. Harrison / Oxysarcodexia chaetopygialis (Will.) det. Lopes (ZMUC) // Majorca

1500‟ St. Vicent, W. I., VII–VIII.1972 / Malaise trap, A. D. Harrison / Oxysarcodexia

chaetopygialis ♂ (Will.) Det. Lopes (MNRJ).

56

Oxysarcodexia cingarus (Aldrich, 1916)

Sarcophaga cingarus Aldrich, 1916

Sarcophaga cingaris Hall (1929), incorrect subsequent spelling of cingarus Aldrich,

1916

(Appendix 12-A)

Type locality. Panama, Natrona. Aldrich, 1916: 288.



well-developed. Thorax with pale golden microtomentum more evident laterally, same for the

abdomen. Dorsocentral post-sutural setae with 2 well differentiated and 1–3 anterior setae

smaller. Apical scutellar seta absent. Legs brownish. T3 with 3 lateral marginal setae. T4 with

1 median marginal and 2 lateral marginal setae. T5 with golden microtomentum only laterally.

ST5 with parallel cleft edges and with setosity and setae on the apex of arms. Cerci, in lateral

view, straight with pointed apexes of oblique upwards edges. Setae ventrally on the cerci

(lateral view) present on apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the

cercus; in posterior view) smaller than the middle part. Conformation of the cerci in posterior

view is parallel. Presence of a remarkable constriction at middle portion of the cerci, also in

posterior view. In lateral view, distiphallus with ventroapical concavity of smooth margins;

distiphallic apical shape square/oblong and straight distiphallic dorsal shape. Pregonite with

same color along the entire extension and expanded base narrowing smoothly until the apex.

Postgonite with expanded base and sudden narrowing at apex. Vesica with symmetrical

branches; terminal lobes reduced, with filamentous at most tapering to apex shape and

sclerotized texture; without spines; and presence of an angular median projection of the main

vesica branch.

Remarks. Vesica of O. cingarus is one of the most simple shape and least ornamented

one among the species included in the genus. Phallus resemble somehow that seen in

Oxysarcodexia wygodzinsky Lopes & Tibana, 1987, although apex seems more angulated on

O. cingarus, and vesica and cerci shape are completely different for each species. Female was

57


undivided.

Distribution. NEARTIC. Canada (Quebec, Ontario), USA (Arkansas, Colorado,

Florida, Iowa, Kansas, New Jersey, New York, Pennsylvania, South Carolina).

Biology. Adults of this fly has been considered scavengers according to their food

habitats, once they were collected associated to pig carcasses exposed on a tree as much as in

the water (Payne & King 1972). Predatism of O. cingarus by the wasp Oxybelus uniglumis

(Linnaeus, 1758) (Hymenoptera: Sphecidae) has been recorded in the literature (Peckham et

al. 1973). This fly was one of the species used to study the phylogeny of Sarcophagidae based

on a molecular approach (mtDNA fragments) (Stamper et al. 2012).

Material examined. ♂♂: Great Falls, Oct. 3, 15 Va / W. L. McAfee, Collector /

Sarcophaga cingarus Ald., Det. J. B. Malloch (ZMUC) // La Fayette, La. Det. 9-11-1947

Follow by trap USPHN: LaF 2 / Oxysarcodexia cingarus Ald. Det. E. R. Dodge 1957

(MNRJ).

Oxysarcodexia comparilis (Reinhard, 1939)

Sarcophaga comparilis Reinhard, 1939

Oxysarcodexia omissa Lopes, 1946

(Appendix 12-B)

Type locality. USA, Texas, Donna. Reinhard, 1939: 66.

Depository of type material. Unknwon.


well-developed. Thorax and abdomen with pale golden microtomentum. Dorsocentral post-

sutural setae with 2 well differentiated and 3 anterior setae smaller. Apical scutellar seta

absent. Legs blackish. T3 with 3 lateral marginal setae. T4 with 1 median marginal and 3

lateral marginal setae. T5 with golden microtomentum along the entire extension. ST5 with

parallel cleft edges and with setosity and setae on the apex of arms. Cerci straight with normal

apexes (i.e. same size as median area) of oblique upwards edges, in lateral view. Setae

ventrally on the cerci (lateral view) present on apical half portion. Apical shape of the cerci

(last ⅓ portion of the cercus; posterior view) smaller than the middle part. Conformation of the

cerci in posterior view is parallel. In lateral view, distiphallus with smooth margins

58

ventroapically; distiphallic apical shape rounded and sinuous distiphallic dorsal shape.

Pregonite with expanded base narrowing smoothly until the apex, which is darker than the

base. Postgonite broad with expanded base and sudden narrowing at apex same color along the

entire extension (darker than pregonite). Vesica with symmetrical branches; terminal lobes

reduced, with filamentous at most tapering to apex shape and sclerotized texture; without

presence of spines; and presence of a rounded median projection of the main vesica branch.

Remarks. This species is considered similar to Oxysarcodexia confusa Lopes, 1946

and Oxysarcodexia parva Lopes, 1946, which the most evident difference is seen on cerci

apex (Lopes 1975c). Reinhard pointed out similarities of O. comparilis with O. culminata,

stating characteristics of genitalia and the lighter parafacialia and parafrontalia

microtomentum as differences between them (Lopes 1946b). Terminalia of O. comparilis

resemble Oxysarcodexia occulta Lopes, 1946, although vesica of O. comparilis present

terminal lobes laterolaterally directed and characteristics as an angular median projection of

the main vesica branch, elongation of basal area of terminal lobes, bigger vesica size with

terminal lobes ventroapically directed, surstylus conic and cerci sinuous are seen only in the

last species.

Distribution. NEARTIC. Mexico (Districto Federal, Morelos), USA (Texas).

NEOTROPICAL. Ecuador, El Salvador, Mexico (Jalisco, Veracruz).

Biology. Unknown. Information available is only about the type of collection, active,

using an entomological net (Lopes 1946b).

Material examined. ♂: Acatlipa Morelos July 29 1950 Coll. N. G. Dorns /

Oxysarcodexia comparilis ♂ (Reinh.) Det. H. S. Lopes (MNRJ).

Oxysarcodexia conclausa (Walker, 1861)

Sarcophaga conclausa Walker, 1861 [female]

Sarcophaga ochripyga Wulp, 1895

Sarcophaga australis Aldrich, 1916

Sarcophaga ochropyga Mattos (1919), incorrect subsequent spelling of ochripyga

Wulp, 1895

Sarcophaga ochrypyga Lopes (1933), incorrect subsequent spelling of ochripyga

Wulp, 1895

59

(Appendix 12-C)

Type locality. Mexico. Walker, 1861: 309.



well-developed. Thorax grayish with pale golden microtomentum. Dorsocentral post-sutural


Legs brownish. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral

marginal setae. T5 with golden microtomentum along the entire extension. ST5 with parallel

cleft edges and with setosity. Cerci, in lateral view, straight with expanded apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) present on apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; posterior view) with same size as the middle

part. Conformation of the cerci in posterior view is parallel. In lateral view, distiphallus with

serrated margins ventroapically; distiphallic apical shape rounded and straight distiphallic

dorsal shape. Presence of a dorsoapical swelling (“finger-like”) on the distiphallus, in lateral

view. Pregonite with expanded base and sudden narrowing at apex, which is darker than the

base. Postgonite with expanded base and sudden narrowing at apex. Vesica with symmetrical

branches; terminal lobes well-developed, with square shape and sclerotized texture; presence

of spines only along the edges; and presence of an angular median projection of the main

vesica branch.

Remarks. Dorsoapical swelling “finger-like” of distiphallus, distiphallus apical

conformation and vesica with oblong shape and serrated edges resemble to those

characteristics of Oxysarcodexia morretesi Tibana & Mello, 1983 and O. thornax, despite of

apical shape of the cerci differing among O. conclausa and these other species. See also

“remarks” section on O. afficta. Female was described by Lopes (1975b) and classified by


two plates. Larvae are known and were described by Knipling (1936).

Distribution. NEARTIC. Canada (Quebec), Mexico (Baja California Sur, Guajanuto,

Morelos, San Luis Potosí), USA (Arizona, California, Illinois, Indiana, Louisiana, Missouri,

New York, Oklahoma, Texas). NEOTROPICAL. Chile (La Araucanía, Patagonia, Tarapacá),

Colombia (Antioquia), Costa Rica (Alajuela, Guanacaste, San José), Ecuador, El Salvador,

Guatemala, Honduras (Tegucilgalpa), Mexico (Chiapas, Guerrero, Jalisco, Nayarít, Sinaloa,

60

Tabasco, Veracruz, Yucatán), Panama, Peru, Trinidad and Tobago (Trinidad), Venezuela

(Aragua).

Biology. This species is close related to bovine dung (Blume 1985), and due to its

sazonality (more active with the onset of the first spring which coincides with the emergence

of Haematobia irritans (Linnaeus, 1758) (Diptera: Muscidae)) and for being rearing on cattle

dung, it was considering a possible candidate for the natural control of the horn fly (Kunz

1978). Strong necrophagous behavior has been indicated by Yepes-Gaurisas et al. (2013),

further a high synanthropic index, pointing out this species preference for human settlements.

Collections with entomological net and with traps for collecting fruit flies are mentioned by

Lopes (1946b). Attractiveness is marked by fish, liver, chicken viscera and pig carcass (Flores

& Dale 1995; Garcés et al. 2004; Ramírez-Mora et al 2012; Yepes-Gaurisas et al. 2013).

Oxysarcodexia conclausa is also considered a floral visitor of Bdallophyton bambusarum

Liebm. (Rafflesiaceae) (Gacía-Franco & Rico-Gray 1997).

Material examined. ♂♂: Hidalgo Co. Tx., 4.X.1986; dysentery fly trap; USPUS Mi /

Oxysarcodexia ochripyga (ZMUC) // COLOMBIA Palmira, Valle 1 Mar 1971 1006m 165 G.

P. Waldbauer / Oxysarcodexia ochripyga Wulp, H. S. Lopes det. (MNRJ).

Oxysarcodexia confusa Lopes, 1946

(Appendix 13-A)

Type locality. BRASIL, RIO DE JANEIRO, Miguel Pereira. Lopes, 1946b:96.




microtomentum of the abdomen. Dorsocentral post-sutural setae with 2 well differentiated and

1–3 anterior setae smaller. Apical scutellar seta absent. Legs brownish. T3 with 2 lateral

marginal seta. T4 with 1 median marginal and three lateral marginal setae. T5 normally

without golden microtomentum, in a few cases a pale golden microtomentum can be present

laterally. ST5 with parallel cleft edges and with setosity. Cerci, in lateral view, straight with

expanded apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view) absent

only in middle portion. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior

view) same size as the middle part. Conformation of the cerci in posterior view is parallel.

61

Presence of a remarkable constriction at middle portion of the cerci, also in posterior view. In

lateral view, distiphallus with ventroapical concavity of smooth margins; distiphallic apical

shape rounded and straight distiphallic dorsal shape. Pregonite and postgonite with same color

along the entire extension and expanded base narrowing smoothly until the apex. Vesica with

symmetrical branches; terminal lobes reduced, with rounded shape and partially membranous

texture; presence of spines on both ventral and dorsal surfaces; and presence of a rounded

median projection of the main vesica branch.

Remarks. A good and detailed male terminalia comparison of the sympatric species

not unusually misidentified, O. avuncula, O. confusa, O. diana and O. parva, is given by Silva

& Mello-Patiu (2008). Distiphallus of O. confusa, in lateral view, is very similar to

Oxysarcodexia molitor (Curran & Walley, 1934), differing by the internal plates (spinous on

O. molitor) and the composition of the glans (Lopes 1975c). Female was classified by Tibana

& Mello (1985), according to the shape of the T6+7, as syntergite undivided. Larvae are

characterized by the presence of sinuous ribbons of festoon on psedocephalon and

conspicuous teeth on maxillae, same as O. thornax, differing from it by the more scattered

teeth of the maxillae and by the festoon arms (Leite & Lopes 1987; Lopes & Leite 1986;

1987). See also “remarks” section on O. comparilis for further discussion.

Distribution. NEOTROPICAL. Argentina (Misiones), Brazil (Mato Grosso, Mato

Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, Santa Catarina, São Paulo).

Biology. Attractiveness by human feces, chicken viscera, mouse and fish carcasses and

banana plus brown sugar is reported (Lopes 1973b; Dias et al. 1984c; Mendes & Linhares

1993). This fly has been found rearing on human feces, under natural conditions, and at

laboratory, it has been reared on agar plus powder milk for 24h, then transferred to meat to

complete the development (Lopes 1973b). Studying synanthropy of flesh flies from Curitiba,

Brazil, Ferreira (1979) collected O. confusa using a trap baited with fish, chicken liver and

human feces, observing this species wasn‟t often associated with inhabited areas.

Material examined. ♂♂: Petrópolis, Tq; E. do Rio, Brasil, H. S. Lopes, 2.69 /

Oxysarc. confusa, Lop., ♂, Det. H. S. Lopes / NRM-DIPT 0014285 (NRM) // IGUASSÚ,

Paraná XII-941, Com. E. N. V. / NRM-DIPT 0014287 (NRM) // Taquara, Petrópolis, E. do

Rio, Brasil / H. S. Lopes, 16.II.75 / Oxysarcodexia confusa, Lopes. Det. H. S. Lopes (ZMUC)

62

// S. Paulo – Cantareira Serra L. Trav. F. Q. 30.VIII.935 / Paratype / Oxysarcodexia confusa

sp.n. Lopes – det 1944 (MNRJ).

Oxysarcodexia corolla Dodge, 1965

(Appendix 13-B)

Type locality. Jamaica, Saint Andrews, Second Breakfast Spring. Dodge, 1965: 508.

Depository of type material. WSU.

Diagnosis. Male. Postocular plate yellowish gray. Ocellar setae weak. Thorax gray and

abdomen with silvery microtomentum. Dorsocentral post-sutural setae with 3 well

differentiated, a small seta can be present among them. Apical scutellar seta present, although

considered small or weak. Legs blackish. Abdomen grayish with pale golden microtomentum

laterally. T4 with long marginal setosity, but without setae. ST5 with a deep cleft. Pregonite

with darker apex, stouter and curved. Postgonite is slender and nearly straight. Cerci brownish

red with black apexes, not divergent apically and a conspicuous preapical “tooth”. Distiphallus

straight with compressed apex and an anterior elongation pointed inward, and bearing a

preapical flap around dorsal and lateral sides. Vesica is black, broad, with an asymmetric

finger-shaped process on the left side. This is a synopsis of the original description (Dodge

1965) and of a redescription (Lopes 1978).

Remarks. See “remarks” section on O. augusta. Female was classified by Tibana &


Distribution. NEOTROPICAL. Jamaica (St. Andrews).

Biology. Unknown.

Oxysarcodexia cuernavaca Dodge, 1966

(Appendix 13-C)

Type locality. Mexico, Morelos, Cuernavaca. Dodge, 1966: 685.


Diagnosis. Male. Postocular plate with silvery microtomentum. Weak ocellar setae.

Thorax gray, slightly yellow laterally. Dorsocentral post-sutural setae with 2 well

differentiated and 2 anterior setae smaller. Apical scutellar seta is absent. Legs blackish.

Abdomen black and brown apically, with silvery microtomentum. T3 with median marginal

63

seta. T4 yellow with a marginal row of setae. T5 with golden microtomentum. Cerci yellow

with divergent apexes (posterior view), and straight shape and a strong preapical “tooth”, in

lateral view. Pregonite long and curved. Postgonite without seta. Large vesica with flattened

and spined lobes, presenting broad and thin margins. This is a synopsis of original description

given by Dodge (1966).

Remarks. It is considered similar to O. ventricosa, by the similar cerci with divergent

apexes (Dodge 1966). See also “remarks” section of O. berlai for further discussion.

Distribution. NEARTIC. Mexico (Jalisco, Morelos, San Luis Potosí).

Biology. Unknown.

Oxysarcodexia culmiforceps Dodge, 1966

Oxysarcodexia culminiforceps Lopes (1969), incorrect subsequent spelling of

culmiforceps Dodge, 1966

(Appendix 14-A)

Type locality. Brazil, Rio de Janeiro, Rio de Janeiro. Dodge, 1966: 687.

Depository of lectotype species. MNRJ.


developed. Thorax with golden microtomentum laterally. Dorsocentral post-sutural setae with

2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta absent. Legs

blackish. Abdomen with silvery microtomentum. T3 with 2 lateral marginal setae. T4 with 1

median marginal and 2 lateral marginal setae. T5 with golden microtomentum laterally

(“stain-like”). ST5 with parallel cleft edges and with setosity and setae along the edges of the

arms. Cerci, in lateral view, bent backwards (very conspicuous) with pointed apexes of

oblique upwards edges. Setae ventrally on the cerci (lateral view) present on apical ⅓ portion.

Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) with same size as the

middle part. Conformation of the cerci in posterior view is divergent. Presence of a remarkable

constriction at middle portion of the cerci, also in posterior view. In lateral view, distiphallus

with a ventroapical concavity of serrated margins ventroapically; presence of a lateroapical

furrow; distiphallic apical shape conic and sinuous distiphallic dorsal shape. Presence of a

large dorsoapical swelling on the distiphallus, in lateral view. Lateroapical expansions present

on distiphallus, in dorsal view. Pregonite with expanded base narrowing smoothly until the

64

apex, which is darker than the base. Postgonite with expanded base and sudden narrowing at

apex. Vesica with symmetrical branches; terminal lobes reduced, with filamentous at most

tapering to apex shape and partially membranous texture; presence of spines only along the

edges; and presence of an angular median projection of the main vesica branch.

Remarks. Initially it was misidentified as O. culminata although morphological

variations, especially at male genitalia, were pointed out by Lopes (1946b), reason of it is

considered a very variable species. Dodge (1966) confirmed these differences as belonging to

another species, proposing the new name O. culmiforceps. Differentiation of O. culmiforceps

from O. culminta is based on distiphallus apex and vesica details (Lopes 1975c). See

“remarks” section on O. aurata for further discussion. Female was classified by Tibana &

Mello (1985), according to the shape of the T6+7, as syntergite membranous.

Distribution. NEOTROPICAL. Argentina (Buenos Aires, Entreríos, Misiones), Brazil

(Distrito Federal, Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul,

Santa Catarina, São Paulo), Paraguay.

Biology. This species is attracted by human and dog feces, chicken viscera, rotten

bovine liver, fish, mouse and pig carcasses and banana plus brown sugar (Ferreira 1979; Dias

et al. 1984c; Mendes & Linhares 1993; Carvalho & Linhares 2001; Leandro & D‟Almeida

2005; Mariluis et al. 2007; Mulieri et al. 2008; Mulieri et al. 2010; Mulieri et al. 2011; Vairo

et al. 2011). Despite of this variety of substrates, it has been observed no differences in

attractiveness among human feces, chicken viscera and mouse carcasses (Linhares 1981; Dias

et al. 1984c), besides no preference for sunlight or shaded areas (Linhares 1981). However,

Ferreira (1979) recorded this species preference by human feces, linking this fact to a possible

negative relationship to the preference for inhabited areas. Grassland, woodland (Mulieri et al.

2008), urban, suburban and rural areas (Mulieri et al. 2011), forest, rural and urban areas (in

order of abundance in each site) at Brazilian South (Ferreira 1979), a Brazilian zoological

garden (Oliveira et al. 2002) and a forest area (our own data) are the environments where O.

culmiforeceps has already been collected on. This fly is also recognized as flower visitor of

Sebastiania brasiliensis Spreng. (Euphorbiaceae) and of Scutia buxifolia Reissek

(Rhamnaceae) (Mulieri et al. 2010).

65

Material examined. ♂: BRAZIL: Minas Gerais, Extrema, Serra do Lopo, Pedra das

Flores, 27.II.2012; A. G. Savino, M. P. Nassu, M. D. Grella / O. culmiforceps, Extrema-MG;

27/02/2012 (L2B-DBA).

Oxysarcodexia culminata (Aldrich, 1916)

Sarcophaga culminata Aldrich, 1916

(Appendix 14-B)

Type locality. Puerto Rico, Mayaguez. Aldrich, 1916: 289.



well-developed. Thorax grayish with golden microtomentum at humeral portion. Dorsocentral


seta absent. Legs blackish. T3 with 2 lateral marginal setae. T4 with 1 median marginal and 3

lateral marginal setae. T5 with exclusively silvery microtomentum. ST5 with parallel cleft

edges and with setosity. Cerci, in lateral view, bent backwards with pointed apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) with the same size as the

middle part. Conformation of the cerci in posterior view is parallel. Presence of a remarkable

constriction at middle portion of the cerci, in posterior view. In lateral view, distiphallus with

ventroapical concavity with serrated margins ventroapically; distiphallic apical shape conic

and sinuous distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until

the apex, which is darker than the base, as well as the postgonite, except for the color which is

the same along the entire extension. Vesica with symmetrical branches; terminal lobes

reduced, with rounded shape and partially membranous texture; presence of spines only on

ventral surface; and presence of an angular median projection of the main vesica branch.

Remarks. See “remarks” section on O. aurata, O. comparilis and O. culmiforceps.

Distribution. NEOTROPICAL. Argentina (Misiones), Brazil (Rio de Janeiro), Puerto

Rico.

Biology. This species has already been reared under laboratory conditions and the

attraction by human feces, rotten Syagrus comosa (Mart.) Mart. (Arecaceae), a species of a

palmtree, and rotten banana plus brown sugar (the last two at coastal area) has been reported

66

on the literature (Lopes 1973b). This author also noted the distribution of O. culminta in

highlands at Rio de Janeiro, Brazil.

Material examined. ♂: PUERTO RICO: Jayuya Bosque Estatal del Toro Negro,

0,7km SE, Cerro de Punta, Cordillera Central / 18-10-09N, 66-35-16W, 1195m, 9 June 1966,

J. Rawlins. C. Young, R. Davidson, W. Zanol, S. Thompson, M. Klinger / Carnegie Museum,

Specimen Number CMNH-67351 / NRM-DIPT 0014285 (NRM).

Oxysarcodexia cyanea Lopes, 1975

(Appendix 14-C)

Type locality. Dominica, Traglafar Falls. Lopes, 1975b: 475.

Depository of type material. NMNH.

Diagnosis. Male. Back of head gray. Ocellar setae well-developed. Thorax gray,

dorsocentral post-sutural setae with 2 well differentiated and 2 anterior smaller setae. Apical

scutellar seta is absent. Legs blackish. Abdomen grayish with silvery microtomentum. T4

without median marginal setae. ST5 reddish brown. Cerci yellow, slightly curved on apexes,

which are blackish. Parallel conformation of cerci and enlarged cerci apexes in posterior view.

Pregonite with expanded apex. Distiphallus sinuous on apex and with a small membranous

area dorsally. Vesica large with apical lobes bearing a few spines. This is a synopsis of the

original description given by Lopes (1975b).

Remarks. Distiphallus similar to Oxysarcodexia zayasi Dodge, 1956, although

differences are notably on cerci which is near to O. orbitalis, but differing from O. cyanea by

the distinct distiphallus. It is also distinguished from O. aurata due to the absence of a

dorsoapical swelling (called by Lopes as a “large membrane on penis apex”) (Lopes 1975b).


syntergite undivided.

Distribution. NEOTROPICAL. Dominica.

Biology. Unknown. There is only a mention in the original description paper about

some specimens collected on human feces, however without mention which species they refer

to (Lopes 1975b).

Oxysarcodexia cyaniforceps (Hall, 1933)

67

Sarcophaga cyaniforceps Hall, 1933

(Appendix 15-A)

Type locality. Panama, Barro Colorado Island. Hall, 1933: 282.


Diagnosis. Male. Postocular plate golden microtomentum. Thorax gray and with

golden microtomentum on humeral region and laterally. Dorsocentral post-sutural setae with 2

well differentiated and 3 anterior smaller setae. Apical scutellar seta is absent. Abdomen

grayish with pale golden microtomentum laterally on T3 and on entire tergites T4 and T5. T4

with median marginal setae. T5 completely black. ST5 reddish with a deep cleft. Cerci with

sinuous shape (lateral view), enlarged and curved apexes. Distiphallus mostly membranous,

but with a pair of spinous plates internally. Large vesica with 2 pair of spinous lobes. This is a

synopsis of diagnosis given by Lopes (1946b; 1975c).

Remarks. Accordingly to drawings presented by Lopes (1975c), cerci seem similar to

the one presented by O. chaetopygialis and vesica resembles Oxysarcodexia floricola Lopes,

1975, although terminal lobes are ventroapically directed and basal portion is placed slightly

below distiphallus apex. See also “remarks” section of O. adunca for further discussion.

Distribution. NEOTROPICAL. Colombia (Antioquia), Costa Rica, Panama (Barro

Colorado Island, Corozal Canal Zone).

Biology. This fly is considered very rare and it was collected only one specimen in

urban area of Antioquia, Colombia, using chicken viscera as baits (Yepes-Gaurisas et al.

2013).

Oxysarcodexia diana (Lopes, 1933)

Sarcophaga diana Lopes, 1933

(Appendix 15-B)

Type locality. Brasil, Rio de Janeiro, Rio de Janeiro. Lopes, 1933:154.



developed. Thorax with golden microtomentum more intense at humeral portion. Dorsocentral


seta absent. Legs brownish. Abdomen grayish with some shades of golden microtomentum

68

laterally. T3 with 3 lateral marginal setae. T4 with 1 median marginal and 1 lateral marginal

seta. T5 with golden microtomentum along the entire extension. ST5 with parallel cleft edges

and with setosity and setae on the apex of the arms. Cerci, in lateral view, sinuous with

expanded apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view) present

in apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; posterior view)

smaller than the middle part. Conformation of the cerci in posterior view is parallel. In lateral

view, distiphallus with ventroapical concavity with smooth margins ventroapically;

distiphallic apical shape rounded and sinuous distiphallic dorsal shape. Pregonite with

expanded base narrowing smoothly until the apex, which is darker than the base. Postgonite

with expanded base and sudden narrowing at apex. Vesica with symmetrical branches;

terminal lobes well-developed, with rounded shape and membranous texture; presence of

spines only on ventral surface; and presence of a rounded median projection of the main

vesica branch.


not unusually misidentified, O. avuncula, O. confusa, O. diana and O. parva, is given by Silva

& Mello-Patiu (2008). See O. admixta “remarks” section for further discussion. Female was


undivided.

Distribution. NEARTIC. Mexico (Morelos, San Luis de Potosí). NEOTROPICAL.

Argentina (Misiones), Brazil (Distrito Federal, Ceará, Goiás, Mato Grosso, Mato Grosso do

Sul, Minas Gerais, Paraná, Rio de Janeiro, Roraima, Santa Catarina, São Paulo), Colombia

(Antioquia), Ecuador, El Salvador, Mexico (Chiapas), Paraguay, Trinidad and Tobago

(Trinidad).

Biology. This fly has already been reared on human feces (D‟Almeida 1989;

D‟Almeida 1994; Lopes 1973b; Lopes 1975c) – and it has great preference for this kind of

substrate for rearing (Mendes & Linhares 1993) –, on bovine feces of pasture (Marchiori

2000; Marchiori et al. 2001), cattle shed (Marchiori 2000), on fish (D‟Almeida 1986) and also

under laboratory conditions (Lopes 1973b). D‟Almeida (1984) noticed this species preference

for inhabited areas, collecting it with human feces, fish, bovine liver and banana plus brown

sugar as baits (in decreasing scale of abundance). Attractiveness has been reported by human

feces, fish, chicken viscera, bovine liver and lung, mouse and pig carcasses, crab, squid,

69


palmtree (at coastal area) (Lopes 1973b; Lopes 1975a; Linhares 1981; D‟Almeida 1984; Dias

et al. 1984c; Mendes & Linhares 1993; D‟Almeida & Lima 1994; Pamplona et al. 2000;

Carvalho & Linhares 2001; Leandro & D‟Almeida 2005; Barros et al. 2008; Barbosa et al.

2009; Ramírez-Mora et al. 2012; Yepes-Gaurisas et al. 2013). Analyzing the attractiveness of

three different baits, Linhares (1981) observed no differences for mouse carcass, human feces

and chicken viscera, although Mendes & Linhares (1993), in a study using the same baits,

pointed out O. diana preference for human feces. Sunlight instead of shaded areas is also

referred as a preference of this species (Linhares 1981). The higher frequency of occurrence of

adult females on chicken viscera and rodent carcasses baits suggest the use of these substrates

as proteic resources for the development of ovarian folicules (Mendes & Linhares 1993).

Besides actively, this fly has been collected using malaise Shannon and W.O.T. traps baited

with different substrates mentioned above (Lopes & Tibana 1991; Pamplona et al. 2000). At

Rio de Janeiro Zoological Garden, it was the third most abundant species among flesh flies

collected, showing a peak of occurrence in January and February (summer) and, consequently,

positive correlation with temperature and preference for urban areas and feces (available

abundantly at the zoo) (Oliveira et al. 2002).

Material examined. ♂♂: Oxysarcodexia diana sp 4 / TdeA 831 (CE-TdeA) [from

Colombia] // BRAZIL: Minas Gerais, Belo Horizonte, Est. Ecológica, UFMG, Campus, 2–

22.vii.1993, S. D. Gaiman / NRM-DIPT 0014598 (NRM) // GRAJAÚ Rio de Janeiro Lopes

10.XI.40 (MNRJ).

Oxysarcodexia eberti Lopes & Tibana, 1987

(Appendix 15-C)

Type locality. Brazil, Minas Gerais, Paracatu. Lopes & Tibana, 1987: 336.



developed. Thorax and abdomen with silvery microtomentum and slightly pale golden

microtomentum shades. Dorsocentral post-sutural setae with 2 well-differentiated and 3

anterior setae smaller. Apical scutellar seta absent. Legs blackish. T3 with 2 lateral marginal

setae. T4 without median marginal and with 2 lateral marginal setae. T5 with slightly pale

70

golden microtomentum, although silvery microtomentum more intense. ST5 with parallel cleft

edges and setosity. Cerci, in lateral view, bent backwards with expanded apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) absent only in middle portion.

Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) smaller than the

middle part. Conformation of the cerci in posterior view is parallel. In lateral view,

distiphallus with smooth margins ventroapically; distiphallic apical shape square/oblong and

sinuous distiphallic dorsal shape. Presence of a large dorsoapical swelling on the distiphallus,

in lateral view. Lateroapical expansions on distiphallus, in dorsal view, present. Pregonite and

postgonite with expanded base and sudden narrowing at apex, which is darker than the base

only in pregonite. Vesica symmetric; terminal lobes well-developed, with square shape and

mostly sclerotized texture; presence of spines along the edges; and presence of an angular


Remarks. See “remarks” section on O. aurata.

Distribution. NEOTROPICAL. Brazil (Distrito Federal, Mato Grosso, Mato Grosso

do Sul, Minas Gerais).

Biology. This species has been collected associated to pig carcass in areas of Brazilian

Cerrado (“savanna-like”) vegetation (Barros et al. 2008; Rosa et al. 2011).

Material examined. ♂: BRASIL: MT: Chap. Dos Guimarães Véu da Noiva – cerrado

S15°24‟ W5550‟ Shannon – peixe 16.i.2012 Mello-Patiu & Patiu col.

SISBIOTA/CNPq/FAPESP / Oxysarcodexia eberti Lopes & Tibana, 1987 Det. C. A. Mello-

Patiu (MNRJ).

Oxysarcodexia edwardsi Lopes, 1946

(Appendix 16-A)

Type locality. Colombia, Gorgona Island. Lopes, 1946b: 123.



well-developed. Thorax grayish and with pale golden microtomentum. Dorsocentral post-

sutural setae with 2 well differentiated and 3 anterior smaller setae. Apical scutellar seta is

absent. Legs blackish. Abdomen grayish with golden microtomentum laterally. T4 with one

pair of median marginal setae. T5 with golden microtomentum along the entire extension. ST5

71

with a deep cleft of V-shaped edges. Cerci reddish with blackish and “arrow-shaped” apexes.

Distiphallus reddish. Vesica of notably asymmetrical branches. This is a synopsis of the

original description given by Lopes (1946b).

Remarks. Comparison among drawings from O. edwardsi terminalia presented in the

original description (Lopes 1946b: Figs. 146–149) and other Oxysarcodexia species enables

recognize some similarities with O. parva and Oxysarcodexia trivialis (Wulp, 1895). Cerci is

very similar to that seen on O. parva, nevertheless vesica is completely different and more

similar to that of O. trivialis. In O. edwardsi the vesica presents an angular projection of the

main vesica branch, a more enlarged basal portion of terminal lobes and the apex of terminal

lobes ventroapical directed. For O. trivialis, the main vesica branch presents a rounded

projection, narrower basal portion of terminal lobes and the apex of terminal lobes dorsoapical

directed.

Distribution. NEOTROPICAL. Brazil (Rio de Janeiro), Colombia (Gorgona Island).

Biology. Collected using traps baited with cow liver and lung, fish and squid and

actively with feces (Pamplona et al. 2000).

Oxysarcodexia favorabilis (Lopes, 1935)

Sarcophaga favorabilis Lopes, 1935

(Appendix 16-B)





microtomentum of the abdomen. Three well differentiated dorsocentral post-sutural setae are

present. Apical scutellar seta absent. Legs dark brownish. T3 with 2 lateral marginal setae. T4

with 1 median marginal and 3 lateral marginal setae. T5 without golden microtomentum, only

silvery microtomentum is present. ST5 with parallel cleft edges and with setosity. Cerci, in

lateral view, bent backwards with pointed apexes of straight edges. Setae ventrally on the cerci

(lateral view) absent only in middle portion. Apical shape of the cerci (last ⅓ portion of the


view is parallel. In lateral view, distiphallus with smooth margins ventroapically; distiphallic

72

apical shape square/oblong and sinuous distiphallic dorsal shape. In lateral view, presence of a

large dorsoapical swelling on the distiphallus, ventroapical projections and lateral lobes.

Lateroapical expansions on the distiphallus, in dorsal view, present. Pregonite with base and

apex with almost same size and same color along the entire extension. Postgonite with

expanded base and sudden narrowing at apex and same color along the entire extension.

Vesica symmetric; terminal lobes well-developed, with filamentous at most tapering to apex

shape, membranous texture and presence of spines.

Remarks. See “remarks” section on Oxysarcodexia 4 n. sp.

Distribution. NEOTROPICAL. Brazil (Rio de Janeiro).

Biology. Unknown.

Material examined. ♂: Angra dos Reis, Est. do Rio, Dario Mendes 8-934 / Holotype /

Typus / Sarcophaga favorabilis Lopes H. S. Lopes DET. – 935 / MNRJ/2307 (MNRJ).

Oxysarcodexia festiva Lopes & Tibana, 1987

(Appendix 16-C)

Type locality. Costa Rica, Cartago, Turrialba. Lopes & Tibana, 1987, 338.

Depository of type material. CAS.

Diagnosis. Male. Postocular plate with golden microtomentum. Ocellar setae with

about the size of smaller frontal setae. Thorax with golden microtomentum dorsally and on

mesonotum. Dorsocentral post-sutural setae with 2 well differentiated and 3 anterior smaller

setae. Apical scutellar seta is absent. Legs brown. Abdomen with yellowish gray

microtomentum. T4 with median marginal setae. ST5 with slender and elongated apical arms

covered with setosity on apexes. Cerci with pre-apical dorsal recurrent lobe and remarkable

setosity on apexes. Distiphallus curved forwards, with setosity anteroapically. Robust vesica

with setosity ventrally; apical lobes elongated, membranous and with a pair of spines. This is a

synopsis of the original description given by Lopes & Tibana (1987).

Remarks. Species easily distinguished from others by the peculiar cerci (Lopes &

Tibana 1987).

Distribution. NEOTROPICAL. Costa Rica.

Biology. Unknown.

73

Oxysarcodexia flavipes Lopes & Tibana, 1987

(Appendix 17-A)

Type locality. Argentina, Tucumán, Cerro, San Javier. Lopes & Tibana, 1987: 355.



developed. Thorax with pale golden microtomentum. Three well-differentiated dorsocentral

post-sutural setae. Apical scutellar seta absent. Legs yellowish. Abdomen with silveryish

microtomentum. T3 with one lateral marginal seta. T4 with 1 median marginal and 1 lateral

marginal seta. T5 without golden microtomentum, only yellowish on ventral edges. ST5 with

parallel cleft edges and with setosity and setae along the edges of the arms. Cerci, in lateral

view, straight with normal (i.e. same size as median area) apexes of oblique upwards edges.

Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical shape of the cerci

(last ⅓ portion of the cercus; in posterior view) smaller than the middle part. Conformation of

the cerci in posterior view is divergent. In lateral view, distiphallus with smooth margins

ventroapically; distiphallic apical shape conic and sinuous distiphallic dorsal shape. Presence

of a small dorsoapical swelling on the distiphallus, in lateral view. Lateroapical expansions on

distiphallus, in dorsal view, present. Pregonite with base and apex with the same size and same

color along the entire extension. Postgonite with expanded base and sudden narrowing at apex

and same color along the entire extension. Vesica symmetric; with presence of lateral lobes

(i.e. division of the vesica coming from or close to the basal branch of the vesica, placed

laterally to phallic tube); terminal lobes reduced, with rounded shape, sclerotized texture and

presence of spines on both ventral and dorsal surfaces.

Remarks. This species is distinctly recognized by the yellow legs, 3 dorsocentral post-

sutural setae, apical distiphallic apex and conspicuous vesica (Lopes & Tibana 1987). See also

“remarks” section on O. augusta.

Distribution. NEOTROPICAL. Argentina (Tucumán).

Biology. Unknown.

Material examined. ♂: Cerro San Javier Tucuman, Arg. 8.X.53 800m P.

Wygodzinsky / Holótipo / Oxysarcodexia flavipes n.sp. holotypus ♂ Det. H. S. Lopes & R.

Tibana / MNRJ/2239 (MNRJ).

74

Oxysarcodexia floricola Lopes, 1975

(Appendix 17-B)

Type locality. Costa Rica, Guanacaste, 24 km NW Canas. Lopes, 1975e: 487.

Depository of type material. Unknown [original description only mentioned material

examined belonging to Dr. E. R. Heithaus].


developed. Thorax and abdomen with pale golden microtomentum. Dorsocentral post-sutural

setae with 2 well differentiated and 2 anterior setae smaller. Apical scutellar seta absent. Legs

blackish. T3 with 1 lateral marginal seta. Cerci, in lateral view, sinuous with normal (i.e. same

size as median area) apexes of oblique upwards edges. Setae ventrally on the cerci (lateral

view) present on apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in

posterior view) with the same size as the middle part. Conformation of the cerci in posterior

view is parallel. Presence of a remarkable constriction at middle portion of the cerci, in

posterior view. In lateral view, distiphallus with smooth margins ventroapically; distiphallic

apical shape conic and straight distiphallic dorsal shape. Pregonite with expanded base

narrowing smoothly until the apex. Postgonite with expanded base and sudden narrowing at

apex. Vesica symmetric; terminal lobes well-developed, with filamentous at most tapering to

apex shape, sclerotized texture and presence of spines only along ventral edge; and presence of


Remarks. Specimen examined presents a damaged abdomen, lacking T4 (partially),

T5 and ST5 (both completely). Terminalia is on a microtube with glycerin. In original

description is mentioned yellow color for T5; T4 without median marginal seta presence and

there is no remarks for ST5 (Lopes 1975e). See “remarks” section on O. cyaniforceps for

further discussion.

Distribution. NEOTROPICAL. Brazil (Santa Catarina), Costa Rica (Guanacaste).

Biology. Unknown. Information available is only about the collection, in flowers

(Lopes 1975e).

Material examined. ♂: Nova Teutonia 27°11‟S, 52°23‟W Brazil, 300–500m 15-III

1968 Fritz Plaumann / Oxysarc. floricola ♂ Lopes Det. H. S. Lopes (MNRJ).

Oxysarcodexia fluminensis Lopes, 1946

75

(Appendix 17-C)

Type locality. Brazil, Rio de Janeiro, Guanabara, Grajaú. Lopes, 1946b: 104.



developed. Thorax as well as abdomen with silvery microtomentum. Dorsocentral post-sutural

setae with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta present.

Legs blackish. T4 with 1 median marginal and 1 lateral marginal seta. T5 with golden

microtomentum in part. ST5 with parallel cleft edges and with setosity. Cerci, in lateral view,

straight with expanded apexes of oblique upwards edges. Setae ventrally on the cerci (lateral

view) present in apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in

posterior view) smaller than the middle part. Conformation of the cerci in posterior view is

divergent. In lateral view, distiphallus with ventroapical concavity with serrated margins

ventroapically; presence of a lateroapical furrow. Distiphallic apical shape rounded and

sinuous distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until the


apex. Vesica with symmetrical branches; terminal lobes well-developed, filamentous at most

tapering to apex shape, sclerotized texture and with presence of lateral lobes (i.e. division of

the vesica coming from or close to the basal branch of the vesica, placed laterally to phallic

tube); presence of spines only along the edges; and presence of a rounded median projection

of the main vesica branch.

Remarks. Distiphallus with a noteworthy concavity of serrated edges ventroapically.


syntergite partially divided into two plates.

Distribution. NEOTROPICAL. Brazil (Distrito Federal, Mato Grosso, Mato Grosso

do Sul, Minas Gerais, Pernambuco, Rio de Janeiro, São Paulo), Colombia (Antioquia).

Biology. Species already reared under laboratory conditions (Lopes 1973b), using dog

and primate feces, mouse carcass, crab (D‟Almeida 1989) and fish (D‟Almeida 1986) as

rearing substrates. Adults have been collected associated to pig carcass (Barros et al. 2008;

Cruz 2008; Barbosa et al. 2009) and cadavers (Oliveira-Costa et al. 2001). D‟Almeida (1984)

collected this fly using fish, bovine liver and human feces (decreasing scale of abundance) as

baits and pointed out a high synanthropic index for it. Baits as human feces, pig carcass, cow

76

liver and lung, chicken viscera, fish, crab, squid, rotten banana plus brown sugar and rotten

Syagrus comosa (Mart.) Mart. (Arecaceae), a species of a palmtree (at coastal area) has been

used for collect this species (Lopes 1973b; D‟Almeida & Lima 1994; Pamplona et al. 2000;

Oliveira et al. 2002; Leandro & D‟Almeida 2005; Vasconcelos & Araújo 2012; Ramírez-Mora

et al. 2012). Adults were already collected actively or using baited or W.O.T. traps at a forest

environment, an urban area and at Rio de Janeiro Zoological Garden (Pamplona et al. 2000;

Oliveira et al. 2002; Vasconcelos & Araújo 2012).

Material examined. ♂: Oxysarcodexia fluminensis sp 7 / COLOMBIA: Antioquia,

Copacabana, Ankon, 06°22‟1”N 75°29‟22,3”W 1.417m 4.viii.2010 Tr VSR 09:35 M. A.

Ramírez, I. Cadavid, C. Rave, Leg M08CP1 sp 7 (CE-TdeA).

Oxysarcodexia fraterna (Lopes, 1946)

Oxysarcodexia titubata ssp. fraterna Lopes, 1946

(Appendix 18-A)

Type locality. Mexico, Morelos, Cuernavaca. Lopes, 1946c: 143.


Diagnosis. Male. Postocular plate with golden microtomentum. Ocellar setae small,

but well-differentiated. Thorax grayish. Dorsocentral post-sutural setae with 3 well-

differentiated setae. Apical scutellar seta is present. Legs blackish. Abdomen grayish with

golden microtomentum more intense on T5. T4 with 1 pair of median marginal setae. ST5

reddish with a deep cleft. Cerci strongly bent backwards. Distiphallus with a large ventral lobe

and numerous pre-apical spines. This is a synopsis of the original description given by Lopes

(1946c).

Remarks. Cerci, in posterior view, convergent and with enlarged apexes, ventral

spines more developed and apex structural conformation of distiphallus are characteristics

used for differentiation of the very close related species O. peruviana and O. vittata (Lopes

1946c). See also “remarks” section on Oxysarcodexia n. sp. 4 for further discussion.

Distribution. NEARTIC. Mexico (Morelos).

Biology. Unknown. Information available is only about the type of collection, using

traps for collecting fruit flies (Lopes 1946c).

77

Material examined. ♂ terminalia mounted on a slide: O. titubata fraterna Lop №

8.101 cx: 55 Det. H. S. Lopes (MNRJ).

Oxysarcodexia fringidea (Curran & Walley, 1934)

Sarcophaga fringidea Curran & Walley, 1934

Oxysarcodexia lopesi Dodge, 1966

(Appendix 18-B)

Type locality. Guyana, Georgetown. Curran & Walley, 1934: 488.



well-developed. Thorax with pale golden microtomentum more evident at humeral portion.


Apical scutellar seta absent. Legs blackish. Abdomen with pale golden microtomentum. T3

with 3 lateral marginal setae. T4 with 1 median marginal and 3 lateral marginal setae. T5 with

golden microtomentum along the entire extension. ST5 with parallel cleft edges and with setae

on the apex of the arms. Cerci, in lateral view, straight with expanded apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) with same size as the middle

part. Conformation of the cerci in posterior view is parallel. In lateral view, distiphallus with

ventroapical concavity with smooth margins ventroapically; distiphallic apical shape conic and

straight distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until the

apex, which is darker than the base, as well as the postgonite, except for the color which is the

same along the entire extension. Vesica with asymmetrical branches; terminal lobes well-

developed, with rounded shape and partially membranous texture; presence of spines only

along the edges; and presence of an angular median projection of the main vesica branch.

Remarks. Lopes (1938) redescribed this species and pointed out the occurrence of

intraspecific variation on the size of the vesica (reduced sometimes), although the asymmetry

is always observed. See “remarks” section on O. aurata for further discussion. Tibana &

Mello (1985) classified O. fringidea female, according to the shape of the T6+7, as syntergite

membranous.

78

Distribution. NEOTROPICAL. Bolivia (Cochabamba), Brazil (Amapá, Amazonas,

Bahia, Colombia, Espírito Santo, Maranhão, Mato Grosso, Minas Gerais, Pará, Pernambuco,

Rio de Janeiro), Colombia, Guyana (Georgetown, Kartabo), Peru (Camchamayo, Tingo

Maria), Venezuela (Caracas, Aragua).

Biology. Human feces are indicated as a rearing substrate under natural conditions. At

laboratory, O. fringidea was reared on agar plus powder milk, developing from 1st instar to

adults in 14–17 days (Lopes 1973b). Human feces, cow liver and lung, fish and squid baits

have already been used for collect (Curran & Walley 1934; Couri et al. 2000; Sousa et al.

2011). Methods of collection reported are actively, with nets, or using baited or Shannon traps

(Couri et al. 2000; Pamplona et al. 2000).

Material examined. ♂: COLOMBIA: Amazonas, PNN Amacayacu Camino a San

Martin, 3°41‟N 70°15‟W, 1-10.iii.2004, sweepnet, T. Pape & D. Arias. Id#4325 / NRM-DIPT

0014659 (NRM).

Oxysarcodexia galeata (Aldrich, 1916)

Sarcophaga galeata Aldrich, 1916

(Appendix 18-C)

Type locality. USA, Indiana, Lafayette. Aldrich, 1916: 280.



developed. Thorax blackish with pale golden microtomentum more evident at humeral

portion. Dorsocentral post-sutural setae with 2 well differentiated and 1–3 anterior setae

smaller. Apical scutellar seta absent. Legs brownish. Abdomen blackish with some shades of

pale golden microtomentum. T3 with 1 lateral marginal seta. T4 with no median marginal and

1 lateral marginal seta. T5 with golden microtomentum along the entire extension, although

more intense at posterodorsal edge. ST5 with V-shaped cleft edges and with setosity and setae

on the apex of the arm. Cerci, in lateral view, sinuous with pointed apexes of oblique upwards

edges. Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical shape of

the cerci (last ⅓ portion of the cercus; in posterior view) with the same size as the middle part.

Conformation of the cerci in posterior view is parallel. Presence of a remarkable constriction

at middle portion of the cerci, in posterior view. In lateral view, distiphallus with smooth

79

margins ventroapically; distiphallic apical shape conic and straight distiphallic dorsal shape.

Presence of a large dorsoapical swelling on the distiphallus in lateral view. Lateroapical

expansions on distiphallus are present, in dorsal view. Pregonite with expanded base

narrowing smoothly until the apex, which is darker than the base. Postgonite with expanded

base and sudden narrowing at apex. Vesica with symmetrical branches; terminal lobes

reduced, with filamentous at most tapering to apex shape and sclerotized texture; presence of

spines along the edges; and presence of a rounded median projection of the main vesica

branch.

Remarks. See “remarks” section on O. aurata. Larvae of 1st, 2

nd and 3

rd instars were

described by Knipling (1936).

Distribution. NEARTIC. USA (Arkansas, Florida, Georgia, Illinois, Indiana, Iowa,

Kansas, New York, Tenessee, Texas).

Biology. Oxysarcodexia galeata has been collected in high lands (Dodge & Seago

1954) and in a wooded area, associated to a dog carcass (Reed 1958). Immatures normally

feed on excrement, however larvae obtained from one female were reared on decaying

hamburger meat and completed the development from larva to adult in 14 days (Knipling

1936).

Material examined. ♂: ALABAMA: Madson Co. 10km NE Maysville, Sharp Core,

Sneed Spg. Bottomland, 240m, 6 oct 1992, Acciavatti / NRM-DIPT 0014309 (NRM).

Oxysarcodexia grandis Lopes, 1946

(Appendix 19-A)

Type locality. Brazil, Rio de Janeiro, Angra dos Reis. Lopes, 1946b: 82.



developed. Thorax with intense golden microtomentum, as well as the abdomen. Dorsocentral

post-sutural setae with 3 well differentiated and 1 smaller seta among these can be present.

Apical scutellar seta present. Legs blackish. T4 with 1 lateral marginal seta. T5 with golden

microtomentum along the entire extension. ST5 with parallel cleft edges and with setosity.

Cerci, in lateral view, straight with pointed apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) present in apical ⅓ portion. Apical shape of the cerci (last ⅓ portion

80

of the cercus; in posterior view) with the same size as the middle part. Conformation of the

cerci in posterior view is parallel. In lateral view, distiphallus with ventroapical concavity with

smooth margins ventroapically; presence of a lateroapical furrow; distiphallic apical shape

rounded and sinuous distiphallic dorsal shape. Pregonite with expanded base and sudden

narrowing at apex with same color along the entire extension. Postgonite with expanded base

narrowing smoothly until the apex. Vesica with symmetrical branches; presence of lateral

lobes (i.e. division of the vesica coming from or close to the basal branch of the vesica, placed

laterally to phallic tube); terminal lobes well-developed, with filamentous at most tapering to

apex shape and sclerotized texture; presence of spines only on ventral surface; and presence of

a rounded median projection of the main vesica branch.

Remarks. Lopes (1946b) reported that dorsocentral post-sutural setae can, even for

species of a same colony, present variable number: 3 or 4 setae being the first 3 equidistant

and a smaller seta can be present among them. See also “remarks” section on O. augusta for

further discussion. Female was classified by Tibana & Mello (1985), according to the shape of

the T6+7, as syntergite undivided.

Distribution. NEOTROPICAL. Brazil (Paraná, Rio de Janeiro, São Paulo), Colombia

(Antioquia), Ecuador, Peru (Arequipa, San Miguel).


1973b), although further information of the development was not specified. Attraction by

human feces, chicken viscera and fish head is documented (Lopes 1973b; Ramírez-Mora et al.

2012). In a study of synanthropy of Sarcophagidae from Colombia carried out by Yepes-

Gaurisas et al. (2013), it is reported the attraction of O. grandis mainly by chicken viscera,

collecting specimens both in rural and in forest areas, although this fly has been considered

infrequent.

Material examined. ♂♂: O. grandis SP 3; Oxysarcodexia, Det. Alejandro Ramírez M

2010 / COLOMBIA: Antioquia, Medellín, Cola del Zorro, 06°22‟19,7”N 75°32‟43,9”W

1.943m 10.iii.2010 Tr VSR 10:05 Col. M. A. Ramírez / T. de A. 1132 (CE-TdeA) //

COLOMBIA: Antioquia, Medellín, Cola del Zorro, 06°22‟19,7”N 75°32‟43,9”W 1.943m

10.iii.2010 Tr VSR 10:05 Col. M. A. Ramírez / T. de A. 1132 / O. grandis (CE-TdeA).

Oxysarcodexia inflata Lopes, 1975

81

(Appendix 19-B)

Type locality. Brazil, Maranhão, Rosário, Igarapé, Paraqueú. Lopes 1975c: 470.


Diagnosis. Male. Postocular plate with silvery microtomentum. Ocellar setae not well-

developed. Thorax and abdomen with pale golden microtomentum, slightly more intense on

the abdomen. Dorsocentral post-sutural setae with 2 well differentiated and 2 anterior setae

smaller. Apical scutellar seta present. Legs brownish. T3 with 1 lateral marginal seta. T4 with

1 median marginal and 2 lateral marginal setae. T5 with golden microtomentum along the

entire extension. ST5 with parallel cleft edges and with setosity and setae on the apex of the

arms. Cerci, in lateral view, sinuous with expanded apexes of oblique upwards edges. Setae

ventrally on the cerci (lateral view) present in full extension. Apical shape of the cerci (last ⅓

portion of the cercus; in posterior view) smaller than the middle part. Conformation of the

cerci in posterior view is parallel. In lateral view, distiphallus with serrated margins

ventroapically and ventroapical projections; distiphallic apical shape rounded and straight

distiphallic dorsal shape. In dorsal view, presence of lateroapical expansions on distiphallus.

Pregonite with expanded base and sudden narrowing at apex, which is darker than the base.

Postgonite with expanded base narrowing smoothly until the apex and same color along the

entire extension. Vesica with symmetrical branches and presence of lateral lobes (i.e. division

of the vesica coming from or close to the basal branch of the vesica, placed laterally to phallic

tube); terminal lobes well-developed, with filamentous at most tapering to apex shape and

sclerotized texture; the median area of terminal lobes more expanded than basal and apical

areas; presence of spines only on ventral surface and presence of a rounded median projection

of the main vesica branch.

Remarks. See “remarks” section on O. amorosa. Female was described by Lopes

(1975c) as presenting ocellar setae well-developed, absence of apical scutellar setae and, as

seen in O. amorosa female, T7 weakly sclerotized on the center and with black setosity.

Distribution. NEOTROPICAL. Brazil (Maranhão, Pará, Pernambuco).

Biology. Unknown.

Material examined. ♂: Igarapé PARAQUEÚ Rosário, MARANHÃO BRASIL /

20/22-XI-70 / H. Berla / Paratype / Oxysarcodexia inflata n.sp. ♂ Det. H. S. Lopes (MNRJ).

82

Oxysarcodexia injuncta (Walker, 1857)

Sarcophaga injuncta Walker, 1857

Oxysarcodexia plaumanni Lopes, 1946

(Appendix 19-C)

Type locality. Brazil, São Paulo, Eugênio Lefevre. Lopes, 1946b: 102.



developed. Thorax with intense golden microtomentum. Dorsocentral post-sutural setae with 2

well differentiated and 1–3 anterior setae smaller. Apical scutellar seta present. Legs blackish.

Abdomen with silvery microtomentum. T3 with 2 lateral marginal setae. T4 with 1 median

marginal and 2 lateral marginal setae. T5 without golden microtomentum, only silvery

microtomentum is seen. ST5 with parallel cleft edges and with setae scattered on arm‟s

surface. Cerci, in lateral view, straight with expanded apexes of oblique upwards edges. Setae



cerci in posterior view is divergent. In lateral view, distiphallus with ventroapical concavity

with serrated margins ventroapically; presence of a lateroapical furrow; distiphallic apical

shape rounded and straight distiphallic dorsal shape. Lateroapical expansions, in dorsal view,

present on the distiphallus. Pregonite with expanded base and sudden narrowing at apex and

same color along the entire extension. Postgonite with expanded base narrowing smoothly

until the apex. Vesica with symmetrical branches; presence of lateral lobes (i.e. division of the

vesica coming from or close to the basal branch of the vesica, placed laterally to phallic tube);

terminal lobes well-developed, with filamentous at most tapering to apex shape and

sclerotized texture; presence of spines on both ventral and dorsal surfaces; and presence of a

rounded median projection of the main vesica branch.

Remarks. Oxysarcodexia injuncta is close related to O. paulistanensis, differing by the

golden microtomentum in the thorax, the straight dorsal shape of the phallus, the ventroapical

distiphallus shape, the vesica constitution and the apical cerci shape (Lopes 1946b). Female

was described by Lopes (1975b) and was classified by Tibana & Mello (1985), according to

the shape of the T6+7, as syntergite undivided. See also “remarks” section on O. bikini for

further discussion.

83

Distribution. NEOTROPICAL. Brazil (Mato Grosso, Mato Grosso do Sul, Minas

Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo).

Biology. This fly has already been reared under laboratory conditions, although no

further information about its development was found on the literature, except the

attractiveness by human feces (Lopes 1973b). Occurrence of this species is basically

associated to highlands, as pointed out by Lopes (1973b) and seen in our collection, once O.

injuncta was only collected in high hills areas (Serra do Japi, in São Paulo state, and Serra do

Lopo, in Minas Gerais state, both in Brazil).

Material examined. ♂♂: Rio Japura / Amazon Roman / april / Oxysarcodexia pr.

plaumanni, Lopes, Det. H. S. Lopes / NRM-DIPT 0014313 (NRM) // BRAZIL: São Paulo /

Jundiaí, Serra do Japi, / 06.I.2012 / M. D. Grella (L2B-DBA) // BRAZIL: Minas Gerais, /

Extrema, Serra do Lopo, / Pedra das Flores, / 27.II.2012 / A. G. Savino, M. P. Nassu, M. D.

Grella (L2B-DBA).

Oxysarcodexia insolita Lopes, 1946

(Appendix 20-A)

Type locality. Guuyana, Esequibo River, Moraballi Creek. Lopes, 1946b: 89.



well-developed. Thorax and abdomen with golden microtomentum more evident laterally.


Apical scutellar seta present. Legs brownish. T3 with 2 lateral marginal setae. T4 with 1

median marginal and 2 lateral marginal setae. T5 with golden microtomentum in part. ST5

with parallel cleft edges and with setosity and setae scattered on arm‟s surface. Cerci, in lateral

view, sinuous with expanded apexes of straight edges. Setae ventrally on the cerci (lateral

view) present in apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in

posterior view) bigger than the middle part. Conformation of the cerci in posterior view is

parallel. Presence of a remarkable constriction at middle portion of the cerci, in posterior view.

In lateral view, distiphallus with serrated margins ventroapically; distiphallic apical shape

rounded and straight distiphallic dorsal shape. Pregonite with expanded base narrowing

smoothly until the apex, which is darker than the base, as well as the postgonite, except for the

84

color which is the same along the entire extension. Vesica with symmetrical branches;


sclerotized texture; basal area of terminal lobes more expanded than the apical area; presence

of spines only on ventral surface; and presence of a rounded median projection of the main

vesica branch.

Remarks. Cerci, surstylus and phallus enable distinguish O. insolita from

Oxysarcodexia major Lopes, 1946, considered a close species. On distiphallus, ventroapically,

there is a serrated edge on O. insolita and a small group of spines on O. major, besides the

vesica, which is not long as seen in O. major, and presents long spines and a long and curved

basal area, absent on O. major whose spines are shorter (Lopes 1946b). Oxysarcodexia

petropolitana Lopes, 1975 is considered similar to O. insolita and O. major due to the vesica

shape, differing mostly by distiphallus apex and cerci shape (Lopes 1975c). Female was


undivided.

Distribution. NEOTROPICAL. Brazil (Pará), Guyana (Moraballi Creek), Mexico

(Chiapas, Veracruz), Trinidad and Tobago (Trinidad).

Biology. Under laboratory conditions, O. insolita was reared on agar plus powder milk,

developing from first instar to adults in 14–17 days. Under natural conditions, it has been

reared on human feces (Lopes 1973b).

Material examined. ♂: ECUADOR: Napo Povince; Yasuní National Park; Yasuní

Research Station; 76°36‟W 00°38‟S; 3–20 XI 1998: T. Pape & B. Viklund / NRM-DIPT

0014317 (NRM).

Oxysarcodexia intona (Curran & Walley, 1934)

Sarcophaga intona Curran & Walley, 1934

Sarcophaga intonsa: Lopes (1969), incorrect subsequent spelling of intona Curran &

Walley, 1934

(Appendix 20-B)

Type locality. Guyana, Kartabo. Curran & Walley, 1934: 489.


85




brownish. T4 with no median marginal and 3 lateral marginal setae. T5 with golden

microtomentum, although not seen along the entire extension. ST5 with parallel cleft edges

and with setae on the apex of the arms. Cerci, in lateral view, sinuous with expanded apexes of

concave edges. Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) with same size as the middle

part. Conformation of the cerci in posterior view is divergent. In lateral view, distiphallus with

smooth margins ventroapically; presence of ventroapical projections and lateral lobes;

distiphallic apical shape rounded and sinuous distiphallic dorsal shape. A large dorsoapical

swelling in the distiphallus (lateral view) is present. Pregonite with expanded base and sudden

narrowing at apex, which is darker than the base, as well as the postgonite, except for the color

which is the same along the entire extension. Vesica with symmetrical branches; terminal

lobes reduced, with filamentous at most tapering to apex shape and partially membranous

texture; presence of spines on both ventral and dorsal surfaces; and presence of an angular


Remarks. See “remarks” section on O. aurata. Female was classified by Tibana &


Distribution. NEOTROPICAL. Brazil (Amazonas, Amapá, Ceará, Espírito Santo,

Maranhão, Minas Gerais, Pará, Pernambuco, Rio de Janeiro), Guyana.

Biology. This fly has been collected associated to pig and rat carcasses, in Brazilian

Cerrado (“savanna-like”) vegetation, forest environments and an urban area (Cruz 2008;

Barbosa et al. 2009; Vasconcelos & Araújo 2012). Baits as feces, banana plus brown sugar,

rotting liver, cow liver and lung, fish and squid has been used with traps (W.O.T. or “can”

traps) (Lopes 1975a; Pamplona et al. 2000; Oliveira et al. 2002).

Material examined. ♂♂: RIO DE JANEIRO, BRASIL / R. Tibana / NRM-DIPT

0014318 (NRM) // Guarapari, Esp. Santo, Brasil / H. S. Lopes, 9.I.75 / Oxysarcodexia intona

♂ (C. et w.), det. H. S. Lopes (ZMUC).

Oxysarcodexia jamesi Dodge, 1968

86

(Appendix 20-C)

Type locality. Panama, Canal Zone, Barro Colorado Island. Dodge, 1968: 433.

Depository of type material. KU.


developed. Thorax and abdomen with golden microtomentum. Dorsocentral post-sutural setae

with 3 well differentiated setae and 1 anterior seta smaller. Apical scutellar seta present. Legs

brownish. T3 with 2 lateral marginal setae. T4 with 2 median marginal and 3 lateral marginal

setae. ST5 with parallel cleft edges and setosity. Cerci, in lateral view, straight with expanded

apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view) present in full

extension. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) smaller

than the middle part. Conformation of the cerci in posterior view is parallel. In lateral view,

distiphallus with membranous ventroapical concavity of smooth margins; distiphallic apical

shape conic and sinuous dorsal shape. Presence of a notable dilatation on the distiphallus

lateroapically. Pregonite with base and apex with same size and same color along the entire

extension. Postgonite with expanded base and sudden narrowing at apex and same color along

the entire extension. Vesica symmetric; terminal lobes reduced, with filamentous at most

tapering to apex shape, partially membranous texture and with micro-spines only on ventral

surface; presence of a rounded median projection of the main vesica branch.

Remarks. Cerci and somehow the distiphallus apical concavity are similar to those

present in Oxysarcodexia peculiaris Lopes, 1975, although vesica is very unlike. See also

“remarks” section on O. augusta for further discussion.

Distribution. NEOTROPICAL. Panama (Barro Colorado Island), Mexico.

Biology. Unknown. This fly specimen was collected from over a swarm raid of Eciton

burchelli (Westwood, 1842), an ant species (Dodge 1968).

Material examined. ♂: Santa Rosa National Park, Guanacaste Prov. COSTA RICA

Nov. 1998 300m D. H. Jansen & W. Hallwachs / Oxysarcodexia jamesi Dodge Det. H. S.

Lopes (MNRJ).

Oxysarcodexia major Lopes, 1946

(Appendix 21-A)

Type locality. Brazil, Rio de Janeiro, Guanabará, Grajaú. Lopes, 1946b: 88.

87



developed. Thorax and abdomen with intense golden microtomentum more evident laterally.


Apical scutellar seta present. Legs blackish. T3 with 2 lateral marginal setae. T4 with 1


with parallel cleft edges and with setosity and setae scattered on arm‟s surface. Cerci, in lateral

view, sinuous with normal apexes (i.e. same size as median area) of oblique upwards edges.

Setae ventrally on the cerci (lateral view) absent only in the middle portion. Apical shape of

the cerci (last ⅓ portion of the cercus; in posterior view) with the same size as the middle part.


at middle portion of the cerci, in posterior view. In lateral view, distiphallus with serrated

margins ventroapically; distiphallic apical shape rounded and straight distiphallic dorsal shape.

Pregonite with expanded base and sudden narrowing at apex, which is darker than the base, as

well as the postgonite, except for the color which is the same along the entire extension.

Vesica with symmetrical branches; terminal lobes well-developed, elongated, with

filamentous at most tapering to apex shape and sclerotized texture; presence of spines only on

ventral surface; and presence of an angular median projection of the main vesica branch.

Remarks. See “remarks” section on O. insolita. Female was classified by Tibana &


Distribution. NEOTROPICAL. Brazil (Amazonas, Amapá, Mato Grosso, Rio de

Janeiro, Roraima), Colombia (Antioquia), Ecuador, El Salvador, Mexico (Veracruz), Peru,

Trinidad and Tobago (Trinidad).

Biology. Collections of this species have been done with human feces, rotten Syagrus

comosa (Mart.) Mart. (Arecaceae), a species of a palmtree (at coastal area), rotten banana and

brown sugar, rotting liver, rotting beef lung, chicken viscera and fish head as baits (Lopes

1973b; Oliveira et al. 2002; Sousa et al. 2011; Ramírez-Mora et al. 2012). The methods of this

collections included flytraps, W.O.T., malaise and Shannon traps (Lopes & Tibana 1991;

Oliveira et al. 2002; Sousa et al. 2011). Oxysarcodexia major has already been reared under

laboratory conditions (Lopes 1973b), even though no detailed information about its

development had been reported.

88

Material examined. ♂: Oxysarcodexia bakeri? SP 8 / TdeA 1311 [from Colombia]

(CE-TdeA).

Oxysarcodexia marina (Hall, 1938)

Apelophyla marina Hall, 1938

(Appendix 21-B)

Type locality. Argentina, Mendoza. Hall, 1938: 257.

Depository of type material. DEI.


developed. Thorax and abdomen with silvery microtomentum. Three well differentiated

dorsocentral post-sutural setae. Apical scutellar seta absent. Legs blackish. T4 with presence

of small median marginal setae. ST5 with parallel cleft edges. Syntergosternite 7+8 black.

Cerci, in lateral view, bent backwards with expanded apexes of oblique upwards edges. Setae


portion of the cercus; posterior view) smaller than the middle part. Conformation of the cerci

in posterior view is divergent. Presence of a remarkable dilatation at middle portion of the

cerci, in posterior view. In lateral view, distiphallus with smooth margins ventroapically;

presence of lateral lobes and a large dorsoapical swelling; distiphallic apical shape

square/oblong and straight dorsal shape. Presence of lateroapical expansions on the

distiphallus, in dorsal view. Pregonite and postgonite with expanded base narrowing smoothly

until the apex and the same color along the entire extension. Vesica symmetric, with presence

of lateral lobes (i.e. division of the vesica coming from or close to the basal branch of the

vesica, placed laterally to phallic tube); terminal lobes well-developed, with square shape and

sclerotized texture; ventroapical area of terminal lobes notably elongated; presence of spines

along the edges (not continuously). This diagnosis is based on the examination of one

specimen (specified below) presenting some conservation issues (“oily” aspect compromising

original microtomentum and color, T5 partially broke and terminalia on glycerin lacking ST5)

and on diagnosis provided by Lopes (1975c).

Remarks. Male morphology and coloration of O. marina are considered similar to O.

paulistanensis, although O. marina presenting black color on syntergosternite 7+8 and

absence of apical scutellar setae (Mulieri et al. 2010) and we also pointed out the peculiarity

89

of O. marina terminalia seen on cerci shape and constitution of the vesica, especially the

ventroapical elongation. A good and description of female was given by Mulieri et al. (2010),

that also stress the sternites of female terminalia resemble in general shape and coloration

those of O. varia.

Distribution. NEOTROPICAL. Argentina (Buenos Aires, Mendoza), Brazil (Rio

Grande do Sul).

Biology. Attractiveness is reported by rotten bovine and cow liver and dog feces

(Mariluis et al. 2007; Mulieri et al. 2008; Mulieri et al. 2010; Mulieri et al. 2011). In Buenos

Aires, Argentina, O. marina has been collected in grassland and in rural areas (Mulieri et al.

2008; Mulieri et al. 2011). Adults of this fly are also considered flower visitors of Ludwigia

spp., a water primrose of Onagraceae family (Mulieri et al. 2010).

Material examined. ♂: Pelotas 7 Maio 1963 R. G. do Sul – Brasil C. M. Biezanko leg.

/ Apelophyla marina ♂ Hall Det. H. S. Lopes (MNRJ).

Oxysarcodexia meridionalis (Engel, 1931)

Sarcophaga (Dasyproctia) ventricosa var. meridionalis Engel, 1931

Sarcophaga illata Lopes, 1938

(Appendix 21-C)

Type locality. Argentina, Chaco, San José. Engel, 1931: 146.

Depository of lectotype species. SMN.


developed. Thorax and abdomen with golden microtomentum, although a slightly silvery

microtomentum is seen ventrally on the abdomen. Dorsocentral post-sutural setae with 2 well

differentiated and 2 anterior setae smaller. Apical scutellar seta absent. Legs blackish. T3 with

2 lateral marginal setae. T4 with 1 median marginal and 3 lateral marginal setae. T5 with

golden microtomentum along the entire extension. ST5 with parallel cleft edges and with

setosity and setae on the apex of the arms. Cerci, in lateral view, straight with pointed apexes

of straight edges. Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.

Conformation of the cerci in posterior view is parallel. In lateral view, distiphallus with

smooth margins ventroapically; distiphallic apical shape rounded and sinuous distiphallic

90

dorsal shape. Pregonite and postgonite with expanded base narrowing smoothly until the apex

and same along the entire extension. Vesica with symmetrical branches; terminal lobes well-

developed, mediolateral expanded, with rounded shape, sclerotized texture and presence of

spines only on ventral surface.

Remarks. It is considered probably as a member of “ventricosa” group, although the

lack of female study forbids this statement (Lopes 1975c). Phallus of O. meridionalis is

similar to that of O. parva, although cerci present slender and more pointed shape and vesica

terminal lobes are bigger, completely sclerotized and adorned with spines.

Distribution. NEOTROPICAL. Argentina (Chaco, Delta Del Paraná, San José),

Bolivia, Brazil (Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, São Paulo).

Biology. Unknown. The only information of O. meridionalis is the occurrence in

Cerrado (“savanna-like”) environment (Lopes 1973b) and on an urban area of Campinas city,

São Paulo state, Brazil (data from our research group).

Material examined. ♂: BRAZIL: São Paulo, Campinas, UNICAMP, 16.VII.2012, D.

Brancoli, A. G. Savino / II – 16/7 ? O. meridionalis (L2B-DBA).

Oxysarcodexia mitifica Lopes, 1953

(Appendix 22-A)

Type locality. Venezuela, Caracas. Lopes, 1953: 50.





blackish. T3 with lateral marginal seta. T4 with 1 pair of median marginal and 2 lateral

marginal setae. T5 with golden microtomentum. ST5 with parallel cleft edges and with

setosity. Cerci, in lateral view, straight with pointed apexes of oblique upwards edges. Setae

ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical shape of the cerci (last

⅓ portion of the cercus; in posterior view) with same size as the middle part. Conformation of

the cerci in posterior view is parallel. Presence of a remarkable constriction at middle portion

of the cerci, in posterior view. In lateral view, distiphallus with smooth margins

ventroapically; presence of a lateroapical furrow, distiphallic apical shape rounded and

91

sinuous distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until the


apex. Vesica with symmetrical branches; terminal lobe well-developed, with filamentous at

most tapering to apex shape, flattened laterally and with partially membranous texture;

presence of spines on both dorsal and ventral surfaces; and presence of an angular median


Remarks. The main difference in comparison to other Oxysarcodexia species is the

vesica, which presents a great development of median lobe (ventroapically directed) while

lateral lobes (laterally directed) are small.

Distribution. NEOTROPICAL. Colombia (Antioquia), Costa Rica, Ecuador, El

Salvador, Panama (Barro Colorado Island), Peru, Venezuela (Caracas).

Biology. This fly specimen was collected from over a swarm raid of Eciton burchelli

(Westwood, 1842), an ant species (Dodge 1968). Chicken viscera and fish head were baits

used to attract and collect them in Antioquia, Colombia (Ramírez-Mora et al. 2012).

Material examined. ♂: COLOMBIA: SP21 4CP2 / O. ramosa? (CE-TdeA) [from

Antioquia].

Oxysarcodexia modesta Lopes, 1946

(Appendix 22-B)

Type locality. Brazil, Rio de Janeiro, Guanabara, Gávea. Lopes, 1946b: 129.





absent. Legs brownish. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2

lateral marginal setae. T5 with golden microtomentum in part. ST5 with parallel cleft edges

and with setae on the apical half of the arms. Cerci, in lateral view, straight with pointed



than the middle part. Conformation of the cerci in posterior view is divergent. In lateral view,

distiphallus with the presence of a ventroapical concavity with serrated margins

92

ventroapically; distiphallic apical shape rounded and sinuous distiphallic dorsal shape;

presence of ventroapical projections. Pregonite with expanded base and sudden narrowing at

apex and same along the entire extension. Postgonite with expanded base narrowing smoothly

until the apex. Vesica with symmetrical branches; terminal lobes well-developed, with

filamentous at most tapering to apex shape and partially membranous texture; presence of

spines on both dorsal and ventral surfaces; and presence of a rounded median projection of the

main vesica branch.

Remarks. See “remarks” section on O. bakeri. Female was classified by Tibana &

Mello (1985), according to the shape of the T6+7, as syntergite partially divided into two

plates.

Distribution. NEOTROPICAL. Brazil (Minas Gerais, Rio de Janeiro, Pará,

Pernambuco, São Paulo), Peru.

Biology. Under laboratory conditions, O. modesta was reared on agar plus powder

milk, developing from 1st instar to adults in 14–17 days (Lopes 1973b). Adults attractiveness

has been observed by human feces, pig and mouse carcasses, chicken viscera (including liver),

rotting cow liver, fish, banana plus brown sugar and rotten Syagrus comosa (Mart.) Mart.

(Arecaceae), a species of a palmtree (at coastal area) (Lopes 1973b; Linhares 1981; Dias et al.

1984c; Mendes & Linhares 1993; Oliveira et al. 2002; Cruz 2008; Barbosa et al. 2009;

Oliveira & Vasconcelos 2010; Rosa et al. 2011). Comparing the attractiveness of human feces,

chicken viscera and mouse carcass baits, Linhares (1981) observed no differences among

these three substrates for O. modesta. Moreover, a lower occurrence in urban zone in contrast

to rural and forest areas analyzed, pointing out a low correlation with the anthropobiocenosis.

Sunlight instead of shaded areas is also referred as a preference of this species (Linhares

1981). This species has been caught on the premises of an institute of legal medicine, attracted

by chicken liver and classified as a necrophagous/omnivore species (Oliveira & Vasconcelos

2010). Adults have also been collected associated to a zoological garden in Rio de Janeiro

state, Brazil (Oliveira et al. 2002) and in urban and forest environments of Brazilian

northeastern (Vasconcelos & Araújo 2012).

Material examined. ♂♂: Quinta da Bôa Vista, São Cristóvão, Rio, Brasil / R. Tibana

21. VI.73 / NRM-DIPT 0014319 (NRM) // Rio das Ostras, E. do Rio, Brasil / 15-I-96, E. S.

93

Conrado col. / Oxysarcodexia modesta Lopes Det. R. Tibana (MNRJ) // Brasil: Amapá,

Mazagão, v–vi 1993 J. M. D‟Almeida col (MNRJ).

Oxysarcodexia molitor (Curran & Walley, 1934)

Sarcophaga molitor Curran & Walley, 1934

(Appendix 22-C)

Type locality. Guyana, Kaieteur. Curran & Walley, 1934: 488.


Diagnosis. Male. Ocellar setae not well-developed. Thorax and abdomen grayish with

golden microtomentum more intense laterally on thorax and on T4. Dorsocentral post-sutural

setae with 2 well differentiated and 2 anterior smaller setae. Apical scutellar setae present,

although very small. Legs blackish. T5 yellowish. ST5 small, with setosity, parallel cleft edges

and with small triangular projection on the posterior part of the arms. Cerci robust with

truncated apex. Anterior area of the apical distiphallic plate largely membranous, with a pair

of spinous internal plates. Vesica very sclerotized with an apical membranous lobe very

reduced. This is a synopsis of the original description given by Curran & Walley (1934) and of

the redescription given by Lopes (1975c).

Remarks. See “remarks” section on O. confusa. Female was described by Curran &

Walley (1934), presenting the first genital segment reddish with golden microtomentum,

sparse black setosity and an apical fringe of setae creased above.

Distribution. NEOTROPICAL. Guyana (Kartabo).

Biology. Unknown. The only information is about the substrate, human excrement, in

which the type species was collected on (Curran &Walley 1934).

Oxysarcodexia morretesi Tibana & Mello, 1983

(Appendix 23-A)

Type locality. Brazil, Paraná, São José dos Pinhais, Castelhanos. Tibana & Mello,

1983b: 279.



developed. Thorax and abdomen with grayish microtomentum. Dorsocentral post-sutural setae

94

with 2 well differentiated and 3 anterior setae smaller. Apical scutellar seta absent. Legs


setae. T5 with golden microtomentum along the entire extension. ST5 with parallel cleft edges

and with setosity along the edges of the arms. Cerci, in lateral view, straight with normal

apexes (same size as median area) of oblique upwards edges, due to a slight curvature and

with the mediolateral margin coming outwards. Setae ventrally on the cerci (lateral view)

present in apical half. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view)

with the same size as the middle part. Conformation of the cerci in posterior view is parallel.

Presence of a remarkable constriction at middle portion of the cerci, in posterior view. In

lateral view, distiphallus with presence of a ventromedial elongation; with smooth margins

ventroapically; distiphallic apical rounded shape and straight dorsal shape; presence of a small

“finger-like” swelling dorsoapically. Pregonite with expanded base narrowing smoothly until

the apex, which is darker than the base. Postgonite with expanded base and sudden narrowing

at apex. Vesica with symmetrical branches; terminal lobes well-developed, with oblong shape

and sclerotized texture; presence of spines only along the edges; and presence of an angular


Remarks. Similar to O. thornax due to the presence of the “finger-like” dorsoapical

swelling and vesica constitution. Differences are observed on cerci (with mediolateral margin

coming outwards and slight curved apex), distiphallus structures (with membranous

expansions extended ventrally and enlarged median process with apical spines) and vesica

(without presence of two enlarged lateral plates with serrated edges) (Tibana & Mello 1983b).

See also “remarks” section on O. conclausa for further discussion.

Distribution. NEOTROPICAL. Brazil (Rio de Janeiro, Paraná, São Paulo).

Biology. Unknown.

Material examined. ♂: Brazil, Município de Bertioga, S.P., 27–30 Nov. 72, B. V.

Peterson / Paratype / Oxysarcodexia ♂ n.sp. near thornax Det. H. S. Lopes (MNRJ).

Oxysarcodexia nitida Soares & Mello-Patiu, 2010

(Appendix 23-B)

Type locality. Peru: Avispas, Madre de Dios. Soares & Mello-Patiu, 2010: 72.


95


well-developed. Thorax and abdomen with golden microtomentum. Dorsocentral post-sutural

setae with 3 well differentiated setae, although a small seta among these 3 can be present.



with parallel cleft edges and with setosity and setae on the apex of the arm. Cerci, in lateral

view, straight with pointed apexes of oblique upwards edges. Setae ventrally on the cerci

(lateral view) present at basal half portion. Apical shape of the cerci (last ⅓ portion of the


view is divergent. In lateral view, distiphallus with smooth margins ventroapically; distiphallic

apical shape conic; straight distiphallic dorsal shape; and presence of lateral lobes. Presence of

lateroapical expansions on distiphallus in dorsal view. Pregonite with base and apex with same

size and the apex is darker than the base. Postgonite with expanded base narrowing smoothly

until the apex. Vesica with symmetrical branches; terminal lobes well-developed, with square

shape and sclerotized texture; without presence of spines.

Remarks. This species is considered very close morphologically to O. vittata, with

differences evident on vesica shape and adornment, distiphallus apex and pregonite (Soares &

Mello-Patiu 2010). In original description the apical scutellar is absent, although it is present

in the material here examined. See “remarks” section on Oxysarcodexia n. sp. 4 and O.

augusta for further discussion.

Distribution. NEOTROPICAL. Ecuador (Napo), Peru (Avispas).

Biology. Unknown.

Material examined. ♂: ECUADOR: Napo Province, Yasuní Research Station;

76°36‟W 00°38‟S; 3–20.XI.1998; T. Pape & B. Viklund / NRM-DIPT 0014471 (NRM).

Oxysarcodexia notata Soares & Mello-Patiu, 2010

(Appendix 23-C)

Type locality. Peru: Avispas, Madre de Dios. Soares & Mello-Patiu, 2010: 74.




96

with 3 well differentiated setae, although a small seta among these 3 can be present. Apical

scutellar seta present. Legs brownish. T3 with 1 lateral marginal seta. T4 with 1 median

marginal and 2 lateral marginal setae. T5 with golden microtomentum along the entire

extension. ST5 with parallel cleft edges and with setosity and setae scattered on arm‟s surface.

Cerci, in lateral view, straight with pointed apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) absent only in the middle portion. Apical shape of the cerci (last ⅓


cerci in posterior view is divergent. In lateral view, distiphallus with serrated margins

ventroapically; distiphallic apical shape square, sinuous distiphallic dorsal shape; and presence

of ventroapical projections and lateral lobes. Presence of lateroapical expansions on

distiphallus in dorsal view. Pregonite and postgonite with expanded base narrowing smoothly

until the apex and same color along the entire extension. Vesica with symmetrical branches;


sclerotized texture; presence of micro-spines both on dorsal and ventral surfaces; and presence

of a rounded median projection of the main vesica branch.

Remarks. Male terminalia is considered similar to that presented by O. xon, with

differences observed on vesica and distiphallus morphologies (Soares & Mello-Patiu 2010).

See “remarks” section on Oxysarcodexia n. sp. 4 and O. augusta for further discussion.

Biology. Unknown.

Distribution. NEOTROPICAL. Ecuador (Napo), Peru (Avispas).

Material examined. ♂: ECUADOR: Napo Province, Yasuní National Park, Yasuní

Research Station; 76°36‟W 00°38‟S; 3–20.XI.1998; T. Pape & B. Viklund / NRM-DIPT

0014466 (NRM).

Oxysarcodexia occulta Lopes, 1946

(Appendix 24-A)

Type locality. Brazil, Rio de Janeiro, Guanabara, Corcovado. Lopes, 1946b: 112.


Diagnosis. Male. Postocular plate golden microtomentum. Ocellar setae not well-

developed. Thorax and abdomen with intense golden microtomentum. Dorsocentral post-


97

absent. Legs blackish. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral

marginal setae. T5 with golden microtomentum in part. ST5 with parallel cleft edges and with

setosity. Cerci, in lateral view, bent backwards with normal apexes (same size as median area)

of oblique upwards edges. Setae ventrally on the cerci (lateral view) present in apical ⅓

portion. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) with the

same size as the middle part. Conformation of the cerci in posterior view is divergent. In

lateral view, distiphallus with smooth margins ventroapically; distiphallic apical shape

rounded and sinuous distiphallic dorsal shape. Pregonite with expanded base and sudden

narrowing at apex, which is darker than the base, as well as the postgonite, except for the color

which is the same along the entire extension. Vesica with symmetrical branches; terminal

lobes well-developed, with filamentous at most tapering to apex shape, flattened

anteroposteriorly and partially membranous texture; without spines; and with the presence of


Remarks. See “remarks” section on O. comparilis. Female was classified by Tibana &


Biology. In laboratory, O. occulta was reared on agar plus powder milk, developing

from first instar to adults in 14–17 days (Lopes 1973b). In natural environment, adults has

been caught with banana and brown sugar as bait and larvae, obtained from one female of the

collected specimens, were reared in human feces, taken 4 days to reach 3rd

larval instar plus 13

days to emerge adults (Lopes 1975a). This fly has also been collected from over a swarm raid

of Eciton burchelli (Westwood, 1842), an ant species (Dodge 1968). Attractiveness is reported

by human feces and rotten banana plus brown sugar (Lopes 1973b). Methods of collection

used for this species include malaise and Shannon traps (Lopes & Tibana 1991), traps for

collecting butterflies (Lopes 1975a) and actively, using baits to attract them and sweepnet for

catching (our collection).

Distribution. NEOTROPICAL. Brazil (Ceará, Mato Grosso, Mato Grosso do Sul,

Pará, Rio de Janeiro, Roraima, São Paulo), Colombia, Ecuador, Panama (Barro Colorado

Island).

Material examined. ♂♂: BRAZIL: São Paulo, Mogi Guaçu, Campininha, 18.IX.2011,

C. G. P. Lima, M. D. Grella, N. M. Jimenez / Oxysarcodexia sp III, Mogi Guaçu-SP,

18/11/2011, 16 (L2B-DBA); BRAZIL: São Paulo, Mogi Guaçu, Campininha, 18.IX.2011, C.

98

G. P. Lima, M. D. Grella, N. M. Jimenez / Oxysarcodexia sp III, Mogi Guaçu-SP, 18/11/2011,

16 (L2B-DBA).

Oxysarcodexia orbitalis Dodge, 1966

(Appendix 24-B)

Type locality. Martinique, Mont Pélé, 2 km N Morne Range. Dodge, 1966: 686.


Diagnosis. Male. Postocular plate with golden microtomentum. Ocellar seta not well-

developed (“hair-like”). Presence of 1 proclinate frontorbital seta. Thorax black with

yellowish-gray microtomentum on humeral region and laterally. Four dorsocentral post-sutural

setae. Apical scutellar seta is absent. Legs black. Abdomen black with grayish

microtomentum. T3 without median marginal row of seta. ST5 with arms crossed internally.

Cerci elongated, fused to apexes, which are shining black bosses and with sinuous shape

(lateral view). Pregonite strongly curved and postgonite straighter. Phallus with usual lobes

and distally strongly compressed. Vesica symmetrical, short, “ear-shaped”, with bulbous base

(in lateral view, a hole in the middle) and spinous arms. This is a synopsis of original

description given by Dodge (1966).

Remarks. Oxysarcodexia orbitalis is the unique species of the entire genus referred

presenting proclinate frontorbital seta (Dodge 1966). See “remarks” section on O. cyanea for

further discussion.

Distribution. NEOTROPICAL. Martinique.

Biology. Unknown. The only information is about the collection of the type species,

made on guava (Dodge 1966).

Oxysarcodexia pallisteri Dodge, 1966

(Appendix 24-C)

Type locality. Peru, Machu Picchu, Pueblo Cusa. Dodge, 1966: 686.



well-developed. Thorax with pale golden microtomentum. Dorsocentral post-sutural setae

with 2 well differentiated and 2 anterior setae smaller. Apical scutellar seta present, although

99

small. Legs brownish. Abdomen with silvery microtomentum. T3 with 1 lateral marginal seta.

T4 with 1 median marginal and 2 lateral marginal setae. T5 without golden microtomentum,

only silvery microtomentum is present. ST5 with parallel cleft edges and with setosity and

setae on the apex of the arms. Cerci, in lateral view, sinuous with pointed apexes of straight

edges. Setae ventrally on the cerci (lateral view) absent in the middle portion. Apical shape of

the cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.


ventroapical concavity of smooth margins and presence of serrated margins ventroapically;

presence of lateral lobes, ventroapical projections and medium dorsoapical swelling;

distiphallic apical shape square/oblong and sinuous distiphallic dorsal shape. Presence of

lateroapical expansions, in dorsal view. Pregonite and postgonite with expanded base

narrowing smoothly until the apex and same color along the entire extension. Vesica

symmetric; terminal lobes well-developed, with rounded shape, sclerotized texture and

presence of spines only on ventral surface.

Remarks. See “remarks” section on Oxysarcodexia n. sp. 4.

Distribution. NEOTROPICAL. Peru (Machu Pichu), Brazil (Paraná).

Biology. Unknown.

Material examined. ♂: IGUASSÚ Paraná XII - 941 Com. E. N. V. / Xarcophaga /

pallisteri (MNRJ).

Oxysarcodexia parva Lopes, 1946

(Appendix 25-A)

Type locality. BRASIL, RIO DE JANEIRO, Rio de Janeiro, Grajaú. Lopes, 1946b: 97.



developed. Thorax grayish with pale golden microtomentum only laterally. Dorsocentral post-


absent. Legs blackish. Abdomen with silvery microtomentum. T3 with 1 lateral marginal seta.

T4 with 1 median marginal and 1 lateral marginal setae. T5 with golden microtomentum in

part. ST5 with parallel cleft edges and with setae scattered on arm‟s surface. Cerci, in lateral

view, sinuous with pointed apexes of oblique upwards edges. Setae ventrally on the cerci

100

(lateral view) absent only in the middle portion. Apical shape of the cerci (last ⅓ portion of the


view is parallel. Presence of a remarkable constriction at middle portion of the cerci, in

posterior view. In lateral view, distiphallus with presence of a ventroapical concavity with

smooth margins; distiphallic apical shape rounded, sinuous distiphallic dorsal shape. Presence

of lateroapical expansions on distiphallus in dorsal view. Pregonite with expanded base

narrowing smoothly until the apex, which is darker than the base, as well as the postgonite,

except for the color which is the same along the entire extension. Vesica with symmetrical

branches; terminal lobes reduced, with rounded shape and membranous texture; without

spines; and with presence of an angular median projection of the main vesica branch.


not uncommonly misidentified, O. avuncula, O. confusa, O. diana and O. parva, is given by

Silva & Mello-Patiu (2008). Female was classified by Tibana & Mello (1985), according to

the shape of the T6+7, as syntergite undivided. See also “remarks” section on O. comparilis,

O. edwardsi and O. meridionalis for further discussion.

Distribution. NEOTROPICAL. Argentina (Jujuy, Misiones), Brazil (Ceará, Mato

Grosso, Minas Gerais, Rio de Janeiro, Paraná, São Paulo).

Biology. This species has been reared on human feces (Lopes 1973b; D‟Almeida

1994). Agar plus powder milk has been used successfully as artificial rearing source for O.

parva under laboratory conditions (Lopes 1973b). In our laboratory, it was reared on minced

bovine meat with larval stage during 6–7 days and pupal stage, until adult emergence, taking

9–11 days. Attractiveness has been already recorded by human feces, chicken viscera, fish,

mouse and pig carcasses, banana plus brown sugar and rotten Syagrus comosa (Mart.) Mart.

(Arecaceae), a species of a palmtree (at costal area) (Lopes 1973b; Lopes 1975a; Dias et al.

1984a; Barbosa et al. 2009; Vairo et al. 2011). Dias et al. (1984a) collected this species in

urban, rural and forest areas, although higher occurrence was observed only for the last

environment, considering O. parva as an asynanthropic species. It was also observed higher

preference for human feces comparing to the other substrates (chicken viscera, raw fish, mice

carcass and banana plus brown sugar) analyzed (Dias et al. 1984c).

101

Material examined. ♂: BRAZIL: São Paulo, Campinas, Sousas, 13.IV.2011, C. M.

Souza, D. L. Brancoli, F. Rezende / ??? 4 Sousas, Campinas-SP, 13/04/2011 / Oxy sp 1 (L2B-

DBA).

Oxysarcodexia paulistanensis (Mattos, 1919)

Sarcophaga paulistanensis Mattos, 1919

Oxysarcodexia amarali Prado & Fonseca, 1932

Oxysarcodexia delpontei Blanchard, 1939

Oxysarcodexia artigasi Dodge, 1966

Oxysarcodexia artegasi: Lopes (1973), incorrect subsequent spelling of artigasi

Dodge, 1966

(Appendix 25-B)

Type locality. Brazil, São Paulo. Mattos, 1919: 72.

Depository of type material. MZUSP.



with 2 well differentiated and 1–3 anterior setae smaller. Apical scutellar seta present. Legs


setae. T5 without golden microtomentum. ST5 with parallel cleft edges and with setosity.

Cerci, in lateral view, sinuous with expanded apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) present in full extension. Apical shape of the cerci (last ⅓ portion of

the cercus; in posterior view) smaller than the middle part. Conformation of the cerci in

posterior view is divergent. In lateral view, distiphallus with ventroapical concavity with

serrated margins; distiphallic apical shape rounded, sinuous distiphallic dorsal shape; and a

small distiphallic dorsoapical swelling. Pregonite and postgonite with expanded base

narrowing smoothly until the apex and same color along the entire extension. Vesica with

symmetrical branches; terminal lobes well-developed, filamentous at most tapering to apex

shape, sclerotized texture and with presence of lateral lobes (i.e. division of the vesica coming

from or close to the basal branch of the vesica, placed laterally to phallic tube); presence of

102

spines only on ventral surface; and presence of a rounded median projection of the main

vesica branch.

Remarks. See “remarks” section on O. bikini, O. injuncta and O. marina. Female was


partially divided into two plates. Larval morphology of 1st, 2

nd and 3

rd instars was described by

Lopes (1943) and studied ultrastructurally, using scanning electron microscope, by Lopes &

Leite (1987).

Distribution. NEOTROPICAL. Argentina (Buenos Aires, Córdoba, Entre Ríos),

Brazil (Distrito Federal, Goiás, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul,

Santa Catarina, São Paulo), Chile (Santiago).

Biology. Oxysarcodexia paulistanensis has been reared under laboratory conditions on

agar plus powder milk for 24h, then transferred to meat; and under natural conditions, on

human feces – which is the preferred substrate for rearing (Lopes 1973b; Mendes & Linhares

1993) –, rat carcas (Mendes & Linhares 1993; Moura 2004; Moura et al. 2005) and chicken

viscera (Mendes & Linhares 1993). Time of development reported in the literature for this fly

is 8 days of larval stage and 6–9 days in pupa, until the emergence of adults (Lopes 1943). At

our laboratory, it was reared on minced bovine meat, although larviposition was observed also

on rotten fish, with larval stage during 5–7 days and pupal stage, until adult emergence, taking

8–12 days. Baits as human and dog feces, chicken viscera, rotten bovine liver, fish, mouse, rat

and pig carcasses and banana plus brown sugar have been reported as attractive for this

species (Lopes 1973b; Ferreira et al. 1980; Dias et al. 1984c; Mendes & Linhares 1993;

Carvalho & Linhares 2001; Moura et al. 2005; Barros et al. 2008; Rosa et al. 2011; Vairo et

al. 2011; Beuter et al. 2012). In a study of synanthropy of flesh flies, Linhares (1981)

observed higher attraction of O. paulistanensis by human feces and mouse carcasses than to

chicken viscera, and also pointed out this species preference for sunlight instead of shaded

areas. The higher frequency of adult females on chicken viscera and rodent carcasses baits

suggested the use of these substrates as proteic resources for the development of ovarian

folicules (Mendes & Linhares 1993). Oxysarcodexia paulistanensis occurring at the coastline

of Buenos Aires was analysed taking into account its seasonal trends in abundance, habitat and

bait attractiveness, evidencing this species positive correlation with mean temperature,

preference by grassland instead of woodland and by dog feces instead of bovine liver (Mulieri

103

et al. 2008). This fly has been found both in urban, suburban and rural areas (Mulieri et al.

2011; Beuter et al. 2012). It is also considered a flower visitor of Coriandrum sativum L.

(Apiaceae), known as coriander or cilantro, and of Condalia spp., Discaria Americana Gillies

& Hook, Scutia buxifolia Reissek, shrubs of Rhamnaceae family (Mulieri et al. 2010).

Material examined. ♂♂: BRAZIL: São Paulo, Campinas, Sousas, 13.IV.2011, C. M.

Souza, D. L. Brancoli, F. Rezende / O. paulistanensis, Sousas, Campinas-SP, 13/04/2011

(L2B-DBA) // BRAZIL: São Paulo, Campinas, UNICAMP, IX.2010, C. M. Souza / O.

paulistanensis, Campinas-SP, SET/2010 (L2B-DBA).

Oxysarcodexia peculiaris Lopes, 1975

(Appendix 25-C)

Type locality. Brazil, Espírito Santo Linhares. Lopes, 1975c: 477.



developed. Thorax with pale golden microtomentum. Dorsocentral post-sutural setae with 2

well differentiated and 2 anterior setae smaller. Apical scutellar seta present. Legs blackish.

Abdomen with silvery microtomentum, although some shades of pale golden microtomentum

is seen especially on T4 and T5. T3 with 2 lateral marginal setae. T4 with 1 median marginal

and 3 lateral marginal setae. T5 with golden microtomentum in part. ST5 with parallel cleft

edges and with setae on the apex of the arms. Cerci, in lateral view, sinuous with expanded

apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view) present in apical

⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) smaller


distiphallus with ventroapical concavity with smooth margins ventroapically; presence of

ventroapical projections; distiphallic apical shape conic and sinuous distiphallic dorsal shape.

Presence of a notable dilatation on the distiphallus lateroapically and membranous dilatations

medially on the phallus. Pregonite with expanded base narrowing smoothly until the apex and

same color along the entire extension. Postgonite with expanded base and sudden narrowing at

apex and same color along the entire extension. Vesica with symmetrical branches; terminal

lobes well-developed, with rounded shape and sclerotized texture; presence of spines only on


104

Remarks. See “remarks” section on O. jamesi.

Distribution. NEOTROPICAL. Brazil (Espírito Santo, Rio de Janeiro).

Biology. The paratype species was reared from human feces (Lopes 1975c), however

without mention any further information about its development.

Material examined. ♂: Linhares, E. Santo, Brasil / P. C. Elias VI-72 / Holotype /

Oxysarcodexia peculiaris ♂ n.sp. Det. H. S. Lopes / MNRJ/2249 (MNRJ).

Oxysarcodexia peltata (Aldrich, 1916)

Sarcophaga peltata Aldrich, 1916

(Appendix 26-A)

Type locality. Puerto Rico, Mayaguez. Aldrich, 1916: 216.





brownish. T3 with 1 lateral marginal seta. T4 without median marginal and with 1 lateral

marginal seta. T5 with golden microtomentum in part. ST5 with parallel cleft edges and with

setae scattered on the apex of the arms. Cerci, in lateral view, bent backwards with normal

apexes (same size as the median area) of concave edges. Setae ventrally on the cerci (lateral

view) present on apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in


divergent. In lateral view, distiphallus with smooth margins ventroapically; distiphallic apical

conic shape, straight distiphallic dorsal shape; presence of a large dorsoapical swelling and

lateral lobes on distiphallus. Pregonite with expanded base and sudden narrowing at apex,

which is darker than the base. Postgonite with expanded base narrowing smoothly until the



texture; presence of spines on both dorsal and ventral surfaces; and presence of an angular


Remarks. See “remarks” section on O. aurata. Female was classified by Tibana &


105

Distribution. NEARTIC. USA (Florida). NEOTROPICAL. Bahamas (Andros, Cat I,

Eleuthera, Grand Bahama, Great Inagua, New Providence), Cuba (Havana), Dominica,

Guadalupe, Jamaica, Mexico (Jalisco, Tabasco), Panama, Puerto Rico, San Andrés Islands,

Santa Lucia, San Vicente.

Biology. Despite of the type-species status for the genus Oxysarcodexia, the only

information about O. peltata biology is the relationship with Laguncularia racemosa (L.) c.f.

Gaertn (Combretaceae), being a probable highly effective pollinator of this mangrove species

(Sánchez-Núñez & Mancera-Pineda 2012).

Material examined. ♂: Est. Cent. Agr. De Cuba 8089 / Sarcophaga peltata Ald. / R.

Panke/1/9/1920 (ZMUC).

Oxysarcodexia perneta (Walker, 1861)

Sarcophaga perneta Walker, 1861

(Appendix 26-B)

Type locality. Mexico. Walker, 1861: 308.





brownish. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral marginal

setae. T5 with golden microtomentum along the entire extension. ST5 with parallel cleft edges

and with setosity and setae on the apex of the arms. Cerci, in lateral view, straight with pointed



than the middle part. Conformation of the cerci in posterior view is parallel. Presence of a

remarkable constriction at middle portion of the cerci, in posterior view. In lateral view,

distiphallus with presence of a ventroapical concavity of smooth margins; distiphallic apical

rounded shape; straight distiphallic dorsal shape; and presence of a small dorsoapical swelling

on distiphallus. Pregonite and postgonite with expanded base narrowing smoothly until the


lobes well-developed, with rounded shape and membranous texture; presence of spines on

106

both dorsal and ventral surfaces; and presence of a rounded median projection of the main

vesica branch.

Remarks. Lopes (1946b) indicated differences on vesica appearance due to specimen

preparation. In dry pinned flies, it can seem juxtaposed to the distiphallus, whereas in wet

preparation or dried properly, the vesica become detached from distiphallus and examination

of distiphallus apical shape and vesica terminal lobes can be easier done. See “remarks”

section of Oxysarcodexia n. sp. 3.

Distribution. NEARTIC. Mexico (Districto Federal, Morelos). NEOTROPICAL.

Honduras, Mexico (Oxaca).

Biology. Unknown. Information available is only about the type of collection, active,

using an entomological net, or with traps for collecting fruit flies (Lopes 1946c).

Material examined. ♂♂: MEX. Chiapas, San Cristobal 7000m, 4 May 1969, H. J.

Teskey / Oxysarc.perneta (Wlk.), det. Lopes (ZMUC) // Col. Inst. O. Cruz N. 9.361 / Mexico

D.F., A. Dampf [unreadable number] / Oxysarcodexia perneta Wulp ♂ xi-44 Det. H. S. Lopes

(MNRJ) // Cuernavaca Est. Morelos Mexico 1800m Dampf 1.IX a 5.XII / Oxysarcodexia

perneta (Walker) Det. H. S. Lopes ♂ (MNRJ).

Oxysarcodexia peruviana (Lopes, 1975)

Xarcophaga titubata ssp. peruviana Lopes, 1975

(Appendix 26-C)

Type locality. Peru, Lambayeque, La Beatita. Lopes, 1975f: 575.

Depository of type material. UNPRG.

Diagnosis. Male. Postocular plate with intense golden microtomentum. Ocellar setae

well-developed. Thorax gray with golden microtomentum more intense on humeral region.

Dorsocentral post-sutural setae with 3 well-differentiated setae. Presence of apical scutellar

seta, small though. Abdomen with golden microtomentum more intense laterally on T4 and

T5. T4 without median marginal setae. ST5 reddish. Cerci curved backwards (lateral view).

Phallus similar to the typical form, but apical and lateral plates besides vesica with different

shapes. This is a synopsis of original description given by Lopes (1975f).

Remarks. See “remarks” section on Oxysarcodexia n. sp. 4, O. augusta and O.

fraterna.

107

Distribution. NEOTROPICAL. Colombia (Antioquia), Peru (Lambayeque).

Biology. Unknown.

Oxysarcodexia petropolitana Lopes, 1975

(Appendix 27-A)

Type locality. BRAZIL: Rio de Janeiro. Lopes, 1975c: 471.




differentiated and 3 anterior setae smaller. Apical scutellar seta absent. Legs brownish.


is seen laterally. T3 with 2 lateral marginal setae. T4 with 1 median marginal and 2 lateral

marginal setae. T5 with golden microtomentum in part. ST5 with parallel cleft edges,

triangular apical shape of the arms and with setosity scattered on arms‟ surface. Cerci, in

lateral view, straight with normal apexes (same size as the median area) of concave edges.

Setae ventrally on the cerci (lateral view) present in apical ⅓ portion. Apical shape of the cerci

(last ⅓ portion of the cercus; in posterior view) with the same size as the middle part.


at middle portion of the cerci, in posterior view. In lateral view, distiphallus with smooth

margins ventroapically; distiphallic apical shape rounded and sinuous distiphallic dorsal

shape. Pregonite with a mediobasal projection (“thorn-like”) and expanded base narrowing

smoothly until the apex and same color along the entire extension. Postgonite with expanded

base and sudden narrowing at apex and same color along the entire extension. Vesica with

symmetrical branches; presence of lateral lobes (i.e. division of the vesica coming from or

close to the basal branch of the vesica, placed laterally to phallic tube); terminal lobes well-

developed, with sclerotized texture, filamentous at most tapering to apex shape, with basal

portion enlarged and apexes pointed backwards; presence of spines only on ventral surface;

and presence of a rounded median projection of the main vesica branch.

Remarks. See “remarks” section on O. insolita. Female was described by Lopes

(1975c) with golden microtomentum on T6+7, which is composed by two plates; sternites

108

reddish and anal tergite represented by a pair of setae. It was classified by Tibana & Mello

(1985), according to the shape of the T6+7, as syntergite partially divided into two plates.

Biology. Species already reared on human feces and on meat (Lopes 1973b; Lopes

1975c), although no other details of its development are reported. Attractiveness is reported

only for human feces (Lopes 1973b).

Distribution. NEOTROPICAL. Brazil (Minas Gerais, Rio de Janeiro, Santa Catarina,

São Paulo), Ecuador.

Material examined. ♂: 6 / Estr. S. Paulo – Santos, Brasil 3.II.68 R. Kano / Paratype /

Oxysarcodexia petropolitana Lopes H. S. Lopes det. (MNRJ).

Oxysarcodexia plebeja Lopes, 1946

(Appendix 27-B)

Type locality. Mexico, Morelos, Cuernavaca. Lopes, 1946c: 142.



developed. Thorax with golden microtomentum more intense laterally. Dorsocentral post-

sutural setae with 3 well differentiated setae, although a small seta among these 3 can be

present. Apical scutellar seta present. Legs blackish. Abdomen with intense golden

microtomentum. T4 with 2 lateral marginal setae. T5 with golden microtomentum along the

entire extension. ST5 with parallel cleft edges and with setosity and setae on the apex of the

arms. Cerci, in lateral view, straight with expanded apexes of oblique upwards edges. Setae

ventrally on the cerci (lateral view) present on apical half. Apical shape of the cerci (last ⅓


cerci in posterior view is parallel. In lateral view, distiphallus with presence of a ventroapical

concavity of serrated margins; distiphallic apical rounded shape; straight distiphallic dorsal

shape. Pregonite and postgonite with expanded base narrowing smoothly until the apex and

same color along the entire extension. Vesica with symmetrical branches; terminal lobes

reduced, with lateral expansions of filamentous at most tapering to apex shape and sclerotized

texture; presence of spines only on ventral surface of the median lobe; and presence of a


109

Remarks. Vesica shape is very peculiar and different from the typical shape seen into

the genus. See also “remarks” section on O. augusta.

Distribution. NEARTIC. Mexico (Morelos, San Luis Potosí), USA (Texas).

NEOTROPICAL. Colombia (Antioquia), Costa Rica, Mexico (Chiapas).

Biology. Original description gives information only about the type of collection, using

traps for collecting fruit flies, without further specification (Lopes 1946c). At Antioquia,

Colombia, this species was collected using chicken viscera and fish head as attractive baits

(Ramírez-Mora et al. 2012).

Material examined. ♂♂: SP 25 / Pajar. Mar 12 / O. plebeja ? (CE-TdeA) [from

Colombia] // MEX. Chis. 32mi W. San Cristobal Jct 190–195 Hwys. 12, V, 1969 H. J. Teskey

/ Oxysarc. plebeja ♂ Lopes Det. H. S. Lopes (MNRJ).

Oxysarcodexia ramosa (Reinhard, 1939)

Sarcophaga ramosa Reinhard, 1939

(Appendix 27-C)

Type locality. USA, Texas, Donnas. Reinhard, 1939: 64.


Diagnosis. Male. Head yellowish-gray. Ocellar setae not well-developed. Thorax

grayish. Four dorsocentral post-sutural setae. Apical scutellar seta is absent. Legs blackish.

Abdomen yellowish-gray. T5 reddish. ST5 reddish and with a U-shaped cleft. Phallus with

vesica of asymmetrical branches; presence of flattened squamas on the smaller branch. This is

a synopsis of diagnoses given by Lopes (1946b).

Remarks. Oxysarcodexia ramosa is one of the species of this genus that have a

noteworthy asymmetrical vesica.

Distribution. NEARTIC. USA (Texas).

Biology. Unknown.

Oxysarcodexia riograndensis Lopes, 1946

(Appendix 28-A)

Type locality. Brazil, Rio Grande do Sul, São João de Montenegro. Lopes, 1946b:

103.

110






setae. T5 without golden microtomentum. ST5 with parallel cleft edges and with setosity.

Cerci, in lateral view, straight with expanded apexes of oblique upwards edges. Setae ventrally

on the cerci (lateral view) absent only on the middle portion. Apical shape of the cerci (last ⅓


cerci in posterior view is divergent. In lateral view, distiphallus with ventroapical concavity of

serrated margins; distiphallic apical square/oblong shape, sinuous distiphallic dorsal shape;

presence of a medium dorsoapical swelling on distiphallus. Pregonite and postgonite with

expanded base narrowing smoothly until the apex and same color along the entire extension.

Vesica with symmetrical branches; presence of lateral lobes (i.e. division of the vesica coming

from or close to the basal branch of the vesica, placed laterally to phallic tube); terminal lobes

well-developed, with rounded shape, sclerotized texture and spines present only on ventral

surface; and presence of a rounded median projection of the main vesica branch.

Remarks. See “remarks” section on O. bikini. Female was classified by Tibana &


Distribution. NEOTROPICAL. Argentina (Jujuy), Brazil (Minas Gerais, Paraná,

Pernambuco, Rio de Janeiro, Rio Grande do Sul, São Paulo).

Biology. This species has been reared from a corpse on bloated stage and collected on

the premises of an institute of legal medicine using chicken liver as bait and, for this,

considered as necrophagous/omnivore species (Oliveira & Vasconcelos 2010). At our

laboratory, it was reared only once, on minced bovine meat with larval stage during 6 days and

pupal stage, until adult emergence, taking 9 days. In a study comparing the attractiveness of

baits, Linhares (1981) observed that human feces and mouse carcasses were more attractive

than chicken viscera for O. riograndensis, pattern corroborated by Mendes & Linhares (1993).

Sunlight instead of shaded areas is also referred as a preference of this species (Linhares

1981). This fly has also been collected using mice, rat, rabbit and pig carcasses, fish, chicken

liver and human feces as baits (Ferreira 1979; Mendes & Linhares 1993; Carvalho & Linhares

111

2001; Cruz 2008; Moretti et al. 2008; Rosa et al. 2011; Vairo et al. 2011; Vasconcelos &

Araújo 2012). Adult specimens have been found both in urban, rural or forest environments

(Linhares 1981; Mendes & Linhares 1993; Rosa et al. 2011; Vairo et al. 2011; Vasconcelos &

Araújo 2012).

Material examined. ♂: BRAZIL: São Paulo, Campinas, UNICAMP, 18.VII.2010, C.

M. Souza / O. riograndensis, Campinas-SP, 18/08/2010 (L2B-DBA).

Oxysarcodexia sarcinata Lopes, 1953

(Appendix 28-B)

Type locality. Colombia, Medellín. Lopes, 1953: 48.





blackish. T3 with 2 lateral marginal setae. T4 with 1 median marginal and with 2 lateral


setae on the apical half of the arms. Cerci, in lateral view, sinuous with expanded apexes of

oblique upwards edges. Setae ventrally on the cerci (lateral view) absent only on the middle

portion. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior view) bigger than

the middle part. Conformation of the cerci in posterior view is divergent. Presence of a

remarkable constriction at middle portion of the cerci, in posterior view. In lateral view,

presence of a ventroapical concavity on distiphallus with smooth margins; distiphallic apical

rounded shape and sinuous distiphallic dorsal shape. Pregonite with expanded base and

sudden narrowing at apex and same color along the entire extension. Postgonite with expanded

base narrowing smoothly until the apex. Vesica with notably asymmetrical branches; terminal

lobes well-developed, with filamentous at most tapering to apex shape and sclerotized texture;

presence of spines only on ventral surface; and presence of an angular median projection of

the main vesica branch.

Remarks. Oxysarcodexia sarcinata is one of the species of this genus that have a

noteworthy asymmetrical vesica.

112

Biology. This fly specimen has already been collected from over a swarm raid of

Eciton burchelli (Westwood, 1842), an ant species (Dodge 1968). Chicken viscera and fish

head were baits used to attract and collect them in Antioquia, Colombia (Ramírez-Mora et al.

2012).

Distribution. NEARTIC. Mexico (San Luis Potosí). NEOTROPICAL. Colombia

(Antioquia, Calí, Medellín), Costa Rica, Mexico (Alajuela, Chiapas, Guanacaste, Heredia),

Panama (Barro Colorado Island), Trinidad and Tobago (Trinidad).

Material examined. ♂♂: PANAMA: Veraguas Pr. Calovébora, Rio Guazarito 14–19

Feb 2000 Col. S. Bermudez C. / 54 / NRM-DIPT 0014326 (NRM) // COLOMBIA: Antioquia,

Caldas, La Clara, 06°03‟06,9”N 75°37‟19,2”W 1.640m 7.ii.2010 Tr VSR 15:22 Cols. M. A.

Ramírez, J. Durango, H. Areiza / Dexosarcophaga, Det. M. A. Ramírez / Oxysarcodexia

sarcinata (CE-TdeA) // COSTA RICA: Alajuela prov. Atenas-Orotina road, Cerro del

Aguacate, 869m, 14.viii.2010, T. Pape, 9°58‟25,12”N 84°24‟52,24”W (ZMUC) // Costa Rica:

Cartago Turialba 2000 23 July 1965 H. .G Real / Oxysarc. sarcinata ♂ Lop. Det. H. S. Lopes

(MNRJ).

Oxysarcodexia similata Lopes & Tibana, 1987

(Appendix 28-C)

Type locality. Trinidad and Tobago, Trinidad, Aripo. Lopes & Tibana, 1987: 335.



well-developed. Thorax and abdomen with not very intense golden microtomentum.



median marginal and with 1 lateral marginal seta. T5 with golden microtomentum in part. ST5

with parallel cleft edges and with setosity. Cerci, in lateral view, straight with expanded




distiphallus with smooth margins ventroapically; distiphallic apical rounded shape and straight

distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until the apex and

113

same color along the entire extension. Postgonite with expanded base and sudden narrowing at

apex. Vesica with symmetrical branches; presence of lateral lobes (i.e. division of the vesica

coming from or close to the basal branch of the vesica, placed laterally to phallic tube),


sclerotized texture; presence of spines only on ventral surface; and presence of a rounded


Remarks. Oxysarcodexia similata can be misidentified as O. amorosa and O.

xanthosoma due to morphological similarities seen on the male terminalia. See “remarks”

section on Oxysarcodexia n. sp. 1 and O. amorosa for further discussion. Female was

described together with the male; it presents a blackish membrane between T5 and T6+7,

golden microtomentum between plates of genital tergite and red genital sternites (Lopes &

Tibana 1987).

Distribution. NEARTIC. Mexico (Morelos, San Luis Potosí). NEOTROPICAL.

Colombia (Antioquia), Costa Rica, Guyana, Mexico (Jalisco), Panama, Trinidad and Tobago

(Trinidad).

Biology. Oxysarcodexia similata showed strong attractiveness for chicken viscera

despite of human feces, which attracted no specimens. In Antioquia, Colombia, occurrence of

O. similata was reported for urban and rural areas and only a low abundance in the forest

environment, pointing out the preference of this species for human settlements (Yepes-

Gaurisas et al. 2013).

Material examined. ♂♂: PANAMA: Panama prov. Punta Charme, 19.ix.2002, J.

Méndez 79°45‟W 8°40‟N red mangrove stand / NRM-DIPT 0014339 (NRM) // Costa Rica:

Cartago Turrialba 2000‟ 23 July 1965 H. G. Real / Herman G. Real Collection / Paratype /

Oxysarcodexia similata n.sp. ♂ Paratipo Tibana & Det. H. S. Lopes.

Oxysarcodexia simplicoides (Lopes, 1933)

Sarcophaga simplicoides Lopes, 1933

(Appendix 29-A)

Type locality. Brazil, Rio de Janeiro, Guanabara. Lopes, 1933: 156.


114



with 2 well differentiated and 3 anterior setae smaller. Apical scutellar seta absent. Legs


setae. T5 without golden microtomentum, only silvery microtomentum is seen. ST5 with

parallel cleft edges and with setosity. Cerci, in lateral view, sinuous with pointed apexes of

straight edges. Setae ventrally on the cerci (lateral view) present in full extension. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) with the same size as the

middle part. Conformation of the cerci in posterior view is divergent. Presence of a remarkable

constriction at middle portion of the cerci, in posterior view. In lateral view, distiphallus with

smooth margins ventroapically and presence of ventroapical projections; distiphallic apical

shape rounded and straight distiphallic dorsal shape. Presence of a notable dilatation on the

distiphallus lateroapically. Pregonite and postgonite with expanded base narrowing smoothly

until the apex and same color along the entire extension. Vesica with symmetrical branches;

presence of lateral lobes (i.e. division of the vesica coming from or close to the basal branch of

the vesica, placed laterally to phallic tube); terminal lobes well-developed, with rounded

shape, sclerotized texture and without spines; basal branch enlarged.

Remarks. This fly is considered a rare species. The presence of slender setulae on the

gena and the postocullar plate with same color as the occiput are its more peculiar

characteristics (Lopes 1933). Female was classified by Tibana & Mello (1985), according to

the shape of the T6+7, as syntergite partially divided into two plates.

Distribution. NEOTROPICAL. Brazil (Ceará, Mato Grosso, Mato Grosso do Sul,

Minas Gerais, Rio de Janeiro).

Biology. Information is concerned about collection methods and attractiveness. Baits

as banana plus brown sugar, cow liver and lung, fish and squid has been used with traps

(“can” traps), whereas feces has been an attractive substrate for collecting actively (Lopes

1975a; Pamplona et al. 2000). Adults of O. simplicoides have also been associated to pig and

rodents carcasses in urban and natural areas of Brazilian Cerrado (“savanna-like”) vegetation

(Barbosa et al. 2009; Rosa et al. 2011; Vasconcelos & Araújo 2012).

Material examined. ♂: P. C. Elias VI-72 / Linhares, E. Santo, Brasil / simplicoides

(MNRJ).

115

Oxysarcodexia terminalis (Wiedemann, 1830)

Sarcophaga terminalis Wiedemann, 1830

Amesothyrsus chilensis Enderlein, 1928

Sarcophaga complicata Hall, 1937

Hybopygia pseudovaria Blanchard, 1939

(Appendix 29-B)

Type locality. Brazil. Wiedemann, 1830: 366.



developed. Thorax and abdomen with golden microtomentum more evident laterally.

Dorsocentral post-sutural setae with 3 well differentiated setae, although a small seta among

these 3 can be present. Apical scutellar seta absent. Legs brownish. T3 with 2 lateral marginal

setae. T4 with 1 median marginal and 1 lateral marginal setae. T5 with golden microtomentum

in part. ST5 with V-shaped cleft edges and setosity and setae along the edges of the arms.

Cerci, in lateral view, bent backwards with pointed apexes of oblique upwards edges. Setae

ventrally on the cerci (lateral view) present at apical ⅓ portion. Apical shape of the cerci (last

⅓ portion of the cercus; in posterior view) smaller than the middle part. Conformation of the

cerci in posterior view is divergent. In lateral view, distiphallus with smooth margins

ventroapically; distiphallic apical square/oblong shape and sinuous distiphallic dorsal shape.

Pregonite with base and apex with same size and same color along the entire extension.


branches; presence of lateral lobes (i.e. division of the vesica coming from or close to the basal

branch of the vesica, placed laterally to phallic tube); terminal lobes well-developed, with

filamentous at most tapering to apex shape, partially membranous texture, and presence of

spines only along the edges.

Remarks. Cerci bent backwards (lateral view) and with slender shape (dorsal view)

associated to the apical distiphallus shape and explanated vesica enable the easy recognition of

116

this species. See also “remarks” section on O. augusta. Female presents genital segments

retracted, reddish and with golden microtomentum (Blanchard 1939).

Distribution. NEOTROPICAL. Argentina (Buenos Aires, Córdoba, La Rioja), Brazil

(Goiás, Minas Gerais, Paraná, Rio de Janeiro, São Paulo), Chile (Coquimbo, Nuble,

Patagonia), Easter Islands, Peru.

Biology. Oxysarcodexia terminalis is listed as a coprophagous species, reared in cattle

dung and shed (Marchiori & Linhares 1999; Marchiori 2000; Marchiori et al. 2001; Mendes &

Linhares 2002; Mulieri et al. 2010). This species has already been reared under laboratory

conditions (Lopes 1973b), although no further information about its development was

documented. Besides bovine, dog and human feces, which are the preferential baits (Ferreira

1979; Linhares 1981; Dias et al. 1984c; Mulieri et al. 2010), attractiveness has also been

observed by chicken viscera, bovine liver, mouse, rat and pig carcasses, fish, crab, fermented

grapes, rotten banana plus brown sugar and rotten Syagrus comosa (Mart.) Mart. (Arecaceae),

a species of a palmtree (at coastal area) (Lopes 1973b; Dias et al. 1984c; Flores & Dale 1995;

Mariluis et al. 2007; Mulieri et al. 2008; Mulieri et al. 2011; Rosa et al. 2011; Beuter et al.

2012). Sunlight instead of shaded areas are also referred as a preference of this species

(Linhares 1981). Adults of O. terminalis are also among the Oxysarcodexia species already

collected associated to cadavers (Oliveira-Costa et al. 2001). Another ecological role of this

fly is the association as flower visitor with shrubs species (Baccharis spp.) of Asteraceae

family (Mulieri et al. 2010). This fly has been collected on grassland, urban, suburban and

rural areas (Mulieri et al. 2008; Mulieri et al 2011; Beuter et al. 2012), showing synanthropic

habits and populational peaks from September to December (Linhares 1981; Dias et al. 1984b;

Mendes & Linhares 1993; Mulieri et al. 2010).

Material examined. ♂♂: Brasília, D.F., Brasil, XI-60, A. B. Guimarães / NRM-DIPT

0014357 (NRM) // On human feces / H. S. Lopes 16.V.85 / Retiro Petrópolis E. do Rio, Brasil

/ O. terminalis (MNRJ).

Oxysarcodexia thornax (Walker, 1849)

Musca auriflua Wiedemann, 1830. Unavailable name. [Originally proposed in

synonymy with Musca tessellata Fabricius, 1805 and not validated by subsequent usage.]

Sarcophaga thornax Walker, 1849

117

Sarcophaga pudica Rondani, 1850

Sarcophaga aurifinis Walker, 1853

Hybopygia auricauda Enderlein, 1928

Oxysarcodexia neotropicale Prado & Fonseca, 1932

(Appendix 29-C)

Type locality. Unknown. Walker, 1849: 814.





absent. Legs blackish. T3 with 2 lateral marginal setae. T4 with 1 median marginal and 2


and with setosity and setae on the apex of the arms. Cerci, in lateral view, straight with normal

(same size as median area) apexes of concave edges. Setae ventrally on the cerci (lateral view)

absent only on the middle portion. Apical shape of the cerci (last ⅓ portion of the cercus; in


view is divergent. In lateral view, distiphallus with serrated margins ventroapically;

distiphallic apical rounded shape and straight dorsal shape; presence of a “finger-like”

swelling dorsoapically. Pregonite and postgonite with expanded base and sudden narrowing at

apex, which, only in pregonite, is darker than the base. Vesica with symmetrical branches;

terminal lobes well-developed, with square shape and sclerotized texture; presence of spines

only along the edges; and presence of an angular median projection of the main vesica branch.

Remarks. See “remarks” section on O. afficta, O. conclausa and O. morretesi. Female

was classified by Tibana & Mello (1985), according to the shape of the T6+7, as syntergite

partially divided into two plates. Larvae are characterized by the presence of sinuous ribbons

of festoon on psedocephalon and conspicuous teeth on maxillae, same as O. confusa, differing

from it by the teeth of the maxillae and the festoon arms (Leite & Lopes 1987; Lopes & Leite

1986; 1987). This species has been intercepted in imported container with cargoes from

Americas in Shanghai Port, China (Deng et al. 2011).

Distribution. NEOTROPICAL. Argentina (Bahía Blanca, Buenos Aires, Catamarca,

Corrientes, Jujuy, Misiones, Tucumán), Bolivia, Brazil (Amapá, Amazonas, Ceará, Distrito

118

Federal, Espírito Santo, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará,

Paraíba, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Roraima, Santa Catarina,

São Paulo), Colombia (Antioquia), Ecuador (Pastaza), Guyana, Paraguay, Peru. PALEARTIC.

China (Shangai).

Biology. Oxysarcodexia thornax is one of the most widespread Oxysarcodexia species.

It is very abundant among flesh flies collected, especially during summer and autumn in Brazil

(Oliveira et al. 2002; Mulieri et al. 2010). Human and bovine feces are the preferred substrates

of O. thornax for rearing and also for feeding, especially for females in order to complete

ovarian development (Mendes & Linhares 1993; Mendes & Linhares 2002). Larviposition is

documented for human, dog and anteater feces, dung, shrimp, bovine kidney, chicken viscera

and dead fish (Lopes 1973b; D‟Almeida 1986; 1988; 1989; 1994; Mendes & Linhares 1993;

Marchiori & Linhares 1999; Marchiori et al. 2001; Marchiori et al. 2002; Mendes & Linhares

2002; Mulieri et al. 2010). Under laboratory conditions, O. thornax has been reared on agar

plus powder milk for 24h, and then transferred to meat in order to complete the development

until the emergence of adults (Lopes 1973b). At our laboratory, it was reared on minced

bovine meat with adults emerging after 17–21 days (8–12 days from larva to pupa and 5–11

days until the emergence of adults). This species has already been collected associated to pig

carcass (Carvalho & Linhares 2001; Barros et al. 2008; Barbosa et al. 2009; Vairo et al.

2011). Besides these, baits as raw pork, mice, rat and rodent carcasses, chicken viscera

(especially liver), fresh and rotten bovine liver and lung, fish, crab, squid, fermented banana,

banana plus brown sugar, rotten Syagrus comosa (Mart.) Mart. (Arecaceae) (a species of a

palmtree – at coastal area) have been reported as attractive for collecting this fly (Lopes

1973b; Lopes 1975a; Ferreira 1979; Ferreira et al. 1980; Dias et al. 1984c; D‟Almeida &

Lima 1994; Couri et al. 2000; Pamplona et al. 2000; Oliveira et al. 2002; Leandro &

D‟Almeida 2005; Costamagna et al. 2007; Marchiori 2007; Mariluis et al. 2007; Moretti et al.

2008; Mulieri et al. 2008; Mulieri et al. 2011; Sousa et al. 2011; Beuter et al. 2012;

Vasconcelos & Araújo 2012; Ramírez-Mora et al. 2012; Moretti & Godoy 2013). Adults have

also been collected associated to cadavers (Oliveira-Costa et al. 2001). Sunlight instead of

shaded areas is also referred as a preference of this species (Linhares 1981). The presence of

O. thornax is known for urban areas, rural areas, pasture, cattle shed, slaughterhouse,

zoological garden, highlands (at 1,000m of altitude), grassland, woodland, archipelago,

119

deforestation area (Couri et al. 2000; Marchiori 2000; Costamagna et al. 2007; Marchiori et

al. 2007a; Mariluis et al. 2007; Couri et al. 2008; Mulieri et al. 2008; Mulieri et al. 2011;

Beuter et al. 2012; Vasconcelos & Araújo 2012). D‟Almeida (1984) pointed out O. thornax

synanthropy related to inhabited areas, which seems a pattern changed along the time, once the

presence of this fly is reported for a wide variety of sites. Other ecological interactions

reported for this species are its role as host of parasitoids such as Trybliographa sp.

(Hymenoptera: Figitidae) (Marchiori et al. 2005; Marchiori et al. 2007a; Marchiori et al.

2007b); as an eventual or rare flower visitor on Metrodorea stipularis Mart. (Rutaceae), a

riparian forest tree species (Pombal & Morellato 2000); and as host of the rove beetles

Aleochara verberans Erichson, 1839 and A. bimaculata Gravenhorst, 1802 (Coleoptera:

Staphilinidae) (Walsh & Posse 2003).

Material examined. ♂: BRAZIL: São Paulo, Campinas, Sousas, 13.IV.2011, C. M.

Souza, D. L. Brancoli, F. Rezende / O. thornax, Sousas, Campinas-SP, 13/04/2011 (L2B-

DBA).

Oxysarcodexia timida (Aldrich, 1916)

Sarcophaga timida Aldrich, 1916

Sarcophaga sanguisuga Hall, 1933

(Appendix 30-A)

Type locality. Guatemala, Porto Barrios. Aldrich, 1916: 283.



developed. Thorax and abdomen grayish sometimes with slight pale golden microtomentum.


Apical scutellar seta absent. Legs blackish. T3 with 2 lateral marginal setae. T4 with 1 median

marginal and 1 lateral marginal seta. T5 with golden microtomentum in part. ST5 with parallel

cleft edges and with setosity and setae on the apex of the arms. Cerci, in lateral view, straight

with expanded apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view)

present in full extension. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior

view) with the same size as the middle part. Conformation of the cerci in posterior view is

divergent. In lateral view, distiphallus with smooth margins ventroapically; distiphallic apical

120

conic shape, straight dorsal shape and presence of a small dorsoapical swelling. Pregonite with

expanded base and sudden narrowing at apex, which is darker than the base. Postgonite with

expanded base narrowing smoothly until the apex. Vesica asymmetric; terminal lobes reduced,

with filamentous at most tapering to apex shape and sclerotized texture; presence of spines

only along the edges; and presence of an angular median projection of the main vesica branch.

Remarks. See “remarks” section on O. afficta. Female was classified by Tibana &

Mello (1985), according to the shape of the T6+7, as syntergite partially divided into two

plates.

Distribution. NEOTROPICAL. Bolivia, Brazil (Ceará, Maranhão, Mato Grosso, Mato

Grosso do Sul, Pará, Rio de Janeiro), Colombia (Antioquia), Costa Rica, El Salvador,

Guatemala, Honduras, Panama, Peru, Venezuela.

Biology. This species has been reared on human feces and under laboratory conditions

(Lopes 1973b). Attractiveness by human feces, cow liver and lung, chicken viscera, pig

carcass, fish, crab, squid, rotten banana plus brown sugar, rotten Syagrus comosa (Mart.) Mart.

(Arecaceae) (a species of a palmtree – at coastal area) is reported for this species (Lopes

1973b; Lopes 1975a; Pamplona et al. 2000; Garcés et al. 2004; Barbosa et al. 2009; Ramírez-

Mora et al. 2012; Yepes-Gaurisas et al. 2013). As consequence of its higher frequency in

urban area, a high synanthropic index was obtained for O. timida in Antioquia, Colombia

(Yepes-Gaurisas et al. 2013).

Material examined. ♂: VENEZUELA: Aragua, Mouth of Rio Ocumare, costal flats

nr. La Boca, 17–20.xi.1997. T. Pape (ZMUC).

Oxysarcodexia trivialis (Wulp, 1895)

Sarcophaga trivialis Wulp, 1895

(Appendix 30-B)

Type locality. Mexico, Guerrero, Tierra Colorada, Amula, Xucumanatlan; Mexico,

Morelos, Cuernavaca; Mexico, Veracruz, Atoyac; Mexico, Tabasco, Teapa. Wulp, 1895: 268.



developed. Thorax and abdomen with golden microtomentum more evident laterally.


121

Apical scutellar seta present. Legs blackish. T3 with 3 lateral marginal seta. T4 with 1 median

marginal and 3 lateral marginal setae. T5 with golden microtomentum in part. ST5 with

parallel cleft edges and with setosity and setae on the apex of the arms. Cerci, in lateral view,

sinuous with expanded apexes of oblique upwards edges. Setae ventrally on the cerci (lateral

view) present at apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in


view is parallel. In lateral view, distiphallus with presence of a ventroapical concavity of

smooth margins; distiphallic apical rounded shape and straight distiphallic dorsal shape.

Pregonite with expanded base and sudden narrowing at apex, which is darker than the base.


branches; terminal lobes well-developed, with filamentous at most tapering to apex shape and

partially membranous texture; presence of spines only on ventral surface; and presence of a


Remarks. Vesica slender, without ornamentation and partially membranous is the

main characteristic which enables identification of O. trivialis with no doubt. See also

“remarks” section on O. edwardsi for further discussion. Female was classified by Tibana &


Distribution. NEARTIC. Mexico (Districto Federal, Michoacán, Morelos, Puebla),

USA (Arizona, New Mexico). NEOTROPICAL. Costa Rica (Alajuela, Puntaneras, San José),

El Salvador, Guatemala, Mexico (Chiapas, Guerrero, Michoacán, Nuevo León, Sinaloa,

Tabasco, Veracruz), Panama.

Biology. Oxysarcodexia trivialis has been found breeding on human feces (Howard

1900). This species can also be an occasional flower visitor of avocado Persea americana Mill

(Castañeda-Vildózola et al. 1999).

Material examined. ♂♂: COSTA RICA: Puntaneras, Las Alturas, Cerro Chai, 2100m,

14.viii.1995, Th. Pape leg. (ZMUC) // COSTA RICA: San José, Rio Savegre, 9km SW route

2, San Gerardo de Dota 2200–2500m, 7–11.viii.1995, Th. Pape leg. (ZMUC).

Oxysarcodexia varia (Walker, 1836)

Sarcophaga varia Walker, 1836

Sarcophaga milleri Johnston & Tiegs, 1922

122

Sarcophaga chilensis Macquart, 1843

Sarcophaga flavicostata Macquart, 1843

Sarcophaga truncata Schiner, 1868

Sarcophaga nobilis Thomson, 1869

(Appendix 30-C)

Type locality. Uruguay, Gorrite Island, off Maldonado. Walker, 1836: 353.



developed. Thorax and abdomen with pale golden microtomentum more evident laterally.

Dorsocentral post-sutural setae with 3 well differentiated although a small bristle among these

3 can be present. Apical scutellar seta absent. Legs blackish. T3 with 1 lateral marginal seta.

T4 without median marginal and with 2 lateral marginal setae. T5 with golden microtomentum

in part. ST5 with V-shaped cleft edges and with setosity and setae along the edges of the arms.

Cerci, in lateral view, sinuous with expanded apexes of concave edges. Setae ventrally on the

cerci (lateral view) present in full extension. Apical shape of the cerci (last ⅓ portion of the


view is divergent. Presence of a remarkable constriction at middle portion of the cerci, in

posterior view. In lateral view, distiphallus with smooth margins ventroapically; distiphallic

apical conic shape; sinuous distiphallic dorsal shape; and presence of a large dorsoapical

swelling on distiphallus. Pregonite with expanded base and sudden narrowing at apex and

same color along the entire extension. Postgonite with expanded base narrowing smoothly

until the apex. Vesica with asymmetrical branches; terminal lobes well-developed, with

rounded shape and sclerotized texture; presence of spines only along the edges; and presence

of a rounded median projection of the main vesica branch.

Remarks. See “remarks” section on O. augusta, O. aurata and O. marina.

Oxysarcodexia varia male and female were redescribed by Lopes & Albuquerque (1955);

characteristics as T7 interrupted medially, T8 compound by two plates without setosity,

ST7+8 small, with marginal setae and connected to ST8 which is well-developed, with golden

microtomentum and, on posterior margin, a deep longitudinal furrow were ascribed for

females.

123

Distribution. NEOTROPICAL. Argentina (Buenos Aires, Mendoza, Neuquén,

Tucumán), Bolivia, Brazil (Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul), Chile

(Bío Bío, Concepción, Coquimbo, Juan Fernandez Islands La Araucanía, Nuble, Patagonia,

Santiago, Valdivia), Uruguay (Corrite Island, Maldonado). AUSTRALASIAN/OCEANIAN.

Australia (New South Wales, Tasmania, Victoria), Fiji, French Polynesia (Society Islands),

New Zealand (North Island, South Island), Norfolk Island.

Biology. This species has been reared on dung (Mendes & Linhares 2002), dead

insects (Blanchard 1939) and in alive and dead lobsters (Blanchard 1939; Blanchard 1942;

Mulieri et al. 2010). This fly is attracted preferentially by feces, especially of dog (Mariluis et

al. 2007; Mulieri et al. 2010; Mulieri et al. 2011), but also by rotten minced kangaroo meat,

sheep‟s liver, carcasses (Meiklejohn et al. 2012) and rotten bovine liver (Mulieri et al. 2011).

Oxysarcodexia varia was more abundant during the spring and summer in Argentina and

Brazilian Southern and also considered asynanthropic due to higher abundance in rural areas

and grassland instead of woodland (Ferreira 1979; Mariluis et al. 2007; Mulieri et al. 2008;

Mulieri et al. 2010). Other ecological relationships of this species are the association as flower

visitor with Eryngium horridum (Spreng) Less (Apiaceae), Baccharis juncea (Cass) Desf,

Helianthus annuus L. (Asteraceae), Sebastiania brasiliensis Spreng (Euphorbiaceae), and

Colletia paradoxa (Spreng) Escal., Discaria americana Gillies & Hook, Scutia buxifolia

Reissek (Rhamnaceae) (Mulieri et al. 2010); and as host of Aleochara notula Erichson, 1839,

A. verberans Erichson, 1839 and A. puberula Klug, 1832 (Coleoptera: Staphilinidae) (Walsh

& Posse 2003). It has been collected in urban, suburban and rural areas (Mulieri et al. 2011).

This species has also a medical importance once it‟s considered a possible mechanical vector

of rabbit hemorrhagic disease virus (Henning et al. 2005).

Material examined. ♂: ARGENTINA, Chubut 47: Tecka, Corcovado, 750m

17.ii.1979, Mision Cientifica Danesa / NRM-DIPT 0014413 (NRM).

Oxysarcodexia ventricosa (Wulp, 1895)

Sarcophaga ventricosa Wulp, 1895

Sarcophaga tenuiventris Wulp, 1895

(Appendix 31-A)

124

Type locality. Mexico, Guerrero, Chilpancingo; Mexico, Tabasco, Teapa. Wulp, 1895:

268.


Diagnosis. Male. Postocular plate with silveryish microtomentum. Ocellar setae not

well-developed. Thorax and abdomen with golden microtomentum more evident laterally.


Apical scutellar seta absent. Legs yellowish. T3 with 3 lateral marginal setae. T4 with 1

median marginal and 2 lateral marginal setae. T5 with golden microtomentum partially. ST5

with parallel cleft edges and with setosity and setae on the apex of the arms. Cerci, in lateral

view, straight with expanded apexes of oblique upwards edges. Setae ventrally on the cerci

(lateral view) present on apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the

cercus; in posterior view) bigger than the middle part. Conformation of the cerci in posterior

view is divergent. Presence of a remarkable constriction at middle portion of the cerci, in

posterior view. In lateral view, distiphallus with presence of a ventroapical concavity of

serrated margins; distiphallic apical conic shape and straight distiphallic dorsal shape.

Pregonite with expanded base narrowing smoothly until the apex, which is darker than the

base; same as postgonite except for the color which is the same along the entire extension.

Vesica with symmetrical branches; terminal lobes reduced, with filamentous at most tapering

to apex shape and sclerotized texture; presence of spines on both dorsal and ventral surfaces;

and presence of a rounded median projection of the main vesica branch.

Remarks. See O. admixta “remarks” section. Ravinia assidua (Walker, 1853) and O.

ventricosa have already been involved in a taxonomical issue as pointed out by Dodge

(1956b). Female was classified by Tibana & Mello (1985), according to the shape of the T6+7,

as syntergite undivided. Larvae are known and were described by Knipling (1936) and

Wharton & Moon (1979). For further comments about the larvae, see “remarks” section on O.

galeata. This species has been intercepted in imported container with cargoes from Americas

in Shanghai Port, China (Deng et al. 2011).

Distribution. NEARTIC. Bermuda, Canada (Ontario, Quebec), Mexico (Districto

Federal, Morelos, San Luis Potosí), USA (Alabama, Arkansas, District of Columbia, Florida,

Georgia, Illinois, Indiana, Iowa, Louisiana, Maryland, Massachusetts, Missouri, Nebraska,

New Jersey, New York, Ohio, Oklahoma, Pennsylvania, South Carolina, Texas, West

125

Virginia). NEOTROPICAL. Argentina (Córdoba), Costa Rica, Honduras, Mexico (Chiapas,

Guerrero, Tabasco, Veracruz, Yucatán). PALEARTIC. China (Shangai).

Biology. This species has been rearing mainly on bovine dung (Knipling 1936; Sanders

& Dobson 1966; Blume 1986). Larvae were reared on cow manure and, in one case, on

decomposing hamburger meat, took 14 days to complete the developmental period, from larva

to adult (Knipling 1936). When immatures were reared from a bovine pad, they took 15–25

days to complete the development until the emergence of adults (Sanders & Dobson 1966).

Larvae were also recorded feeding on eggs and larvae of Oxybelus wasp, probably as an

alternative choice due to scarcity of the preferential resource (Bohart et al. 1966; Peckham

1991), thus it‟s considered a species probably capable of prey-parasitism (Peckham 1991).

Oxysarcodexia ventricosa is considered a scavenger species, being collected associated to pig

carcass exposed on tree and water environments (Payne & King 1972) and in the active decay

stage of decomposition (Tabor et al. 2004; Tabor et al. 2005). This dung-breeding species has

been collected perching on pitcher plants (Dahlem & Naczi 2006), on high lands (Dodge &

Seago 1954) and in a bog (Judd 1970). Pupae originated from larvae bred on dung were

parasitized by Aphaereta pallipes (Say) (Hymenoptera: Braconidae) (Figg et al. 1983). This

fly was one of the species used to study the phylogeny of Sarcophagidae based on a molecular

approach (mtDNA fragments) (Stamper et al. 2012).

Material examined. ♂♂: ALABAMA: Madison Co. 10km NE Maysville, Sharp

Cove, Sneed Spg. bottomland, 240m, 6 oct 1992, Acciavath / NRM-DIPT 0014456 (NRM) //

USA, Fla, Highlands Co., Archbold Biol. St., 15.vi.1989, T. Pape leg. (ZMUC).

Oxysarcodexia villosa Lopes, 1946

(Appendix 31-B)

Type locality. Brazil, Pará. Lopes, 1946a: 458.



developed. Thorax with pale golden microtomentum. Dorsocentral post-sutural setae with 2

well differentiated and 2 anterior setae smaller. Apical scutellar seta absent. Legs blackish.


is seen laterally. T3 with 1 lateral marginal seta. T4 with 1 median marginal and 2 lateral

126


setae scattered on arm‟s surface. Cerci, in lateral view, straight with normal apexes (same size

as the median area) of oblique upwards edges. Setae ventrally on the cerci (lateral view)

present in apical ⅓ portion. Apical shape of the cerci (last ⅓ portion of the cercus; in posterior

view) with the same size as the middle part. Conformation of the cerci in posterior view is

parallel. Presence of a remarkable constriction at middle portion of the cerci, in posterior view.

In lateral view, distiphallus with smooth margins ventroapically; distiphallic apical shape

rounded and straight distiphallic dorsal shape. Pregonite and postgonite with expanded base

narrowing smoothly until the apex and same color along the entire extension. Vesica with

asymmetrical branches; terminal lobes well-developed, with oblong shape and sclerotized

texture; presence of spines on basal half of ventral surface and on the edge of the right lobe

(more developed than the left lobe); and presence of an angular median projection of the main

vesica branch.

Remarks. This is another species of this genus that have a noteworthy asymmetrical

vesica. In O. villosa, vesica is very developed, surpassing substantially the distiphallus apex.

Distribution. NEOTROPICAL. Brazil (Pará).

Biology. Unknown.

Material examined. ♂: Inst. Agronômico do Norte, Belém, Est. Do Pará – Brasil XI-

959. L. Trav. D. Lacombe J. Evangelista E. Lobato / Oxysarc villosa ♂ Lop det. H. S. Lopes

(MNRJ).

Oxysarcodexia vittata (Walker, 1836)

Sarcophaga vittata Walker, 1836

Oxysarcodexia titubata Lopes, 1946

(Appendix 31-C)

Type locality. South America. Walker, 1836: 353.




differentiated, a small bristle among these 3 can be present. Apical scutellar seta present. Legs

blackish. Abdomen with silvery microtomentum. T4 with 2 lateral marginal setae. T5 with

127

golden microtomentum in part. ST5 with parallel cleft edges and with setosity and setae on the

apex of the arms. Cerci, in lateral view, sinuous with pointed apexes of oblique upwards

edges. Setae ventrally on the cerci (lateral view) present in full extension. Apical shape of the

cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.


smooth margins ventroapically; distiphallic apical shape square/oblong; sinuous distiphallic

dorsal shape; presence of ventroapical projections, lateral lobes, and a medium dorsoapical

swelling on distiphallus. Presence of lateroapical expansions on distiphallus, in dorsal view.

Pregonite with base and apex with same size and apex darker than the base. Postgonite with

expanded base and sudden narrowing at apex and same color along the entire extension.

Vesica symmetric; terminal lobes well-developed, with square shape and sclerotized texture;

and without spines.

Remarks. See “remarks” section on Oxysarcodexia n. sp. 4, O. augusta and O. nitida.

Female is characterized by T7 entire, sclerotized dorsally and with the presence of a large

membrane connecting it to T5; T8 constituted by two elongated plates without setae; ST6

larger than ST5 and with apical setae; ST7 larger than long and connected to ST8 by an

extensive membranous region (Lopes 1946a).

Distribution. NEOTROPICAL. Brazil (Mato Grosso, Mato Grosso do Sul, Rio de

Janeiro, Paraná, Santa Catarina, São Paulo).

Biology. This species is attracted by human feces and had already been reared on

laboratory (Lopes 1973b), although no further information was reported.

Material examined. ♂♂: BRAZIL: São Paulo, Jundiaí, Serra do Japi; 06.I.2012; M.

D. Grella / Oxysarcodexia 01; Jundiaí-SP; 06/01/2012 / O. titubata (L2B-DBA) // Brasilien,

Nova Teutonia, 27°11‟B 52°23‟L, Fritz Plaumann / Xarcophaga titubata (Lopes) /

Oxysarcodexia (ZMUC).

Oxysarcodexia wygodzinskyi Lopes & Tibana, 1987

(Appendix 32-A)

Type locality. Argentina, Salta, San Martin. Lopes & Tibana, 1987: 336.


128


developed. Thorax with pale golden microtomentum on humeral region. Dorsocentral post-

sutural setae with 2 well differentiated and 3 anterior setae smaller. Apical scutellar seta

absent. Legs blackish. Abdomen with silvery microtomentum. T3 with 2 lateral marginal

setae. T4 with 1 median marginal and 2 lateral marginal setae. T5 with silvery

microtomentum, although some pale golden microtomentum on anterior margin is seen. ST5

with parallel cleft edges and with setae on the apex of the arms. Cerci, in lateral view, sinuous

with pointed apexes of oblique upwards edges. Setae ventrally on the cerci (lateral view)

present in apical half portion. Apical shape of the cerci (last ⅓ portion of the cercus; in


divergent. In lateral view, distiphallus with ventroapical concavity with smooth margins

ventroapically; distiphallic apical shape rounded and straight distiphallic dorsal shape.

Pregonite with base and apex with same size and sudden narrowing at apex and same color

along the entire extension. Postgonite with expanded base narrowing smoothly until the apex

and same color along the entire extension. Vesica symmetric; terminal lobes well-developed,

with rounded shape and sclerotized texture; presence of spines only apically along the edges.

Remarks. See “remarks” section on O. cingarus.

Distribution. NEOTROPICAL. Argentina (Salta), Brazil (Mato Grosso, Mato Grosso

do Sul).

Biology. Unknown.

Material examined. ♂: Oxysarcodexia wygodzinsky Lopes & Tibana, 1987, det. C. A.

Mello-Patiu / BRASIL, MS, Porto Murtinho, Fazenda Retiro Conceição S21°41‟18.8”

W057°45‟53.7” Coleta manual (rede) 11.xii.2011 Patiu & Patiu col. (MNRJ).

Oxysarcodexia xanthosoma (Aldrich, 1916)

Sarcophaga xanthosoma Aldrich, 1916

(Appendix 32-B)

Type locality. Guatemala, Los Amates. Aldrich, 1916: 274.



well-developed. Thorax and abdomen with intense golden microtomentum. Dorsocentral post-

129


present. Legs blackish. T3 with 3 lateral marginal setae. T4 with 1 median marginal and 2


and setosity. Cerci, in lateral view, straight with expanded apexes of oblique upwards edges.

Setae ventrally on the cerci (lateral view) absent only in the middle portion. Apical shape of

the cerci (last ⅓ portion of the cercus; in posterior view) smaller than the middle part.


presence of a ventroapical concavity of smooth margins; distiphallic apical rounded shape;

sinuous distiphallic dorsal shape. Pregonite with expanded base and sudden narrowing at apex,

which is darker than the base. Postgonite with expanded base narrowing smoothly until the

apex and same color along the entire extension. Vesica with symmetrical branches; presence

of lateral lobes (i.e. division of the vesica coming from or close to the basal branch of the

vesica, placed laterally to phallic tube); terminal lobes well-developed, with filamentous at

most tapering to apex shape and sclerotized texture; presence of spines only on ventral

surface; and presence of a rounded median projection of the main vesica branch.

Remarks. This species can exhibit high morphological variation, especially in cerci

and phallus apical portions and also on microtomentum, as pointed out by Lopes (1975c),

leading to misidentification of O. xanthosoma as O. amorosa. See “remarks” section on

Oxysarcodexia n. sp. 1. Female was classified by Tibana & Mello (1985), according to the

shape of the T6+7, as syntergite undivided.

Distribution. NEARTIC. Mexico (San Luis Potosí, Sonora). NEOTROPICAL.

Argentina (Misiones), Brazil (Amapá, Amazonas, Ceará, Espírito Santo, Mato Grosso, Minas

Gerais, Pará, Paraná, Rio de Janeiro, Roraima, São Paulo), Colombia, Costa Rica, Ecuador, El

Salvador, Guatemala (Los Amates), Guyana (Kartabo), Mexico (Jalisco, Veracruz), Panama,

Peru.


1973b) and on human feces (D‟Almeida 1994). This fly has been collected using human feces,

peccary entrails, rotting liver, fish, pig carcass, banana plus brown sugar, rotten Syagrus

comosa (Mart.) Mart. (Arecaceae), a species of a palmtree (at coastal area) as baits (Curran &

Walley 1934; Lopes 1973b; Lopes 1975a; Oliveira et al. 2002; Leandro & D‟Almeida 2005;

Barbosa et al. 2009; Rosa et al. 2011; Vairo et al. 2011). The presence of O. xanthosoma is

130

known for urban (zoological garden), Cerrado (“savanna-like”) vegetation, peri-urban and

forest areas (Lopes & Tibana 1982; Oliveira et al. 2002; Rosa et al. 2011; Vairo et al. 2011).

Material examined. ♂♂: ECUADOR: Napo Province: Yasuní National Park: Yasuní

Research Station: 76°36‟W 00°38‟S 3–20.XI.1998 T. Pape & B. Viklund / NRM-DIPT

0014476 / O. xanthosoma (NRM) // Oxysarcodexia xanthosoma / Campinas, SP VII/2012

Brancoli DL (L2B-DBA) // Oxysarcodexia xanthosoma / Campinas, SP VII/2012 Brancoli DL

(L2B-DBA) // Oxysarcodexia xanthosoma / Campinas, SP VII/2012 Brancoli DL (L2B-DBA).

Oxysarcodexia xon (Dodge, 1968)

Xarcophaga xon Dodge, 1968

(Appendix 32-C)

Type locality. Panama, Canal Zone, Barro Colorado Island. Dodge, 1968: 449.

Depository of type material. KU.

Diagnosis. Male. Postocular plate pale golden microtomentum. Ocellar setae not well-

developed. Thorax and abdomen with silveryish microtomentum. Three well differentiated

dorsocentral post-sutural setae. Apical scutellar seta is absent. T3 with median marginal setae.

T5 with yellowish microtomentum more evident laterally. T4 with median marginal setae. ST5

with V-shaped cleft edges and short apical prolongations. Cerci with straight shape (lateral

view) with a small anterior point. Phallus entire with the presence of an expanded apical

serrated plate and lateral serrated plates. Small vesica. This is a synopsis of the original

description given by Dodge (1968) and of the diagnosis given by Lopes (1975c).

Remarks. See “remarks” section on Oxysarcodexia n. sp. 4 and O. notata.

Distribution. NEOTROPICAL. Brazil (Rio de Janeiro), Panama (Barro Colorado

Island, Canal Zone).

Biology. Unknown.

Oxysarcodexia zayasi Dodge, 1956

(Appendix 32-D)

Type locality. Cuba, Habana, Lomas de Camoa. Dodge, 1956: 100.


131


well-developed. Thorax with slightly pale golden microtomentum. Dorsocentral post-sutural


Legs yellowish. Abdomen with slightly silveryish microtomentum. T3 with 3 lateral marginal

setae. T4 without median marginal and with 2 lateral marginal setae. T5 without golden

microtomentum, but silveryish microtomentum shades are present. ST5 with V-shaped cleft

edges and with setosity. Cerci, in lateral view, straight with expanded apexes of oblique

upwards edges. Setae ventrally on the cerci (lateral view) present in full extension. Apical

shape of the cerci (last ⅓ portion of the cercus; in posterior view) bigger than the middle part.


at middle portion of the cerci, in posterior view. In lateral view, distiphallus with presence of a

ventroapical concavity of smooth margins; distiphallic apical square/oblong shape; and

straight distiphallic dorsal shape. Pregonite with expanded base narrowing smoothly until the




texture; presence of spines only on ventral surface; and presence of a rounded median


Remarks. See “remarks” section on O. cyanea. Female was classified by Tibana &


Distribution. NEOTROPICAL. Bahamas (Andros, Eleuthera, Grand Bahama, New

Providence), Cuba (Havana, Lomas de Camoa).

Biology. Unknown. Type specimens were collected with fish head exposed in a

wooded area (Dodge 1956).

Material examined. ♂♂: KUBA: Provinz Santiago de Kuba; Cordillera de La Gran

Piedra [unreadable information] 75°38‟W, 20°02‟N, [unreadable information] 19–26.XII.1997

Leg. Stark / Oxysarcodexia zayasi Dodge, det. T. Pape 1998-08-27 / NRM-DIPT 0014463

(NRM) // Cuba prov. Las Villas Trinidad 100m 29.VI.196[0 or 6 – damaged by the pinhole]

leg. F. Gregor / Oxysarcodexia zayasi Dodge B. Rohdendorf det 1970. I (MNRJ).

132

Final considerations

Similarities on male terminalia of very close related species, the notable number of

species presenting asymmetry on the vesica, in comparison to other genus of Sarcophagidae,

and a better comprehension of “Xarcophaga”, “paulistanensis”, “ventricosa” and “thornax”

groups call for a thorough investigation, in order to provide a phylogenetic approach for

clarify these issues. Furthermore, it is also needed a deeper study of females and larval

morphology, besides ultrastructural analysis of male terminalia (e.g. scanning electron

microscopy) and molecular approaches for a wider and more complete understanding of

relationships inside the genus Oxysarcodexia.

Thus, this taxonomic conspectus is an initial wider step towards Oxysarcodexia

comprehension, contributing with an update of the male knowledge of species within this

genus, their morphological peculiarities and geographical distribution, also gathering

information about their biology, which is still incipient. Also, pictorial information here

documented, together the presented diagnosis, enable a comparison and, consequently, an

accurate identification until the level of species, particularly important in the case of close

related species and living in sympatry.

Acknowledgments

To Dr. Cátia Antunes de Mello-Patiu for provided the means for the examination of

specimens from the Museu Nacional/Universidade Federal do Rio de Janeiro; MsC. Eliana

Buenaventura for provided the means for the examination of specimens from the Tecnológico

de Antioquia, Institución Universitaria; and Dr. Kjell Arne Johanson for provided the means

for the examination of specimens from the Swedish Museum of Natural History for loan of

material.

This research was supported by CAPES Foundation, Ministry of Education of Brazil

(grant #0906/12-3).

References

Aldrich, J.M. (1916) Sarcophaga and allies in North America. Thomas Say Foundation, Entomological

Society of America, La Fayette, Indiana, 301 pp. + 16 pls.

133

Aldrich, J.M. (1930) Notes on the types of American two-winged flies of the genus Sarcophaga and a

few related forms, described by the early authors. Proceedings of the United States National

Museum, 78, 1–39 + 3 pls. http://dx.doi.org/10.5479/si.00963801.78-2855.1

Amat, E. (2010) Notes on necrophagous flies (Diptera: Calyptratae) associated to fish carrion in

Colombian Amazon. Acta Amazonica, 40 (2), 397–400.

Barbosa, R.R., Mello-Patiu, C.A., Mello, R.P. & Queiroz, M.M.C. (2009) New records of calyptrate

dipterans (Faniidae, Muscidae and Sarcophagidae) associated with the decomposition of domestic

pigs in Brazil. Memórias do Instituto Oswaldo Cruz, 104 (6), 923–926.

Barros, R.M. de, Mello-Patiu, C.A. de & Pujol-Luz, J.R. (2008) Sarcophagidae (Insecta, Diptera)

associados à decomposição de carcaças de Sus scrofa Linnaeus (Suidae) em área de Cerrado do

Distrito Federal, Brasil. Revista Brasileira de Entomologia, 52 (4), 606–609. [In Portuguese]

http://dx.doi.org/10.1590/S0085-56262008000400011

Beuter, L., Fernandes, P.A., Barros, P.B., Souza, C.R. & Mendes, J. (2012) Insetos de potencial

importância forense e na saúde pública em região urbana de Minas Gerais: frequência relativa e

variação sazonal de fauna atraída e criada em carcaças de roedores. Revista de Patologia Tropical,

41 (4), 480–490. [In Portuguese]

Blanchar, E.E. (1935) Contribución al estúdio de los sarcofágidos argentinos. Physis, 11, 508. [In

Spanish]

Blanchard, E.E. (1939) Los sarcofágidos argentinos. Contribución a su conocimiento. Physis, 17, 791–

856. [In Spanish]

Blanchard, E.E. (1942) Parasitos de Alabama argillacea Hbn. en la República Argentina. Estudio

preliminar. Anales de la Sociedad Científica Argentina, 134, 54–63. [In Spanish]

Blume, R.R. (1985) A checklist, distribution record, and annotated bibliography of the insects

associated with bovine droppings on pastures in America north of Mexico. The Southwestern

Entomologist, 9, 1–28.

Blume, R.R. (1986) Parasites of Diptera associated with bovine droppings on a pasture in east central

Texas. The Southwestern Entomologist, 11 (3), 215–222.

Boff, S., Graciolli, G., Boaretto, A.G. & Marques, M.R. (2008) Insetos visitantes de gomas exsudadas

por Terminalia argentea Marct & Zucc (Combretaceae). Revista Brasileira de Entomologia, 52

(3), 477–479. [In Portuguese]

Bohart, R.M., Lin, C.S. & Holland, J.F. (1966) Bionomics of Oxybelus sparideus at Lake Texoma,

Oklahoma (Hymenoptera: Sphecidae: Craboninae). Annals of the Entomological Society of

America, 59 (4), 818–820.

134

Brown, B.V. (2009) Introduction. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M.,

Woodley, N.E. & Zumbado, M.A. (eds.), Manual of American Central Diptera. Vol. 1. NCR

Research Press, Ottawa, Ontario, Canada, pp. 1–7.

Carvalho, C.J.B. & Mello-Patiu, C.A. (2008) Key to the adults of the most common forensic species of

Diptera in South America. Revista Brasileira de Entomologia, 52, 390–406.

http://dx.doi.org/10.1590/S0085-56262008000300012

Carvalho, C.J.B., Rafael, J.A., Couri, M.S. & Silva, V.C. (2012) Diptera. In: Rafael, J.A., Melo,

G.A.R., Carvalho, C.J.B., Casari, S. & Constantino, C. (eds.), Insetos do Brasil: diversidade e

taxonomia. Holos Editora, Ribeirão Preto, São Paulo, Brasil, pp. 701–743. [In Portuguese]

Carvalho, L.M.L. & Linhares, A.X. (2001) Seasonality of insect succession and pig carcass

decomposition in a natural forest area in southeastern Brazil. Journal of Forensic Sciences, 46 (3),

604–608.

Castañeda-Vildózola, A., Equihua-Martínez, A., Valdés-Carrasco, J., Barrientos-Priego, A.F., Ish-Am,

G. & Gazit, S. (1999) Insectos polinizadores del aguacareto em los estados de México y

Michoacán. Revista Chapingo Serie Horticultura, 5, 129–136. [In Spanish]

Costamagna, R., Visciarelli, E.C, Lucchi, L.D., Basabe, N.E., Esteban, M.P. & Oliva, A. (2007)

Aportes al conocimiento de los dípteros ciclorrafos em el área urbana de Bahía Blanca (província

de Buenos Aires), Argentina. Revista del Museo Argentino de Ciencias Naturales, 9 (1), 1–4.

Cornaby, B.W. (1974) Carrion reduction by animals in contrasting tropical habitats. Biotropica, 6 (1),

51–63.

Couri, M.S., Barros, G.P.S., Orsini, M.P. (2008) Dipterofauna do Arquipélago de Fernando de

Noronha (Pernambuco, Brasil). Revista Brasileira de Entomologia, 52 (4), 588–590. [In

Portuguese]

Couri, M.S., Lamas, C.J.E., Aires, C.C.C., Mello-Patiu, C.A., Pamplona, D.M. & Magno, P.D. (2000)

Diptera from Serra do Navio (Amapá, Brasil): Asilidae, Bombyliidae, Calliphoridae,

Micropezidae, Muscidae, Sarcophagidae, Stratiomyiidae, Syrphidae, Tabanidae and Tachinidae.

Revista Brasileira de Zoociências, 2 (1), 91–100.

Cruz, T. M. (2008) Diversidade e sucessão ecológica de insetos associados à decomposição animal em

um fragmento de Mata Atlântica de Pernambuco. Dissertation. Recife, Pernambuco, Brasil.

102pp. [In Portuguese]

Curran, C.H. & Walley, G.S. (1934) Sarcophagidae. In: Curran, C.H. The Diptera of Kartabo, Bartica

District, British Guiana, with descriptions of new species from other British Guiana localities.

Bulletin of the American Museum of Natural History, 66, pp. 287–532.

135

D‟Almeida, J.M. & Lima, S.F. (1994) Atratividade de diferentes iscas e sua relação com as fases de

desenvolvimento ovariano em Calliphoridae e Sarcophagidae (Insecta, Diptera). Revista

Brasileira de Zoologia, 11(2), 177–186. [In Portuguese]

D‟Almeida, J.M. (1984) Sinantropia de Sarcophagidae (Diptera) na região metropolitana do Estado do

Rio de Janeiro. Arquivos da Universidade Federal Rural do Rio Janeiro, 7, 101-110. [In

Portuguese]

D‟Almeida, J.M. (1986) Substratos para criação de dípteros caliptratos em área rural do Estado do Rio

de Janeiro. Arquivos da Universidade Federal Rural do Rio de Janeiro, 9 (1–2): 13–22. [In

Portuguese]

D‟Almeida, J.M. (1988) Substratos utilizados para a criação de dípteros caliptratos em uma área urbana

do município do Rio de Janeiro. Memórias do Instituto Oswaldo Cruz, 83, 201–206. [In

Portuguese] http://dx.doi.org/10.1590/S0074-02761988000200009

D‟Almeida, J.M. (1989) Substratos utilizados para a criação de dípteros caliptratos no jardim zoológico

do Rio de Janeiro. Memórias do Instituto Oswaldo Cruz, 84, 257–264. [In Portuguese]

http://dx.doi.org/10.1590/S0074-02761989000200016

D‟Almeida, J.M. (1994) Ovipositional substrates used by Calyptrate Diptera in Tijuca Forest, Rio de

Janeiro. Memórias do Instituto Oswaldo Cruz, 89 (2), 261–264.

Dahlem, G.A. & Naczi, R.F.C. (2006) Flesh flies (Diptera: Sarcophagidae) associated with North

American pitcher plants (Sarraceniaceae), with descriptions of three new species. Annals of the

Entomological Society of America, 99 (2), 218–240.

Deng, Y.H., Wang, G.P., Chen Z.Z & Fan Z.D. (2011) Three species of the genus Oxysarcodexia

Townsend, 1917 (Diptera: Sarcophagidae) intercepted in imported container with cargoes from

Americas in Shanghai Port. Acta Parasitologica et Medica Entomologica Sinica, 18 (1), 42–47.

doi:10.3969/j.issn.1005-0507.201.1.01.009 [In Chinese]

Dias, E.S., Neves, D.P. & Lopes, H.S. (1984a) Estudos sobre a fauna de Sarcophagidae (Diptera) de

Belo Horizonte – Minas Gerais I – Levantamento taxonômico e sinantrópico. Memórias do

Instituto Oswaldo Cruz, 79 (1), 83–91. [In Portuguese]

Dias, E.S., Neves, D.P. & Lopes, H.S. (1984b) Estudos sobre a fauna de Sarcophagidae (Diptera) de

Belo Horizonte – Minas Gerais II – Variação Sazonal. Memórias do Instituto Oswaldo Cruz, 79

(4), 409–412. [In Portuguese]

Dias, E.S., Neves, D.P. & Lopes, H.S. (1984c) Estudos sobre a fauna de Sarcophagidae (Diptera) de

Belo Horizonte – Minas Gerais II – Atratividade das Iscas. Memórias do Instituto Oswaldo Cruz,

79 (4), 413–417. [In Portuguese]

Dodge, H.R. & Seago, J.M. (1954) Sarcophagidae and other Diptera taken by trap and net on Georgia

Mountain Summits in 1952. Ecology, 35 (1), 50–59.

136

Dodge, H.R. (1956) Two new Sarcophagid flies from Cuba (Diptera). Memórias de la Sociedad

Cubana de Historia Natural, 23 (1), 97–103.

Dodge, H.R. (1965) The Sarcophagidae (Diptera) of the West Indies. II. Jamaica. Annals of the

Entomological Society of America, 58, 497–517.

Dodge, H.R. (1966) Some new or little-known Neotropical Sarcophagidae (Diptera), with a review of

the genus Oxysarcodexia. Annals of the Entomological Society of America, 59, 674–701.

Dodge, H.R. (1968) The Sarcophagidae of Barro Colorado Island, Panama (Diptera). Annals of the


Enderlein, G. (1928) Klassifikation der Sarcophagiden. Sarcophagiden-Studien I. Archiv für

Klassifikatorische und Phylogenetische Entomologie, Wien, 1, 1–56. [In German]

Engel, O. (1931) Die Ausbeute der deutschen Chaco –Expedition 1925/1926 – Diptera XXVI.

Anthomyidae, XXVII. Muscidae und XXVIII. Sarcophagidae. Konowia, 10 (2), 133–154. [In

German]

Evenhuis, N.L., Pape, T., Pont, A.C. & Thompson, F.C. (eds.) (2007) Biosystematic Database of

World Diptera, Version 10. Available at http://www.diptera.org/biosys.htm. apud Pape, T.,

Bickel, D. & Meier, R. (2009) Diptera Diversity: status, challenges and tools. Brill, Portland,

459pp.

Ferreira, M.J.M. (1979) Sinantropia de Dipteros Muscoides de Curitiba. II: Sarcophagidae. Revista

Brasileira de Biologia, 39 (4), 773–781. [In Portuguese]

Ferreira, M.J.M., Rizzo, J.A., Rasmussen, G., Pereira, I.F. (1980) Ocorrência e frequência de

califorídeos em formações de mata e cerrado no município de Goiânia-Goiás. Anais da E.A.V., 1,

13–26. [In Portuguese]

Figg, D.E., Hall, R.D. & Thomas, G.D. (1983) Host range and eclosion success of the parasite

Aphaereta pallipes (Hymenoptera: Braconidae) among dung-breeding Diptera in Central

Missouri. Enviromental Entomology, 12 (3), 993–995.

Figueiredo, R.R., Dorf, S., Couri, M.S., Azevedo, A.A. & Mossumez, F. (2002) Corpos estranhos

animados em otorrinolaringologia. Revista Brasileira de Otorrinolaringologia, 68 (5), 722–728.

Flores, V.I. & Dale, W.E. (1995) Um estúdio sobre ecologia de las moscas Sarcophagidae em la costa

central peruana. Revista Peruana de Entomología, 38, 13–17. [In Spanish]

Garcés, P.A., Bermudes, S. & Quintero, G. (2004) Determinación de la entomofauna associada a

carcaças de cerdos domésticos vestidos (Sus scrofa), en el Puerto de Vacamonte, Prov. de

Panamá. Tecnociencia, 6 (2), 59–74. [In Spanish]

137

García-Franco, J.G. & Rico-Gray, V. (1997) Reproductive biology of the holoparasitic endophyte

Bdallophyton bamburarum (Rafflesiaceae). Botanical Journal of the Linnean Society, 123, 237–

247.

Giroux, M., Pape, T. & Wheeler, T.A. (2010) Towards a phylogeny of the flesh flies (Diptera:

Sarcophagidae): morphology and phylogenetic implications of the acrophallus in the subfamily

Sarcophaginae. Zoological Journal of the Linnean Society, 158, 740–778.

http://dx.doi.org/10.1111/j.1096-3642.2009.00561.x

Hall, D.G. (1933) The Sarcophaginae of Panama (Diptera: Calliphoridae). Bulletin of the American

Museum of Natural History, 66, 251–285.

Hall, D.G. (1937) New muscoid flies (Diptera) in the United States National Museum. Proceedings of

the United States National Museum, 84, 201–216. http://dx.doi.org/10.5479/si.00963801.84-

3011.201

Hall, D.G. (1938) New genera and species of South American Sarcophagidae (Diptera). Arbeiten über

Morphologische und Taxonomische Entomologie aus Berlin-Dahlem, 5, 253–259.

Henning, J., Schnitzler, F.R., Pfeiffer, D.U. & Davies, P. (2005) Influence of weather conditions on fly

abundance and its implications for transmission of rabbit hemorrhagic disease virus in the North

Island of New Zealand. Medical and Veterinary Entomology, 19, 251–262.

Horenstein, M.B., Linhares, A.X., Ferradas, B.R. & García, D. (2010) Decomposition and dipteran

succession in pig carrion in central Argentina: ecological aspects and their importance in forensic

science. Medical and Veterinary Entomology, 24, 16–25.

Howard, L.O. (1900) A Contribution to the study of the insect fauna of human excrement. Proceedings

of the Washington Academy of Science, 2, 541–604.

Johnston, T.H. & Tiegs, C.W. (1922) Sarcophagidae flies in the Australian Museum collection.

Records of the Australian Museum. 13: 175–188.

Judd, W.W. (1970) Studies on the Byron Bog in Southwestern Ontario XLIV flies (Sarcophagidae,

Muscidae and Tachinidae) trapped in the Bog. Entomology News, 81, 189–190.

Knipling, E.F. (1936) A comparative study of the first-instar larvae of the genus Sarcophaga

(Calliphoridae, Diptera), with notes on the biology. The Journal of Parasitology, 22 (5), 417–454.

Kunz, S.E. (1978) Notes on the seasonal activity of dung infesting Diptera in Central Texas. The

Southwestern Entomologist, 3 (3), 167–170.

Leandro, M.J.F. & D‟Almeida, J.M. (2005) Levantamento de Calliphoridae, Fanniidae, Muscidae e

Sarcophagidae em um fragmento de mata na Ilha do Governador, Rio de Janeiro, Brasil.

Iheringia, Série Zoologia, 95 (4) 377–381. [In Portuguese]

138

Lehrer, A.Z. & Barbet, A. (2008) Sarcophagides de la faune de Nouvelle-Calédonie (Diptera,

Sarcophagidae). Fragmenta Dipterologica, 17: 1–30. [In French]

Leite, A.C.R. & Lopes, H.S. (1987) Second contribution to the knowledge of the larvae of the

Raviniini (Diptera, Sarcophagidae) based on observations using scanning electron microscope.

Memórias do Instituto Oswaldo Cruz, 82 (2), 219–226.

Linhares, A.X. (1981) Synanthropy of Calliphoridae and Sarcophagidae (Diptera) in the city of

Campinas, São Paulo, Brazil. Revista Brasileira de Entomologia, 25, 189–215.

Lopes, H.S. & Albuquerque, D.O. (1955) Los insectos de las Islas Juan Fernandez. Calliphoridae et

Sarcophagidae (Diptera). Revista Chilena de Entomología, 4, 3–20.

Lopes, H.S. & Leite, A.C.R. (1986) Studies on some features of the first instar larvae of Oxysarcodexia

(Diptera, Sarcophagidae) based on scanning electron microscope observations. Revista Brasileira

de Biologia, 46 (4), 741–746.

Lopes, H.S. & Leite, A.C.R. (1987) Third contribution to the knowledge of the Raviniini (Diptera,

Sarcophagidae), based on observations of the larvae, using scanning electron microscope.


Lopes, H.S. & Tibana, R. (1982) Sarcophagid flies (Diptera) from Sinop, State of Mato Grosso, Brazil.


Lopes, H.S. & Tibana, R. (1987) On Oxysarcodexia (Diptera, Sarcophagidae), with descriptions of five

new species, key, list and geographic distribution of the species. Revista Brasileira de Biologia,

47 (3), 329–347.

Lopes, H.S. & Tibana, R. (1991) Sarcophagidae (Diptera) de Roraima, Brasil. Acta Amazonica, 21,

151–157. [In Portuguese]

Lopes, H.S. (1933) Sobre algumas espécies de Sarcophaga do Brasil, com a descripção de cinco

espécies novas (Dipt. Sarcophagidae). Revista de Entomologia, 3 (2), 153–158. [In Portuguese]

Lopes, H.S. (1935) Notas sobre Sarcophagidae com descripção de tres espécies novas do Brasil (Dipt.).

Revista de Entomologia, 5, 315–322. [In Portuguese]

Lopes, H.S. (1938) Notas sobre Sarcophagidae neotrópicos. Um novo gênero e algumas novas

espécies. Memórias do Instituto Oswaldo Cruz, 33, 333–348 + 6 pls. [In Portuguese]

Lopes, H.S. (1939) Sobre alguns Sarcophagídeos de Misiones (Argentina). Physis, 17, 117–123. [In

Portuguese]

Lopes, H.S. (1943) Contribuição ao conhecimento das larvas dos Sarcophagidae com especial

referência ao esqueleto cefálico (Diptera). Memórias do Instituto Oswaldo Cruz, 38, 127–163. [In

Portuguese]

139

Lopes, H.S. (1946a) Um novo gênero e três novas espécies de “Sarcophagidae” do Brasil (Diptera).

Revista Brasileira de Biologia, 5 (4), 453–462. [In Portuguese]

Lopes, H.S. (1946b) Contribuição ao conhecimento das espécies do gênero Oxysarcodexia Towsend,

1917 (Diptera Sarcophagidae). Boletim Escolar Nacional de Vetores, 1, 62–134. [In Portuguese]

Lopes, H.S. (1946c) Sarcophagidae do México, capturados pelo professor A. Dampf. (Diptera).

Memórias do Instituto Oswaldo Cruz, 44 (1), 119–146. [In Portuguese]

Lopes, H.S. (1953) Seis novos “Sarcophagidae” neotrópicos (Diptera). Revista Brasileira de Biologia,

13, 41–51. [In Portuguese]

Lopes, H.S. (1969) A Catalogue of the Diptera of the Americas South of the United States: Family

Sarcophagidae. Departamento de Zoologia, Secretaria da Agricultura, 103, 88p.

Lopes, H.S. (1973a) Notes on some Neotropical Sarcophagidae studied by H. R. Dodge (Diptera).

Anais da Academia Brasileira de Ciências, 45, 293–299.

Lopes, H.S. (1973b) Collecting and rearing sarcophagid flies (Diptera) in Brazil, during 40 years.

Anais da Academia Brasileira de Ciências, 45, 279–291.

Lopes, H.S. (1975a) Sarcophagid flies Diptera from Pacatuba, State of Ceará, Brazil. Revista Brasileira

de Biologia, 34 (2), 271–294.

Lopes, H.S. (1975b) On female holotypes of some American species described by Francis Walker and

J. Macquart (Diptera, Sarcophagidae, Calliphoridae). Revista Brasileira de Biologia, 34 (2), 535–

550.

Lopes, H.S. (1975c) New or little known Oxysarcodexia (Diptera, Sarcophagidae). Revista Brasileira

de Entomologia, 35 (3), 461–483.

Lopes, H.S. (1975d) Bredin-Archbold Smithsonian Biological Survey of Dominica: The Sarcophagidae

of Dominica (Diptera). Anais da Academia Brasileira de Ciências, 45, 467–487.

Lopes, H.S. (1975e) On some new species of Sarcophagidae from Costa Rica (Diptera). Revista

Brasileira de Entomologia, 35 (3), 485–489.

Lopes, H. S. (1975f) Some new Sarcophagidae from Peru (Diptera). Revista Brasileira de Biologia, 34

(4), 573 – 580.

Lopes, H.S. (1976) On the holotypes, mostly females, of some Sarcophagidae (Diptera) described by

Francis Walker. Revista Brasileira de Biologia, 36, 629–641.

Lopes, H.S. (1978) On the Types of Some Neotropical Sarcophagidae described by H. R. Dodge

(Diptera). Revista Brasileira de Biologia, 38 (2), 501–507.

140

Lopes, H.S. (1980) On some Sarcophagidae (Diptera) from Pirapora, State of Minas Gerais, Brazil.

Revista Brasileira de Biologia, 40 (1), 5–8.

Lopes, H.S. (1985) On types of Sarcophagid flies (Diptera) described by Francis Walker, Camilo

Rondani and Van der Wulp. Revista Brasileira de Entomologia, 29 (3/4), 555–558.

Macquart, J. (1843) Diptères exotiques nouveaux ou peu connus. Tome deuxième, 3e partie. Memoires

de la Société Royale des sciences, de l’Agriculture et des Arts de Lille, 1842, 162–460. [In French]

Macquart, J. (1846) Diptéres exotiques nouveaux ou peu connus. Supplément. Memoires de la Société

Royale des sciences, de l’Agriculture et des Arts de Lille, 1844, 133–364 + pls. 1–20. [Last four

pages numbered 363, 364, 363, 364. Reprinted with pagination 5–238, Roret, Paris] [In French]

Macquart, J. (1851) Diptères exotiques nouveaux ou peu connus. Suite de 4.e supplément publié dans

les Mémoirs de 1849. Memoires de la Société Royale des sciences, de l’Agriculture et des Arts de

Lille, 1850, 134–294 + pls. 1–20. [In French]

Macquart, J. (1855) Diptères exotiques nouveaux ou peu connus. 5.e supplément. Memoires de la

Société Royale des sciences, de l’Agriculture et des Arts de Lille, 2 (1), 25–156 + pls. 1–7.

[Reprinted with pagination 5–136, Roret, Paris, 1855] [In French]

Marchiori, C.H. & Linhares, A.X. (1999) Constância, dominância e frequência de dípteros muscóides e

seus parasitóides (Hymenoptera e Coleoptera), associados a fezes frescas de bovinos, em

Uberlândia, MG. Anais da Sociedade de Entomologia do Brasil, 28, 275–387. [In Portuguese]

Marchiori, C.H. (2000) Parasitóides de Sarcophagula occidua Fabricius (Diptera, Sarcophagidae)

coletados em fezes bovinas em Goiás. Pesquisa Agropecuária Tropical, 30 (1), 17–21. [In

Portuguese]

Marchiori, C.H. (2007) Gnathopleura quadridentata Wharton, 1986 (Hymenoptera; Braconidae;

Alysiinae) and their hosts collected in different substrates in Caldas Novas, Goiás. Brazilian

Journal of Biology, 67 (1), 101–103.

Marchiori, C.H., Barbaresco, L.F. & Miranda, M.F. (2007a) Parasitóides de dípteros coletados em um

matadouro de Tupaciguara, Minas Gerais, Brasil. Semina: Ciências agrárias, 28 (4), 695–700. [In

Portuguese]

Marchiori, C.H., Filho, O.M.S., Borges, M.P. & Alvarenga, V.A. (2005) Ocorrência de Trybliographa

sp. (Hymenoptera: Figitidae: Eucoilinae) como parasitóide de Oxysarcodexia thornax (Walker,

1849) (Diptera: Sarcophagidae) no Brasil. Arquivos do Instituto de Biologia, 72 (2), 265–266. [In

Portuguese]

Marchiori, C.H., Filho, O.M.S., Borges, M.P. & Alvarenga, V.A. (2007b) Parasitóides de dípteros

coletados usando armadilha ptifall em Itumbiara, Goiás. Biotemas, 20 (1), 115–118. [In

Portuguese]

141

Marchiori, C.H., Oliveira, A.T. & Linhares, A.X. (2001) Artrópodes associados a massas fecais

bovinas no sul do estado de Goiás. Neotropical Entomology, 30 (1), 19–24. [In Portuguese]

Marchiori, C.H., Pereira, L.A., Filho, S.M.O. (2002) Parasitóides de Oxysarcodexia thornax (Walker,

1849) (Diptera: Sarcophagidae) coletados no estado de Goiás, Brasil. Revista de Patologia

Tropical, 31 (1), 134–137. [In Portuguese]

Marilius, J.C., Schnack, J.A., Mulieri, P.R. & Torretta, J.P. (2007) The Sarcophagidae (Diptera) of the

Coastline of Buenos Aires City, Argentina. Journal of the Kansas Entomological Society, 80 (3),

243–251.

Mattos, W.B. (1919) As Sarcophagas de São Paulo. Thesis, Faculdade de Medicina e Cirurgia de São

Paulo, iii + 116 + xii pp. [In Portuguese]

McAlpine, J.F. (1981) Morphology and terminology. In: McAlpine, J.F., Peterson, B.V., Shewell,

G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (eds.), Manual of Nearctic Diptera. Vol. 1.

Monograph 27, Agriculture Canada, pp. 9–63.

Meiklejohn, K.A., Dowton, M. & Wallman, J.F. (2012) Notes on the distribution of 31 species of

Sarcophagidae (Diptera) in Australia, including new records in Australia for eight species.

Transactions of the Royal Society of South Australia, 136 (1), 56–64.

Mello-Patiu, C.A. & Pape, T. (2000) Definitions of Dexosarcophaga Townsend 1917 and

Sarcofahrtiopsis Hall 1933, including two new species and a redescription of Sarcofahrtiopsis

cuneata (Townsend 1935) (Diptera: Sarcophagidae). Boletín de Entomología Venezolana, 15 (2):

181–194.

Mendes, J. & Linhares, A.X. (1993) Sazonalidade, preferência por iscas e desenvolvimento ovariano

em várias espécies de Sarcophagidae (Diptera). Revista Brasileira de Entomologia, 37 (2), 355–

364. [In Portuguese]

Mendes, J. & Linhares, A.X. (2002) Cattle dung breeding Diptera in pastures in Southeastern Brazil:

diversity, abundance and seasonality. Memórias do Instituto Oswaldo Cruz, 97 (1): 37–41.

Moretti, T.C. & Godoy, W.A.C. (2013) Spatio-temporal dynamics and preference for type of bait in

necrophagous insects, particularly native and introduced blow flies (Diptera: Calliphoridae).

Journal of Medical Entomology, 50 (2), 415–424.

Moretti, T.C., Ribeiro, O.B., Thyssen, P.J. & Solis, D.R. (2008) Insects on decomposing carcasses of

small rodents in a secondary forest in Southeastern Brazil. European Journal of Entomology, 105,

691–696.

Moura, M.O. (2004) Variação especial como mecanismo promotor da coexistência em comunidades de

insetos necrófagos. Revista Brasileira de Zoologia, 21 (3), 409–419. [In Portuguese]

142

Moura, M.O., Carvalho, C.J.B. & Monteiro-Filho, E.L.A. (2005) Estrutura de comunidades necrófagas:

efeito da partilha de recursos na diversidade. Revista Brasileira de Zoologia, 22 (4), 1134–1140.

[In Portuguese]

Mulieri, P.R., Mariluis, J.C. & Patitucci, L.D. (2010) Review of the Sarcophaginae (Diptera:

Sarcophagidae) of Buenos Aires Province (Argentina), with a key and description of a new

species. Zootaxa, 2575, 1–37.

Mulieri, P.R., Patitucci, L.D., Schnack, J.A. & Mariluis, J.C. (2011) Diversity and seasonal dynamic of

an assemblage of Sarcophagid Diptera in a gradient of urbanization. Journal of Insect Science, 11

(91) 1–15.

Mulieri, P.R., Schnack, J.A., Mariluis, J.C. & Torretta, J.C. (2008) Flesh flies species (Diptera:

Sarcophagidae) from a grassland and a woodland in a nature reserve of Buenos Aires, Argentina.

Revista Biologia Tropical, 56 (3), 1287–1294.

Oliveira, T.C. & Vasconcelos, S.D. (2010) Insects (Diptera) associated with cadavers at the Institute of

Legal Medicine in Pernambuco, Brazil: implications for forensic entomology, Forensic Science

International, 198, 97–102.

Oliveira, V.C., D‟Almeira, J.M., Paes, M.J & Sanavria, A. (2002) Population dynamics of caliptrate

Diptera (Muscidade and Sarcophagidae) at the Rio-Zoo Foundantion, Rio de Janeiro, RJ, Brazil.

Brazilian Journal of Biology, 62 (2), 191–196.

Oliveira-Costa, J., Mello-Patiu C.A. & Lopes, S.M. (2001) Dípteros muscóides associados com

cadáveres humanos na cena da morte no estado do Rio de Janeiro - Brasil. Boletins do Museu

Nacional de Zoologia, 464, 1–6. [In Portuguese]

Pamplona, D., Maia, V.C., Couri, M.S., Lamas, C.J.E. & Aires, C.C.C. (2000) A survey of diptera on

Paquetá Island, Rio de Janeiro, Brazil. Entomologist’s Monthly Magazine, 136, 169–175.

Pape, T. & Dahlem, G.A. (2010) Sarcophagidae. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood,

D.M., Woodley, N.E. & Zumbado, M. (eds), A Manual of Central American Diptera. Vol. 2. NRC

Research Press, Ottawa, pp. 1313–1335.

Pape, T. (1996) Catalogue of the Sarcophagidae of the world (Insecta: Diptera). Memoirs on

Entomology, International, 8, 1–558.

Pape, T. (2008) Nominal species of Australasian/Oceanian Sarcophagidae proposed by P. J. M.

Macquart (Insecta: Diptera). Zootaxa, 1702, 41–51.

Payne, J.A. & King, E.W. (1972) Insect succession and decomposition of pig carcasses in water.

Journal of the Georgia Entomological Society, 7 (3), 153–162.

143

Peckham, D.J. (1991) Consequences relating to the inclusion of female Sarcophagids (Diptera) as prey

of Oxybelus sparideus (Hymenoptera, Sphecidae). Annals of the Entomological Society of

America, 84 (2), 170–173.

Peckham, D.J., Kurczewski, F.F. & Peckham, D.B. (1973) Nesting behavior of Neartic species of

Oxybelus (Hymenoptera: Sphecidae). Annals of the Entomological Society of America, 66 (3),

647–661.

Pérez, S.P., Duque, P. & Wolff, M. (2005) Successional behavior and occurrence matrix of carrion-

associated arthropods in the urban area of Medellfn, Colombia. Journal of Forensic Sciences, 50

(2), 448–453.

Pombal, E.C.P. & Morellato, P.C. (2000) Differentiation of floral color and odor in two fly pollinated

species of Metrodorea (Rutaceae) from Brazil. Plant System Evolution, 221, 141–156.

Prado, A. & Fonseca, F. (1932) Algumas espécies novas de Sarcophagas da cidade de São Paulo

(Diptera. Stephanostomatidae). Revista Medico-Cirúrgica do Brasil, 40, 35–39. [In Portuguese]

Ramírez-Mora, M.A., Buenaventura, E., Gómez-P, L.M. & Amat, E. (2012) Uptadet cheklist and new

records of Calyptrate carrion flies (Diptera, Schizophora) from Valle de Aburrá and other

localities in Colombia. Entomotropica, 27 (1), 27–35.

Reed, H. B. (1958) A study of dog carcass communities in Tennessee, with special reference to the

insects. American Midland Naturalist, 59 (1), 213–245.

Reinhard, H.J. (1939) New genera and species of muscoid Diptera. Bulletin of Brooklyn Entomological

Society, 34: 61–74.

Roback, S.S. (1954) The evolution and taxonomy of the Sarcophaginae (Diptera, Sarcophagidae).

Illinois Biological Monographs, 23, 1–181. http://dx.doi.org/10.1086/401540

Rondani, C. (1850) Osservazioni sopra alquante specie di Esapodi ditteri del Museu Torinense. Nuovi

annali dele scienze naturali, 3 (2), 165–197. [In Italian]

Rosa, T.A., Babata, M.L.Y., Souza, C.M., Sousa, D., Mello-Patiu, C.A., Vaz-de-Mello, F.Z. &

Mendes, J. (2011) Arthropods associated with pig carrion in two vegetation profiles of Cerrado in

the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, 55 (3), 424–434.

Salazar-Ortega, J.A., Amat, E. & Gomez-Piñerez, L.M. (2012) A check list of necrophagous flies

(Diptera, Calyptrate) from urban area in Medellín, Colombia. Revista Mexicana de Biodiversidad,

83, 502–505.

Sánchez-Núñez, D.A. & Pineda-Mancera, J.E. (2012) Pollination and fruit set in the main neotropical

mangrove species from Southwestern Caribbean. Aquatic Botany, 103, 60–65.

Sanders, D.P. & Dorson, R.C. (1966) The insect complex associated with bovine manure in Indiana.

Annals of the Entomological Society of America, 59 (5), 955–959.

144

Schiner, I.R. (1868) Diptera. In: Reise der österreichischen Fregatte Novara um die Erde in den

Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B, von Wüllerstorf-Urbair.

Zoologischer Theil. Vol. 01. Zweiter Band. Abtheilung, Wien, pp. 388–392. [In German]

Silva, K.P. & Mello-Patiu, C.A. (2008) Morfologia comparada da terminalia masculina de quatro

espécies de Oxysarcodexia Towsend, 1917 (Diptera, Sarcophagidae). Arquivos do Museu

Nacional, 66 (2), 363–372. [In Portuguese]

Soares, W.F. & Mello-Patiu, C.A. (2010) Two new Neotropical species of the genus Oxysarcodexia

Towsend (Diptera, Sarcophagidae). Revista Brasileira de Entomologia, 54 (1), 72–75.

Sousa, J.R.P., Esposito, M.C. & Carvalho-Filho, F.S. (2011) Composition, abundance and richness of

Sarcophagidae (Diptera: Oestroidea) in forests and forest gaps with different vegetation cover.

Neotropical Entomology, 40 (1), 20–27.

Stamper, T., Dahlem, G.A., Cookman, C., & Debry R.W. (2012) Phylogenetic relationships of flesh

flies in the subfamily Sarcophaginae based on three mtDNA fragments (Diptera: Sarcophagidae).

Systematic Entomology, 38 (1): 35–44. 10.1111/j.1365-3113.2012.00646.x

Tabor, K.L., Brewster, C.C. & Fell, R.D. (2004) Analysis of the successional patterns of insect on

carrion in Southwest Virginia. Journal of Medical Entomology, 41 (4), 785–795.

Tabor, K.L., Fell, R.D. & Brewster, C.C. (2005) Insect fauna visiting carrion in Southwst Virginia.

Forensic Science International, 150, 73–80.

Thomson, C.G. (1869) Diptera. Species nova descripsit. In: Kongliga Svenska Fregatten Eugenies resa

omkring jorden under befäl af C.A. Virgin, åren 1851–1853. Vetenskapliga Iakttagelser på H.M.

Konung Oscar den förstes befallning, af K. Svenska Vetenskaps-Akademien. Vol. 2. Zoologi. Part

1. Insecta. P.A. Norstedt & Soner, Stockholm, [1868], 617 pp. + 9 pls. [In German]

Tibana, R. & Mello, C.A. (1983a) Estudo sobre as fêmeas de Oxysarcodexia do grupo Peltata (Diptera,

Sarcophagidae). Revista Brasileira de Biologia, 43 (3), 241–250. [In Portuguese]

Tibana, R. & Mello, C.A. (1983b) Redescrição de Oxysarcodexia thornax (Walker, 1849) e descrição

de O. morretesi, sp. n. (Diptera, Sarcophagidae). Revista Brasileira de Entomologia, 27 (3), 277–

283. [In Portuguese]

Tibana, R. & Mello, C.A. (1985) O sintergito 6+7 nas fêmeas de Oxysarcodexia Towsend, 1917

(Diptera, Sarcophagidae). Revista Brasileira de Biologia, 45 (4), 439–445. [In Portuguese]

Towsend, C.H.T. (1917) Genera of the dipterous tribe Sarcophagini. Proceedings of the Biological

Society of Washington, 30, 189-198.

Vairo, K.P., Mello-Patiu, C.A. & Carvalho, C.J.B. (2011) Pictorial identification key for species of

Sarcophagidae (Diptera) of potential forensic importance in Southern Brazil. Revista Brasileira de

Entomologia, 55 (3), 333–347.

145

Vasconcelos, S.D. & Araujo, M.C.S. (2012) Necrophagous species of Diptera and Coleoptera in

northeastern Brazil: state of art and challenges for the Forensic Entomologist. Revista Brasileira

de Entomologia, 56 (1), 7–14.

Walker, F. (1836) Diptera. In: Curtis, J., Haliday, A. H. & Walker, F. (eds), Descriptions of insects

collected by Captain P. P. King, R. N., F. R. S., in the survey of the Straights of Magellan. Vol.

17. Trans. Linn. Soc. London, pp. 315–369.

Walker, F. (1849) Part IV. In: List of the specimens of dipterous insects in the collection of the British

Museum. Vol. 01. British Museum, London, pp. 689–1172.

Walker, F. (1853) Diptera. Part IV. In: Saunders, W.W. (ed.), Insecta Saundersiana: or characters of

undescribed insects in the collection of William Wilson Saunders, Esq., F.R.S., F.L.S., Vol. 1, Van

Voorst, London, 474 pp. + 8 pls.

Walker, F. (1857) Characters of undescribed Diptera in the collection of W. W. Saunders, Esq., F. R.

S. [part]. Transactions of the Entomological Society of London, London, 260–280 pp.

Walker, F. (1861) Characters of undescribed Diptera in the collection of W. W. Saunders, Esq., F. R.

S. [part]. Vol. 2. Transactions of the Entomological Society of London, London, 297–334 pp.

Walsh, G.C. & Posse, M.C. (2003) Abundance and seasonal distribution of predatory coprophilous

argentine rove beetles (Coleoptera: Staphylinidae), and their effects on dung breeding flies. The

Coleopterists Bulletin, 57 (1), 43–50.

Wharton, R.A. & Moon, R.D. (1979) Puparia of cyclorrhaphous Diptera from bovine dung in open

pasture and rangeland in the transition zone of Western North America. Annals of the


Wiedemann, C.R.W. (1830) Aussereuropäische zweiflügelige Insekten. Als Fortsetzung des

Meigenschen Werkes. Zweiter Theil., Schulz, Hamm, xii + 684 pp. +5 pls. [In German]

Williston, S.W. (1896) On the Diptera of St. Vincent (West Indies). Transactions of the Entomological

Society of London, Part 3, 253–446. http://dx.doi.org/10.1111/j.1365-2311.1896.tb00965.x

Wolff, M., Uribe, A., Ortiz, A. & Duque, P. (2001) A preliminar study of forensic entomology in

Medellín, Colombia. Forensic Sicence International, 120, 53–59.

Wulp, F.M. (1895) Fam. Muscidae [part]. In: Godman, F. D. & Salvin, O. (eds). Biologia Centrali-

Americana. Vol. 2. Class Insecta. London, 289 pp.

Yepes-Gaurisas, D., Sánchez-Rodríguez, J.D., Mello-Patiu, C.A. & Echeverri, M.W. (2013)

Synantropy of Sarcophagidae (Diptera) in La Pintada, Antioquia-Colombia. Revista de Biologia

Tropical, 61 (3), 1275–1287.

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Figure 1 – Genital structures of Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae).

Abbreviations: b- basiphallus; d- distiphallus; C- cercus; Ep- epandrium; J- juxta; P- Phallus;

Po- postgonite; Pr- pregonite; Su- Surstylus; St- syntergosternite 7+8; V- vesica.

147

Appendices

Appendix 1 - Oxysarcodexia n. sp. 1. A) Habitus. B) Genitalia, lateral view. C) Genitalia,

anterior view. D) Genitalia, posterior view.

148



149



150



A

B

C

D

151



A

B

C D

152



153

Appendix 7 – A) Oxysarcodexia admixta (Lopes, 1933); B) Oxysarcodexia adunca Lopes,

1975; C) Oxysarcodexia afficta (Wulp, 1895). Habitus; genitalia lateral view; genitalia

posterior view.

A

B

C

154

Appendix 8 – A) Oxysarcodexia amorosa (Schiner, 1868); B) Oxysarcodexia angrensis

(Lopes, 1933); C) Oxysarcodexia augusta Lopes, 1946. Habitus; genitalia lateral view;

genitalia posterior view.

A

A

B

C

155

Appendix 9 – A) Oxysarcodexia aura (Hall, 1937); B) Oxysarcodexia aurata (Macquart,

1851); C) Oxysarcodexia avuncula (Lopes, 1933). Habitus; genitalia lateral view; genitalia

posterior view.

A

C

B

156

Appendix 10 – A) Oxysarcodexia bakeri (Aldrich, 1916); B) Oxysarcodexia berlai Lopes,

1975; C) Oxysarcodexia bicolor Lopes, 1946. Habitus; genitalia lateral view; genitalia

posterior view.

A

B

C

157

Appendix 11 – A) Oxysarcodexia bikini Dodge, 1966; B) Oxysarcodexia carvalhoi Lopes,

1946; C) Oxysarcodexia chaetopygialis (Williston, 1896), lateral view with and without the

presence of the pregonite. Habitus; genitalia lateral view; genitalia posterior view.

A

C

B

158

Appendix 12 – A) Oxysarcodexia cingarus (Aldrich, 1916); B) Oxysarcodexia comparilis

(Reinhard, 1939); C) Oxysarcodexia conclausa (Walker, 1861). Habitus; genitalia lateral

view; genitalia posterior view.

A

C

B

159

Appendix 13 – A) Oxysarcodexia confusa Lopes, 1946; B) Oxysarcodexia corolla Dodge,

1965, presenting the genitalia in lateral view, adapted from Lopes 1978; C) Oxysarcodexia

cuernavaca Dodge, 1966, presenting the genitalia in lateral and posterior views, adapted from

Dodge, 1966.

A

C

B

160

Appendix 14 – A) Oxysarcodexia culmiforceps (Dodge, 1966); B) Oxysarcodexia culminata

(Aldrich, 1916), both presenting the habitus; genitalia in lateral view; and genitalia in posterior

view; C) Oxysarcodexia cyanea Lopes, 1975, presenting the genitalia in lateral view, adapted

from Lopes 1975b.

B

C

A

161

Appendix 15 – A) Oxysarcodexia cyaniforceps (Hall, 1933), presenting the genitalia in lateral

view, adapted from Lopes 1975c; B) Oxysarcodexia diana (Lopes, 1933); B) Oxysarcodexia

eberti Lopes & Tibana, 1987. Habitus; genitalia lateral view; genitalia posterior view.

A

C

B

162

Appendix 16 – A) Oxysarcodexia edwardsi Lopes, 1946; B) Oxysarcodexia favorabilis

(Lopes, 1935) with habitus; genitalia lateral view; genitalia posterior view, and, for O.

favorabilis, also anterior view; C) Oxysarcodexia festiva Lopes & Tibana, 1987, presenting

the genitalia in lateral view and cerci in posterior view , adapted from Lopes & Tibana 19875.

A

B

C

163

Appendix 17 – A) Oxysarcodexia flavipes Lopes & Tibana, 1987; B) Oxysarcodexia floricola

Lopes, 1975; C) Oxysarcodexia fluminensis Lopes, 1946. Habitus; genitalia lateral view;


A

B

C

164

Appendix 18 – A) Oxysarcodexia fraterna (Lopes, 1946); B) Oxysarcodexia fringidea

(Curran & Walley, 1934); B) Oxysarcodexia galeata (Aldrich, 1916). Habitus; genitalia lateral

view; genitalia posterior view, except for O. fraterna which is presented only lateral view.

A

B

C

165

Appendix 19 – A) Oxysarcodexia grandis Lopes, 1946; B) Oxysarcodexia inflata Lopes,

1975; C) Oxysarcodexia injuncta (Walker, 1858). Habitus (except for O. grandis); genitalia

lateral view; genitalia posterior view.

A

C

B

166

Appendix 20 – A) Oxysarcodexia insolita Lopes, 1946; B) Oxysarcodexia intona (Curran &

Walley, 1934); C) Oxysarcodexia jamesi Dodge, 1968. Habitus; genitalia lateral view;


A

C

B

167

Appendix 21 – A) Oxysarcodexia major Lopes, 1946; B) Oxysarcodexia marina (Hall, 1938);

C) Oxysarcodexia meridionalis (Engel, 1931). Habitus; genitalia lateral view; genitalia

posterior view.

A

C

B

168

Appendix 22 – A) Oxysarcodexia mitifica Lopes, 1953; B) Oxysarcodexia modesta Lopes,

1946; C) Oxysarcodexia molitor (Curran & Walley, 1934). Habitus; genitalia lateral view;

genitalia posterior view, except for O. molitor with only lateral view, adapted from Lopes

1946f.

A

C

B

169

Appendix 23 – A) Oxysarcodexia morretesi Tibana & Mello, 1983; B) Oxysarcodexia nitida

Soares & Mello-Patiu, 2010; B) Oxysarcodexia notata Soares & Mello-Patiu, 2010. Habitus;

genitalia lateral view; genitalia posterior view.

A

C

B

170

Appendix 24 – A) Oxysarcodexia occulta Lopes, 1946; B) Oxysarcodexia orbitalis Dodge,

1966, adopted from Dodge, 1966; C) Oxysarcodexia pallisteri Dodge, 1966. Habitus, except

for O. orbitalis; genitalia lateral view; genitalia posterior view.

A

C

B

171

Appendix 25 – A) Oxysarcodexia parva Lopes, 1946; B) Oxysarcodexia paulistanensis

(Mattos, 1919); C) Oxysarcodexia peculiaris Lopes, 1975. Habitus; genitalia lateral view;


A

C

B

172

Appendix 26 – A) Oxysarcodexia peltata (Aldrich, 1916); B) Oxysarcodexia perneta

(Walker, 1861); C) Oxysarcodexia peruviana (Lopes, 1975), with only genitalia lateral view,

adapted from Lopes 1975f. Habitus; genitalia lateral view; genitalia posterior view.

A

B

C

173

Appendix 27 – A) Oxysarcodexia petropolitana Lopes, 1975; B) Oxysarcodexia plebeja

Lopes, 1946; C) Oxysarcodexia ramosa (Reinhard, 1939), adapted from Dodge 1966. Habitus,

except for O. ramosa; genitalia lateral view; genitalia posterior view.

A

B

C

174

Appendix 28 – A) Oxysarcodexia riograndensis Lopes, 1946; B) Oxysarcodexia sarcinata

Lopes, 1953; C) Oxysarcodexia similata Lopes & Tibana, 1987. Habitus, except for O.

ramosa; genitalia lateral view; genitalia posterior view.

A

B

C

175

Appendix 29 – A) Oxysarcodexia simplicoides (Lopes, 1933); B) Oxysarcodexia terminalis

(Wiedemann, 1830); C) Oxysarcodexia thornax (Walker, 1849). Habitus; genitalia lateral

view; genitalia posterior view.

A

B

C

176

Appendix 30 – A) Oxysarcodexia timida (Aldrich, 1916); B) Oxysarcodexia trivialis (Wulp,

1895); C) Oxysarcodexia varia (Walker, 1836). Habitus; genitalia lateral view; genitalia

posterior view, and also for O. varia, genitalia anterior view.

A

B

C

177

Appendix 31 – A) Oxysarcodexia ventricosa (Wulp, 1895); B) Oxysarcodexia villosa Lopes,

1946; C) Oxysarcodexia vittata (Walker, 1836). Habitus, except for O. vittata; genitalia lateral

view; genitalia posterior view, and also, for O. villosa genitalia in anterior view.

A

C

B

178

Appendix 32 – A) Oxysarcodexia wygodzinskyi Lopes & Tibana, 1987; B) Oxysarcodexia

xannthosoma (Aldrich, 1916); C) Oxysarcodexia xon (Dodge, 1968), adapted from Lopes

1975c; D) Oxysarcodexia zayasi Dodge, 1956. Habitus; genitalia lateral view; genitalia

posterior view. For O. xon only genitalia in lateral view.

A

B

D

C

179

6. CAPÍTULO II

ABORDAGEM FILOGENÉTICA DO GÊNERO Oxysarcodexia (DIPTERA: SARCOPHAGIDAE) COM

ESPECIAL REFERÊNCIA À GENITÁLIA ASSIMÉTRICA DE ALGUNS MACHOS

CARINA MARA DE SOUZA1, DAVID K-B CHEUNG

2, THOMAS PAPE

3 & PATRICIA

JACQUELINE THYSSEN4

1 Departamento de Biologia Animal, Universidade de Campinas - UNICAMP, CP: 6109,

CEP:13083-970, Campinas, São Paulo, Brasil. E-mail: [email protected]

2 Museu de História Natural da Dinamarca, Universitetsparken 15, DK - 2100 Copenhagen,

Dinamarca. E-mail: [email protected]

3 Museu de História Natural da Dinamarca, Universitetsparken 15, DK - 2100 Copenhagen,

Dinamarca. E-mail: [email protected]

4 Departamento de Microbiologia e Biologia, Universidade Federal de Pelotas - UFPel, CP:

354, CEP: 96010-900, Pelotas, Rio Grande do Sul, Brasil. E-mail: [email protected]

RESUMO

Uma análise filogenética baseada em caracteres morfológicos a partir do exame

machos de 54 espécies de Oxysarcodexia Townsed, 1917 (Diptera: Sarcophagidae) foi

realizada a fim de investigar a assimetria fálica vista em algumas espécies e a ocorrência de

possíveis agrupamentos de espécies dentro do gênero. Para isso, foram realizadas análises de

parcimônia por busca heurística, utilizando as metodologias de busca tradicional e novas

tecnologias do programa TNT, com e sem pesos implicados e valores de suporte de ramos de

Bremer, tanto para a matriz completa quanto para a matriz contendo apenas caracteres da

terminália. Além disso, micrografias rotacionais foram produzidas, após a confecção de

imagens de microscopia eletrônica de varredura para Oxysarcodexia fringidea (Curran &

Walley, 1934), Oxysarcodexia timida (Aldrich, 1916) e Oxysarcodexia varia (Walker, 1836),

utilizadas como modelos, para analisar mais pormenorizadamente as alterações morfológicas

consequentes da assimetria. Foram listados 73 caracteres morfológicos, dos quais 14 foram

referentes a caracteres não localizados na terminália e 59 exclusivos da terminália masculina.

A assimetria detectada foi do tipo direcional sinistral, presente em oito espécies, restringindo-

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se aos lobos terminais da vésica. Esse estado de caráter foi considerado homoplásico, com

base nas topologias obtidas. Quanto aos agrupamentos de espécies, a topologia encontrada

mostrou similaridade com o proposto na literatura, embora sem completa correspondência

para as relações entre as espécies, exceto para o “grupo ventricosa”, para o qual não houve

nenhuma correspondência.

Palavras-chave: Cladística, caracteres morfológicos, moscas coprófagas, microscopia

eletrônica de varredura rotacional.

INTRODUÇÃO

A reconstrução da filogenia de muitos grupos da ordem Diptera, incluindo aqueles

pertencentes à família Sarcophagidae (Arthropoda: Diptera), pode ser satisfatoriamente

acessada por meio do estudo de caracteres morfológicos, especialmente daqueles presentes na

terminália dos machos (e.g. Pape, 1992; 1996; Blackith et al., 1997; Giroux e Wheeler, 2009;

Giroux et al., 2010; Whitmore et al., 2013). Análises de sequências moleculares baseadas em

marcadores mitocondriais como citocromo oxidase subunidade I, 12S, 16S, e citocromo b,

e/ou nucleares como RNA ribossomal 18S e 28S, fator de elongação 1-α e fragmentos do gene

CAD, vão de encontro às abordagens morfológicas (e.g. Wells et al. 2001; Kutty et al., 2008),

corroborando entre si a confiabilidade do resgate filogenético baseado em ambas as fontes de

dados.

A morfologia externa dos sarcofagídeos é considerada, de maneira geral, altamente

homoplásica (Giroux et al., 2010). Contudo, a terminália dos machos é uma importante fonte

de dados filogenéticos, por apresentar estruturas com elevado grau de complexidade, podendo

assim ser utilizada para esclarecer a história filogenética de um determinado grupo, a partir de

análises minuciosas e comparativas (Blackith et al., 1997; Giroux et al., 2010; Whitmore et

al., 2013).

Agrupamentos para algumas espécies do gênero Oxysarcodexia Townsend, 1917

(Diptera: Sarcophagidae) já foram propostos na literatura como detalhado abaixo:

“grupo paulistanensis” (Lopes, 1973; 1975b), incluindo Oxysarcodexia bikini Dodge, 1966,

Oxysarcodexia injuncta (Walker, 1857), Oxysarcodexia paulistanensis Prado & Fonseca, 1932

e Oxysarcodexia riograndensis Lopes, 1946;

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“grupo peltata” (Lopes, 1975a; Tibana & Mello, 1983), incluindo Oxysarcodexia aurata

(Macquart, 1851), Oxysarcodexia culminata (Aldrich, 1916), Oxysarcodexia culmiforceps

Dodge, 1966, Oxysarcodexia fringidea (Curran & Walley, 1934), Oxysarcodexia intona

(Curran & Walley, 1934) e Oxysarcodexia peltata (Aldrich, 1916);

“grupo thornax” (Lopes, 1976), incluindo Oxysarcodexia afficta (Wulp, 1895),

Oxysarcodexia conclausa (Walker, 1861), Oxysarcodexia thornax (Walker, 1849) e

Oxysarcodexia timida (Aldrich, 1916);

“grupo ventricosa” (Lopes, 1975a), incluindo Oxysarcodexia admixta (Lopes, 1933),

Oxysarcodexia avuncula (Lopes, 1933), Oxysarcodexia berlai Lopes, 1975, Oxysarcodexia

carvalhoi Lopes, 1946, Oxysarcodexia diana (Lopes, 1933) e Oxysarcodexia ventricosa

(Wulp, 1895);

“grupo Xarcophaga” (Lopes, 1975a; Soares & Mello-Patiu, 2010), incluindo

Oxysarcodexia favorabilis (Lopes, 1935), Oxysarcodexia fraterna (Lopes, 1946),

Oxysarcodexia nitida Soares & Mello-Patiu, 2010, Oxysarcodexia notata Soares & Mello-

Patiu, 2010, Oxysarcodexia pallisteri Dodge, 1966, Oxysarcodexia peruviana (Lopes, 1973),

Oxysarcodexia vittata (Lopes, 1946) e Oxysarcodexia xon (Dodge, 1968).

Esses grupos foram baseados em caracteres morfológicos externos de machos e/ou de

fêmeas; entretanto, nenhuma abordagem filogenética formal para as relações interespecíficas

foi proposta.

Apesar da homogeneidade na morfologia externa das espécies inclusas em

Oxysarcodexia, é possível observar diversas modificações nas estruturas fálicas,

especialmente na vésica. Em um número considerável de espécies, os ramos da vésica sofrem

“deformações”, tornando-se assimétricos (Souza et al. in prep.). Uma vez que os organismos

do filo Arthropoda apresentam como via de regra a simetria bilateral (Triplehorn & Johnson,

2005), a ocorrência de assimetria torna-se uma característica marcante e instigante. Em

Insecta, a assimetria já foi relatada para Orthoptera, nas linhas laterais que cobrem a asa; para

Thysanoptera, na lateral do estilete mandibular; para Coleoptera, no entalhe lateral presente na

mandíbula; dentre outros insetos, cuja assimetria é observada na genitália dos machos. Os

traços assimétricos de um mesmo grupo podem ser dextrais (deslocados para a direita),

sinistrais (deslocados para a esquerda) ou randômicos (ambas as formas comuns e em

frequências similares) (Palmer, 2009).

182

A assimetria de estruturas morfológicas nos animais, assim como nos protistas e nas

plantas, é considerada um traço extremamente raro que corresponde a uma característica que

evoluiu por várias vezes de forma independente e convergente (Huber et al., 2007; Palmer,

2009). Assim, a partir de uma filogenia baseada em caracteres morfológicos e da análise

ultraestrutral por meio de microscopia eletrônica de varredura das estruturas fálicas de

algumas espécies, a ocorrência de assimetria em Oxysarcodexia foi investigada numa tentativa

de rastrear o surgimento desta característica dentro deste gênero. Além disso, buscou-se

elucidar a relação entre as espécies de Oxysarcodexia com base na filogenia morfológica

apresentada.

MATERIAL E MÉTODOS

O material utilizado para a confecção de uma matriz de caracteres morfológicos

baseada em machos compreendeu 54 espécies de Oxysarcodexia (Apêndice I) disponíveis para

exame. Fêmeas e larvas não foram inclusas nas análises devido à dificuldade de identificação

em nível específico, ao não conhecimento dessas formas para muitas espécies e também, para

as conhecidas, à indisponibilidade de espécimes para estudo. De acordo com uma análise

filogenética prévia para Sarcophaginae (Giroux et al., 2010), as espécies Dexosarcophaga

transita (Townsend, 1917) e Ravinia belforti (Prado & Fonseca, 1932) foram inclusas nas

análises como grupo externo, por pertencerem a gêneros considerados filogeneticamente

próximos à Oxysarcodexia.

A matriz de dados foi compilada usando o programa Mesquite versão 2.75 (Maddison

& Maddison, 2011), incluindo caracteres morfológicos registrados a partir do exame

comparativo entre as espécies. Para os caracteres morfológicos não localizados na terminália,

a terminologia usada seguiu McAlpine (1981) e, para aqueles presente na terminália dos

machos, seguiu-se Mello-Patiu & Pape (2000) e Giroux et al. (2010). A filogenia foi inferida

com auxílio do programa TNT (Goloboff et al., 2008), a partir de análises de parcimônia por

busca heurística, utilizando as metodologias de busca tradicional (“traditional search” com o

algoritmo heurístico “tree-bissection-reconnection” – TBR; RAM 1.000 Mb; memória de

99.999 árvores; “random seed” de 50.000 árvores, 1.000 réplicas; e 1.000 árvores salvas por

replicação) e novas tecnologias de busca (“new technology search” com o algoritmo heurístico

TBR; RAM 1.000 Mb; memória de 99.999 árvores; “ratched search” com 10 sequências

183

adicionadas inicialmente e 10 árvores salvas por replicação). As árvores mais parcimoniosas

foram geradas considerando-se pesos iguais e pesos implicados (k=3). Os valores de suporte

de Bremer para os ramos foram calculados para 50.000 árvores sub-ótimas com até 10 passos

a mais que a árvore mais curta obtida. Foi realizada também uma análise por busca tradicional

e valores de suporte de Bremer (mesmos parâmetros descritos acima) para a matriz

considerando apenas caracteres da terminália, considerados na literatura como mais

informativos.

Para analisar as alterações morfológicas consequentes da assimetria, as espécies

Oxysarcodexia fringidea (Curran & Walley, 1934), Oxysarcodexia timida (Aldrich, 1916) e

Oxysarcodexia varia (Walker, 1836), que apresentam vésicas assimétricas e são mais

frequentes que as demais portadoras de tal característica, foram utilizadas como modelos de

estudo. Para isso, essas espécies foram submetidas à técnica de microscopia eletrônica de

varredura (MEV) para a produção de micrografias rotacionais. A terminália de espécimes

alfinetados foi cuidadosamente seccionada no nível do epândrio, colocada em ácido lático

80% aquecido por aproximadamente 5 minutos, dissecada, lavada em água morna e

desidratada em etanol 96%. O processo de clarificação com ácido lático foi preferido ao

hidróxido de potássio a fim de evitar deformações em estruturas membranosas do falo

(Triplehorn & Johnson, 2005). A confecção de micrografias rotacionais interativas foi

realizada de acordo com a metodologia proposta por Cheung et al. (2013). Cada falo, após

secar naturalmente, foi posicionado em um suporte metálico (stub) com fita adesiva dupla

face, coberto com platina para que, então, as imagens fossem produzidas, sequencialmente,

utilizando um microscópio eletrônico de varredura do tipo “JEOL-JSM-6335F”, localizado no

Museu de Zoologia da Universidade de Copenhagen, Copenhagen, Dinamarca. As imagens

foram submetidas à edição para ajuste de exposição, brilho e contraste, além do ajuste de

tamanho e qualidade, para assegurar que todas as imagens ficassem alinhadas possibilitando

uma transição contínua quando em sequência. As micrografias foram então agrupadas em uma

animação rotacional, disponibilizada na web com o auxílio de um plug-in específico (Magic

360®

).

RESULTADOS

Foram listados 73 caracteres morfológicos, dentre os quais 30 referem-se a caracteres

184

multiestado e 43 a binários. Do total, foram 26 caracteres localizados não na terminália,

distribuídos na cabeça, tórax e abdômen, excluindo a terminália, e 47 restritos à terminália dos

machos, divididos entre sintergosternito 7+8, surstilo, cercos, pênis, pré e pós-gonitos e vésica

(Apêndices II e III). As árvores de consenso resultantes de 12 árvores mais parcimoniosas

geradas pela método de parcimônia e busca heurística inferida por busca tradicional (Fig. 1) e

de três árvores mais parcimoniosas geradas pelas novas tecnologias de busca (Fig. 2),

considerando pesos iguais para os caracteres em ambos os casos, apresentaram as mesmas

ramificações e topologias, com exceção apenas de O. mitifica. As árvores de consenso, quando

associadas ao valor de suporte de Bremer (Figs. 3 e 4), apresentaram as mesmas topologias e

valores associados, exceto para o clado O. aurata, O. intona e O. peltata, melhor acomodado

na árvore resultante da busca por novas tecnologias. A utilização do peso implicado resultou

em árvores com as mesmas ramificações e topologias, tanto para a busca tradicional (Fig. 5)

quanto via novas tecnologias de busca (Fig. 6). A análise sob os critérios de busca tradicional,

pesos iguais e suporte de Bremer, utilizando a matriz com caracteres apenas da terminália,

gerou uma árvore de consenso a partir de 14 árvores mais parcimoniosas (Fig. 7). Sob os

mesmos critérios, porém com pesos implicados, observou-se maior separação entre os clados,

apesar do suporte de Bremer ainda ser considerado baixo (Fig. 8).

A genitália assimétrica foi detectada em oito espécies: Oxysarcodexia corolla Dodge,

1965, Oxysarcodexia edwardsi Lopes, 1946, Oxysarcodexia fringidea, Oxysarcodexia ramosa

(Reinhard, 1939), Oxysarcodexia sarcinata Lopes, 1953, Oxysarcodexia timida,

Oxysarcodexia varia e Oxysarcodexia villosa Lopes, 1946 (Fig. 9), sendo que exemplares de ,

O. edwardsi, O. fringidea, O. sarcinata, O. timida, O. varia e O. villosa foram examinados e

O. corolla e O. ramosa foram determinadas assimétricas com base nas descrições originais e

suas ilustrações (Lopes, 1946; Dodge, 1965). As micrografias rotacionais dos falos de O.

fringidea, O. timida e O. varia foram disponibilizadas no sítio eletrônico

<http://dkbdigitaldesigns.com/semrotate/sem.html>. A análise por MEV da composição fálica

das espécies O. fringidea, O. timida e O. varia (Fig. 10) permitiu verificar que a assimetria

está restrita aos lobos terminais da vésica, com redução e/ou deslocamento do lobo terminal

esquerdo, correspondendo a uma assimetria direcional sinistral.

185

DISCUSSÃO

Os baixos valores de suporte de Bremer alcançados em todas as análises refletem o alto

grau de homoplasia da matriz de caracteres. Isso pode ser em parte decorrente de uma

fragilidade da matriz em resposta à escolha dos caracteres, mas é mais provável que seja

devido ao alto grau de homoplasia já relatado na literatura para a morfologia externa de

Sarcophagidae, assim como de outros dípteros (e.g. Couri & Pont, 2000; O‟Hara, 2002;

Giroux et al., 2010). Apesar das árvores terem sido geradas a partir da matriz com caracteres

homoplásicos, a resolução entre os taxa parece ser razoável.

Um agrupamento inserido em Sarcophaginae e denominado “clado Ravinia”

compreende os gêneros Ravinia Robineau-Desvoidy, 1863 e Oxysarcodexia Townsend, 1917,

considerados grupos próximos ou irmãos em várias filogenias. A presença de ornamentação,

representada por estrias transversais (“festoon-like”) largas e nítidas em torno do anel da

cavidade oral das larvas de primeiro estádio e a presença de ctenídios formados por espinhos

achatados no fêmur médio, são características que corroboram a proximidade destes gêneros

(Roback, 1954; Downes, 1955; Lopes, 1982; Pape, 1996; Giroux et al., 2010). As filogenias

baseadas em caracteres moleculares, no entanto, são divergentes. Stamper et al. (2012),

utilizando análises Bayesianas para avaliar a variação gênica proveniente das regiões

mitocondriais COI, a subunidade II (COII) e a subunidade 4 da desidrogenase (ND4),

encontrou suporte para o clado Ravinia + Oxysarcodexia, embora mais consistente somente

após considerar previamente a politomia. Já Kutty et al. (2010) não encontraram suporte para

uma relação próxima destes dois gêneros ao analisar regiões gênicas mitocondriais (12S, 16S,

COI e citocromo b) e nucleares (18S, 28S, a região carbamoil fosfato sintetase – CAD

(rudimentar) e o fator de elongação 1-α). Vale ressaltar que no estudo de Kutty et al. (2010)

apenas uma espécie de cada gênero foi inclusa nas análises, o que pode ter gerado um viés nos

resultados, culminando na ausência de suporte para a relação entre esses gêneros. Nas

presentes análises, Ravinia enraizou o cladograma, tendo como grupo-irmão um clado

formado por Dexosarcophaga Townsend, 1917 e Oxysarcodexia. Essa divergência na

proximidade desses gêneros, em comparação à literatura, pode ter ocorrido devido à inclusão

de apenas uma espécie de cada um desses gêneros, enviesando a filogenia aqui obtida.

A formação de um clado para as espécies O. nitida, O. notata e O. vittata sob a análise

por busca tradicional com pesos implicados para a matriz com caracteres apenas da terminália

186

corrobora a proximidade destas espécies já apontada por Lopes (1975a) e Soares & Mello-

Patiu (2010). Esses autores nomeiam o grupo formado por tais espécies como “grupo

Xarcophaga”, baseando o agrupamento na forma do falo, alongado e afinado, com um

alargamento apical característico.

As espécies O. injuncta, O. paulistanensis e O. riograndensis, pertencentes ao “grupo

paulistanensis”, caracterizado pela morfologia das fêmeas, com o tergito 8 presente e a

ausência de cerdas representando o tergito anal (Lopes, 1975b), não formaram um clado único,

considerando a árvore recuperada a partir da análise por busca tradicional com pesos

implicados para a matriz com caracteres apenas da terminália. Do “grupo paulistanensis”,

apenas as espécies O. paulistanensis e O. riograndensis se apresentaram como grupos irmãos,

abrigados em um único clado. As demais espécies desse grupo ficaram agrupadas em um

clado maior, que abrigou também outras espécies que apresentam a vésica com o ápice

filamentoso, como visto, por exemplo, em O. carvalhoi e O. amorosa.

Apesar de O. intona e O. aurata formarem um grupo irmão de O. peltata em todas as

árvores geradas, as demais espécies do “grupo peltata” (O. culminata, O. culmiforceps e O.

fringidea), caracterizado pela redução do tergito genital nas fêmeas (Lopes 1975c), não se

apresentaram como irmãs desse clado. Da mesma forma, a dilatação dorsoapical larga do falo

se mostrou como uma característica homoplásica, uma vez que O. galeata e O. varia, que

também apresentam tal estado de caráter, foram acomodadas em clados filogeneticamente

distantes.

O “grupo thornax”, caracterizado pela presença, nas fêmeas, de uma larga região

membranosa entre o tergito 5 e o tergito 6+7, sendo este último reduzido, apresentou-se nas

presentes análises formando um clado que não incluiu O. timida e foi filogeneticamente

próximo a O. occulta. Similarmente ao observado para o “grupo peltata”, a dilatação

dorsoapical média do falo se mostrou como uma característica homoplásica devido à formação

dos clados para O. conclausa, O. riograndensis, O. thornax e O. vittata.

Para as espécies do “grupo ventricosa”, caracterizado pela presença, nas fêmeas, do

tergito 6+7 inteiro e fortemente esclerotizado (Lopes, 1975a), não foi observada a formação de

agrupamentos em nenhuma das árvores obtidas. Isso pode ter ocorrido devido à ausência de

caracteres da morfologia das fêmeas na matriz utilizada e responsáveis pelo agrupamento

proposto na literatura.

187

Em abordagens filogenéticas, a simetria bilateral é considerada um estado padrão

devido à predefinição dos eixos anteroposterior e dorsoventral. As alterações no padrão

simétrico surgem devido a mudanças nos eixos que deslocam o lado oposto, embora os

mecanismos envolvidos sejam ainda desconhecidos para a maioria dos organismos que

apresentam estruturas morfológicas assimétricas. Os caracteres assimétricos podem ser

divididos em dois tipos: antissimétrico ou assimetria aleatória, no qual as formas sinistrais

(deslocamento para a esquerda) e dextrais (deslocamento para a direita) são igualmente

comuns numa mesma espécie; e assimetria direcional, no qual a maioria dos indivíduos

apresenta assimetria no mesmo lado (Palmer, 2004). As assimetrias direcionais fixas permitem

o desenvolvimento de estudos moleculares a fim de investigar se a regulação gênica dos

processos envolvidos na assimetria é feita por alguns ou vários genes. Para assimetrias

aleatórias, a investigação de regulação gênica é praticamente impossível devido à variação

dextral ou sinistral ser considerada uma variação fenotípica quase sempre sem uma base

genética (Palmer, 2009). Além de aranhas (Pholcidae e Theridiidae), Apterygota, Paleoptera,

Orthoptera, Phasmida, Embiidina, Grylloblattodea, Mantophasmatodea, Plecoptera,

Dermaptera, Zoraptera, Dictyoptera, Thysanoptera, Heteroptera, Psocodea, Neuropterida,

Siphonaptera, Mecoptera, Strepsiptera, Coleoptera, Hymenoptera, Trichoptera e Lepidoptera

possuem representantes portadores de caracteres assimétricos já relatados na literatura (Huber

et al., 2007).

Das espécies de Oxysarcodexia com vésica assimétrica, cinco (O. corolla, O. edwardsi,

O. fringidea, O. timida e O. villosa) ocorrem exclusivamente na região Neotropical; uma (O.

ramosa) é exclusiva da região Neártica; uma (O. sarcinata) ocorre nas regiões Neotropical e

Neártica; e outra (O. varia) nas regiões Neotropical e Australásia/Oceânica (Souza et al., in

prep.). A ocorrência de assimetria fálica em Oxysarcodexia já foi associada a deformações

causadas por compressões durante a fase de pupa (Lopes, 1946). Contudo, esta justificativa

parece ser procedente apenas para variações fenotípicas sem conexão com fatores genéticos, o

que parece não ser o caso da genitália assimétrica das espécies anteriormente mencionadas.

Isto porque, no caso de ser fixa (quando o desvio ou modificação da estrutura sempre ocorre

do mesmo lado), a assimetria pode ser considerada independente de fatores extrínsecos, como

compressão, e dependente de fatores genéticos, logo um caráter herdado (Palmer, 2009).

O tipo de assimetria visto em Oxysarcodexia, direcional sinistral com redução e/ou

188

deslocamento do lobo terminal, é similar ao observado em Syrphidae, que geralmente

apresenta deslocamento ou encurtamento das estruturas genitais do lado esquerdo (Metcalf,

1921), embora neste caso a assimetria possa ser decorrente da rotação que as estruturass da

genitália sofrem após a emergência do adulto. Em Tachydromiinae (Diptera: Empididae), por

exemplo, a terminália assimétrica pode ser encontrada em todos os taxa, sendo que o lado

direito apresenta-se mais alargado que o esquerdo (Chvála, 1975; Cumming & Cooper, 1992),

contrapondo o visto em Oxysarcodexia.

A ocorrência de genitália assimétrica em Diptera é considerada esporádica, embora em

Acalyptratae o ápice assimétrico do falo seja recorrente (Huber et al., 2007). Na superfamília

Empidoidea e em Phoridae, por exemplo, considera-se que todos os machos apresentam

genitálias assimétricas em certo grau, apesar da simetria bilateral ser considerada o estado

padrão (Huber et al., 2007; Nakayama, 2008; 2012). Huber et al. (2007) apresentam um

panorama amplo e detalhado sobre a ocorrência de assimetria entre os dípteros e outros

artrópodes, listando a presença desse estado para algumas espécies de Agromyzidae,

Anthomyiidae, Asteiidae, Atelestidae, Bombyliidae, Brachystomatidae, Ceratopogonidae,

Diopsidae, Dolichopodidae, Drosophilidae, Dryomyzidae, Empididae, Ephydridae, Hybotidae,

Lauxaniidae, Lonchaeidae, Micropezidae, Mycetophilidae, Periscelididae, Phoridae,

Pipunculidae, Scatopsidae, Sepsidae, Sphaeroceridae, Syrphidae, Tachinidae, Tephritidae,

Thaumaleidae, Tipulidae (Diptera: Arthropoda). As estruturas morfológicas da terminália dos

machos que podem ser acometidas pela assimetria são os pós e pré-gonitos, o surstilo, o

epândrio, o hipândrio, os gonostilos, os cercos e o falo (Huber et al., 2007).

A análise das árvores recuperadas com base em caracteres morfológicos indica que a

assimetria na genitália de Oxysarcodexia possa ser um caráter homoplásico, que surgiu várias

vezes e de maneira independente no grupo. Situação semelhante ocorre em Phoridae e

Pipunculidae, cujos caracteres assimétricos da genitália dos machos parecem ter evoluído

várias vezes dentro da família, sendo filogeneticamente informativos para o estudo cladístico

de determinados grupos, gêneros ou mesmo subfamílias (Rafael & De Meyer, 1992;

Nakayama, 2012).

Várias hipóteses são relatadas na literatura na tentativa de esclarecer a evolução da

assimetria em estruturas genitais. Dentre elas, as pressões espaciais causadas pela disposição

das estruturas internas (favorecendo a fixação assimétrica e morfológica de órgãos internos,

189

refletindo secundariamente na genitália), a seleção sexual antagônica via coevolução (partindo

do pressuposto de divisão de funções entre os lados direito e esquerdo) e da seleção sexual por

meio da alteração de posições de acasalamento (a assimetria morfológica como uma

compensação mecânica para mudanças evolutivas e comportamentais da posição de

acasalamento) (Huber et al., 2007; Huber, 2010). Cada uma se suporta em casos específicos,

porém a última parece ser a mais recorrente explicação para o surgimento da assimetria na

genitália de insetos (Huber, 2010).

Quanto às fêmeas, a assimetria nas estruturas genitais é muito pouco estudada.

Considera-se que seja um estado raro, menos infrequente apenas em Phoridae, em que a

presença da vagina e escleritos vaginais assimétricos é provavelmente associada à assimetria

distifálica dos machos (Buck, 2001; Buck & Disney, 2001; Huber et al., 2007). Assim, estudos

futuros da morfologia externa e interna das fêmeas de Oxysarcodexia, cujos machos

apresentam assimetria podem vir a corroborar tal hipótese também para este gênero de

dípteros, uma vez que a modificação nos machos está associada diretamente ao falo. Da

mesma forma, uma análise da posição de cópula pode facilitar a compreensão de como e se as

estruturas assimétricas podem modular a interação entre machos e fêmeas de Oxysarcodexia.

Apesar da dificuldade de reconhecimento e desconhecimento de muitas espécies, o estudo

morfológico das fêmeas de Oxysarcodexia à luz da filogenia, assim como das larvas, poderá

contribuir significativamente para um melhor entendimento da relação entre as espécies.

AGRADECIMENTOS

À Dra. C.A. Mello-Patiu por possibilitar o exame dos espécimes depositados no Museu

Nacional/Universidade Federal do Rio de Janeiro; à MsC. E. Buenaventura por possibilitar o

exame dos espécimes depositados no Centro Tecnológico de Antioquia, Instituição

Universitária; e ao Dr. K.A. Johanson por possibilitar o exame dos espécimes depositados no

Museu de História Natural Sueco.

Esta pesquisa foi apoiada pela Fundação CAPES, Ministério da Educação do Brasil

(processo #0906/12-3).

190

REFERÊNCIAS BIBLIOGRÁFICAS

Blackith, R.; Blackith, R.; Pape, T. 1997. Taxonomy and systematics of Helicophagella Enderlein,

1928 (Diptera: Sarcophagidae) with the description of a new species and a revised catalogue.

Studia Dipterologica 4: 383–434.

Buck, M. & Disney, R.H.L. 2001. Revision of the Megaselia giraudii and M. densior species

complexes of Europe, including ecological notes (Diptera, Phoridae). Beiträge zur Entomologie,

51: 73–154.

Buck, M. 2001. Identification of females of European Conicera Meigen, with a discussion of certain

features of the reproductive anatomy of female Phoridae (Diptera). Deutsche Entomologische

Zeitschrift, 48: 69–81.

Cheung, D.K.-B., Brunke, A.J., Akkari, N., Souza, C.M., Pape, T. 2013. Rotational scanning electron

micrographs (rSEM): A novel and accessible tool to visualize and communicate complex

morphology. ZooKeys, 328: 47–57.

Chvála, M. 1975. The Tachydromiinae (Dipt. Empididae) of Fennoscandia and Denmark. Fauna

Entomologica Scandinavica, 3: 1–336.

Couri, M.S., Pont, A.C. 2000. Cladistic analysis of Coenosiini (Diptera: Muscidae: Coenosiinae).

Systematic Entomology, 25: 373–392.

Cumming, J.M. & Cooper, B.E. 1992. A revision of the Nearctic species of the Tachydromiinae fly

genus Stilpon Loew (Diptera: Empidoidea). The Canadian Entomologist, 124: 951–998.

Dodge, H.R. 1965. The Sarcophagidae (Diptera) of the West Indies. II. Jamaica. Annals of the

Entomological Society of America, 58: 497–517.

Dodge, H.R. 1966. Some new or little-known Neotropical Sarcophagidae (Diptera), with a review of

the genus Oxysarcodexia. Annals of the Entomological Society of America, 59: 674–701.

Downes, W.L. 1955. Notes on the morphology and classification of the Sarcophagidae and other

calyptrates (Diptera). Iowa Academy of Science, 62 (5): 514–538.

Giroux, M. & Wheeler, T.A. 2009. Systematics and phylogeny of the subgenus Sarcophaga

(Neobellieria) (Diptera: Sarcophagidae). Annals of the Entomological Society of America, 102 (4):

567–587.

Giroux, M., Pape, T. & Wheeler, T.A. 2010. Towards a phylogeny of the flesh flies (Diptera:


Sarcophaginae. Zoological Journal of the Linnean Society, 158: 740–778.

Goloboff, P.A.; Farris, J.S.; Nixon, K.C. 2008. TNT, a free program for phylogenetic analysis.

Cladistics, 24: 774–786.

191

Huber, B.A. 2010. Mating positions and the evolution of asymmetric insect genitalia. Genetica, 138:

19–25.

Huber, B.A.; Sinclair, B.J.; Schmitt, M. 2007. The evolution of asymmetric genitalia in spiders and

insects. Biological Reviews, 82: 647–698.

Kutty, S.N.; Pape, T.; Pont, A.; Wiegmann, B.M.; Meier, R. 2008. The Muscoidea (Diptera:

Calyptratae) are paraphyletic: evidence from four mitochondrial and four nuclear genes. Molecular

Phylogenetics and Evolution, 49: 639–652.

Kutty, S.N.; Pape, T.; Pont, A.; Wiegmann, B.M.; Meier, R. 2010. Molecular phylogeny of the

Calyptratae (Diptera: Cyclorrhapha) with an emphasis on the superfamily Oestroidea and the

position of Mystacinobiidae and McAlpine‟s fly. Systematic Entomology, 35: 614–635.

Lopes, H.S. 1946. Contribuição ao conhecimento das espécies do gênero Oxysarcodexia Towsend,

1917 (Diptera Sarcophagidae). Boletim Escolar Nacional de Vetores, 1: 62–134.

Lopes, H.S. 1973. Notes on some Neotropical Sarcophagidae studied by H. R. Dodge (Diptera). Anais

da Academia Brasileira de Ciências, 45: 293–299.

Lopes, H.S. 1975a. New or little known Oxysarcodexia (Diptera, Sarcophagidae). Revista Brasileira de

Entomologia, 35 (3), 461–483.

Lopes, H.S. 1975b. On female holotypes of some American species described by Francis Walker and J.

Macquart (Diptera, Sarcophagidae, Calliphoridae). Revista Brasileira de Biologia, 34 (2): 535–550.

Lopes, H.S. 1976. On the holotypes, mostly females, of some Sarcophagidae (Diptera) described by

Francis Walker. Revista Brasileira de Biologia, 36: 629–641.

Lopes, H.S. 1978. On the Types of Some Neotropical Sarcophagidae described by H. R. Dodge

(Diptera). Revista Brasileira de Biologia, 38 (2): 501–507.

Lopes, H.S. 1982. The importance of the mandible and clypeal arch of the first instar larvae in the

classification of the Sarcophagidae (Diptera). Revista Brasileira de Entomologia, 26: 293–326.

Maddison, W.P. & Maddison, D.R. 2011. Mesquite: A modular system for evolutionary analysis.

Version 2.75. http://mesquiteproject.org.

McAlpine, J.F. 1981. Morphology and terminology. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E.,

Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (eds.), Manual of Nearctic Diptera. Vol. 1.

Monograph 27, Agriculture Canada, pp. 9–63.

Mello-Patiu, C.A. & Pape, T. 2000. Definitions of Dexosarcophaga Townsend 1917 and

Sarcofahrtiopsis Hall 1933, including two new species and a redescription of Sarcofahrtiopsis

cuneata (Townsend 1935) (Diptera: Sarcophagidae). Boletín de Entomología Venezolana, 15 (2):

181–194.

192

Metcalf, C. L. 1921. The genitalia of male Syrphidae: their morphology, with special reference to its

taxonomic significance. Annals of the Entomological Society of America, 14: 169–225.

Nakayama, H. 2008. Systematic and morphological evaluations of a new species of Kuenburgia

Schmitz (Diptera: Phoridae) with remarkable asymmetry of the male genitalia, including a

bifurcated left surstylus. Entomological Science, 11: 179–188.

Nakayama, H. 2012. Complex asymmetric male genitalia of Anevrina Lioy (Diptera: Phoridae).

Arthropod Structure & Development, 41: 35–49.

O‟Hara, J.E. 2002. Revision of the Polideini (Tachinidae) of America north of Mexico. Studia

dipterologica, Supplement 10: 1–170.

Palmer, A.R. 2004. Symmetry breaking and the evolution of development. Science, 306: 828–833.

Palmer, A.R. 2009. Animal asymmetry. Current Biology, 19 (12): 473–477.

Pape, T. 1992. Phylogeny of the Tachinidae family-group (Diptera: Calyptratae). Tijdschrift voor

Entomologie, 135: 43–86.

Pape, T. 1996. Catalogue of the Sarcophagidae of the world (Insecta: Diptera). Memoirs on

Entomology, International, 8, 1–558.

Rafael, J.A. & De Meyer, M. 1992. Generic classification of the family Pipunculidae (Diptera): a

cladistic analysis. Journal of Natural History, 26: 637–658.

Roback, S.S. 1954. The evolution and taxonomy of the Sarcophaginae (Diptera, Sarcophagidae).

Illinois Biological Monographs, 23: 1–181.

Soares, W.F. & Mello-Patiu, C.A. 2010. Two new Neotropical species of the genus Oxysarcodexia

Towsend (Diptera, Sarcophagidae). Revista Brasileira de Entomologia, 54 (1): 72–75.

Souza, C.M.; Pape, T. & Thyssen, P.J. In prep. On Oxysarcodexia (Diptera: Sarcophagidae): a

taxonomic conspectus with the description of six new species.

Stamper, T., Dahlem, G.A., Cookman, C., & Debry R.W. 2012. Phylogenetic relationships of flesh

flies in the subfamily Sarcophaginae based on three mtDNA fragments (Diptera: Sarcophagidae).

Systematic Entomology, 38 (1): 35–44.

Tibana, R. & Mello, C.A. 1983. Estudo sobre as fêmeas de Oxysarcodexia do grupo Peltata (Diptera,

Sarcophagidae). Revista Brasileira de Biologia, 43 (3), 241–250.

Triplehorn, C.A. & Johnson, N.F. 2005. Collecting, preserving, and studying insects. In: Triplehorn,

C.A. & Johnson, N.F. 2005. Borror and DeLong's introduction to the study of insects. 7th ed.

Thomson Brooks/Cole, California, USA, pp. 745–778.

193

Wells, J.D.; Pape, T.; Sperling, F. 2001. DNA-based identification and molecular systematics of

forensically important Sarcophagidae (Diptera). Journal of Forensic Sciences, 46 (5): 1098–1102.

Whitmore, D., Pape, T. & Cerretti, P. 2013. Phylogeny of Heteronychia: the largest lineage of

Sarcophaga (Diptera: Sarcophagidae). Zoological Journal of the Linnean Society, 169: 604–639.

194

Figura 1 – Árvore de consenso resultante das 12 árvores mais parcimoniosas recuperadas da

análise da matriz de caracteres completa para espécies de Oxysarcodexia Townsend, 1917

(Diptera: Sarcophagidae), por meio do programa TNT, considerando pesos iguais para os

caracteres (análise “traditional search” com RAM 1.000 Mb; memória de 99.999 árvores;

“random seed” de 50.000 árvores, 1.000 réplicas; e 1.000 árvores salvas por replicação).

195

Figura 2 – Árvore mais parcimoniosa recuperada da análise da matriz de caracteres completa

para espécies de Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae), por meio do

programa TNT, considerando pesos iguais para os caracteres (análise “new technology search”

com RAM 1.000 Mb; memória de 99.999 árvores; “ratched search” com 1.000 sequências

adicionadas inicialmente e 1.000 árvores salvas por replicação).

196

Figura 3 – Árvore de consenso resultante das 12 árvores mais parcimoniosas recuperada da



caracteres e suporte Bremer calculado para 50.000 árvores sub-ótimas com até 10 passos a

mais que a árvore mais curta obtida (análise “traditional search” com RAM 1.000 Mb;

memória de 99.999 árvores; “random seed” de 50.000 árvores, 1.000 réplicas; e 1.000 árvores

salvas por replicação).

197




caracteres e suporte Bremer calculado para 50.000 árvores sub-ótimas com até 10 passos a

mais que a árvore mais curta obtida (análise “new technology search” com RAM 1.000 Mb;

memória de 99.999 árvores; “random seed” de 50.000 árvores, 1.000 réplicas; e 1.000 árvores

salvas por replicação).

198



programa TNT, considerando pesos diferenciados para os caracteres (k=3) (análise “traditional

search” com “implied weight” (k=3), RAM 1.000 Mb; memória de 99.999 árvores; “ratched

search” com 1.000 sequências adicionadas inicialmente e 1.000 árvores salvas por replicação).

199



programa TNT, considerando pesos diferenciados para os caracteres (k=3) (análise “new

technology search” com “implied weight” (k=3), RAM 1.000 Mb; memória de 99.999 árvores;

“ratched search” com 1.000 sequências adicionadas inicialmente e 1.000 árvores salvas por

replicação).

200


análise da matriz de caracteres apenas da terminália, para espécies de Oxysarcodexia

Townsend, 1917 (Diptera: Sarcophagidae), por meio do programa TNT, considerando pesos

iguais para os caracteres e suporte Bremer calculado para 50.000 árvores sub-ótimas com até

10 passos a mais que a árvore mais curta obtida (análise “traditional search” com RAM 1.000

Mb; memória de 99.999 árvores; “ratched search” com 1.000 sequências adicionadas

inicialmente e 1.000 árvores salvas por replicação).

201

Figura 8 – Árvore mais parcimoniosas recuperadas da análise da matriz de caracteres apenas

da terminália, para espécies de Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae), por

meio do programa TNT, considerando pesos implicados (k=3) para os caracteres e suporte

Bremer calculado para 50.000 árvores sub-ótimas com até 10 passos a mais que a árvore mais

curta obtida (análise “traditional search” com RAM 1.000 Mb; memória de 99.999 árvores;

“ratched search” com 1.000 sequências adicionadas inicialmente e 1.000 árvores salvas por

replicação).

202

Figura 9 – Terminália assimétrica dos machos de Oxysarcodexia Townsend, 1917. A)

Oxysarcodexia corolla Dodge, 1965 (reproduzido a partir de Lopes, 1978); B) Oxysarcodexia

edwardsi Lopes, 1946; C) Oxysarcodexia fringidea (Curran & Walley, 1934); D)

Oxysarcodexia ramosa (Reinhard, 1939) (reproduzido a partir de Dodge, 1966); E)

Oxysarcodexia sarcinata Lopes, 1953; F) Oxysarcodexia timida (Aldrich, 1916); G)

Oxysarcodexia varia (Walker, 1836); H) Oxysarcodexia villosa Lopes, 1946.

A

C

B

|

D

E

F

G

H

203

Figura 10 – Falos assimétricos submetidos à técnica de microscopia eletrônica de varredura.

A) Oxysarcodexia fringidea (Curran & Walley, 1934) (7,0kV; x65; 41,7mm); B)

Oxysarcodexia timida (Aldrich, 1916) (7,0kV; x65; 41,2mm); C) Oxysarcodexia varia

(Walker, 1836) (7,0kV; x65; 39,8mm). Ver também <http://dkbdigitaldesigns.com/

semrotate/sem.html>.

A B

C

204

APÊNDICE

APÊNDICE I. Espécies do gênero Oxysarcodexia Townsend, 1917, Dexosarcophaga Townsend,

1917 e Ravinia Robineau-Desvoidy, 1863 (Diptera: Sarcophagidae) examinadas para o

desenvolvimento da matriz de caracteres morfológicos, com indicação do local de origem

(estado ou província e país) e depositário dos espécimes examinados entre parênteses.

Onde: CE-TdeA = Tecnológico de Antioquia, Institución Universitaria, Medellín, Colômbia; L2B-DBA =

Laboratório de Entomologia do Departamento de Biologia Animal, Universidade Estadual de Campinas,

Campinas, São Paulo, Brasil; MNRJ = Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro,

Brasil; NMR = Swedish Museum of Natural History, Estocolmo, Suécia; ZMUC = Zoological Museum of

Copenhagen, Copenhagen, Dinamarca.

Espécie Local de origem e depositário

Dexosarcophaga transita (Townsend, 1917)

Mogi Guaçu, SP, Brasil (L2B-DBA)

Ipameri, GO, Brasil (L2B-DBA)

Uberlândia, MG, Brasil (L2B-DBA)

Oxysarcodexia admixta (Lopes, 1933) Jundiaí, SP, Brasil (L2B-DBA)

Oxysarcodexia adunca Lopes, 1975 Linhares, ES, Brasil (NRM)

Província de Napo, Equador (NRM)

Oxysarcodexia afficta (Wulp, 1895) Otavalo, Equador (NRM)

Chiapa de Corzo, México (ZMUC)

Oxysarcodexia amorosa (Schiner, 1868) Angra dos Reis, RJ, Brasil (NMR)

Angra dos Reis, RJ, Brasil (MNRJ)

Oxysarcodexia angrensis (Lopes, 1933) Mogi Guaçu, SP, Brasil (L2B-DBA)

Campinas, SP, Brasil (L2B-DBA)

Oxysarcodexia augusta Lopes, 1946 Belo Horizonte, MG, Brasil (NRM)

Oxysarcodexia aura (Hall, 1937)

Campinas, GO, Brasil (NRM)

Pirapora, MG, Brasil (MNRJ)

Uberlândia, MG, Brasil (L2B-DBA)

Oxysarcodexia aurata (Macquart, 1851) Província Nord, Nova Caledônia (NRM)

Society Is., Tahiti (ZMUC)

Oxysarcodexia avuncula (Lopes, 1933) Campinas, SP, Brasil (L2B-DBA)

Extrema, MG, Brasil (L2B-DBA)

Oxysarcodexia bakeri (Aldrich, 1916) Antioquia, Colômbia (CE-TdeA)

Cuba (MNRJ)

205

Continuação Apêndice I.



desenvolvimento da matriz de caracteres morfológicos, com indicação do local de origem (estado

ou província e país) e depositário dos espécimes examinados entre parênteses.



Campinas, São Paulo, Brasil; MNRJ = Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de

Janeiro, Brasil; NMR = Swedish Museum of Natural History, Estocolmo, Suécia; ZMUC = Zoological Museum

of Copenhagen, Copenhagen, Dinamarca.


Oxysarcodexia berlai Lopes, 1975 Avispas, Madre de Dios, Peru (NRM)

Igarapé, Paraqueú Rosário, MA, Brasil (MNRJ)

Oxysarcodexia bicolor Lopes, 1946 Petrópolis, RJ, Brasil (MNRJ; NMR)

Campos do Jordão, SP, Brasil (MNRJ)

Oxysarcodexia carvalhoi Lopes, 1946 Campinas, SP, Brasil (L2B-DBA)

Mogi Guaçu, SP, Brasil (L2B-DBA)

Oxysarcodexia chaetopygialis (Williston, 1896) Majorca, St. Vicent (MNRJ; ZMUC)

Oxysarcodexia cingarus (Aldrich, 1916) Great Falls, USA (ZMUC)

La Fayette, La., USA (MNRJ)

Oxysarcodexia conclausa (Walker, 1861) Hidalgo Co., Texas, USA (ZMUC)

Palmira, Colombia (MNRJ)

Oxysarcodexia confusa Lopes, 1946

Petrópolis, RJ, Brasil (NRM; ZMUC)

Iguassú, PR, Brasil (NRM)

São Paulo, SP, Brasil (MNRJ)

Oxysarcodexia culmiforceps Dodge, 1966 Extrema, MG, Brasil (L2B-DBA)

Oxysarcodexia culminata (Aldrich, 1916) Cerro de Punta, Porto Rico (NRM)

Oxysarcodexia diana (Lopes, 1933)

Antioquia, Colômbia (CE-TdeA)

Belo Horizonte, MG, Brasil (NRM)

Grajaú, RJ, Brasil (MNRJ)

Oxysarcodexia fluminensis Lopes, 1946 Antioquia, Colômbia (CE-TdeA)

Oxysarcodexia fringidea (Curran & Walley, 1934) Amazonas, Colômbia (NMR)

Oxysarcodexia galeata (Aldrich, 1916) Madson Co., Alabama, USA (NMR)

206












Oxysarcodexia grandis Lopes, 1946 Antioquia, Colômbia (CE-TdeA)

Oxysarcodexia injuncta (Walker, 1857)

Extrema, MG, Brasil (L2B-DBA)

Jundiaí, SP, Brasil (L2B-DBA)

Rio Japura, AM, Brasil (NRM)

Oxysarcodexia insolita Lopes, 1946 Província de Napo, Equador (NRM)

Oxysarcodexia intona (Curran & Walley, 1934) Rio de Janeiro, RJ, Brasil (NRM)

Guarapari, ES, Brasil (ZMUC)

Oxysarcodexia major Lopes, 1946 Antioquia, Colômbia (CE-TdeA)

Oxysarcodexia mitifica Lopes, 1953 Antioquia, Colômbia (CE-TdeA)

Oxysarcodexia modesta Lopes, 1946 Rio de Janeiro, RJ, Brasil (MNRJ; NRM)

Mazagão, AP, Brasil (MNRJ)

Oxysarcodexia nitida Soares & Mello-Patiu, 2010 Província de Napo, Equador (NRM)

Oxysarcodexia notata Soares & Mello-Patiu, 2010 Província de Napo, Equador (NRM)

Oxysarcodexia occulta Lopes, 1946 Mogi Guaçu, SP, Brasil (L2B-DBA)

Oxysarcodexia parva Lopes, 1946 Campinas, SP, Brasil (L2B-DBA)

Oxysarcodexia paulistanensis (Mattos, 1919) Campinas, SP, Brasil (L2B-DBA)

Oxysarcodexia peltata (Aldrich, 1916) Cuba (ZMUC)

Oxysarcodexia perneta (Walker, 1861)

San Cristobal, Chiapas, México (ZMUC)

México D.F., México (MNRJ)

Cuernavaca, Morelos, México (MNRJ)

Oxysarcodexia plebeja Lopes, 1946 Antioquia, Colômbia (CE-TdeA)

San Cristobal, Chiapas, México (MNRJ)

207












Oxysarcodexia riograndensis Lopes, 1946 Campinas, SP, Brasil (L2B-DBA)

Oxysarcodexia sarcinata Lopes, 1953

Calovébora, Província de Veraguas, Panamá (NRM)

Caldas, Antioquia, Colômbia (CE-TdeA)

Cerro del Aguacate, Província de Alajuela,

Costa Rica (ZMUC)

Turialba, Cartago, Costa Rica (MNRJ)

Oxysarcodexia similata Lopes & Tibana, 1987

Punta Charme, Província do Panamá, Panamá

(NRM)

Turialba, Cartago, Costa Rica (MNRJ)

Oxysarcodexia terminalis (Wiedemann, 1830) Brasília, DF, Brasil (NMR)

Petrópolis, RJ, Brasil (MNRJ)

Oxysarcodexia thornax (Walker, 1849) Campinas, SP, Brasil (DBA-L2B)

Oxysarcodexia timida (Aldrich, 1916) Aragua, Venezuela (ZMUC)

Oxysarcodexia trivialis (Wulp, 1895) Puntaneras, Costa Rica (ZMUC)

San José, Costa Rica (ZMUC)

Oxysarcodexia varia (Walker, 1836) Chubut, Argentina (NRM)

Oxysarcodexia ventricosa (Wulp, 1895) Alabama, USA (NRM)

Flórida, USA (ZMUC)

Oxysarcodexia vittata (Walker, 1836) Jundiaí, SP, Brasil (L2B-DBA)

Nova Teutonia, MG, Brasil (ZMUC)

Oxysarcodexia xanthosoma (Aldrich, 1916) Província de Napo, Equador (NRM)

Campinas, SP, Brasil (L2B-DBA)

208








Campinas, São Paulo, Brasil; MNRJ = Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro,

Brasil; NMR = Swedish Museum of Natural History, Estocolmo, Suécia; ZMUC = Zoological Museum of

Copenhagen, Copenhagen, Dinamarca.


Oxysarcodexia zayasi Dodge, 1956 Santiago, Cuba (NRM)

Las Villas Trinidad, Cuba (MNRJ)

Oxysarcodexia n. sp. 1 Amazonas, Colômbia (ZMUC)

Província de Napo, Equador (ZMUC)

Oxysarcodexia n. sp. 2 Montsinery, Guiana Francesa (ZMUC)

Oxysarcodexia n. sp. 3 Puntaneras, Monteverde, Costa Rica (ZMUC)

Oxysarcodexia n. sp. 6 Província de Napo, Equador (ZMUC)

Ravinia belforti (Prado & Fonseca, 1932) Campinas, SP, Brasil (L2B-DBA)

209

APÊNDICE II. Lista de caracteres morfológicos baseada no exame de machos de Oxysarcodexia

Townsend, 1917 (Diptera: Sarcophagidae).

1. Cabeça, área postocular, polinosidade: (0) prateada; (1) dourada; (2) dourada

esbranquiçada.

2. Cabeça, cerda ocelar: (0) bem desenvolvida (= maior ou mesmo tamanho que as menores

cerdas frontais); (1) não bem desenvolvidas (= menor que as menores cerdas frontais).

3. Tórax, cerda apical escutelar: (0) presente; (1) ausente.

4. Tórax, asa, setosidade na veia R4+5: (0) presente na ½ proximal; (1) presente nos ⅔

proximais; (2) presente nos ¾ proximais.

5. Tórax, cerdas pós-suturais dorsocentrais: (0) 2 bem diferenciadas+1–2–3 pequenas; (1) 3

bem diferenciadas (uma pequena cerda pode estar presente).

6. Tórax, pernas, cor: (0) enegrecidas; (1) marrons; (2) amareladas.

7. Tórax, pernas, fêmur médio, número de espinhos no ctenídio: (0) 4; (1) 5; (2) 6; (3) 7; (4)

9.

8. Tórax, pernas, fêmur posterior, cerdas anteroventrais: (0) presentes somente na metade

apical; (1) presentes em toda a extensão.

9. Abdômen, tergito 3, número de cerdas laterais marginais: (0) 1; (1) 2; (2) 3.

10. Abdômen, tergito 4, número de cerdas laterais marginais: (0) 1; (1) 2; (2) 3.

11. Abdômen, tergito 4, número de cerdas medianas marginais: (0) 0; (1) 1; (2) 2.

12. Abdômen, tergito 5, polinosidade: (0) completamente dourada; (1) parcialmente dourada;

(2) não dourada.

13. Abdômen, esternitos 2–4, formato: (0) retangular; (1) quadrado.

14. Abdômen, esternitos 2–4, membrana entre os esternitos: (0) visível; (1) não visível.

15. Abdômen, esternito 5, cor: (0) completamente amarelo; (1) parcialmente amarelo; (2) não

amarelo.

16. Abdômen, esternito 5, bordas da fenda: (0) paralelas; (1) divergentes.

17. Abdômen, esternito 5, ornamentação nos ramos: (0) setosidade; (1) cerdas; (2) setosidade

+ cerdas.

18. Abdômen, esternito 5, posição das cerdas: (0) no ápice do ramo; (1) na metade apical dos

ramos; (2) ao longo da borda dos ramos; (3) espalhados pela superfície dos ramos.

210

19. Abdômen, esternito 5, tamanho dos ornamentos (cerdas ou sétulas): (0) micro (menor que

ou cerca de 0,1 x o comprimento da menor sétula na superfície do tergito 4); (1) pequeno

(cerca de 0,2–0,5 x o comprimento da menor sétula na superfície do tergito 4); (2) grande

(praticamente o mesmo comprimento da menor sétula na superfície do tergito 4).

20. Abdômen, esternito 5, espaço entre os ramos: (0) uniforme; (1) não uniforme.

21. Abdômen, esternito 5, espaço entre os ramos: (0) convergindo para o ápice; (1) não

convergindo para o ápice.

22. Abdômen, esternito 5, espaço entre os ramos: (0) alargado na base; (1) não alargado na

base.

23. Abdômen, esternito 5, comprimento dos ramos: (0) curtos; (1) longos.

24. Abdômen, esternito 5, largura dos ramos: (0) estreitos; (1) medianos; (2) largos.

25. Abdômen, esternito 5, invaginação da margem distal (visível lateralmente aos ramos): (0)

presente; (1) ausente.

26. Abdômen, esternito 5, “janela” (abertura visível entre as bordas internas dos ramos): (0)

estreita; (1) intermediária; (2) larga.

27. Abdômen, sintergosternito 7+8, cor: (0) marrom amarelado; (1) marrom enegrecido.

28. Abdômen, sintergosternito 7+8, formato dorsal: (0) linear; (1) sinuoso.

29. Abdômen, sintergosternito 7+8, cerdas na margem distal: (0) 6; (1) 7; (2) 8; (3) 10; (4) 4;

(5) 9.

30. Terminália, surstilo, formato: (0) triangular, (1) retangular.

31. Terminália, surstilo, cerda discal: (0) presente; (1) ausente.

32. Terminália, cerco, formato (vista lateral): (0) curvado para trás; (1) sinuoso; (2) reto.

33. Terminália, cerco, ápice (vista lateral): (0) pontiagudo; (1) expandido; (2) normal (mesmo

tamanho que a área mediana).

34. Terminália, cerco, expansão apical (vista lateral): (0) reto; (1) oblíquo para cima; (2)

côncavo.

35. Terminália, cerco, cerdas na face ventral (vista lateral): (0) presente na metade apical; (1)

presente na metade basal; (2) presente em toda a extensão; (3) ausente somente na porção

mediana; (4) presente no ⅓ apical.

211

36. Terminália, cerco, tamanho das cerdas dorsais (vista lateral): (0) mesmo tamanho em toda

a extensão; (1) mais longas na metade basal; (2) mais longas no ⅓ basal; (3) mais longas nos

⅔ basais.

37. Terminália, cerco, formato basal (vista posterior): (0) arredondado; (1) não arredondado.

38. Terminália, cerco, formato apical (vista posterior): (0) mesmo tamanho que a porção

mediana; (1) menor que a porção mediana; (2) maior que a porção mediana.

39. Terminália, cerco, conformação (vista posterior): (0) paralela; (1) divergente.

40. Terminália, cerco, espaço entre os cercos no nível apical (vista posterior): (0) pequeno; (1)

médio; (2) grande; (3) sem espaço entre os cercos.

41. Terminália, cerco, tufos de sétulas lateromedialmente no ápice (vista posterior): (0)

presente; (1) ausente.

42. Terminália, cerco, notável constrição na porção mediana (vista posterior): (0) presente; (1)

ausente.

43. Terminália, pênis; tamanho da expansão triangular (“tooth-like”) localizada acima da

vésica (vista lateral): (0) pequena (não ultrapassa a margem distal do tubo fálico); (1) grande

(ultrapassa a margem distal do tubo fálico).

44. Terminália, pênis/distifalo, concavidade ventroapical (vista lateral): (0) presente; (1)

ausente.

45. Terminália, pênis/distifalo, borda ventroapical (vista lateral): (0) serrada; (1) reta.

46. Terminália, pênis/distifalo, sulco lateroapical (vista lateral): (0) presente; (1) ausente;

47. Terminália, pênis/distifalo, formato apical (vista lateral): (0) cônico; (1) arredondado; (2)

quadrado/retangular.

48. Terminália, pênis/distifalo, formato da margem dorsal (vista lateral): (0) reta; (1) sinuosa.

49. Terminália, pênis/distifalo, projeções ventroapicais (vista lateral): (0) presente; (1)

ausente.

50. Terminália, pênis/distifalo, dilatação dorsoapical (vista lateral): (0) presente; (1) ausente.

51. Terminália, pênis/distifalo, tamanho da dilatação dorsoapical (vista lateral): (0) pequena;

(1) média; (2) grande.

52. Terminália, pênis/distifalo, lobos laterais (vista lateral): (0) presente; (1) ausente.

53. Terminália, pênis/distifalo, expansões lateroapicais (vista dorsal): (0) presente; (1) ausente.

54. Terminália, pênis/distifalo, fenda apical (vista dorsal): (0) presente; (1) ausente.

212

55. Terminália, pênis/distifalo, tamanho da fenda apical (vista dorsal): (0) pequena; (1)

grande.

56. Terminália, pênis, sulco dorsal (vista dorsal): (0) presente; (1) ausente.

57. Terminália, pênis, comprimento do sulco dorsal (vista dorsal): (0) ao longo de toda a

extensão; (1) restrita aos ⅔ basais.

58. Terminália, pregonito, formato: (0) base expandida estreitando repentinamente no ápice;

(1) base expandida estreitando suavemente até o ápice; (2) base e ápice com praticamente a

mesma largura.

59. Terminália, pregonito, cor: (0) mesma cor em toda a extensão da estrutura; (1) ápice mais

escuro do que a base.

60. Terminália, pós-gonito; formato: (0) base expandida estreitando repentinamente no ápice;

(1) base expandida estreitando suavemente até o ápice.

61. Terminália, vésica, ramos: (0) simétricos; (1) assimétricos.

62. Terminália, vésica, lobos terminais: (0) bem desenvolvidos; (1) reduzidos.

63. Terminália, vésica, lobos laterais: (0) presentes; (1) ausentes.

64. Terminália, vésica, forma da margem ventral dos lobos laterais: (0) côncava; (1) convexa.

65. Terminália, vésica, formato dos lobos terminais: (0) filamentoso, ou pelo menos afilando

no ápice; (1) arredondada; (2) quadrada.

66. Terminália, vésica, textura dos lobos terminais: (0) membranosa; (1) parcialmente

membranosa e parcialmente esclerotizada; (2) esclerotizada.

67. Terminália, vésica, espinhos: (0) presentes; (1) ausentes.

68. Terminália, vésica, tamanho dos espinhos: (0) micro; (1) pequeno; (2) longo.

69. Terminália, vésica, posição dos espinhos: (0) presente nas faces dorsal e ventral; (1)

presente somente ao longo da borda dos lobos terminais; (2) presente somente na face ventral.

70. Terminália, vésica, projeção mediana do ramo principal: (0) presente; (1) ausente.

71. Terminália, vésica, formato do pico da projeção mediana do ramo principal: (0) angular,

(1) arredondada.

72. Terminália, vésica, comprimento da projeção mediana do ramo principal: (0) comprimento

horizontal igual ao vertical; (1) comprimento horizontal maior que o vertical; (2) comprimento

vertical maior que o horizontal.

213

73. Terminália, vésica, comprimento da margem apical do lobo terminal: (0) ultrapassa o

ponto de conexão com o ramo principal; (1) não ultrapassa o ponto de conexão com o ramo

principal.

214

APÊNDICE III. Matriz de dados provenientes dos caracteres morfológicos levantados a partir

do exame de machos de Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae) ( _ = não se

aplica; ? = “missing data”).

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36

D. transita 1 0 0 1 0 0 2 1 0 1 0 1 0 0 2 1 0 - 1 1 1 1 0 2 0 2 0 1 2 0 0 2 0 - 0 3

R. belforti 1 1 1 1 0 1 4 1 2 2 1 1 0 0 2 1 2 0 0 1 1 1 1 2 1 2 0 0 0 1 1 1 0 - 0 3

O. admixta 1 0 0 0 0 0 1 1 0 1 1 1 1 0 2 0 0 - 0 0 1 1 1 2 1 1 0 1 1 0 0 2 1 1 2 2

O. adunca 2 1 1 1 0 0 2 1 0 0 1 0 0 0 0 0 2 0 1 1 1 0 1 1 0 0 0 0 2 0 0 1 1 1 3 1

O. afficta 2 1 1 1 0 0 2 0 1 1 2 1 1 1 0 0 0 - 1 1 0 1 1 0 0 1 0 1 2 0 0 1 1 1 2 2

O. amorosa 2 1 0 1 0 1 1 1 2 1 1 1 1 1 2 0 0 - 1 0 1 1 1 1 0 1 0 1 2 0 1 2 1 1 2 3

O. angrensis 1 0 1 1 0 0 2 1 2 1 1 0 1 0 1 0 2 0 0 0 1 1 1 1 0 0 0 0 2 0 1 2 0 1 3 3

O. augusta 1 1 0 1 1 0 0 1 1 1 1 2 0 1 1 0 2 0 0 0 1 1 1 1 0 1 0 0 0 0 0 2 1 1 2 3

O. aura 0 0 1 2 0 1 2 0 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 1 1 2 1 1 1 0 0 2 0 2 1 2

O. aurata 1 0 1 1 0 0 3 1 2 2 1 1 0 0 2 0 1 0 1 0 1 1 1 2 0 0 0 0 2 0 0 1 2 2 4 1

O. avuncula 1 0 0 2 0 0 0 1 2 2 1 1 1 0 1 0 2 0 0 1 1 0 1 1 0 1 0 0 2 0 0 1 0 2 4 3

O. bakeri 1 0 0 1 0 0 0 1 ? 2 ? 0 1 1 2 0 0 - 0 0 1 1 0 1 1 0 0 0 4 1 1 2 0 1 2 3

O. berlai 2 1 0 2 0 1 1 1 0 1 1 0 1 1 1 0 2 1 0 0 1 1 1 1 0 1 0 0 0 0 0 2 1 0 3 3

O. bicolor 1 1 1 1 0 1 2 0 0 1 1 2 0 0 2 0 0 - 0 1 0 1 0 0 0 0 0 0 5 0 0 1 2 1 3 1

O. carvalhoi 1 1 0 2 0 0 1 1 1 1 1 0 1 0 0 0 0 - 0 0 0 1 1 1 1 1 0 1 2 1 0 2 1 1 4 3

O. chaetopygialis 1 0 1 2 0 1 0 1 1 0 ? 2 1 1 0 0 2 2 2 0 1 1 0 1 1 1 0 1 0 0 0 1 0 1 4 1

O. cingarus 2 1 1 1 0 1 0 1 0 1 1 1 1 1 0 0 2 0 0 0 1 0 0 0 0 2 0 0 0 0 1 2 0 1 4 3

O. conclausa 2 1 1 1 0 1 1 1 2 1 1 0 0 0 0 0 0 - 0 0 0 1 1 1 1 1 0 0 3 0 0 2 1 1 4 3

O. confusa 1 0 1 1 0 1 1 1 1 2 1 2 1 1 0 0 0 - 1 0 0 1 0 1 1 1 0 1 2 0 0 2 1 1 3 0

O. culmiforceps 1 0 1 1 0 0 3 1 1 1 1 1 0 0 1 0 2 2 1 0 1 1 0 2 1 0 1 0 2 0 1 0 0 1 4 2

O. culminata ? 0 1 1 0 0 2 ? 1 2 1 2 1 1 1 0 0 - 1 0 1 1 0 2 1 2 0 0 1 0 0 0 0 1 4 1

O. diana 1 1 1 2 0 1 3 1 2 0 1 0 1 1 0 0 2 0 0 0 1 1 0 0 1 1 0 0 0 0 1 1 1 1 4 3

O. fluminensis 1 0 0 2 0 0 0 1 ? 0 ? 1 1 1 1 0 0 - 1 0 1 1 1 1 1 1 0 0 3 0 0 2 1 1 4 3

O. fringidea 2 1 1 1 0 0 3 1 2 2 1 0 1 0 0 0 1 0 1 1 1 0 1 2 1 0 0 0 1 0 0 2 1 1 4 1

O. galeata 0 0 1 1 0 1 1 1 0 0 0 0 0 1 0 1 2 0 0 1 1 1 0 1 1 2 0 0 1 0 0 1 0 1 4 2

O. grandis 1 0 0 2 1 0 1 1 ? 0 ? 0 1 1 2 0 0 - 1 1 1 0 1 0 0 1 0 0 2 0 0 2 0 1 4 3

O. injuncta 1 1 0 1 0 0 0 1 ? 1 ? 2 1 1 1 0 1 3 1 0 1 1 1 1 1 1 0 1 2 0 0 2 1 1 2 3

O. insolita 2 1 0 0 0 1 1 1 1 1 1 1 1 0 2 0 2 3 1 1 0 1 0 2 1 1 0 1 1 0 0 1 1 0 4 3

O. intona 1 1 1 2 0 1 3 1 ? 2 ? 1 0 1 2 0 1 0 1 0 1 1 1 2 1 0 0 0 1 0 0 1 1 2 4 1

O. major 1 1 0 0 0 0 2 1 1 1 1 1 1 1 0 0 2 3 1 0 1 1 1 2 1 2 0 0 1 1 0 1 2 1 3 1

O. mitifica 1 0 1 0 0 0 1 1 ? 1 ? 0 1 1 0 0 0 - 0 1 1 0 0 2 1 0 0 0 0 0 1 2 0 1 4 0

O. modesta 2 0 1 0 0 1 1 1 0 1 1 1 1 1 0 0 1 1 2 0 1 1 1 2 1 0 1 0 - 0 0 2 0 1 2 0

O. nitida 2 0 0 1 1 0 0 1 1 1 1 1 1 0 2 0 2 0 1 1 1 0 1 1 1 0 0 0 1 0 0 2 0 1 1 0

O. notata 1 0 0 1 1 1 0 1 0 1 1 0 1 0 0 0 2 3 2 0 1 1 1 1 1 1 0 0 4 0 0 2 0 1 3 3

O. occulta 1 1 1 2 0 0 1 1 0 1 1 1 1 0 0 0 0 - 1 1 0 1 1 1 1 0 0 0 2 0 1 0 2 1 4 3

O. parva 1 0 1 1 0 0 1 1 0 0 1 1 1 1 2 0 1 3 1 0 1 1 0 1 1 1 0 0 0 0 0 1 0 1 3 3

O. paulistanensis 1 1 0 1 0 0 1 1 1 1 1 2 1 0 1 0 0 - 1 0 1 1 1 1 1 2 0 1 5 0 0 1 1 1 2 3

O. peltata 1 0 1 1 0 1 1 0 0 0 0 1 0 1 1 0 1 0 1 0 1 1 1 2 0 0 0 0 0 0 0 0 2 2 4 2

O. perneta 1 1 0 0 0 1 1 1 0 1 1 0 1 1 1 0 2 0 1 0 1 1 0 1 1 0 0 0 2 0 0 2 0 1 2 1

O. plebeja 1 1 0 1 1 0 0 1 ? 1 ? 0 1 0 1 0 2 0 1 0 1 1 1 1 1 0 0 0 2 1 1 2 1 1 0 0

O. riograndensis 1 0 0 1 0 0 0 1 1 1 1 2 1 0 2 0 0 - 2 0 1 1 1 1 1 2 0 1 1 0 0 2 1 1 3 3

O. sarcinata 1 0 0 2 0 0 1 1 1 1 1 1 1 0 1 0 1 1 1 0 1 1 1 0 1 0 0 1 0 0 0 1 1 1 3 3

O. similata 2 0 0 2 0 0 1 1 1 0 1 1 1 1 2 0 0 - 0 1 1 0 0 1 1 1 0 0 2 0 0 2 1 1 2 1

O. terminalis 0 0 1 1 1 1 1 0 1 0 1 2 1 1 1 1 2 2 0 1 1 1 0 2 1 2 0 0 1 0 0 0 0 1 4 1

O. thornax 2 1 1 1 0 0 1 1 1 1 1 1 0 0 1 0 2 0 0 1 0 1 1 1 1 1 0 0 2 1 0 2 2 0 3 1

O. timida 1 1 1 2 0 0 2 1 1 0 1 1 0 1 0 0 2 0 1 0 1 1 1 1 1 0 0 1 3 0 0 2 1 1 2 2

O. trivialis 1 1 0 1 0 0 2 1 2 2 1 1 1 1 0 0 2 0 2 0 1 1 0 2 1 1 0 0 2 0 0 1 1 1 4 3

O. varia 1 0 1 0 1 0 2 1 0 1 0 1 0 1 2 1 2 2 2 1 1 1 0 2 1 2 1 1 2 1 0 1 1 2 2 3

O. ventricosa 0 1 1 2 0 2 1 1 2 1 1 1 1 1 0 0 2 0 0 0 1 1 0 0 1 1 0 1 2 0 0 2 1 1 4 2

O. vittata 1 0 0 2 1 0 1 ? 2 1 1 1 0 1 0 0 2 0 0 1 1 0 0 2 1 1 0 1 2 1 0 1 0 1 2 1

O. xanthosoma 2 1 0 1 0 0 1 1 2 1 1 1 1 1 1 0 0 - 1 1 1 0 1 1 1 1 0 0 1 0 0 2 1 1 3 3

O. zayasi 2 1 1 2 0 2 1 1 2 1 0 2 0 1 0 1 0 - 0 1 1 1 0 2 1 2 0 1 2 0 0 2 1 1 2 2

Oxysarcodexia n. sp. A 1 0 0 1 0 0 1 1 0 1 1 0 1 1 2 0 0 - 1 0 1 1 1 2 1 1 0 1 2 0 0 2 1 1 2 3

Oxysarcodexia n. sp. B 1 1 0 1 0 0 1 1 1 0 1 1 1 1 1 0 2 0 1 1 0 1 1 1 0 1 0 ? 5 0 0 2 1 1 2 0

Oxysarcodexia n. sp. C 1 1 1 0 0 0 2 1 1 2 0 2 0 1 2 1 2 2 1 1 1 1 0 2 0 2 1 1 3 0 0 1 0 1 1 0

Oxysarcodexia n. sp. F 2 0 0 2 0 1 2 1 1 0 1 0 1 1 1 0 0 - 0 0 1 1 1 1 1 1 0 1 2 0 0 2 1 1 2 3

215

Continuação apêndice III. Matriz de dados provenientes dos caracteres morfológicos levantados a

partir do exame de machos de Oxysarcodexia Townsend, 1917 (Diptera: Sarcophagidae) ( _ = não se

aplica; ? = “missing data”).

37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73

D. transita 1 1 0 0 1 1 - 0 1 1 1 1 1 0 0 1 1 1 - 1 - 1 0 1 0 0 1 - 1 1 1 - - - - - 0

R. belforti 1 1 1 2 1 1 - 1 1 1 1 1 1 1 - 1 1 1 - 1 - 1 0 0 0 0 1 - 1 2 1 - - - - - 0

O. admixta 1 1 1 1 1 1 0 1 1 1 0 0 1 1 - 1 1 1 - 0 1 1 0 1 0 1 0 0 0 2 0 0 1 0 1 2 1

O. adunca 1 0 1 0 1 1 1 1 1 1 1 0 1 1 - 1 1 1 - 0 0 0 1 0 0 0 1 - 1 0 0 0 2 0 0 2 0

O. afficta 1 0 1 1 1 1 1 1 1 1 0 0 1 0 0 1 1 1 - 0 1 1 1 1 0 0 1 - 0 2 0 0 2 1 - - 1

O. amorosa 1 1 1 1 1 1 1 0 1 1 1 0 1 1 - 1 1 1 - 0 1 1 0 0 0 0 0 0 0 2 0 1 2 0 1 0 0

O. angrensis 0 0 0 3 1 1 1 0 1 1 0 1 1 1 - 1 1 0 0 0 0 1 1 1 0 0 1 - 0 2 0 1 0 0 0 1 0

O. augusta 0 1 1 1 1 1 1 0 0 0 1 1 1 1 - 1 0 1 - 0 1 1 0 0 0 0 0 1 0 2 0 0 2 0 1 1 0

O. aura 0 0 1 2 1 1 0 0 1 1 1 0 1 1 - 1 1 1 - 0 1 2 0 1 0 0 1 - 1 2 1 - - 1 - - 1

O. aurata 0 0 1 1 0 1 1 1 1 1 1 0 1 0 2 0 1 1 - 0 1 2 1 1 0 0 1 - 0 1 0 0 2 0 0 0 0

O. avuncula 1 0 1 0 1 1 1 0 1 1 1 0 1 1 - 1 1 1 - 0 0 1 1 1 0 0 1 - 1 1 0 1 1 0 0 1 0

O. bakeri 1 0 1 1 1 1 0 1 0 1 0 0 0 1 - 1 1 1 - 1 - 0 1 1 0 0 1 - 0 2 0 1 1 0 1 1 1

O. berlai 1 1 1 1 1 1 1 0 0 1 1 1 1 1 - 1 1 1 - 0 1 0 0 1 0 0 1 - 0 2 0 1 2 0 1 2 0

O. bicolor 1 1 0 3 1 0 0 1 1 1 1 1 1 0 2 1 0 0 0 0 1 1 0 0 0 1 1 - 1 0 0 0 0 0 1 0 0

O. carvalhoi 1 1 1 0 1 1 1 1 1 0 0 0 0 1 - 1 1 1 - 0 0 1 0 1 0 1 0 0 0 2 0 0 2 0 1 0 0

O. chaetopygialis 0 0 0 3 1 1 1 1 1 1 2 1 1 1 - 1 1 0 1 0 0 2 0 1 0 1 1 - 1 0 1 - - 0 1 1 1

O. cingarus 1 1 0 3 1 0 0 0 1 1 2 0 1 1 - 1 1 1 - 0 0 1 0 0 0 1 1 - 0 2 1 - - 0 0 0 1

O. conclausa 0 0 0 0 1 1 0 1 0 1 1 0 1 0 1 1 1 1 - 0 1 0 1 0 0 0 1 - 2 2 0 1 1 0 0 1 1

O. confusa 1 0 0 0 1 0 0 0 1 1 1 0 1 1 - 1 1 1 - 0 0 1 0 1 0 1 1 - 1 1 0 0 0 0 1 1 0

O. culmiforceps 0 0 1 1 1 0 1 0 0 0 0 1 1 1 - 1 0 1 - 0 0 1 1 0 0 1 1 - 0 1 0 0 1 0 0 0 0

O. culminata 1 0 0 0 0 0 0 0 0 1 0 1 1 1 - 1 1 1 - 0 0 1 1 1 0 1 1 - 1 1 0 0 2 0 0 1 1

O. diana 1 1 0 0 1 1 1 0 1 1 1 1 1 1 - 1 1 1 - 0 0 1 1 0 0 0 1 - 1 0 0 0 2 0 1 0 1

O. fluminensis 1 1 1 0 1 1 1 0 0 0 1 1 1 1 - 1 1 1 - 0 1 1 1 0 0 0 0 1 0 2 0 1 1 0 1 1 0

O. fringidea 0 0 0 0 1 1 1 0 1 1 0 0 1 1 - 1 1 1 - 0 1 1 1 1 1 0 1 - 1 1 0 1 1 0 0 1 0

O. galeata 1 0 0 3 1 0 0 1 1 1 0 0 1 0 2 1 0 1 - 0 1 1 1 0 0 1 1 - 0 2 0 0 1 0 1 1 1

O. grandis 1 0 0 0 1 1 1 0 1 0 1 1 1 1 - 1 1 0 0 0 0 0 0 1 0 0 0 1 0 2 0 1 2 0 1 1 0

O. injuncta 1 1 1 2 1 1 1 0 0 0 1 0 1 1 - 1 0 0 0 0 1 0 0 1 0 0 0 1 0 2 0 2 0 0 1 0 0

O. insolita 1 2 0 0 1 0 1 1 0 1 1 0 1 1 - 1 1 0 0 0 1 1 0 1 0 0 1 - 0 2 0 1 2 0 1 0 0

O. intona 0 0 1 2 0 1 1 1 1 1 1 1 0 0 2 0 1 1 - 0 1 0 1 0 0 1 1 - 0 1 0 0 0 0 0 0 1

O. major 1 0 0 0 1 0 1 1 0 1 1 0 1 1 - 1 1 0 0 0 1 0 1 0 0 0 1 - 0 2 0 1 2 0 0 0 1

O. mitifica 1 0 0 3 1 0 0 1 1 0 1 1 1 1 - 1 1 1 - 0 0 1 1 0 0 0 1 - 0 1 0 1 0 0 0 1 1

O. modesta 1 1 1 0 1 1 0 0 0 1 1 1 0 1 - 1 1 1 - 0 0 0 0 1 0 0 1 - 0 1 0 0 0 0 1 2 0

O. nitida 0 1 1 1 1 1 0 1 1 1 0 0 1 1 - 0 0 1 - 0 0 2 1 1 0 0 1 - 2 2 1 - - 1 - - 1

O. notata 1 1 1 0 1 1 0 1 0 1 2 1 0 1 - 0 0 1 - 0 1 1 0 1 0 0 1 - 0 2 0 0 2 0 1 2 1

O. occulta 1 0 1 0 1 1 1 1 1 1 1 1 1 1 - 1 1 0 0 0 0 0 1 0 0 0 1 - 0 1 1 - - 0 0 2 1

O. parva 1 1 0 3 1 0 0 0 1 1 1 1 1 1 - 1 0 1 - 0 0 1 1 1 0 1 1 - 1 0 1 - - 0 0 0 1

O. paulistanensis 1 1 1 1 1 1 1 0 0 1 1 1 1 0 0 1 1 1 - 0 1 1 0 1 0 0 0 0 0 2 0 2 2 0 1 1 0

O. peltata 1 1 1 0 0 1 1 1 1 1 0 0 1 0 2 0 1 1 - 0 1 0 1 1 0 1 1 - 0 1 0 0 0 0 0 1 1

O. perneta 1 1 0 3 1 0 1 0 1 1 1 0 1 0 0 1 1 1 - 0 0 1 0 1 0 0 1 - 1 0 0 0 0 0 1 0 0

O. plebeja 1 1 0 0 1 1 0 0 0 1 1 0 1 1 - 1 1 1 - 0 0 1 0 1 0 1 1 - 0 2 0 0 2 0 1 1 1

O. riograndensis 1 1 1 0 1 1 0 0 0 1 2 1 1 0 1 1 1 1 - 0 0 1 0 1 0 0 0 0 1 2 0 1 2 0 1 0 0

O. sarcinata 1 2 1 0 1 0 1 0 1 1 1 1 1 1 - 1 1 1 - 0 0 0 0 1 1 0 1 - 0 2 0 1 2 0 0 1 1

O. similata 1 1 1 2 1 1 0 1 1 1 1 0 1 1 - 1 1 1 - 0 0 1 0 0 0 0 0 0 0 2 0 1 2 0 1 1 0

O. terminalis 0 1 1 0 1 1 1 1 1 1 2 1 1 1 - 1 1 1 - 0 0 2 0 1 0 0 0 0 0 1 0 0 1 1 - - -

O. thornax 1 0 1 1 1 1 1 1 0 1 1 0 1 0 1 1 1 1 - 0 1 0 1 0 0 0 1 - 2 2 0 0 1 0 0 1 0

O. timida 1 0 1 1 0 1 1 1 1 1 0 0 1 0 0 1 1 1 - 0 1 0 1 1 1 1 1 - 0 2 0 1 1 0 0 1 0

O. trivialis 1 0 0 0 1 1 1 0 1 1 1 0 1 1 - 1 1 1 - 0 1 0 1 1 0 0 1 - 0 1 0 0 2 0 1 2 1

O. varia 0 1 1 1 1 0 1 1 1 1 0 1 1 0 2 1 1 1 - 0 0 0 0 1 1 0 1 - 1 2 0 1 1 0 1 1 0

O. ventricosa 1 2 1 1 1 0 1 0 0 1 0 0 1 1 - 1 1 0 0 0 0 1 1 1 0 1 1 - 0 2 0 0 0 0 1 1 0

O. vittata 1 1 1 2 1 1 ? 1 1 1 2 1 0 0 1 0 0 1 - 0 0 2 1 0 0 0 1 - 2 2 1 - - 1 - - 1

O. xanthosoma 1 1 1 1 1 1 1 0 1 1 1 1 1 1 - 1 1 1 - 0 1 0 1 1 0 0 0 0 0 2 0 1 2 0 1 1 0

O. zayasi 1 2 0 3 1 0 0 0 1 1 2 0 1 1 - 1 1 1 - 0 0 1 1 0 0 1 1 - 0 1 0 0 2 0 1 2 1

Oxysarcodexia n. sp. A 1 1 1 1 1 1 0 1 1 1 1 1 1 1 - 1 1 0 0 0 1 1 0 1 0 0 0 0 0 2 0 1 2 0 1 1 1

Oxysarcodexia n. sp. B 1 1 1 1 1 1 0 0 0 1 1 1 1 1 - 1 1 0 0 0 0 1 1 1 0 0 1 - 0 2 0 0 2 0 1 1 1

Oxysarcodexia n. sp. C 1 1 1 2 1 1 0 0 1 1 1 1 0 1 - 1 1 0 0 0 0 0 0 1 0 0 1 - 0 2 0 0 2 0 1 1 -

Oxysarcodexia n. sp. F 1 1 1 1 1 1 0 1 0 1 0 0 1 1 - 1 1 0 0 0 1 1 0 1 0 1 1 - 0 2 0 0 2 0 1 1 1

217

7. CONSIDERAÇÕES FINAIS

O conspecto apresentado para Oxysarcodexia, visou contribuir para uma melhor

compreensão da variedade morfológica dos machos deste gênero, assim como fornecer uma

atualização acerca da nomenclatura válida e sinonimização, distribuição geográfica e aspectos

biológicos das espécies pertencentes a esse gênero. Espera-se que o banco de dados pictórico

elaborado para os machos de Oxysarcodexia, associado às diagnoses, possa auxiliar no

processo de identificação de espécies, em especial daquelas morfologicamente próximas e

simpátricas, sendo um facilitador nos mais diversos estudos, os quais dependem da acurácia na

identificação, como, por exemplo, a entomologia forense.

A abordagem filogenética proposta representa um passo inicial para a compreensão das

relações dentro desse gênero, apontando paralelos nas variações morfológicas encontradas

entre as diferentes espécies, sugerindo agrupamentos interespecíficos, apesar do alto grau de

homoplasia presente nas espécies. Além disso, a assimetria na genitália dos machos revelou-se

uma característica independente que surgiu várias vezes durante a diversificação das espécies.

Vale ressaltar que o estudo morfológico das fêmeas de Oxysarcodexia à luz da

filogenia, assim como das larvas, poderá contribuir significativamente para um melhor

entendimento da relação entre as espécies. Da mesma forma, a análise morfológica de fêmeas

e a observação do comportamento de cópula de casais de espécies cujos machos apresentam a

assimetria na genitália possibilitará esclarecer como a assimetria evolui dentro desse gênero.

219

8. REFERÊNCIAS BIBLIOGRÁFICAS

Amat, E. 2010. Notes on necrophagous flies (Diptera: Calyptratae) associated to fish carrion in

Colombian Amazon. Acta Amazonica, 40 (2): 397–400.

Amorim, D.S. 2002. Fundamentos de Sistemática Filogenética. Holos Editora, Ribeirão Preto, São

Paulo, Brasil. 154pp.

Amorim, D.S. 2009. Neotropical Diptera diversity: richness, patterns, and perspectives. In: Pape, T.,

Bickel, D. & Meier, R. 2009. Diptera Diversity: status, challenges and tools. Brill, Portland, 71–

97.

Amorim, D.S. 2012. Biogeografia da Região Neotropical. In: Rafael, J.A., Melo, G.A.R., Carvalho,

C.J.B., Casari, S. & Constantino, C. (eds.), Insetos do Brasil, diversidade e taxonomia. Holos

Editora, Ribeirão Preto, São Paulo, Brasil, pp. 112–132.

Battán-Horenstein, M., Linhares, A.X., Ferradas, B.R. & Garcia, D. 2010. Decomposition and dipteran

succession in pig carrion in central Argentina: ecological aspects and their importance in forensic

science. Medical and Veterinary Entomology, 24: 16–25.

Battán-Horenstein, M., Rosso, B. & García, M.D. 2012. Seasonal structure and dynamics of

sarcosaprophagous fauna on pig carrion in a rural area of Cordoba (Argentina): their importance in

forensic science. Forensic Science International, 217: 146–156.

Beuter, L., Fernandes, P.A., Barros, P.B., Souza, C.R. & Mendes, J. 2012. Insetos de potencial

importância forense e na saúde pública em região urbana de Minas Gerais: frequência relativa e

variação sazonal de fauna atraída e criada em carcaças de roedores. Revista de Patologia Tropical,

41 (4): 480–490.

Brown, B. V. 2009. Introduction. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M.,

Woodley, N.E. & Zumbado, M.A. (eds.), Manual of American Central Diptera. Vol. 1. NCR

Research Press, Ottawa, Ontario, Canada, pp. 1–7.

Campobasso, C.P.; Di Vella, G. & Introna, F. 2001. Factors affecting decomposition and Diptera

colonization. Forensic Science International, 120: 18–27.

Carrera, M. 1944. Relação de alguns dípteros capturados em Monte Alegre, Estado de São Paulo.

Papéis avulsos do Departamento de Zoologia, 6 (5): 37–50.

Carvalho, C.J.B., Rafael, J.A., Couri, M.S. & Silva, V.C. 2012. Diptera. In: Rafael, J.A., Melo, G.A.R.,

Carvalho, C.J.B., Casari, S. & Constantino, C. (eds.), Insetos do Brasil, diversidade e taxonomia.

Holos Editora, Ribeirão Preto, São Paulo, Brasil, pp. 701–743.

Carvalho, L.M.L. & Linhares, A.X. 2001. Seasonality of insect succession and pig carcass

decomposition in a natural forest area in southeastern Brazil. Journal of Forensic Sciences, 46 (3):

604–608.

220

Carvalho, L.M.L., Thyssen, P.J., Goff, M.L. & Linhares, A.X. 2004. Observations on the succession

patterns of necrophagous insects onto a pig carcass in an urban area of Southeastern Brazil.

Aggrawal’s Internet Journal of Forensic Medicine and Toxicology, 5 (1): 33–39.

Carvalho, L.M.L., Thyssen, P.J., Linhares, A.X. & Palhares, F.B. 2000. A checklist of arthropods

associated with carrion and human corpses in southeastern Brazil. Memórias do Instituto Oswaldo

Cruz, 95 (1): 135–138.

Çiftçioğlu, N.; Altintaș, K. & Haberal M. 1997. A case of human orotracheal myiasis caused by

Wohlfahrtia magnifica. Parasitology Research, 83: 34–36.

Constamagna, R., Visciarelli, E.C, Lucchi, L.D., Basabe, N.E., Esteban, M.P. & Oliva, A. 2007.

Aportes al conocimiento de los dípteros ciclorrafos em el área urbana de Bahía Blanca (província

de Buenos Aires), Argentina. Revista del Museo Argentino de Ciencias Naturales, 9 (1): 1–4.

Cornaby, B.W. 1974. Carrion reduction by animals in contrasting tropical habitats. Biotropica, 6 (1),

51–63.

Couri, M.S., Lamas, C.J.E., Aires, C.C.C., Mello-Patiu, C.A., Pamplona, D.M. & Magno, P.D. 2000.

Diptera from Serra do Navio (Amapá, Brasil): Asilidae, Bombyliidae, Calliphoridae, Micropezidae,

Muscidae, Sarcophagidae, Stratiomyiidae, Syrphidae, Tabanidae and Tachinidae. Revista

Brasileira de Zoociências, 2 (1): 91–100.

Cruz, T. M. 2008. Diversidade e sucessão ecológica de insetos associados à decomposição animal em

um fragmento de Mata Atlântica de Pernambuco. Dissertação de Mestrado. 102 fls. Universidade

Federal de Pernambuco, Recife, Pernambuco, Brasil.

Dias, E.S., Neves, D.P. & Lopes, H.S. 1984. Estudos sobre a fauna de Sarcophagidae (Diptera) de Belo

Horizonte – Minas Gerais I- Levantamento Taxonômico e Sinantrópico. Memórias do Instituto

Oswaldo Cruz, 79 (1): 83–91.

Dutto, M. & Bertero, M. 2010. Traumatic myiasis from Sarcophaga (Bercaea) cruentata Meigen, 1826

(Diptera, Sarcophagidae) in a hospital environment: reporting of a clinical case following

polytrauma. Journal of Preventive Medicine and Hygiene, 51 (1): 50–52.

Evenhuis, N.L., Pape, T., Pont, A.C. & Thompson, F.C. (eds.) 2007. Biosystematic Database of World

Diptera, Version 10. Available at http://www.diptera.org/biosys.htm. apud Pape, T., Bickel, D. &

Meier, R. (2009) Diptera Diversity: status, challenges and tools. Brill, Portland, 459pp.

Evenhuis, N.L., Pape, T., Pont, A.C. & Thompson, F.C. (eds.) 2007. Biosystematic Database of World

Diptera, Version 10. Available at http://www.diptera.org/biosys.htm. apud Amorim, D.S. 2009.

Neotropical Diptera diversity: richness, patterns, and perspectives. In: Pape, T., Bickel, D. &

Meier, R. 2009. Diptera Diversity: status, challenges and tools. Brill, Portland, 71–97.

Ferreira, M.J.M. 1978. Sinantropia de dípteros muscóideos de Curitiba, Paraná. I. Calliphoridae.

Revista Brasileira de Biologia, 38: 445–454.

221

Figueiredo, R.R.; Dorf, S.; Couri, M.S.; Azevedo, A.A. & Mossumez, F. 2002. Corpos estranhos

animados em otorrinolaringologia. Revista Brasileira de Otorrinolaringologia, 68 (5): 722–728.

Giroux, M., Pape, T. & Wheeler, T.A. 2010. Towards a phylogeny of the flesh flies (Diptera:


Sarcophaginae. Zoological Journal of the Linnean Society, 158: 740–778.

Greenberg, B. 1971. Flies and diseases. Ecology, classification and biotic association. Vol. 1.

Princeton: Princeton University, 856p.

Greenberg, B. 1973, Flies and diseases. Biology and disease transmission. Vol. 2. Princeton, Princeton

University, 447p.

Guimarães, J.H. & Papavero, N. 1999. Myiasis in man and animals in the Neotropical region:

bibiographic database. São Paulo: Editora Plêaide/FAPESP, 1 ed., 308pp.

Leão, R.N.Q; Fraiha, H.; Cruz, J.P.N. & Tibana, R. 1996. Miíase uretral por Sarcodexia lambens

(Wiedemann, 1830) (Diptera: Sarcophagidae): relato de um caso amazônico. Revista Paraense de

Medicina, 10 (1): 27–29.

Linhares, A.X. & Thyssen, P.J. 2007. Miíases de importância médica: moscas e entomologia forense.

In: De Carli, G.A. (org.) Parasitologia Clínica: seleção de métodos e técnicas de laboratório para o

diagnóstico das parasitoses humanas. 2ª ed., São Paulo: Atheneu, 709–730.

Liu, D. & Greenberg, B. 1989. Immature stages of some flies of forensic importance. Annals of the

Entomological Society of America, 82 (1): 80–93.

Lopes, H.S. 1969. A Catalogue of the Diptera of the Americas South of the United States: Family

Sarcophagidae. Departamento de Zoologia, Secretaria da Agricultura, 103, 88p.

Lopes, H.S. 1982. The importance of the mandible and clypeal arch of the first instar larvae in the

classification of the Sarcophagidae (Diptera). Revista Brasileira de Entomologia, 26 (3/4): 293–

326.

Mégnin, J. La faune des cadavres: application de l'entomologie a la medecine legale. Encyclopedie

Scientifique des Aides Memoires. Masson et Gauthiers-Villars, Paris, 1894. 214p.

Monteiro-Filho, E.L.A. & Pernereiro, J.L. 1987. Estudo de decomposição e sucessão sobre uma

carcaça animal numa área do estado de São Paulo, Brasil. Revista Brasileira de Biologia, 47 (3):

289–295.

Nuorteva, P. 1977. Sarcosaprophagous insects as forensic indicators. In: Forensic medicine: a study in

trauma and environmental hazards. vol. II. W.B. Saunders Company, 1072–1095.

Pamplona, D., Maia, V.C., Couri, M.S., Lamas, C.J.E. & Aires, C.C.C. 2000. A survey of díptera on

Paquetá Island, Rio de Janeiro, Brazil. Entomologist`s Monthly Magazine, 136: 169–175.

222

Pape, T. & Dahlem, G.A. 2010. Sarcophagidae. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood,

D.M., Woodley, N.E. & Zumbado, M. (eds), A Manual of Central American Diptera. Vol. 2. NRC

Research Press, Ottawa, pp. 1313–1335.

Pape, T. 1996. Catalogue of the Sarcophagidae of the world (Insecta: Diptera). Memoirs on

Entomology, International, 8: 1–558.

Pérez, S.P., Duque, P., Wolff, M. 2005. Successional and behavior and occurrence matrix of carrion-

associated arthropods in the urban área of Medellín, Colombia. Journal of Forensic Science, 50 (2):

448–453.

Rafael, J.A.; Aguiar, A.P. & Amorim, D.S. 2009. Knowledge of insect diversity in Brazil: challenges

and advances. Neotropical Entomology, 38 (5) : 565–570.

Ramírez-Mora, M.A., Buenaventura, E., Gómez-P, L.M. & Amat, E. (2012) Uptadet cheklist and new

records of Calyptrate carrion flies (Diptera, Schizophora) from Valle de Aburrá and other localities

in Colombia. Entomotropica, 27 (1), 27–35.

Roback, S.S. 1954. The evolution and taxonomy of the Sarcophaginae (Diptera, Sarcophagidae).

Illinois Biological Monographs, 23: 1–181.

Shewell, G.E. 1987. Sarcophagidae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J.,

Vockeroth, J.R. & Wood, D.M. (eds.), Manual of Nearctic Diptera. Vol. 2. Monograph 27,

Agriculture Canada, pp. 1159–1186.

Smith, K.G.V. 1986. A manual of Forensic Entomology. Cornell University Press, Ithaca, NY, 205 pp.

Sousa, J.R.P., Esposito, M.C. & Carvalho-Filho, F.S. 2011. Composition, abundance and richness of

Sarcophagidae (Diptera: Oestroidea) in forests and forest gaps with different vegetation cover.

Neotropical Entomology, 40 (1): 20–27.

Souza, A.M. & Linhares, A.X. 1997. Diptera and Coleoptera of potential forensic importance in

Southeastern Brazil: relative abundance and sazonality. Medical and Veterinary Entomology, 11:

8–12.

Thyssen, P.J.; Moretti, T.C.; Ueta, M.T. & Ribeiro, O.B. 2004. O papel de insetos (Blattodea, Diptera e

Hymenoptera) como possíveis vetores mecânicos de helmintos em ambiente domiciliar e

peridomiciliar. Cadernos de Saúde Pública, 20: 1096–1102.

Turhan, V.; Ulaş, U.H.; Aydin, A.F.; Eyigün, C.P.; Araz, E.; Tanyüksel, M. & Pahsa, A. 2007. An

intestinal myiasis case of flesh fly (Sarcophaga haemorrhoidalis). The Anatolian Journal of

Clinical Investigation, 1 (4): 270–272.

Tuygun, N.; Taylan-Özkan, A.; Tanir, G. & Mumcuoğlu, K.Y. 2009. Furuncular myiasis in a child

caused by Wohlfahrtia magnifica (Diptera: Sarcophagidae) associated with eosinophilia. The

Turkish Journal of Pediatrics, 51: 279–281.

223

Verves, Y. 1989. Prof. Hugo de Souza Lopes and the modern system of Sarcophagidae (Diptera).

Memórias do Instituto Oswaldo Cruz, 84: 529–545.

Yazgi, H.; Uyanik, M.H.; Yoruk, O. & Aslan, I. 2009. Aural myiasis by Wohlfahrtia magnifica: case

report. The Eurasian Journal of Medicine, 41: 194–196.

Zhang, Z.-Q. 2013. Phylum Arthropoda. In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An outline of

higher-level classification and survey of taxonomic richness (Addenda 2013). Zootaxa, 3703, 1–82.

Zumpt, F. Myiasis in man and animals in the Old World. London: Butterworths, 1965. 267pp.

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