Muscidae (Diptera) of the Neotropical Region: Taxonomy

Claudio Jose Barros de Carvalho is Full Professor at the Department of Zoology of the Universidade Federal do Parana; ...

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Claudio Jose Barros de Carvalho is Full Professor at the Department of

Zoology of the Universidade Federal do Parana; fellow of the CNPq, with many papers published in Systematics, Bionomics and Biogeography of Diptera. A former student in the Museu ~acional,

Rio de Janeiro, he did his

Ph.D. studies in Curitiba, and in the ~arIy

1990's, he spent a year at the

~atural History Museum in London.

To

date. 20 graduate students have obtained their degrees under him. ~Iarda Souto Couri is Professor at the

.\Iuseu Nacional, Universidade Federal do Rio de Janeiro; C~Pq.

fellow of

the

with expressive scientific

contributions on Systematics, Bionomics and Phylogeny of Diptera. \Yith a Ph.D. degree in Parasitology trom the Universidade Federal Rural do Rio de Janeiro and a postdoctoral project developed at the Natural History Museum, London, she has under her leadership a team of researchers and students. She has also supelTised several M.Sc. and Ph.D. candidates in Zoology.

Muscidae (Diptera) of the Neotropical Region:

TAXONOMY

.. UFPR

Reitor Carlos Roberto Antunes dos Santos Vice-Reitor Romolo Sandrini Diretor da Editora da UFPR Luiz Carlos Ribeiro

Conselho Editorial Alberto Pio Fiori Andre de Macedo Duarte Fany Reicher Izaura Hiroko Kuwabara Jose Carlos Cifuentes Vasquez Leilah Santiago Bufrem Manoel Eduardo A. Camargo e Gomes Maria Benigna Martinelli de Oliveira Pedro Ronzelli Junior Sergio Herrero de Moraes Victor Manoel Pelaez Alvarez

Muscidae (Diptera) of the Neotropical Region:

TAXONOMY

Edited by Claudio Jose Barros de Carvalho

~ UFPR

© Claudio Jose Barros de Carvalho

Muscidae (Diptera) of the Neotropical Region: Taxonomy Coordena~ao

editorial: Marildes Rocio Artigas Santos Revisao de texto: Gabriel Augusto Rodrigues de Melo Revisao final: do Autar Editora~ao eletronica: Alquimia Esrudio de Arte Final Fotos: Philippe Wauters Capa: Se9ao de Programa9ao Visual da Universidade Federal do Rio de Janeiro Serie Pesquisa n. 68 Coardena9ao de Processos Tecnicos. Sistema de Bibliotecas, UFPR Muscidae (Diptera) of the Neotropical Region: taxonomy / edited by Claudio Jose Barros de Carvalho. - Curitiba : Ed. UFPR, 2002. 287 p. : il. - (Pesquisa ; 68) ISBN 85-7335-091-1 1. Muscidae. 2. Diptera. 3. Entomologia. I. Carvalho, Claudio Jose Barros de. II. Titulo. III. Serie.

CDD 595.770913 CDU 595.77(213)

ISBN 85-7335-091-1 Ref. 315 Apoio: Funda9ao Araucana Editora UFPR Centro Politecnico - Jardim das Americas Caixa Postal 19 .029 81531-980 - Curitiba - Parana - Brasil tel.Jfax: (41) 361-3380/361-3381/267-5973 e-mail: [email protected]

2002

FOREWORD

iosystematic revisions are vitally essential in biological research as guide and reference work to the continually expanding knowledge of the fauna and flora. They bring together, assimilate and organize scattered information from around the world so that students and researches can quickly ascertain the current status of the taxonomic hierarchy and find reference to many kinds of information. The dipterous of the family Muscidae constitute an important group of human and animal pest, widely distributed in the world fauna. Larvae breed mainly in decaying plant and animal material or manure. Adults of many species bite or passively vector pathogens for disease such as typhoid fever, dysentery, anthrax, etc. Although numerous, ubiquitous, and economically important, muscids have not had the attention they deserve. More than 4000 described species are recorded worldwide, but only 84 genera and about 840 species of them are known from the Neotropical Region and the actual number that must occur in this region is inestimable. No comprehensive monograph has ever published to this area of the world and the present contribution of Claudio Jose Barros de Carvalho and Marcia Souto Couri is well warranted. Subfamilies, tribes and genera are revised, and keys for identification of all genera and species with Neotropical occurrence are proposed. The authors have undertaken a formidable task, and the present work will constitute a significant step in the systematics of N eotropical muscids providing a basic reference for further research.

B

Jose Henrique Guimaraes Associate Professor, rCB Universidade de Sao Paulo February 2001

PREFACE

ow great are God's riches! How deep are his wisdom and knowledge! How impossible to explain his decisions or to understand his methods! As Scripture says, "Who could ever known the mind of the Lord?" Who could ever be his counsellor? ... For all things were created by him and all things exist through him and for him. To him be the glory forever! Amen. [Romans, Ch. 11:33-36] apud McALPINE (1989). The scientific literature shows that there is no consensus about the definition of some words commonly used by a taxonomist or a systematist. Systematics and some related words as taxonomy, classification and biological system have received different definitions by different authors (MINELLI, 1993). Among the definitions still accepted, there are the ones of SIMPSON (1961) for Systematics, Taxonomy and zoological classification. According to this author, Systematics is the scientific study of the groups, of the diversity of the organisms and of all the relations among them. On the other side, Taxonomy is the theoretic study of the classification, including its bases, principles, procedures and rules. Zoological classification is the arrangement of the animals into groups having their similarities as a basis. In his point of view, Taxonomy is part of the Systematic and so, subordinated to it. However, about 30 years ago, NELSON (1970) introduced a new concept of Comparative Biology (WILEY, 1981;AX, 1987; BROOKS & McLENNAN, 1991), where characters of the diverse taxa are analysed comparatively, clearly seeking for similarities and differences among them. This is the study of the diversity patterns, looking for the comprehension of the origin of the patterns of similarities and differences among the groups (AMORIM, 1997). So, Comparative Biology is understood as the study of the biological diversity under a historical perspective (NELSON & PLATNICK, 1981). Its main scope is the reconstruction of the similarity and dissimilarity patterns that exist inside

H

the organic diversity, both concerning the diversity of forms or attributes as well as their geographical distribution. Systematics and Taxonomy are areas that integrate the Comparative Biology. According to some authors (MINELLI, 1993), Taxonomy can be understood as a synonym of Systematics and has the role to produce taxonomic information, including classification, nomenclature, description, keys to identification and to show the biological inter-relations among the analysed taxa. In other words, Systematics has, as one of its main roles, to present a general reference system on the biological diversity (HENNIG, 1966). The scarce know ledge of the dipterofauna of the neotropics has other reasons than the different academic interpretations of Systematics and Taxonomy described above. About 30,000 species ofDiptera in the neotropics are known nowadays (Nelson Papavero, personal communication), what represents a very timid number in relation to the real number of species that live in this region. In Brazil, two main points bring difficulties to the knowledge of the dipterofauna: the abundant richness associated with the reduced number of qualified persons to study it. As it was pointed by THOMPSON (1990), new technologies, research training and also new work conditions are necessary to the comprehensible know ledge of the Diptera of any region. The union of efforts of several levels is necessary to change this reality in a relatively short period of time. But for sure, one of the first steps is the publication of basic information for the identification of the species as well as information on the phylogenetic relations among them and also on the patterns of their distribution (CARVALHO, 1998). Using this know ledge for a more consolidated for a more consolidate study of the biodiversity, as well as, through the development of new theories in Systematics, and a better comprehension of the phylogeny, this scenery should change quickly. Unfortunately, in the threshold of the Third Millennium, the knowledge of the Diptera of the Neotropical Region is still incipient (GRAZIA et ai., 2000). The present publication is a great step to the comprehension of the muscid biodiversity. Before undertaking any study of a basic or practical nature, it is always necessary to know with whom we are dealing, or, in other words, to know who is whothat is the beginning of everything.

References AMORIM, D. de S. 1997. Elementos basicos de Sistematica Filogenetica; 2' edi'tao, revista e ampliada. Holos & Sociedade Brasileira de Entomologia, Ribeirao Preto, 276 p. AX, P. 1987. The Phylogenetic System. The systematization of organisms on the basis of their phylogenesis. John Wiley & Sons, Chichester, 340 p. BROOKS, D.R. & McLENNAN, D.A. 1991. Phylogeny, Ecology, and Behavior. The University of Chicago Press, Chicago, 434 p. CARVALHO, CJ.B. de. 1998. Taxonomista de insetos, uma especie em extin'tao no Brasil. Inf. Soc. Entomol. Bras. 23: 1. GRAZIA, J.; CJ.B. de CARVALHO; L.M. de ALMEIDA; M.M. CASAGRANDE & O.H.H. MIELKE. 2000. A biodiversidade de insetos no Brasil no Terceiro Milenio: dificuldades e perspectivas. Entomol. Vect. 7: 123-141. HENNIG, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, IlL, 263 p. McALPINE, J.E 1989. Phylogeny and classification of the Muscomorpha, p. 1397-1518. In: McAlpine, J.E & D.M. Wood (eds.), Manual ofNearctic Diptera, volume 3. Research Branch, Monograph No. 32, p. 1333-1581. MINELLI, A. 1993. Biological Systematics; the state of the art. Chapman & Hall, London, 387 p. NELSON, GJ. 1970. Outline ofa theory of comparative biology. Syst. Zool. 20: 471-472. NELSON, G.E & PLATNICK, N.L 1981. Systematics and Biogeography, Cladistics and Vicariance. Columbia University Press, New York, 567 p. SIMPSON, G.G. 1961. Principles of Animals Taxonomy. Columbia University Press, New York, 254 p. THOMPSON, EC. 1990. Biosystematic information dipterists ride the third wave, p. 179-201. In: M. Kosztarab & C.W. Schaefer. Systematics on the North American Insects and Arachnids: status and needs. Virginia Agricultural Experiment Station Information series 90-1. Blacksburg: Virginia Polytechnic Institute and State University. WILEY, E.O. 1981. Phylogenetics; the theory and practice of phylogenetic systematics. New York, Wiley Sons, 439 p.

Claudio Jose Barros de Carvalho April 2002

ACKNOWLEDGEMENTS This book is a result of about 25 years dedicated to the study of the muscids flies. During this period an intense scientific exchange and a great friendship grew among us since the beginning of our scientific life in the Museu Nacional, Rio de Janeiro. Along this whole path, many common colleagues and friends have given us valuable help and to all these people we would like to offer our special thanks - Adrian Charles Pont (Oxford University Museum), Oxford, Peter Skidmore, Swansea, and our many colleagues at The Natural History Museum (London), The National Museum ofNatural History (Washington), The Canadian National Collection (Ottawa), Universidade Federal do Parana (Curitiba) and the Museu N acional, Universidade Federal do Rio de Janeiro (Rio de Janeiro), for study facilities and much valuable discussion. We also are grateful to the Conselho Nacional de Desenvolvimento Cientifico e Tecnol6gico, an agency of the Brazilian Government, for scientific and technological development, for the support provided by a grant to CJBC (Proc. Nr. 300043/86-4) and to MSC (Proc. Nr. 300386/80-0). More specifically to the elaboration of this book, we would like to thank Luciane Marinoni and Sonia Maria Noemberg Lazzari (Universidade Federal do Parana, Curitiba), Jose Albertino Rafael (Instituto Nacional de Pesquisas daAmazonia, Manaus), all our present graduates and undergraduates students for their support and critical reviews of earlier versions of the manuscript. To Denise Pamplona (Museu N acional, Rio de Janeiro) and one anonymous referee for their careful and critical review of the final version of the manuscript. To Jose Henrique Guimaraes for the foreword. To Gabriel Augusto Rodrigues de Melo (Universidade Federal do Parana, Curitiba) for revising the English. We owe a special debt to Philippe Wauters, who, with his sensibility, enthusiasm and photography skills, knew how to capture the spirit of the flies. We also thank the Section ofPrograma9ao Visual of Un iversidade Federal do Rio de Janeiro for the cover design. Claudio Jose Barros de Carvalho Marcia Souto Couri

CON'!'ENTS PART I. BASAL GROUPS By Claudio Jose Barros de Carvalho & Marcia Souto Couri INTRODUCTION .................................................................................................... 17 KEY TO NEOTROPICAL GENERA OF MUSCIDAE ............................................. 21 DIAGNOSIS AND KEY TO SPECIES ..................................................................... 37 SUBFAMILY ATHERIGONINAE FAN, 1965 ........................................................... 38 Atherigona Rondani, 1856 ................................................................................. 38 SUBFAMILY MUSCINAE LATREILLE, 1802 ......................................................... 39 Tribe Muscini Latreille, 1802 ................................................................................ .40 Biopyrellia Townsend, 1932 ............................................................................... .40 Cyacyrtoneura Townsend, 1931 ........................................................................ .41 Dasymorellia Malloch, 1923 ............................................................................... .41 Morellia Robineau-Desvoidy, 1830 ..................................................................... .42 Musca Linnaeus, 1758 ....................................................................................... 48 Neomyia Walker, 1859 ....................................................................................... 49 Neorypellia Pont, 1972 ....................................................................................... 49 Parapyrellia Townsend, 1915 .............................................................................. 50 Polietina Schnabl & Dziedzicki, 1911.. .................................................................. 51 Sarcopromusca Townsend, 1927 .......................................................................... 55 Xenomorellia Malloch, 1923 .............................................................................. 57 Tribe Stomoxyini Meigen, 1824 .............................................................................. 58 Haematobia Le Peletier & Serville in Latreille et aI., 1828 .................................... 59 Neivamyia Pinto & Fonseca, 1930 ......................................................................... 59 Stomoxys Geoffroy, 1762 ..................................................................................... 60 SUBFAMILY AZELIINAE ROBINEAU-DESVOIDY, 1830 ...................................... 62 Tribe Azeliini Robineau-Desvoidy, 1830 ..................................................................... 62 Azelia Robineau-Desvoidy, 1830 ........................................................................... 62 Drymeia Meigen, 1826 ....................................................................................... 63 Hydrotaea Robineau-Desvoidy, 1830 ................................................................... 63 Micropotamia Carvalho, 1993 .............................................................................. 68 Ophyra Robineau-Desvoidy, 1830 ....................................................................... 70 Potamia Robineau-Desvoidy, 1830 ....................................................................... 74 Thricops Rondani, 1856 ....................................................................................... 74

Tribe Reinwardtiini Brauer & Bergenstamm, 1889 .................................................... 76 Brachygasterina Macquart, 1851 ........................................................ ,............... 76 Chaetagenia Malloch, 1928 ............................................................................... 78 Correntosia Malloch, 1934 ................................................................................. 79 Dalcyella Carvalho, 1989 ................................................................................. 79 Itatingamyia Albuquerque, 1979 ......................................................................... 80 Muscina Robineau-Desvoidy, 1830 ...................................................................... 80 Palpibracus Rondani, 1863 ................................................................................ 82 Philornis Meinert, 1890 ...................................................................................... 88 Psilochaeta Stein, 1911 ...................................................................................... 96 Reinwardtia Brauer & Bergenstamm, 1889 .......................................................... 97 Synthesiomyia Brauer & Bergenstamm, 1893 ....................................................... 98 SUBFAMILY PHAONIINAE MALLOCH, 1917 .......................................................... 98 Dolichophaonia Carvalho, 1993, stat. rev ........................................................... 99 Helina Robineau-Desvoidy, 1830 ......................................................................... 105 Phaonia Robineau-Desvoidy, 1830 ....................................................................... 119 Souzalopesmyia Albuquerque, 1951 .................................................................. 126

PART II. APICAL GROUPS By Marcia Souto Couri & Claudio Jose Barros de Carvalho INTRODUCTION ..................................................................................................... 133 DIAGNOSIS AND KEY TO SPECIES ..................................................................... 135 SUBFAMILY CYRTONEURININAE SNYDER, 1954 .............................................. 135 Arthurella Albuquerque, 1954 ............................................................................. 135 Cariocamyia Snyder, 1951 ................................................................................. 136 Charadrella Wulp, 1896 ..................................................................................... 137 Chortinus Aldrich, 1932 ..................................................................................... 139 Cyrtoneurina Giglio-Tos, 1893 ............................................................................. 139 Cyrtoneuropsis Malloch, 1925 ............................................................................. 141 Mulfordia Malloch, 1928 ..................................................................................... 149 Neomuscina Townsend, 1919 ................................................................................ 149 Neomuscittiopsis Albuquerque & Lopes, 1992....................................................... 155 Neurotrixa Shannon & Del Ponte, 1926 ................................................................. 155 Pseudoptilolepis Snyder, 1949 .............................................................................. 156 Xenothoracochaeta Malloch, 1921 ....................................................................... 158 SUBFAMILY MYDAEINAE VERRAL, 1888 ............................................................. 160 Brontaea Kowarz, 1873 ..................................................................................... 160

Graphomya Robineau-Desvoidy, 1830 ................................................................. 163 Hemichlora Wulp, 1893 ................................"..................................................... 167 Mydaea Robineau-Desvoidy, 1830 ......................................................................... 167 Myospila Rondani, 1856 ................................................................................... 170 Scenetes Malloch, 1936, stat.rev.................................................... ..................... 171 Scutellomusca Townsend, 1931 ............................................................................ 172 SUBFAMILY COENOSIINAE VERRAL, 1890 .......................................................... 173 Tribe Limnophorini Villeneuve, 1902 ........................................................................... 174 Agenamyia Albuquerque, 1953 ........................................................................... 174 Albertinella Couri & Carvalho, 1997 ..................................................................... 176 Drepanocnemis Stein, 1911 ................................................................................. 177 Limnophora Robineau-Desvoidy, 1830 .................................................................. 178 Lispe Latreille, 1797 ............................................................................................ 182 Lispoides Malloch, 1920 ........................................................................................ 184 Pachyceramyia Albuquerque, 1955 ........................................................................ 187 Rhabdotoptera Stein, 1919 ................................................................................... 189 SpUo gona Schnabl, 1911 ....................................................................................... 190 Syllimnophora Speiser, 1923 ................................................................................. 192 Tetralnerinx Berg, 1898 ........................................................................................ 200 Thaumasiochaeta Stein, 1911 ............................................................................... 20 1 Tribe Coenosiini Verral, 1890 ................................................................................... 205 Apsil Malloch, 1929 ............................................................................................ 205 Bithoracochaeta Stein, 1911 ................................................................................ 211 Coenosia Meigen, 1826 ........................................................................................ 214 Cordiluroides Albuquerque, 1954 .......................................................................... 218 lnsulamyia Couri, 1982 ..................................................................................... 219 Neodexiopsis Malloch, 1920 ................................................................................ 220 Notoschoenomyza Malloch, 1934 .......................................................................... 237 Oxytonocera Stein, 1911 ...................................................................................... 238 Pentacricia Stein, 1898 ...................................................................................... 239 Pilispina Albuquerque, 1954 ................................................................................ 240 Plumispina Albuquerque, 1954 .............................................................................. 241 Reynoldsia Malloch, 1934 ..................................................................................... 242 Schoenomyza Halliday, 1833 ................................................................................. 247 Schoenomyzina Malloch, 1934 .............................................................................. 251 Spathipheromyia Bigot, 1884 .............................................................................. .254 Stomopogon Malloch, 1930 ............................................................................... 257

REFERENCES ...................................................................................................... 263

PART I

BASAL GROUPS By Claudio Jose Barros de Carvalho* & Marcia Souto Couri**

INTRODUCTION The Muscidae occurs in all biogeographical regions and contains more than 4000 species (PONT, 1986a, 1989). In the Neotropical Region, about 800 species are recognised (CARVALHO et al., 1993). Muscidae species have extremely varied habitsthe larvae can be coprophagous, saprophagous or carnivorous (SKIDMORE, 1985) and the adults of many species live on decaying animal or vegetable matter. Muscidae is one of the most interesting families ofDiptera, due to the association of many species with the antropobiocenosis (GREENBERG, 1971). Some species have a great medical and veterinary importance, acting as vectors of many diseases and other species have a considerable ecological importance. The adults are very abundant in rural or semi-rural areas. At high altitudes, they form a very high proportion ofthe fauna, both in number of species as in individuals (PONT, 1986a). The larvae of some species are agriculturally important, being primarily pests of plantations. Some species attack stored products with agricultural and commercial importance. * **

Universidade Federal do Parana, Departamento de Zoologia, C.P. 19020, Curitiba, 81.531-980, Parana, Brazil; email: [email protected] Museu Nacional do Rio de Janeiro, Departamento de Entomologia, Quinta da Boa Vista, Rio de Janeiro, 20.940040, Brazil; e-mail: [email protected]

Muscidae (Diptera) of the Neotropical Region: TAXONOMY

17

Part I - Basal Groups

Muscidae can be recognised by diverse characters among the Muscomorpha (AMORIM & CARVALHO, 1992) as, for example, the absence ofa vertical series of setae on meron and the presence of developed calypteres. These characters identify the Muscoidea (sensu McALPINE, 1989), but the Muscidae shows some exclusive characters, a condition that indicates the monophyly of the family. Some authors (ROBACK, 1951; HENNIG, 1965, 1973; SKIDMORE, 1985; McALPINE, 1989; MICHELSEN, 1991; PONT & CARVALHO, in preparation) consider the family monophyly based on the presence ofthree derived characters: (1) female with the abdomen showing less than 7 pairs of spiracles (spiracles 6 and 7 were lost, but some genera not related amongst themselves, have reacquired the spiracle 6). This reduction in the number of spiracles is a unique apomorphy among the Calyptratae, (2) male reproductive system without the accessory pair of glands. Among the Muscoidea (McALPINE, 1989), this condition is present in the three other families - Fanniidae, Scathophagidae and Anthomyiidae, (3) larval mouth-hooks closely appressed to one another. Although the Muscidae are considered to be a family satisfactorily known worldwide (THOMPSON, 1990), in the Neotropical Region its taxonomy is still in incipient levels (CARVALHO, 1989a). Many classifications of this family were proposed by the authors. Until the beginning of the 1960's, typological classifications were presented (e.g. ROBACK, 1951; HENNIG, 1956, 1957, 1958, 1959, 1960a, 1960b, 1962a, 1962b, 1962c, 1962d, 1963a, 1963b, 1964), but HENNIG (1965) proposed the first classification for Muscidae based on cladistic methodology. Starting about that time, some catalogues (PONT, 1972, 1977, 1980, 1986a, 1989) were published following the classification proposed by HENNIG (1965), with few modifications. In the beginning of the nineties, CARVALHO et al. (1993) published a catalogue for the Neotropical Region following the classification proposed by CARVALHO (1989d). The natural classification of the family, however, still needs improvements (HUCKETT & VOCKEROTH, 1987). Many taxa are known only from the original descriptions or only from the adults forms. In this way, important characters for the understanding of the family were not properly evaluated, considering the family as a whole (CARVALHO, 1989a). A more consistent natural classification will be possible only when the Neotropical genera are fully revised. A fundamental contribution to the studies on the biology of the muscid of the 18


Claudio Jose Barros de Carvalho & Marcia Souto Co uri

world was presented by SKIDMORE (1985), where the immature stages of about 440 muscid species were treated. His classification of the family was proposed mainly based on characters of the immature stages. Four years latter, CARVALHO (1989d) proposed a classification for the muscids, on the basis of a cladistic analysis of characters of the adult morphology. Species representing 27 genera, 12 of them exclusively Neotropical were utilised. The resulting cladogram was used to propose a phylogenetic classification with seven subfamilies and eight tribes: Acanthipterinae, Atherigoninae, Muscinae (Stomoxyini and Muscini), Azeliinae (Azeliini and Reinwardtiini), Phaoniinae, Mydaeinae (Mydaeini and Graphomyini) and Coenosiinae (Coenosiini and Limnophorini). This classification is very similar to the one proposed by SKIDMORE (1985). From 1989 on, many revisions of Neotropical muscid genera were published (CARVALHO, 1989a, 1989b, 1989c, 1993a, 1999; COURI & CARVALHO, 1992, 1993, 1997b;COURI&PAMPLONA, 1992;CARVALHO&COURI, 1993;ARAlJJO & COURI, 1993; COURI, 1995, 1996a, b, 1998a, 1999a, 2000a, b, in press; COURI & LAMAS, 1993; PALKA-ROCHA & CARVALHO, 1994; CARVALHO & PONT, 1998; MOTTA & COURI, 1999; PAMPLONA, 1999; COELHO, 2000). Several papers with descriptions of new genera or partial generic revisions were also recently published (COURI & ALBUQUERQUE, 1979; COURI, 1982a, b, 1989; PAMPLONA, 1983, 1986 a, b, c, 1992; COURI & LOPES, 1985, 1986, 1988a; CARVALHO, 1985a, b, 198ge; PAMPLONA & COURI, 1989, 1995, 1998b; LOPES, 1992; CARVALHO & COELHO, 1993; COURI & CARVALHO, 1995; COELHO, 1998a, b). These complete or partial revisions represent 50% of genera recognised by CARVALHO et al. (1993) for the Neotropical Region. Some keys for the identification of the Neotropical muscid genera are available in the literature (COURI & LOPES, 1985, 1986, 1988b; LOPES & COURI (1989a), VOCKEROTH, 1996; COURI & PONT, 1999). The patterns of distribution of the South American muscid species are scarcely known. A small number of papers on the historical biogeography have been published for subgroups of Muscidae, a few of them dealing with Neotropical groups. HENNIG (1965) indicated that the first muscid invasion in South America occurred from the Northern Hemisphere, during the Upper Cretaceous or at the beginning of the Tertiary Period, under Muscidae (Diptera) of the Neotropical Region: TAXONOMY

19


a clearlydispersionist view. Recently CARVALHO (1999) presented an alternative hypothesis for the origin of the muscids in the Neotropical Region, based on the ideas proposed by AMORIM & PIRES (1996) and PONT & CARVALHO (1997). The transoceanic relations of some genera were also investigated by COURI & CARVALHO (2000). For the southern area of the South America, CARVALHO & COURI (in press) presented a hypothesis for the explanation of the distribution pattern of two small muscid clades. The scope of this chapter is to revise the taxonomy of the basal groups of Neotropical Muscidae (see CARVALHO, 1989d). Diagnoses for the subfamilies, tribes and genera, discussions of the supra-generic relationships, and keys for identification of all genera and species of basal groups are presented. The classification followed here was the one proposed by CARVALHO et al. (1993). The type-species of each genera are cited before the diagnosis. Known synapomorphic and autapomorphic characters were included in the diagnosis, and the monophyly of the subfamilies, tribes and genera are discussed when possible. The terminology used was taken mainly from McALPINE (1981) and COELHO (1997), the last paper in particular for the morphology of the proboscis. Some Latin abbreviations commonly used in taxonomic papers (see PAPAVERO, 1994) were also used. The key to genera was based mainly in the following papers: COURI & LOPES (1985,1986, 1988b), LOPES & COURI (1989a), VOCKEROTH (1996) and COURI & PONT (1999). VOCKEROTH (1996) presented a key for the identification of the genera of Muscidae occurring in Central America. In this key, he included Fannia RobineauDesvoidy and Euryomma Stein, genera of Fanniidae. This family is one of the most basal groups within the Muscoidea, and perhaps among the Calyptratae (VOSSBRINK & FRIEDMAN, 1989; MICHELSEN, 1991; CARVALHO et al., 1993 and BERNASCONI et al., 2000). The keys to species were based on diagnostic characters available in the literature, the study of type-material for some species and adaptations on some published keys. As in the Catalogue of the Neotropical Region by CARVALHO et al. (1993), the geographical coverage is the continent of South America as far north as, and including, Mexico; the islands of the Caribbean and the Antilles, including the Bahamas but excluding Bermuda; the Falklands (Malvinas), Galapagos Islands, Easter Island, and Juan Fernandez Islands, in the southern Atlantic and Pacific Oceans respectively. The known geographical distribution in the Neotropical Region was included in the keys. It was not possible to 20


Claudio Jose Barros de Carvalho & Marcia Souto Couri

include all species in the keys, since some of them are insufficiently known in the literature and the type-material was not examined. Most of the illustrations have been already published, but a few modifications were introduced. Abbreviations for the Museums and Institutions from which material was examined are as follows: AMNH - American Museum ofNatural History, New York, USA BMNH - The Natural History Museum, London, UK CNC- Canadian National Collection, Ottawa, Canada DZUP - Departamento de Zoologia, Universidade Federal do Parana (Cole

R4 +5

3 Figs. 1-3, characters of Atherigona (modified from PONT & MAGPAYO, 1995): 1. head, dorsolateral view; 2, wing, dorsal view; 3, ovipositor, dorsal view. Abbreviations: ar, arista; M, vein; R4+5 , vein; tg, tergite.

Subfamily Muscinae Latreille, 1802 Diagnosis. Arista plumose; anepimeron setulose; most of the genera with vein M strongly curved forward apically; Musca-type egg (according to HERTING, 1957); larvae coprophagous. Monophyly. HENNIG (1965) justified the monophy ly of the subfamily based mainly on characters of the ovipositor -long and tubular and sternite 8 well developed along its whole length. CARVALHO (1989d) justified the monophy ly of the subfamily based on 7 characters of the ovipositor, associated with the shape of the egg (Muscatype) and the presence of setulae in the anepimeron. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

39


Comments. Traditionally, the Muscinae has been divided into two tribesMuscini and Stomoxyini, as presented by CARVALHO (1989d) and CARVALHO et al. (1993).

Tribe Muscini Latreille, 1802 Diagnosis. Most of the genera with lower calypter strongly enlarged, Muscatype, sub-truncate posteriorly, with its anteromedian angle extending at least slightly under base of scutellum; cereal plate, on its ventral surface, with spiny processes. Monophyly. HENNIG (1965) considered the tribe monophyletic, based on the setulose anepimeron, plumose arista and tortuous slits of posterior larval spiracles (see SKIDMORE, 1985). CARVALHO (1989d) indicated the monophyly ofthe tribe based on the enlarged Musca-type lowercalypter (see HENNIG, 1965). Comments. The tribe has 23 genera worldwide, 11 of them occurring in the neotropics, with about 58 species ascribed to this region.

Biopyrellia Townsend, 1932 Type-species. Musca bipuncta Wiedemann (orig. des.). Diagnosis. General body coloration dark metallic blue; male with anterointemal ommatidia very enlarged, with about the same diameter as anterior ocellus; vibrissa weak, inserted above oral margin; dorsocentral setae 0: 1 or 0:2 (the anterior one weak); acrostichal 0:0 or 0: 1 (weak); humerall; intra-alars 0: 1; supra-alars 0:2; intrapostalar seta absent; katepistemals 0: 1 or 0:2 (the postero-anterior one weak); lower calypter enlarged, about twice as long as upper calypter; apical portion of vein M strongly bent forward. Reference. PAMPLONA, 1986a. Monophyly. Not yet established. Geographical record. Mexico, Costa Rica, Panama, Venezuela, Trinidad, Bolivia, Brazil, Paraguay, Argentina. Comments. Monobasic genus: Biopyrellia bipuncta (Wiedemann, 1830), validated by PAMPLONA (1986a).

40



Cyacyrtoneura Townsend, 1931 Type-species. Cyacyrtoneura cyanea Townsend (orig. des.). Diagnosis. General body coloration metallic blue with abdomen metallic green; katepistemal setae 1: 1. Reference. TOWNSEND, 1931. Geographical record. Venezuela, Peru, Brazil. Monophyly. Not yet established. Comments. Monobasic genus: Cyacyrtoneura cyanea Townsend, 1931. This species is known only from the type-material (ZMHU), which has not been examined after of the original description.

Dasymorellia Malloch, 1923 Type-species. Dasymorellia trichops Malloch (orig. des.). Diagnosis. Male holoptic; female with a pair of fronto-orbital proclinate setae; arista plumose; humeral 4; prealar strong; dorsocentrals 2:4; acrostichals 2:2 or 2:3; katepistemals 1:2; wings with cilia at node on both surfaces (except D. benoisti); apical portion of vein M bent forward; lower calypter glossiform, about twice as long as upper calypter; fore tibia on dorsal surface with 2 preapical setae; hind tibia with a strong posterodorsal seta (calcar). Reference. PAMPLONA, 1983. Monophyly. PAMPLONA (1983) did not present unique characters to support the monophyly of the genus. Geographical record. Mexico, Guatemala, Ecuador, Brazil, Colombia, Venezuela, Bolivia, Argentina, Peru. Comments. D.fulvipes (Bigot, 1887) (Mexico), D. saphirina (Seguy, 1935) (Ecuador) and D. spinifera (Wulp, 1883) (Argentina) were not included in the key due to lack of decisive information in the available descriptions (types not seen). PAMPLONA (1983) did not include D. saphirina in this genus, a lapsus corrected by CARVALHO et al. (1993). Nine species are known in the Neotropical Region.


41


Key to species (modified from PAMPLONA, 1983) 1.

Antenna and palpus yellow (Brazil) .......................... D.j1avipalpis Pamplona, 1983 Antenna and palpus brown ..................................................................................... 2

2.

Eye strongly haired in both sexes; parafacial, facial and gena shiny; gena with many hairs; acrostichals 2:3 .............................................................................................. 3 Eye haired in both sexes; hairs short and sparse in female (almost bare in D. callidimera); parafacial, facial and gena with grey pruinescence; gena with weak and

sparse hairs; acrostichals 2:2 .................................................................................. 4 3.

Wing bare on dorsal surface, mid femur with no anterodorsal setae (Ecuador) .................................................................................. .D. benoisti Pamplona, 1983

Wmg with about 5 cilia on dorsal surface, beginning at node, mid femur with 4 anterodorsal setae on middle third (Venezuela, Colombia) .................... .D. seguyi Pamplona, 1983 4.

Hind femur with 2 preapical anterodorsal setae, hind tibia with 5 anteroventral setae on apical half (Colombia, Ecuador, Peru, Bolivia, Brazil, Argentina) .........................................................................................D. trichops Malloch, 1923

Hind femur with a row of anterodorsal setae, hind tibia with less than 5 anteroventral setae on apical half. ................................................................................................. 5 5.

Legs yellow, hind femur with one median posteroventral seta (Mexico, Guatemala) ............................................................................... .... D. callidimera (Bigot, 1887)

Legs brown, hind femur with 2 median posteroventral setae (Colombia) .................................................................................. ... D. nigritibia Snyder, 1949

Morellia Robineau-Desvoidy, 1830 (Figs. 4-22, Plate 1) Type-species. Morellia agilis Robineau-Desvoidy (TOWNSEND, 1916) = hortorum (Fallen). 42



Diagnosis. Male holoptic; anterointemal ommatidia enlarged; eyes bare or with very few cilia; palpus a little spatulate apically; arista plumose on basal two-thirds or threequarters and bare on apically (Fig. 4); acrostichals 0: 1; dorsocentrals 0:2, 0:3 or 1:2; intraalars 1: 1 or 1:2; supra-alars 0:3 or 1:3; katepisternals 1:2; anepimeron setulose; meron bare; metepimeron haired above hind coxa; M strongly curved forward apically (Fig. 5); stemite 1 setulose; ovipositor long with tergites very thin and possessing strong spines (Figs. 6a, 6b, 7a, 7b). Reference. PAMPLONA, 1986b. Monophyly. No revision has been publishing for this genus. Its species are found in all biogeographic regions of the world. Denise Pamplona (personal communication) is finishing a worldwide revision and a cladistic analysis for the species of Morellia. Geographical record. Mexico, Jamaica, El Salvador, Costa Rica, Panama, Venezuela, Colombia, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical and Australian Regions. Comments. There are 19 species recorded to the Neotropical Region, 14 of them occurring in South America.

Key to species (modified from PAMPLONA, 1986b) 1.

Humeral callus yellow............................................................................................. 2 Humeral callus blue or dark metallic green ............................................................. 3

2.

Humeral callus with 4 setae; proepimeron setulose; wings without clouds (Mexico, EI Salvador, Costa Rica, Jamaica, Cuba, Haiti, Puerto Rico, Virgin Is., Venezuela; Nearctic Region) ..................................................... ......................M. basalis (Walker, 1853) Humeral callus with 3 setae; proepimeron bare; wings with clouds (Peru, Bolivia, Brazil) ...........................................................................M. humeralis (Stein, 1918)

3.

Wing without clouds .............................................................................................. 4 Wing with clouds (Figs. 12-22) ............................................................................. 7

4.

Antenna brown ..................................................................................................... 5 Antenna yellow...................................................................................................... 6


43


5.

Palpus yellow; lower half of fronto-orbital plate and upper half of gena grey pruinose; lower calypter brown (Brazil) ..................................... ..M. hirtitibia Pamplona, 1986 Palpus brown; fronto-orbital plate and gena entirely shiny; lowercalypteryellow (South America) ............................................... ................M. diversipalpis (Rondani, 1863)

6.

Intra-alar setae frequently 1:2; supra-alar setae frequently 0:3; mid femur on anteroventral surface without setae; hind femur with a row of dorsal setae; sursty Ius with a median internal seta (Fig. 8); paramere with setulae apically (Figs. 9,10); epiproct with spines on base, closely placed to each other (Fig. 11) (Brazil)

.................................................................................. ....M. dalcyi Pamplona, 1986 Intra-alar setae frequently 1: 1, supra-alar setae frequently 1:3; mid femur with a row of anteroventral setae; hind femur without dorsal setae; surstylus without a median internal seta; paramere with setulae apically; epiproct with spines frequently shorter than in M. dalcyi (Mexico, Costa Rica, Jamaica, Colombia, Ecuador, Venezuela, Peru, Brazil, Paraguay, Argentina) ............. M. violacea (Robineau-Desvoidy, 1830) 7.

Wing with no more than 2 faint, small spots (Figs. 12-16) ...................................... 8 Wing with at least 2 conspicuous spots (Figs. 17-22) .......................................... 13

8.

Flagellomere yellow.............................................................................................. 9 Flagellomere brown ............................................................................................ 10

9.

Wing with a discrete cloud on humeral vein; mid femur on posterodorsal surface with a group of strong and short apical setae, forming a rigid tuft; hind tibia on anteroventral surface with 5-6 setae on apical half (Brazil) .................. .

............................ .............................. M. paulistensis Pamplona & Mendes, 1995 Wing with a discrete brown cloud on humeral vein and crossvein r-m (Fig. 14); no tuft on mid femur; quetotaxy of hind tibia not as above (Mexico, Cuba, Puerto Rico, Galapagos Is., Brazil) ........................................ M. ochricornis (Wiedemann, 1830) 10. Pro sternum setulose ............................................................................................. 11 44



Pro sternum bare ................................................................................................ 12 11. Hind tibia, on anterodorsal surface, with 4 setae evenly distributed from base to apex and with one seta on basal third of posterodorsal surface (Panama) ....................................................................................... .M. affinis Malloch, 1925

Hind tibia, on anterodorsal surface, with 2 setae on apical half and with one seta on apical third of posterodorsal surface (Brazil) ..............M. concacata Pamplona, 1986 12. Dorsocentral setae 1:2, hind tibia with 6 posteroventral setae (Brazil) .................................................................................. .... M. roppai Pamplona, 1986

Dorsocentral setae 0:2, hind tibia without posteroventral setae (Brazil) .............................................................................. .M. sinopensis Pamplona, 1986

13. Wing brown from base to apex along costa, this stripe connected to a dark cloud over crossvein r-m (Cuba, Peru, Bolivia) ......................M. semimarginata (Stein, 1918) Wing not as above ............................................................................................... 14 14. Crossvein dm-cu without spots (Fig. 19) ............................................................. 15 Crossvein dm-cu spotted (Fig. 17) ....................................................................... 16 15. Wing with a long cloud extending from apical portion of vein Sc to wing apex (Costa Rica, Guyana, Peru, Brazil, Paraguay) ............................M. nigricosta Hough, 1900 Wing cloud not as above (Fig. 22) (Brazil) ....................M. lopesae Pamplona, 1986 16. Cell r1 with 1 spot (Fig. 21) (Brazil) ...................... M. xanthoptera Pamplona, 1986 Cell r 1 with 2 spots (Fig. 22) ............................................................................... 17 17. Flagellomere brownish yellow with a basal yellow spot; hind femur with weak and short anteroventral setae only on apical third; male: anteroventral, ventral and posteroventral surfaces with few setae on apical half (Brazil) ........................................................................... ........ M. couriae Pamplona, 1986

Flagellomere brownish yellow but lacking a basal yellow spot; hind femur, on Muscidae (Diptera) of the Neotropical Region: TAXONOMY

45


anteroventral surface, with weak and short setae from base to apex; male: anteroventral, ventral and posteroventral surfaces with many setae on apical 'half (Brazil)

........................................................... .M. dendropanacis Pamplona e Couri, 1995

fl

st7

IO,3mm

6a

6b

-cer

7a

7b

Figs. 4-7, characters de Morellia (modified from PAMPLONA, 1986c; PAMPLONA & COURI, 1995): 4, Morellia sp., antenna, lateral view; 5, M. concacata, wing, dorsal view; 6a, M. concacata, ovipositor, dorsal view; 6b, M. concacata, ovipositor, ventral view; Fig. 7a, M. dendropanacis, ovipositor, dorsal view; 7b, M. dendropanacis, ovipositor, ventral view. Abbreviations: ar, arista; cer, cerci; f1, f1agellomere; M, vein; R I , vein; R2 +3 , vein; Sc, subcostal vein; st, sternite.

46



pal'

2mm

~

~2 2m01

201m

201m

2mm

Figs. 8-22, characters of Morellia (modified from PAMPLONA, 1986c; PAMPLONA & COURI, 1995): 8, M. dalcyi, left surstylus, dorsal view; 9, M. dalcyi, aedeagus, ventral view; 10, M. dalcyi, aedeagus, lateral view; 11, M. dalcyi, ovipositor, dorsal view; 12, M. affinis, wing, dorsal view; 13, M. concacata, wing, dorsal view; 14, M. ochricornis, wing, dorsal view; 15, M. roppai, wing, dorsal view; 16, M. sinopensis, wing, dorsal view; 17, M. couriae, wing, dorsal view; 18, M. humeralis, wing, dorsal view; 19, M. iopesae, wing, dorsal view; 20, M. nigricosta, wing. dorsal view; 21, M. xanthoptera, wing, dorsal view; 22, M. dendropanacis, wing, dorsal view. Abbreviations: dis, distiphallus; dm-cu, crossvein; epi, epiphallus; gon, gonopod; hyp, hypandrium; par, paramere; r-m, crossvein.


47


Musca Linnaeus, 1758 Type-species. Musca domestica Linnaeus (I. C. Z. N., 1925, Opinion 82). Diagnosis. Male holoptic; anterointernal ommatidia enlarged; eyes bare; arista enlarged on basal fourth, plumose; pre sutural acrostichals not developed; dorsocentral 2:4; katepisternals 1:2; anepimeron setulose; prosternum setulose; lower calypter broad, subtruncate posteriorly, with its anteromedian angle extending, at least, to slightly under base of scutellum; mid tibia without ventral setae; vein M strongly curved forward apically; sternite 1 setulose. References. HENNIG, 1963b, 1964. Monophyly. According to HENNIG (1965), the monophyly of this genus is not well established. Geographical record. Mexico, Guadeloupe Is., Guatemala, Belize, San Salvador, Nicaragua, Costa Rica, Panama, Nassau, Jamaica, Puerto Rico, Haiti, Dominica, Cuba, Barbados, Mona Is., St. Lucia, St. Vincent, Virgin Is., Grenada, Trinidad, Venezuela, Guyana, Surinam, Colombia, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is., Easter Is.; Nearctic and Palaearctic Regions; northwestern Oriental; cosmopolitan. Comments. Only M. domestica is recorded for the Neotropical Region. The following key also includes M. autumnalis De Geer, 1776, based on its economic importance and the possibility that it might invade the Neotropical Region. The larvae of both species can be easily distinguished by the posterior end ofthe third instar (STEYSKAL, 1973).

Key to species 1.

Frons totally black in male and grey in female, not shiny (Mexico, Nearctic and Palaearctic Regions, northwestern Oriental) ................M. autumnalis De Geer, 1776 Frons reddish brown around and under antenna in male and silver shiny in female (Mexico, Guadeloupe Is., Guatemala, Belize, San Salvador, Nicaragua, Costa Rica, Panama, Nassau, Jamaica, Puerto Rico, Haiti, Dominica, Cuba, Barbados, Mona Is., St. Lucia, St. Vincent, Virgin Is., Grenada, Trinidad, Venezuela, Guyana, Surinam,

48


Claudio Jose Barros de Carvalho & Marcia Souto Co uri

Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is., Easter Is.; cosmopoIltan) .......M. domestica Linnaeus, 1758

Neomyia Walker, 1859 Type-species. Neomyia gavisa Walker (mon.). Diagnosis. General body coloration metallic green; male holoptic; eyes bare; arista plumose; presutural acrostichals not developed; dorsocentral2:3; katepisternals 1:2; anepimeron setulose; prosternum setulose; lower calypter broad, subtruncate posteriorly, with its anteromedian angle extending, at least, to slightly under base of scutellum; mid tibia with a strong, ventral seta apically; vein M strongly curved forward apically; sternite 1 setulose. Reference. HENNIG, 1963b (as caesarion). Monophyly. Not yet established. Geographical record. Mexico, Cuba, Costa Rica, Brazil; Nearctic and Palaearctic Regions, northwestern Oriental. Comments. Only Neomyia cornicina (Fabricius, 1781) is found in the Neotropical Region.

Neorypellia Pont, 1972 Type-species. Neopyrellia neglecta Townsend (orig. des.). Diagnosis. General body coloration metallic green; male holoptic; anterointernal facets elongated; eyes bare; interfrontal setae absent; arista plumose; prosternum bare; pre sutural acrostichals 1; posterior dorsocentral2 strong setae; intra-alar 1: 1; supra-alars 2; katepisternals 1:2; anepimeron setulose; calypteres glossiform, the upper one with basal half ciliated ventrally; base of veins R 2+3 and R 4+5 with cilia on dorsal surface; stem vein with cilia on both surfaces; M strongly curved forward apically; hind tibia with a small posterodorsal seta (calcar), shorter than the diameter of tibia; sternite 1 setulose. Reference. ALBUQUERQUE, 1955a. Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: Neorypellia neglecta (Townsend, 1939). Muscidae (Diptera) of the Neotropical Region: TAXONOMY

49


Parapyrellia Townsend, 1915 (Figs. 23-25) Type-species. Musca violacea Fabricius of authors (orig. des.) =maculipennis Macquart Diagnosis. General coloration metallic green; male holoptic with anterointernal facets elongated; eyes bare; arista plumose; interfrontal setae absent; prosternum bare; dorsocentrals 1:2; supra-alars 2; katepisternals 1:2; anepimeron bare; calypteres glossiform, the lower one enlarged; vein M strongly curved forward apically (Figs. 23,24); sternite 1 setulose. Reference. ALBUQUERQUE & LOPES, 1979a. Monophyly. Not yet established. Geographical record. Mexico, Cuba, Puerto Rico, Jamaica, Dominica, Trinidad, Guadeloupe Is., Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay. Comments. The two known species occur only in the Neotropical Region.

Key to species 1.

Male: hind tarsi without a strong spine; female: calypter brown; wing not as below (Fig. 23) (Mexico, Cuba, Puerto Rico, Jamaica, Dominica, Trinidad, Guadeloupe Is., Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay)

.......................................................................... P. maculipennis (Macquart, 1846) Male: hind tarsi with a strong spine (Fig. 25); female: calypter yellow; wing as in Fig. 24 (Brazil) ............................................... P. oportuna Albuquerque & Lopes, 1979

50



~ 23

met

Figs. 23-25, characters of Parapyrellia (modified from ALBUQUERQUE & LOPES, 1979a): 23, P. lI1aclllipennis, wing, dorsal view; 24, P. oportllna, wing, ventral view; 25, P. oportllna, hind leg, lateral view. Abbreviation: met, metatarsus.

Polietina Schnabl & Dziedzicki, 1911 (Figs. 26-32, Plate 1) Type-species. Aricia pruinosa Macquart (orig. des.) = Mydaea concinna Wulp. Aricia pruinosa Macquart, 1846, preoc. Zetterstedt, 1845. Diagnosis. Eyes bare or ciliated; male holoptic; arista with long cilia on both surfaces, longer near base, measuring about twice the length of the pedicel; one proclinated orbital seta; palpus filiform or slightly spatulate apically; presutural acrostichals developed or not; dorsocentrals 2:3 or 2:4; prealar strong and long; pro sternum setulose laterally; membrane around prosternum bare; one long proepisternal setae and one long upward directed proepimeral seta; notopleuron with 3 setae, the median one short; postalar wall ciliated; margin of posterior spiracle ciliated; katepisternals 1:2; calypteres bare on dorsal surface, lower calypter about twice as long as upper one; vein R] on dorsal surface ciliated from base to apex; R 4+5 on dorsal surface with cilia from node to apex or almost so, on ventral surface with sparse cilia up to near crossvein r-m or near the apex; hind tibia, on posterodorsal surface, with a strong calcar and a strong pre apical seta; sternite 1 setulose; male: sternite 5 longer than wide; cercal plate on ventral surface with spiny processes (Figs. 26-32); female: ovipositor long, tergites elongated, wider basally. Reference. COURI & CARVALHO, 1997b. Monophyly. Genus monophyletic, based at least, on the presence of setae on Muscidae (Diptera) of the NeotropicaI Region: TAXONOMY

51


postalar wall, a derived character among the Muscidae (COURI & CARVALHO, 1997b). Geographical record. Mexico, Venezuela, Bolivia, Peru, Brazil, Argentina. Comments. COURI & PAMPLONA (1997) transferred Cyrtoneurina ingrata Couri, 1982, Cyrtoneurina prima Couri & Machado, 1990 and Cyrtoneurina stellata Couri, 1992 to this genus. These species were included in the key below. Key to species (modified from COURI & CARVALHO, 1997b)

Male 1.

Postsutural dorsocentral setae 3 ............................................................................ 2 Postsutural dorsocentral setae 4 ............................................................................ 8

2.

Hind tibia, on anteroventral surface, with less than 4 setae on middle-third (usually 3) .......................................................................................................................... 3 Hind tibia, on anteroventral surface, with more than 4 setae on middle-third (usually 5) ......................................................................................................................... 7

3.

Presutural acrostichal setae not developed; spiny lower process of cercal plate with 3 spines similar in length (Fig. 27) (Mexico )............. .l~ wulpi Couri & Carvalho, 1997 Presutural acrostichal setae developed (one or two pairs); spiny lower process of cercal plate with 3 spines, one more developed than the others ............................ ..4

4.

Spines on lower process of cercal plate directed upward (Fig. 28) (Brazil)

.............................................................................. ..... P. minor Albuquerque, 1956 Spines on lower process of cercal plate directed downward (Fig. 29) (Brazil) ......... 5 5.

Lower process of cercal plate with 5 spines; hind tibia on anterodorsal surface with at most 5 setae on middle third (Brazil) ................ P. univittata Couri & Carvalho, 1996 Lower process of cercal plate with less than 5 spines; hind tibia on anterodorsal surface with more than 5 setae on middle third ....................................................... 6

6.

Lower process of cercal plate with 3 spines (Fig. 29) (Brazil) .......................... ..

52



.......................................................................... ....... .P. bicolor Albuquerque, 1956 Lower process of cercal plate with 4 spines (Brazil) ..... ..P. nigra Couri & Carvalho, 1996 7.

Wings not spotted; spiny lower process of cercal plate with 3 downwarddirected spines, 1 more developed than others (Fig. 26) (Peru, Brazil) .......... . ..................................................................................... P. jlavithorax (Stein, 1904)

Wings with brown spots on stigma, apical third of vein R 2+3 and on crossveins r-m and dm-cu (Brazil) .............................................................. ....... P. stellata (Couri, 1982) 8.

Spiny lower process of cercal plate with 4 well developed spines (Fig. 30) (Brazil) ....................................................................................... P. steini (Enderlein, 1927)

Spiny lower process of cercal plate with less than 4 spines ...................................... 9 9.

Spiny lower process of cercal plate with 2 developed spines, one almost vestigial (Fig. 39) (Mexico) .............................................. ............. P. concinna (Wulp, 1896) Spiny lower process of cercal plate with only one developed spine (Fig. 32) (Peru, Bolivia, Brazil, Argentina) ............................................ ..... P. orbitalis (Stein, 1904)

Female 1.

Postsutural dorsocentral setae 3 ............................................................................. 2 Postsutural dorsocentral setae 4 ............................................................................. 7

2.

Tergite 5 paler than remainder of abdomen ............................................................. 3 Tergite 5 concolours with remainder of abdomen .................................................. ..4

3.

Tergite 5 yellow on apical half; hind tibia with about 5 anteroventral setae on middle third (Venezuela, Peru, Bolivia, Brazil) ........................... P.jlavithorax (Stein, 1904) Tergite 5 entirely light brown; hind tibia with about 2 anteroventral setae on middle third (Brazil) .............................................. ......... P. prima (Couri & Machado, 1990)


53


4. Fore leg brown; hind tibia on anterodorsal surface with no more than 5 setae (Brazil)

................................................................................... P minor Albuquerque, 1956 Fore leg yellow or light brown; hind tibia on anterodorsal surface with more than 5 setae ..................................................................................................................... 5 5.

Fore leg light brown (Brazil) ........................... P. univittata Couri & Carvalho, 1996 Fore leg yellow................................................... ,......................................... '" ..... 6

6.

Hind tibia on anterodorsal surface with 7 setae on middle third (Brazil)

............................................................................... ? nigra Couri & Carvalho, 1996 Hind tibia on anterodorsal surface with 8-9 setae on middle third (Brazil)

................................................................................... ? bicolor Albuquerque, 1956 7.

Hind tibia on anteroventral surface with 5 or more setae (Brazil) .............. .

................................................................... .................. P. major Albuquerque, 1956 Hind tibia on anteroventral surface with no more than 4 setae .................................. 8 8.

Abdomen brown, with yellow, translucent areas on tergites 1+2 and 3, tergite 5 yellow; antennal flagellomere brown (Mexico, Venezuela) ............... P. rubella (Wulp, 1896) Abdomen not as above (see below); antennal flagellomere yellow.......................... 9

9.

Hind tibia on anteroventral surface with 4 setae on middle third; legs yellow, mid femur brown apically; hind femur brown on apical half (Brazil) ............ .

................................................................ ....................... P. steini (Enderlein, 1927) Hind tibia on anteroventra1 surface with 2 setae on middle third; leg colour not as below................................................................................................................... 10 10. Legs brown; fore femur, fore tibia and mid femur yellowish brown; abdomen silver pruinose and with yellow lateral spots on basal half of tergites 1-3 (Brazil)

.............................................................................. ? distincta Couri & Lopes, 1987 Legs and abdomen not as above (see below) ......................................................... 11 54



11. Abdomen yellowish with dark brown vittae on apex of tergites 1-4, tergite 5 with yellow apex (Mexico) ....................................................... P. concinna (Wulp, 1896) Abdomen not as above ........................................................................................ 12 12. Legs yellow (Peru, Bolivia, Brazil, Argentina) ......................P. orbitalis (Stein, 1904) Legs brown, with apical portion of fore and hind femora yellow (Brazil)

.......................................................................................... P. ingrata (Couri, 1982) Species of Polietina not included in the key and their respective geographical distribution Species

Geographical distribution

P. basicincta (Stein, 1904)

Peru, Bolivia

P. jlavidicincta (Stein, 1904)

Colombia, Venezuela, Bolivia

P. mellina (Stein, 1904)

Bolivia

Sarcopromusca Townsend, 1927 Type-species. Sarcopromusca arcuata Townsend (mon.) =pruna (Shannon & Del Ponte). Diagnosis. General coloration blackish with grey pruinescence; male holoptic; arista enlarged at basal fourth, plumose; acrostichals 0:2; dorsocentra13 :4; meron setulose;

katepisternals 1:3; anepimeron setulose; vein M strongly curved forward apically; lower calypter with cilia on internal border; mid tibia on ventral surface with a pre apical seta; apex of abdomen yellow gold; sternite 1 setulose. Reference. PAMPLONA, 1991. Monophyly. Not yet established. Geographical record. Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Brazil, Argentina. Comments. Only two species are known, both restricted to the Neotropical Region.


55


Key to species 1.

Costal margin without an elongated brown cloud (Guatemala, Nicaragua, Costa Rica, Brazil, Argentina) ........................................ S. pruna (Shannon & Del Ponte, 1926) Costal margin with an elongated brown cloud (Costa Rica, Panama, Colombia, Brazil, Argentina) ................................................. ................ S. sarcophagina (Wulp, 1896)

1 mm

1mm

1mm

Figs. 26-32, characters of cereal plate, ventral view, of Polietina (modified from COURI & CARVALHO, 1997b): 26, P. flavithorax; 27, P. wUlpi; 28, P. minor; 29, P. bicolor; 30, P. steini; 31, P. concillfla; 32, P. orbitalis. Abbreviation: pr, process.

56



Xenomorellia Malloch, 1923 (Figs. 33-36, Plate 1) Type-species. Xenamarellia haiti Malloch (orig. des.). Diagnosis. Male holoptic; arista plumose, bare on apical fourth; prealar seta long and strong; acrostichals 2:2; dorsocentrals 2:4; anepimeron setulose; M curved forward apically (Fig. 33); stem vein with cilia on dorsal surface and with a long seta near the insertion of the humeral crossvein (Fig. 34); radial node with cilia on both surfaces; hind tibia with 2 posterior setae; hind coxa with posterior cilia; sternite 1 setulose. Reference. ALBUQUERQUE, 1952. Monophyly. Not yet established. Geographical record. Peru, Brazil. Comments. A Neotropical genus with only two described species.

Key to species 1.

Antenna and palpus black; setae on basal half of anteroventral surface of hind femur at least twice as long as the width of the tibia, on apical half with about 5 setae of unequal lengths (Fig. 35) (Brazil) .................. ...... .x. mantanhesa Albuquerque, 1952 Antenna and palpus dark yellow; setae on basal half of anteroventral surface of hind femur less than 1,5 times as long as the width of the tibia, on apical half with a series of long setae (Fig. 36) (Peru, Brazil) ............................................


x. haiti Malloch, 1923

57


hum

O,5mm

1 rnm

35

36

Figs. 33-36, characters of Xenomorellia (modified from ALBUQUERQUE, 1952): 33, X. montanhesa, wing, dorsal view; 34, X. montanhesa, wing, humeral vein, dorsal view; 35, X. montanhesa, hind leg, lateral view; 36, X. holti, hind leg, lateral view. Abbreviations: ad, anterodorsal surface; av, anteroventral surface; hum, humeral vein; sv, stem vein.

Tribe Stomoxyini Meigen, 1924

Diagnosis. Proboscis elongated, adapted to haematophagy, strongly sclerotized; labella reduced and not retractile; arista with long dorsal cilia and generally bare or at most with very few ventral cilia; male: epiphallus absent; female: 2 spermathecae. Monophyly. Tribe undoubtedly monophyletic (HENNIG, 1965, 1973), having Muscini as sister-group (CARVALHO, 1989d). Comments. This tribe have species distributed allover the world, with 10 described genera. Only three genera and six species are recorded to the Neotropical Region.

58


m


Haematobia Le Peletier & Serville in LATREILLE et ai., 1828 Type-species. Conops irritans Linnaeus (WESTWOOD, 1840: 140 and I.C.Z.N., 1974, Opinion 1008). Diagnosis. General coloration dark greyish, with yellowish tinge and often yellow setae; head width 1.25 times its height; arista plumose only on dorsal surface; palpus spatulate, a little narrowed on basal quarter to half, grooved inside, reaching tip ofhaustellum when proboscis flexed; prostemum setulose; propleuron and postalar wall bare; prealar seta barely differentiated or vestigial; lower calypter about twice as long as upper one; vein AJlong. Reference. HENNIG, 1964. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, EI Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., Haiti, S. Domingo, Colombia, Venezuela, Guyana, Brazil, Argentina, Chile; Nearctic Region; Europe, Asia, North Africa, Hawaii. Comments. This genus has species distributed in all biogeographical regions, but only Haematobia irritans (Linnaeus, 1758) occurs in the Neotropical Region. This species probably came from Venezuela to Brazil in the beginning of the eighties, and its widespread occurrence in this country was due to dispersion (VALERIO & GUIMARAES, 1983).

Neivamyia Pinto & Fonseca, 1930 (Figs. 37-44) Type-species. Neivamyia Zutzi Pinto & Fonseca (orig.des.)

=flavicornis

Malloch. Diagnosis. Head slightly wider than high; palpus stout subcylindric and reaching tip ofhaustellum when proboscis flexed; arista plumose on upper surface and with about 3 or 4 shorter cilia below on middle portion (Figs. 37,42,44); prostemum setulose; propleura and postalar wall bare, with at least 2 long postsutural dorsocentral setae; vein Al moderately long; lower calypter about twice as long as upper one; mid and hind tibiae with a submedian seta. Reference. LOPES & MANGABEIRA FILHO, 1938; LOPES, 1955. Monophyly. Not yet established. Adrian Charles Pont (personal communication)


59


understands that Neivamyia could be synonymized with Stomoxys, since there are no significant differences between them. Geographical record. Trinidad, Guyana, Surinam, Peru, Brazil. Comments. A Neotropical genus with only four species.

Key to species (males only) 1.

Male: orbital proclinate setae on frons .................................................................... 2 Male: no proclinate setae on frons ......................................................................... 3

2.

Frons about one-fifth of head width (Figs. 38, 39) (Colombia) ....................... .

...... ........ ...... ........ .... ..... ...... ........ ......... ..... ....................... .N. antenusi Lopes, 1955 Frons less than one-fifth of head width (Peru, Brazil) (Fig. 40) .......................... ..

.................. ......... ..... ......... ......... ....... .................... .........N. latifrons Malloch, 1932 3.

Frons less than one-tenth of head width (Fig. 41); proboscis measuring about the same length as the maximum height of the eye (Fig. 42) (Trinidad, Guyana, Surinam,

Brazil) ..................................................................... .N.Jlavicomis (Malloch, 1928) Frons more than one-tenth of head width (Fig.43); proboscis shorter than the maximum height the eye (Fig. 44) (Brazil) .. ..............N. travassoi Lopes & Mangabeira, 1938

Stomoxys Geoffroy, 1762 Type-species. Conops calcitrans Linnaeus (LC.Z.N., 1957, Opinion 441). Diagnosis. General coloration brownish grey to yellow brown; head slightly wider than high; vertex about one-fourth of head width in male and well over one-third in female; arista plumose; pal pus slender, subcylindric, scarcely one-third as long as haustellum; flagellomere about 2,5 as long as pedicel; prostemum and proepistemum setulose; prealar seta absent. Reference. HENNIG, 1964. Monophyly. Not yet established, but probably monophyletic (see discussion under Neivamyia). 60



O,5mm

1 mm

1 mm 1 mm

prb Figs. 37-44, characters of Neivamyia (modified from LOPES & MANGABEIRA, 1938; LOPES, 1955): 37, N. travassoi, antenna, lateral view; 38, N. antenusi, head, dorsal view; 39, N. antenusi, head, lateral view; 40, N. tatifrons, head, dorsal view; 41, N. ftavicornis, head, dorsal view; 42, N. ftavicornis, head, lateral view; 43, N. travassoi, head, dorsal view; 44, N. travassoi, head, lateral view. Abbreviations: ar, arista; pip, palpus; prb, proboscis.

Geographical record. Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., St. Vincent, Haiti, Dominica, Trinidad, Tobago, Venezuela, Guyana, Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Argentina, Chile, Easter Is.; cosmopolitan. Comments. The genus has species distributed in all biogeographical regions, but only Stomoxys calcitrans (Linnaeus, 1758) occurs in the Neotropical Region.


61


Subfamily Azeliinae Robineau-Desvoidy, 1830 Diagnosis. Arista not plumose, with the cilia shorter than the basal width of arista. Ovipositor with tergites 6 and 7 divided anteriorly; tergite 8 sc1erotized, anchor-type. Monophyly. Not established yet. Comments. SKIDMORE (1985) and CARVALHO (1989d) recognised the subfamily; the first author based on characters of the immature forms and the second, on derived characters of the adults listed in the diagnosis. The subfamily is divided in to two tribes, Azeliini and Reinwardtiini, tentatively indicated as sister-groups by CARVALHO (1989d).

Tribe Azeliini Robineau-Desvoidy, 1830 Diagnosis. Gena, in most species, with strong, upcurved setae; subcostal vein reaching apical margin of wing in a smooth curve; distiphallus with apical part of juxta spinulose (Figs. 53-56); female with a proc1inate fronto-orbital setae. Monophyly. Probably Azeliini is a monophyletic tribe (CARVALHO, 1993b). In most genera 2 character-states are found: a proc1inate fronto-orbital in female, and subcostal smooth. The former character-state is in the ground plan of the Muscidae (HENNIG, 1965). The second character is probably in the ground plan of Muscoidea (sensu McALPINE, 1989). However the most striking character in the tribe is the configuration of the male distiphallus, with the juxta spinulose (CARVALHO, 1993b). Comments. In the world there are nine genera, seven of them with species occurring in the Neotropical Region. 24 species are known to this region.

Azelia Robineau-Desvoidy, 1830 Type-species. Azeliajlorea Robineau-Desvoidy (RONDANI, 1866) =triquetra Wiedemann. Diagnosis. Wing with vein Al ending abruptly, extending at most halfway between the apex of the posterior cubital cell and the margin of the wing; hind coxa with a row of weak posterior setae; male with enlarged ommatidia; female with interfrontal setae. Reference. SNYDER, 1957b.

62



Monophyly. Not yet established. Geographical record. Mexico, Argentina. Comments. Genus with species distributed all over the world, but after CARVALHO (1993b), only one species, Azelia neotropica Snyder, 1957, has been recognised in the neotropics. VOCKEROTH (1996) in his key to the Muscidae of Central America, mentioned this genus to Mexico and Panama, but unfortunately he did not provide specific identification of the material.

Drymeia Meigen, 1826 Type-species. Drymeia obscura Meigen (mon.) =hamata (Fallen). Diagnosis. General coloration blackish; male wing brownish; female wing light brown; arista pubescent; prealar strong; hind tibia usually with 3 or more strong posterodorsal setae which are more or less uniformly distributed along tibial length. References. HUCKETT, 1975; VOCKEROTH, 1996. Monophyly. Not yet established. Geographical record. Mexico; Nearctic, Palaearctic and Oriental regions. Comments: It seems that only Drymeia aterrima Wulp, 1896 occurs in Mexico (CARVALHO et al. 1993), although VOCKEROTH (1996) did not include this species in his key to the Muscidae of Central America. The syntypes males and females are in the BMNH and they are conspecifics.

Hydrotaea Robineau-Desvoidy, 1830 (Figs. 45-70) Type-species. Musca meteorica Linnaeus (des. CURTIS, 1839). Diagnosis. General coloration black or bluish-black, not metallic; gena with strong, upward-directed setae; thorax with no vittae or with 2 or 3 white, pruinose vittae; arista pubescent; dorsocentral setae 2:4; notopleuron covered with setulae, with the 2 setae of similar size; male: fore femur on ventral and posteroventral surfaces with 2 toothed processes; fore tibia flattened along basal half or two-thirds of ventral surface; female: inter-segmentar membrane of the ovipositor with granulations. Reference. PALKA-ROCHA & CARVALHO, 1994. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

63


Monophyly. Probably monophyletic. After the works of SKIDMORE (1985) and PALKA-ROCHA (1992), it has been hypothesized, based on a large number of characters, that Ophyra Robineau-Desvoidy is the sister-group of Australophyra + Hydrotaea Robineau-Desvoidy. PONT (1986a, 1989) considered Ophyra a junior synonym of Hydrotaea, a classification not followed by PAMPLONA & COURI (1989) and CARVALHO et al. (1993). Geographical record. Mexico, Costa Rica, Colombia, Ecuador, Peru, Bolivia, Brazil, Argentina, Chile, Juan Fernandez Is.; Nearctic and Palaearctic Regions; northern India; cosmopolitan. Comments. This genus is found in all biogeographical regions; only eight species occur in the Neotropical Region. Hydrotaea Juscocalyptrata Macquart, 1855 was not included in the key because this species is known only from one male in very bad condition, deposited at UMO (PALKA-ROCHA & CARVALHO, 1994).

Key to species (modified from PALKA-ROCHA & CARVALHO, 1994) 1.

Male: unknown; female: general coloration shiny black; wing darker on apical half (Colombia) ............................................ ..................... H. nubilicosta Malloch, 1923 Male: eyes ciliated or bare; prealar setae 1 or 2; sternite 5 similar or wider in length than in width (Figs. 49-52); surstylus, in posterior view, rounded or elongated (Figs. 45-48); distiphallus as in Figs. 53-56; hypandrium, in dorsal view, triangular or rounded (Figs. 57-60); female: general coloration black to bluish black, not shiny; wing hyaline or uniformly brownish ............................................................................................. 2

2.

Halter with the knob light brown or yellow; male: eyes bare; mid tibia on ventral surface with long and fine setae on apical half; hind femur on ventral surface with 2 strong basal spines (HUCKETT, 1954; ALBUQUERQUE, 1957a); female: prealar absent; mesonotum with 2 white, pruinose vittae that reach the third and fourth pairs of dorsocentral postsutural setae; ovipositor without granulations on the intersegmentar membrane 6-8 (Figs. 61-66) (Chile; North America) .................H. acuta Stein, 1898 Halter with the knob black; male: eye ciliated; mid tibia, on ventral surface with no setae on apical half; hind femur on ventral surface with no spines; female: prealar

64



present; mesonotum with no vittae; ovipositor not as described above (Figs. 62-65; 67-70) ................................................................................................................... 3 3.

General coloration bluish black; calypter dark brown; male: cerci with anterior margin angulated (Fig. 45); female: ovipositor without granulations on membrane of segment 8 (Fig. 67) (Colombia, Bolivia, Argentina, Chile )... .H. cyaneiventris Macquart, 1851 General coloration black; calypter not as described above; male: cerci with the anterior margin not as described above (Figs. 46-48); female: ovipositor with granulations on membrane of segment 8 (Figs. 63-65; 68-70) ........................................................ ..4

4.

Eye with long and dense cilia; calypteres yellow; fore femur, on anterior surface, with setae; male: prealar setae 1; surstyIi on posterior view elongated (Fig. 48); hypandrium in dorsal view rounded (Fig. 60); female: prealar 1; mid tibia, on anterodorsal surface with no setae on limit of the apical third (Colombia) ..................H. villosa Stein, 1904 Eyes with short and sparse cilia; calypteres not yellow; male: prealar 1 or 2; fore femur, on anterior surface, with a row of setae on basal half; surstyli on posterior view rounded (Figs. 45-47); hypandrium in dorsal view, triangular (Figs. 57-59); female: prealar 2; fore femur, on anterior surface, with cilia or with a row of weak setae on basal half; mid tibia, on anterodorsal surface, with one seta on the limit of the apical third ..................................................................................................................... 5

5.

Outer vertical seta similar in length to the inner ones; male: prealar 2; fore tibia on anteroventral surface, with a row of short and strong setae on apical third; stemite 5 longer than wider (Fig. 50); cerci as in Fig. 46; distiphallus with apical spines developed (Fig. 54); hypandrium as in Fig. 58; female: mesonotum with no vittae (Venezuela, Ecuador; North America; Hawaii) .......................................H. houghiMalloch, 1916 Outer vertical seta shorter than the inner ones; male: prealar 1 or 2; fore tibia not as described above; stemite 5 with similar length and width (Fig. 51); cerci as in Fig. 47; distiphallus with apical spines not as described above (Fig. 55); hypandrium as in Fig. 59; female: mesonotum with three polinose vittae (Ecuador, Peru, Bolivia, Brazil, Argentina, Chile) ............................................. ................H. nicholsoni Curran, 1939


7

65


Q,5mm

53

54

55

56

Figs. 45-60, characters de Hydrotaea (modified from PALKA-ROCHA & CARVALHO, 1994): 45, H. cyaneiventris, cerci, epandrium and surstyli, posterior view; 46, H. houghi, cerci, epandrium and surstyli, posterior view; 47, H. nicholsoni, cerci, epandrium and surstyli, posterior view; 48, H. villosa, cerci, epandrium and surstyli, posterior view; 49, H. cyaneiventris, sternite 5, posterior view; 50, H. houghi, sternite 5, posterior view; 51, H. nicholsoni, sternite 5, posterior view; 52, H. villosa, sternite 5, posterior view; 53, H. cyaneiventris, aedeagus, lateral view; 54, H. houghi, aedeagus, lateral view; 55, H. nicholsoni, aedeagus, lateral view; 56, H. villosa, aedeagus, lateral view; 57, H. cyaneiventris, hypandrium, dorsal view; 58, H. houghi, hypandrium, dorsal view; 59, H. nicholsoni, hypandrium, dorsal view; 60, H. villosa, hypandrium, dorsal view. Abbreviations: aed apo, aedeagal apodeme; cer, cerci (cereal plate); dis, distiphallus; epd, epandrium; epi, epiphallus; gon, gonopod; sur, surstylus.

66

~~~~


_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _EW"


66

67

68

69

70

Figs. 61-70, characters of Hydrotaea (modified from PALKA-ROCHA & CARVALHO, 1994): 61, H. acuta, ovipositor, dorsal view and spermathecae; 62, H. cyaneiventris, ovipositor, dorsal view and spermathecae; 63, H. houghi, ovipositor, dorsal view and spermathecae; 64, H. nicholsoni, ovipositor, dorsal view and spermathecae; 65, H. villosa, ovipositor, dorsal view and spermathecae; 66, H. acuta, ovipositor, ventral view; 67, H. cyaneiventris, ovipositor, ventral view; 68, ovipositor, ventral view, H. houghi; 69, ovipositor, ventral view, H. nicholsoni; 70, H. villosa, ovipositor, ventral view. Abbreviation: spr, spermathecae.


67


Micropotamia Carvalho, 1993 (Figs. 71-76) Type-species. Phaonia minuscula Albuquerque (orig. des.). Diagnosis. Parafacial very narrow in both sexes, strongly tapering downward; head of female with proclinate orbitals; interfrontal setae absent, frontal plate with several setae; arista pubescent, with hairs shorter than basal width of the arista; gena with a strong up curved seta; dorsocentral setae 2:3; pre sutural acrostichals developed; prealar short, slightly longer than adjacent ground-setulae; anepimeron bare; katepistemals 1:2, the lower one weak; posterior spiracle small in both sexes, smaller than the knob of halter, bare on margins; wing veins bare, except C; vein Sc smoothly curved on apical portion; mid femur with a preapical anterior seta; mid tibia with two median posterior setae; hind coxa bare on posterior apical margin; hind tibia with a strong calcar on apical third; stemite 1 bare; male: distiphallus with spiny and sclerotized processes; female: ovipositor long (Figs. 71,72). Reference. CARVALHO, 1993b. Monophyly. Probably monophyletic. According to CARVALHO (1993b), Micropotamia share an apomorphic character - absence of interfrontal setae in female with Megophyra Emden, 1965, which has only species occurring in the Oriental Region (PONT, 1977). Geographical record. Mexico, Brazil, Argentina. Comments. This genus was proposed for three South American species, but Phorbia fuscisquama Wulp, considered insertae sedis by CARVALHO et al. (1993), also belongs to this genus and is included in the key below. This species is known only from Mexico.

Key to species (modified from CARVALHO, 1993b) 1.

Antenna dark brown with the pedicel yellowish; thorax dark brown with humeral callus yellowish; halter yellow (Brazil) .... M. amazonica (Albuquerque & Lopes, 1982) Antenna entirely dark brown; thorax black, halter yellow, dark brown at base ......... 2

2.

Male: hind femur enlarged on apical third (AlBUQUERQUE, 1955a:Fig.1);cercalplate as in Fig. 75; stemite 5 as in Fig. 76 (Brazil) ............ M. minuscula (Albuquerque, 1955)

68



Male: hind femur not enlarged on apical third; cereal plate not as above ................. 3 3.

Male: hind femur with short setae on anteroventral face; cereal plate as in Fig. 73; sternite 5 as in Fig. 74; female: ovipositor not as below (Brazil) ........................ .

..............................................................~ .............M. cilitibia (Albuquerque, 1955) Male: hind femur with only long setae on whole anteroventral surface; cereal plate not as above; female: ovipositor with narrow stemites and tergites (Mexico) ..................... .

.............................................................. .... M.fuscisquama (Wulp, 1896), comb.n.

1 mm

73 E E

1

mm/

1mm

Figs. 71-76, characters of Micropotamia (modified from CARVALHO, 1993b): 71, M. amazonica, ovipositor, dorsal view; 72, M. amazonica, ovipositor, ventral view; 73, M. cilitibia, cereal plate, posterior view; 74, M. cilitibia, sternite 5, dorsal view; 75, M. minuscula, cereal plate, posterior view; 76, M. minuscula, sternite 5, dorsal view.


69


Ophyra RobineaugDesvoidy, 1830 (Figs. 77-88) Type-species. Ophyra nitida Robineau-Desvoidy (des. RONDANI, 1866) = ignava Harris. Diagnosis. General coloration metallic black; male holoptic; eyes bare; ocellar triangle shiny, large and long, sometimes reaching lunule; frontal setae 6-11 pairs; proclinate seta and interfrontal setae present in female; arista bare or pubescent; dorsocentral setae 2:4; katepisternals 1: 1; anepimeron bare; vein M straight; ovipositor long with narrow tergites. Reference. PAMPLONA & COURI, 1989. Monophyly. See discussion under genus Hydrotaea. Geographical record. Mexico, Nicaragua, St. Vincent, Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Argentina, Chile; Nearctic and Western Palaearctic Regions; Eastern Pacific). Comments. Besides the seven species recognised in the Neotropical Region, Ophyra ignava was recently identified in Chile (FIGUEIROA, 1999). VOCKEROTH (1996) followed PONT (1986a, 1989) and did not recognise Ophyra as a separate genus (see discussion under genus Hydrotaea).

Key to species (modified from PAMPLONA & COURI, 1989) 1.

Male ................................................................................................................... 2 Female ................................................................................................................ 8

2.

Palpus yellow or light brown; hind trochanter, on ventral surface, with a tuft of fine, hooked setae (Fig. 77) .......................................................................................... 3 Palpus dark brown or black; hind trochanter, on ventral surface, without a tuft of fine setae ..................................................................................................................... 4

3.

Mid femur with 6 short spinules on basal third; mid and hind femur brown to black; pedicel and flagellomere brown (Mexico, Nicaragua, St. Vincent; Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Guyana, Venezuela, Ecuador, Galapagos Is., Peru, Bolivia,

70



Brazil, Uruguay, Argentina, Chile; Nearctic and Palaearctic Regions; Eastern Pacific)

.......................................................................... 0. aenescens (Wiedemann, 1830) Mid femur with 2-3 long spinules on basal third; mid and hind femur yellow; pedicel and flagellomere yellow (Cuba) ........................................... O. cubana Gregor, 1974 4.

Mid femur, on basal half of ventral surface, with a row of 10-12 strong setae (Figs. 78-79) .................................................................................................................. 5 Mid femur, on ventral surface, with a row of 2-6 setae (two rows in O. ignava, Fig. 80) ....................................................................................................................... 6

5.

Wing brown on upper half; calypteres yellow with brown margins (Brazil)

............................................................................... 0. solita ria Albuquerque, 1958 Wings uniformly light brown; calypteres infuscated with dark brown margins (Brazil) ................................................................................ O. albuquerquei Lopes, 1985 6.

Hind tibia strongly curved, its anteroventral, ventral and posteroventral surfaces with many fine and long setae on basal third (Fig. 81) (?Venezuela, Chile; Nearctic and Palaearctic Regions; North-west Oriental) ............................ 'O. ignava (Harris, 1780) Hind tibia not curved, setae on its anteroventral, ventral and posteroventral surfaces not as above ......................................................................................................... 7

7.

Fore tarsomeres yellow on ventral surface; mid femur on ventral surface with 4-6 fine cilia basally (Fig. 82); hind tibia on anteroventral, ventral and posteroventral without long setae on apical two-thirds (Brazil, Chile; Nearctic, Afrotropical, Oriental and Australasian regions ......................................... O. chalcogaster (Wiedemann, 1824) Fore tarsomeres uniformly dark brown to black; mid femur on ventral surface with 2 short and strong setae basally (Fig. 83); hind tibia on anteroventral, ventral and posteroventral with a series of long setae on apical two-thirds (Fig. 84) (Chile; Nearctic, Palaearctic and Afrotropical Regions; North India) ...... O. capensis (Wiedemann, 1818)

8.

Ocellar triangle long, reaching lunule (Fig. 85), strongly sclerotized, reddish-brown


71


(Chile; Nearctic, Palaearctic and Afrotropical Regions; North India) ............... . ................................................................................. 0. capensis (Wiedemann, 1818) Ocellar triangle short to long, not strongly sclerotized, blackish ................................ 9

9.

Ocellar triangle long, reaching lunule or ending very close to it (Figs. 86,88) ......... 10 Ocellar triangle shorter, not reaching lunule and not ending close to it (Fig. 87) ............. 13

10. Palpusyellow........................ " ................................................................................ 11 Palpus dark brown to black .................................................................................. 12

11. Pedicel and flagellomere brown, ocellar triangle broad with a round apex (Fig. 86); calcar long; anterior katepisternal seta hair-like (Mexico, Nicaragua, St. Vincent, Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Argentina, Chile; Nearctic and western Palaearctic Regions; eastern Pacific ) .................................... .0. aenescens (Wiedemann, 1830) Pedicel and flagellomere yellow, ocellar triangle narrow; calcar short; anterior katepisternal seta well developed (Cuba) ............................... O' cubana Gregor, 1974 12. Wing brown on upper half; calypteres yellow with light brown margins (Brazil) ................................................................................ 0. solitaria Albuquerque, 1958 Wing uniformly light brown; calypteres brown with dark brown margins (Brazil)

................................................................................. 0. albuquerquei Lopes, 1985 13. Upper calypter uniformly yellowish; fore tarsi usually with tarsomeres yellow on ventral surface (not so evident as in male); hind tibia on anteroventral surface with one seta (Brazil, Chile; Nearctic, Afrotropical, Oriental and Australasian Regions)

...................................................................... 0. chalcogaster (Wiedemann, 1824) Upper calypter slightly brownish with dark brown margins; fore tarsi usually with tarsomeres uniformly brown; hind tibia on anteroventral surface with 2 setae (?Venezuela, Chile; Nearctic and Palaearctic Regions, northwestern Oriental) ......................................................................................... 0. ignava (Harris, 1780) 72



C::::i'~;;f1Imtr ) 78

c

~

~';1~

81

82

c·····,::·· . ,;·. ? 83

oc tr

85

\rmmNJ 86

87

88

Figs. 77-88, characters of Ophyra (modified from PAMPLONA & CO URI, 1989): 77, O. aenescens, male, trochanter and mid femur; 78, O. salitaria, male, mid femur; 79. O. albuquerquei. male. mid femur; 80, O. ignava. male, mid femur; 81. O. ignava, male, hind tibia; 82, O. chalcagaster, mid femur; 83, O. capensis, male, mid femur; 84, O. capensis, male, hind tibia; 85, O. capensis, female, ocellar triangle; 86, O. aenescens, female, ocellar triangle; 87, O. chalcagaster, female. ocellar triangle; 88, O. albuquerquei, female, ocellar triangle. Abbreviation: oc tr, ocellar triangle.


73


Potamia Robineau-Desvoidy, 1830

(Figs. 89-93) Type-species. Potamia littoralis Robineau-Desvoidy (des. SEGUY, 1937). Diagnosis. Gena with 1-3 differentiated and upward-curved anterior setae; arista with cilia about twice as long as the basal diameter of the arista; katepistemum bare; male: distiphallus with spicules but not strongly sclerotized (Fig. 91); female: fronto-orbital proclinate setae present. References. SNYDER, 1955; CARVALHO, 1981. Monophyly. Probably monophyletic; this genus is one ofthe most basal of the tribe, judging from the presence of a primitive configuration of the male distiphallus (CARVALHO, 1993a), with spicules (Fig. 91), but without the strong sclerotization as found in Micropo tam ia, Ophyra, Australophyra Malloch (Australian genus), Hydrotaea (Figs. 53-56) and in the monophyletic group formed by the genera Azelia and Thrichops. Geographical record. Mexico, Peru, United States of America. Comments. Only two species occur in the Neotropical Region, both in Peru. Key to species

1. Male: cercal plate, in posterior view, without a cleft on posterior margin (Fig. 90); surstylus, in lateral view, as Fig. 89 (Peru) ..................... P plumata (Carvalho, 1981) Male: cercal plate, in posterior view, with a cleft on posterior margin (Fig. 92); surstylus, in lateral view, as Fig. 93 (Mexico, Peru; Nearctic Region) ... ... P scabra (Giglio-Tos, 1893)

Thricops Rondani, 1856

Type-species. Anthomyza hirtula Zetterstedt (orig. des.). Diagnosis. General coloration yellow; more than one inclined frontal setae present; vein Al extending more than halfway to wing margin; hind coxa with hairs on posterior apical margin; hind tibia with one posterodorsal setae. References. HUCKETT, 1975; VOCKEROTH, 1996.

74



apoede

89 90

92

93

Figs. 89-93, characters of Potamia (modified from CARVALHO, 1981; SNYDER, 1955): 89, P. plumata, epandrium, cereal plate and surstyli, lateral view; 90, P. plumata, epandrium, cereal plate and surstyli, posterior view; 91, P. plumata, aedeagus, lateral view; 92, P. scabra, epandrium, cerci and surstyli, posterior view; 93, P. scabra, epandrium, cereal plate and surstyli, lateral view. Abbreviations: apo ede, aedeagal apodeme; dis, distiphallus; epi, epiphallus; gon, gonopod.

Monophyly. Not proposed yet. Geographical record. Mexico; Nearctic, Palaearctic and Oriental regions. Comments. This genus, represented by Thricops diaphanus (Wiedemann, 1817) was reported for the first time in the Neotropical Region (Oaxaca, Mexico) by VOCKEROTH (1996). The male of this species can be recognised by a pronounced strong apical thorn on the ventral surfaces of hind tibia (HUCKETT, 1975).


75


Tribe Reinwardtiini Brauer & Bergenstamm, 1889 Diagnosis. Hind femur, on posterodorsal surface, usually with a developed calcar; haustellum with a ciliated sclerotization; male: distiphallus membranous; female: frontoorbital usually absent; interfrontal setae absent. Monophyly. This tribe is probably not monophyletic (CARVALHO, 1989d), being the genera assembled on basis of primitive characters in relation to the ground plan of the Azeliinae. In the cladistic analysis of the family, CARVALHO (1989d) found only one synapomorphic character for this tribe - haustellum with a posterior ciliated sclerotization, but this character is homoplastic, occurring in many muscid genera. According to PONT (1986a, 1989), this tribe includes the most primitive genera of the family. Comments. A cosmopolitan tribe, with 16 genera, 11 of the them occurring in the Neotropical Region (CARVALHO etaZ., 1993). SKIDMORE (1985) called attention to the high number of Reinwardtiini genera occurring in the Neotropical Region. There are 85 species recorded from the neotropics.

Brachygasterina Macquart, 1851 (Figs. 94-95) Type-species. Brachygasterina vioZaceiventris Macquart (orig. des.). Diagnosis. Male holoptic, eyes with cilia; arista bare; flagellomere strongly dilated (Fig. 94-95); palpus dilated; interfrontal setae present or absent in female; presutural acrostichal developed; prealar seta present or absent; dorsocentrals 2:4; katepimeron bare; katepisternals 1: 1; R node with or without cilia; calcar present and strong; veins R4+5 and M parallel apically; ovipositor long, tergite 6 anchor-shaped, tergite 7 a little larger, sternites 6 and 7 divided posteriorly. Reference. CARVALHO, 1989b. Monophyly. Genus monophyletic, having the strongly dilated antenna as an autapomorphy (CARVALHO, 1989b). Geographical record. Chile, Argentina. Comments. The genus has only three species endemic to southern South America.

76



Key to species (modified from CARVALHO, 1989b) 1.

General coloration metallic violet-blue; humeral callus yellowish; calypteres dark brown; interfrontal setae present in female (Chile) ......... .B.fulvohumeralis (Malloch, 1922) General coloration violet-blue; calypteres brownish with dark brown margins; interfrontal setae absent in female .......................................................................... .2

2.

Wing with Rs node bare on both surfaces; hind tibia on anterodorsal and anteroventral surfaces with at most 3 setae (frequently 2); prealar seta similar in length or much longer than notopleural anterior seta; sternite 1 bare; male: hind femur on posteroventral surface with long setae on apical half, shorter on basal half; female: fore tibia on dorsal surface without setae (Chile, Argentina) .............. .B. violaceiventris Macquart, 1851 Wing with Rs node with cilia on both surfaces; hind tibia on anterodorsal and anteroventral surfaces with at least 4 and 3 setae respectively; prealar seta absent in both sexes; sternite 1 setulose; male: hind femur on posteroventral surface bare on apical half and with longer setae on basal half; female: fore tibia on dorsal surface with 4 short setae on apical half (Chile, Argentina) ..................... .B. major Malloch, 1934

94

95

Figs. 94-95, characters of Brachygasterina (modified from CARVALHO, 1989b) 94, B. major, antenna, lateral view; 95, antenna, lateral view, B. violaceiventris; Abbreviations: ar, arista; ped, pedicel.


77


Chaetagenia Malloch, 1928 (Fig. 96) Type-species. Chaetagenia stigmatica Malloch (orig. des.). Diagnosis. Male dichoptic, eyes bare; facial ridges with short cilic" arista plumose, with long cilia on upper surface and short on lower; female with; directed setae on head; prealar seta present, moderately developed; scutellur;: on its lateral margins; prostemum with few setulae laterally; katepistemal posterior upper one, about twice the length of the others; metathoracic spiraclt: on its lower margin; anepimeron setulose; katepimeron with short and postalar wall setulose; lower calypter about twice as long as the upper one; on apical portion of stem vein and on veins R4+5 and CuA!; hind coxa with Cil::~lC)r surface; calcar absent; stemite 1 setulose. Reference. COURI & LAMAS, 1993. Monophyly. The bifid configuration ofthe distiphallus is very characl:(;":-l:siic and considered here as an apomorphy. This kind of aedeagus is also found in Pseudopri;o2epis Snyder, 1949 (LOPES & CARVALHO, 1985). Geographical record. Brazil. Comments. Monobasic genus: Chaetagenia stigmatica Malloch, 1928. COURI & LAMAS (1993) synonymized the other four described species with the type-species.

Fig. 96, Chaetagenia stigmata, head, lateral view

78

Muscidae (Diptera) of the Neotropicai Region: TAXONOMY


Correntosia Malloch, 1934 Type-species. Correntosia bicolor Malloch (orig. des.). Diagnosis. Male dichoptic, eyes bare; arista pubescent, with cilia shorter than the basal width of the arista; flagellomere weakly dilated apically; interfrontal setae present in male; presutural acrostichals developed; prealar seta strong; wing veins bare; calcar present; abdomen elongated, longer than thorax length; ovipositor long, tergite 6 anchortype; tergite 7 a little larger than 6; stemite 8 entire; microtrichia absent. References. CARVALHO, 1989b; PONT, 2001. Geographical record. Argentina. Monophyly. Correntosia is close related to Palpibracus Rondani, but the presence of interfrontal setae in males is a rare character among the Muscidae, and it is not found in Palpibracus. Comments. This genus was for a long time monobasic (Correntosia bicolor Malloch, 1934), but recently PONT (2001) transferred Psilochaeta camifex Stein, 1911 to this genus. Differently from the type-species, C. camifex has orange antennal scape and pedicel, and yellow palpus (PONT, 2001).

Dalcyella Carvalho, 1989 Type-species. Helina veniseta Dodge (orig. des.). Diagnosis. Male holoptic, eyes with cilia; arista pubescent, with cilia shorter than basal width of the arista; flagellomere weakly dilated apically; interfrontal setae present in female; pre sutural acrostichals developed; prealar seta strong; anepimeron, katepimeron and meron setulose; vein R 4+5 with 2-3 cilia on both surfaces; calcar strong; abdomen oval, shorter than thorax length; ovipositor long, tergite 6 anchor-type; tergite 7 narrow; stemites 6 and 7 divided posteriorly; stemite 8 developed over its whole extension. Reference. CARVALHO, 1989c. Monophyly. DODGE (1967) described the type-species in Helina RobineauDesvoidy, because he could not relate it to any other muscid genera. Many species which would not conveniently fit Helina, have been assigned to it, and consequently this genus, as well other muscid genera (e.g. Phaonia), has been a depository of poorly studied species (SKIDMORE, 1985). Nowadays Helina is seen as an aggregate of distinct genera


79


(HUCKETT & VOCKEROTH, 1987). CARVALHO (1989c) erected Dalcyella for H. veniseta but no autapomorphic character was pointed out. Geographical record. Chile. Comments. Monobasic genus: Dalcyella veniseta (Dodge, 1967).

Itatingamyia Albuquerque, 1979 (Figs. 97-98) Type-species. Itatingamyia bivitatta Albuquerque (orig. des.). Diagnosis. Male dichoptic; antennal axis longer than oral axis; arista with short cilia, longest cilia shorter than twice of the basal width of pedicel; prealar and anterior intra-alar setae absent; dorsocentral setae 2: 3; acrostichals 0: 1, ciliform; pro sternum bare; anepimeron with 2-3 setae; katepisternals 1:2; posterior spiracle with cilia on margin; lower calypter glossiform, about as long as upper one; wing veins bare; veins R 4+s and M divergent at apex prosternum bare; sternite 1 setulose; female: ovipositor with cerci surpassing the hypoproct; tergite 6 anchor-type (Fig. 97), sternite 8 developed all its extension (Fig. 98). References. ALBUQUERQUE, 1979; CARVALHO & COURI, 1993. Monophyly. ALBUQUERQUE (1979) included the genus among the Mydaeinae, based mainly on the ovipositor shape. CARVALHO & COURI (1993), based on the shape of the female cerci (very long, surpassing the hypoproct), the development of sternite 8 and the shape on sternite 6, transferred the genus to Reinwardtiini, in the Azeliinae. Geographical record. Brazil. Comment. Monobasic genus: Itatingamyia bivitatta Albuquerque, 1979.

Muscina Robineau-Desvoidy, 1830 Type-species. Musca stabulans Fallen (des. COQUILLET, 1910). Diagnosis. Male holoptic, eyes bare; arista plumose with cilia on basal threefourths and bare on the apical fourth; interfrontal setae present in female; anterior intra-alar postsutural seta placed posteriorly to level of supra-alar seta; presutural acrostichals developed; prealar seta short, but distinct; anepimeron bare; meron setulose; wing with

80



apical section of vein M slightly curved forward, veins bare; calcar strong, submedian; sternite 1 setulose. Reference. SNYDER, 1955. Monophyly. HENNIG (1965) had indicated the affinity of this genus with the Hydrotaeini based on the morphology of the ovipositor, while SKIDMORE (1985) placed this genus close to Synthesiomyia andPasseromyia based on characters of the immature stages. Geographical record. Mexico, Costa Rica, Guatemala, Peru, Venezuela, Brazil, Uruguay, Argentina, Chile; Nearctic and Palaearctic Regions. Comments. In the Neotropical Region, induding Mexico, there are four species, two of them occurring in South America.

Key to species 1.

Flagellomere dark brown ....................................................................................... 2 Flagellomere entirely or partly yellow...................................................................... 3

2.

Pedicel reddish-brown (Mexico; Nearctic and Palaearctic Regions) ........................ .

....................... .................. ................ ......... ................ ..... .M. levida (Harris, 1780) Apex of pedicel rather reddish; flagellomere lighter at base of pedicel (cosmopolitan)

.............................................................................. .....M. stabulans (Fallen, 1817) 3.

Pedicel and the longitudinal posterior half of flagellomere yellow; abdomen with greyish pruinescence (Mexico; Nearctic Region) ........................ M. dorsilinea (Wulp, 1896) Pedicel reddish and flagellomere rather reddish; abdomen reddish yellow translucent, last segment darker (South America) ........................ .M. principalis (Schiner, 1868)


81


0,4mm 0,4 mm

97

98

Figs. 97-98, characters of ltatingamyia (modified from CARVALHO & COURI, 1993): 97, l. bivitatta, ovipositor, dorsal view; 98, I. bivitatta, ovipositor, ventral view.

Palpibracus Rondani, 1863 (Plate 1) Type-species. BrachypaZpus pilosus Macquart (aut.). Diagnosis. Male holoptic, eyes with cilia; arista pubescent, with cilia shorter than basal width of the arista; flagellomere weakly dilated apically or not; interfrontal setae present or absent in female; pre sutural acrostichals developed; prealar seta strong; katepimeron setulose or bare; wing veins bare; calcar present; abdomen oval, shorter than 82



thorax length; ovipositor long, tergite 6 anchor-type; tergite 7 thin (except in P. veneris); sternites 6 and 7 divided posteriorly (except in P. lancifer); sternite 8 developed over its whole extension. Reference. CARVALHO, 1989a. Monophyly. Probably monophyletic. CARVALHO (1989a) did not indicate putative synapomorphic characters for this genus, but pointed out its uniqueness among the muscids. Geographical record. Chile, Argentina. Comments. The species of Palpibracus have a very restricted distribution, occurring only in Chile and Argentina, in the Andes, southern of 30° S (CARVALHO, 1989a). LOPES & KHOURI (1996) described Palpibracus carvalhoi based on material from Chile. The type-material was not examined but, based on the original description, this species seems to be very close morphologically to P. univittatus (Bigot, 1857). PAMPLONA & COURI (2000) described P. valdiviensis from Valdivia, Chile, based only in female specimens. Based on their description it seems that it is close to P. apicalis (Malloch, 1934). Both species, P. carvalhoi and P. valdiviensis were not included in the key below.

Key to species (modified from CARVALHO, 1989a) 1.

Katepimeron setulose on posterior margin, with black cilia (sometimes yellow) in front of posterior spiracle ....................................................................................... 2 Katepimeron bare ................................................................................................ 8

2.

Antenna and palpus dark brown; mid tibia on posteroventral surface bare; male: head width similar to its height; eyes with anterointemal facets very enlarged; frontal setae limited to the anterior half offrons (Chile, Argentina) ............ P. veneris (Bigot, 1888) Antenna and palpus at least partly yellow; mid tibia on posteroventral surface with one or more setae; male: head wider than high; eyes with anterointernal facets not very enlarged; frontal setae over entire frons, those on upper frons weaker.. ............. 3

3.

Male: fore coxa without a strong apical spur; fore tibia on anteroventral surface with 1


83


seta on apical third; hind tibia on posteroventral surface with a series of long setae; female: interfrontal setae absent; mid tibia on posteroventral surface with only 1 submedian seta (Chile ).................................................. F separatus (Malloch, 1934) Male: fore coxa with a strong apical spur; fore tibia on anteroventral surface with 2-3 setae on apical third; hind tibia on posteroventral surface bare, with small cilia on basal or apical half, never over its whole extension; female: interfrontal setae present; mid tibia on posteroventral surface with 1-3 submedian setae ................................ .4 4.

Male: parafaciallarge, its width, at the level of pedicel, similar to the width of gena at the lower margin of eyes; mid femur with no depression; female: frontal vitta totally reddish; fore tibia on anteroventral surface with 1-2 submedian seta (Chile, Argentina)

...................................................................... .............. F reynoldsi (Malloch, 1934) Male: parafacial, at the level of pedicel, about two-thirds as wide as gena at the lower margin of eyes; mid femur concave, anterior surface with a strong or faint median depression; female: frontal vitta dark brown sometimes with the anterior half reddish; fore tibia on anteroventral surface with only one submedian seta ...................... '" ...... 5 5.

Fore coxa entirely black; male: 10-11 pairs of frontal setae; mid femur on anterior surface without short spines; hind tibia on posterodorsal surface with 5 setae, the calcar submedian; female: frontal vitta dark brown (Chile, Argentina) ....................... .

....................................................................................... F similis (Malloch, 1934) Fore coxa yellow or at most dark brown on apical half; male: at most, 9 pairs of frontal setae (maximum); mid femur on anterior surface with 1-2 series of short spines apically; hind tibia on posterodorsal surface with 3-4 setae, the calcar inserted on the apical third; female: frontal vitta partly reddish ........................................................ 6 6.

Male: mid and hind femora yellowish; mid femur, on the median depression, without ground setulae; female: hind femur yellow, except for a black preapical ring; hind tibia on anteroventral surface with a series of 4-5 setae on apical half; posterodorsal surface with 2 setae on middle third (Chile) ................................... F chilensis (Bigot, 1885)

84



Male: mid and hind femora partly dark brown; mid femur, on the median depression, with the usual ground setulae; female: hind femur yellow on the basal third and black on the apical two-thirds; hind tibia on anteroventral surface with a series of 5-6 setae on apical half; posterodorsal surface with 2-3 setae on middle third ......................... 7 7.

Male: fore tibia on anteroventral surface with 2 setae on apical half; ventral surface with 3-4 weak setae on apical third; posterodorsal surface with 0 or 2 setae on apical fifth; hind femur yellow, black on apical two-thirds; hind tibia black, anteroventral surface with a series of weak setae over its whole extension, longer toward the apex; posteroventral surface with cilia only on apical third, posterodorsal surface with 3-4 short setae evenly spaced along tibial length; mid tibia on posteroventral surface with 1-3 setae on apical half (Chile) ...................................... .l~ confusus (Malloch, 1928) Male: fore tibia on anteroventral surface with a series of very weak setae on apical half; posterodorsal surface with 1 median seta; hind femur yellow with a black ring on middle third; hind tibia yellowish; anteroventral surface with a series of weak setae along its whole length; posteroventral and ventral surfaces with cilia on apical half, longer apically; posterodorsal surface with 3 setae on middle third; female: unknown (Chile, Argentina) ............................................ ............P. pilosus (Macquart, 1851)

8.

General coloration metallic blue; abdomen, at most with tergite 5 partly orange yellow................................................................................................................... 9 General coloration yellow to black, thorax, at least, partly yellow; abdomen never metallic blue ........................................................................................................... 10

9.

Eyes with long and conspicuous cilia; legs black with tarsi partly yellowish white (more evident on fore leg of male); abdomen with the apical half of tergite 5 orange yellow; male: 12-16 pairs of frontal setae; fore coxa without an apical spur; fore tibia on anterior and anteroventral surfaces with fine, long cilia along its whole length; mid tibia on posterior surface with 4-5 setae on basal two-thirds; female: fore tibia on posterodorsal surface bare; mid femur on posteroventral surface with a series of setae


85


on basal half (Chile, Argentina) ......................................... P. apicalis (Malloch, 1934) Eyes with short and sparse cilia; legs yellow, except for coxa, trochanter and tarsi black; abdomen entirely metallic violet blue; male: 8 pairs of frontal setae; fore coxa with a strong apical spur; fore tibia on posterior surface with long and strong setae along its length; female: fore tibia on posterodorsal surface with 2 short setae inserted at the base of the apical third; mid femur on posteroventral surface with one strong seta on basal third (Chile, Argentina) ............................... P. lancifer (Malloch, 1934) 10. Male: mesonotum yellow with a black median vitta; fore coxa with a strong apical spur; female: interfrontal setae present or absent; hind tibia on anteroventral surface with 3-5 setae on apical half. ................................................................................ 11 Male: mesonotum yellow with a black median vitta or black with grey pruinose vittae; fore coxa bare; female: interfrontal setae present; hind tibia on anteroventral surface with 2-3 setae on apical half.. ................................................................................ 13 11. Antenna dark brown, with scape, pedicel and base of flagellomere yellow; palpus yellow; male: hind tibia with a median curvature; anteroventral surface with a series of short setae, shorter than the width of tibia, over its whole length; posteroventral surface with a row of cilia on its whole extension, longer on apical fifth; female: frontal vitta yellow, a little darker posteriorly (Chile ) ........................... P. univittatus (Bigot, 1857) Antenna dark brown, pedicel a little lighter; palpus dark brown; male: hind tibia with or without a curvature; anteroventral surface with a series of long setae, longer than the width of tibia, over its whole length; posteroventral surface with a series of long and fine setae on apical two-thirds; female: frontal vitta dark brown, reddish in middle ornot. .................................................................................................................. 12 12. Gena orange yellow; mesonotum yellow with one narrow median vitta; male: mid tibia on posterodorsal surface with 2-4 setae evenly spaced along tibial length; hind tibia with no median curvature; female: interfrontal setae absent (Chile, Argentina)

.................................................................................... P. spicatus (Malloch, 1934) 86


2


Gena dark brown; mesonotum yellow with one wide median vitta; male: mid tibia on posterodorsal surface with 4-6 setae evenly spaced along tibial length; hind tibia with a median curvature; female: interfrontal setae present (Chile)........... P. albuquerquei Carvalho, 1989 13. Palpus yellow with a brown base; mesonotum black with 3 grey pruinose vittae and small yellow areas; male: antenna inserted at level above the middle of eyes; female: abdomen dark yellow (Chile, Argentina) ..................... P. peruvianus (Malloch, 1929) Palpus dark brown; mesonotum yellow with 1-3 vittae, sometimes fused; male: antenna inserted at a level below the middle of eyes; female: abdomen dark brown .............. 14 14. Male: eyes with anterointernal facets strongly enlarged; fore tibia on anteroventral surface with 1 seta on middle third; ventral surface with no cilia on apical third; posteroventral surface with 1-2 setae on middle third; female: frontal vitta dark brown with only the anterior half reddish (Chile, Argentina) ..... P. nigriventris (Malloch, 1928) Male: eyes with anterointernal facets enlarged; fore tibia on anteroventral surface with 2-3 setae on apical third; ventral surface with 1-4 long cilia on apical third; posteroventral surface with 2-3 setae on middle third; female: frontal vitta reddish on almost all extension ................................................................................................ 15 15. Mesonotum yellow with a large black vitta (sometimes narrow in female); male: fore tibia on ventral surface with 4 long cilia on apical third; hind tibia with a median curvature, posteroventral surface with a series of fine cilia on middle third; female: mid tibia on ventral surface with one submedian seta; hind femur strongly curved and slightly enlarged preapically (Chile, Argentina) ......................................P.fasciculatus (Malloch, 1934) Mesonotum yellow with 3 black vittae (sometimes fused); male: fore tibia on ventral surface with 1-2 long median cilia; hind tibia without a median curvature, posteroventral surface with a series of strong cilia, absent on the basal two-thirds; female: mid tibia on ventral surface with no setae; hind femur only weakly curvature and not enlarged preapically (Chile )...................................................... P. trivittatus (Malloch, 1934)


87


Philornis Meinert, 1890 (Figs. 99-108, Plate 2) Type-species. Philornis molesta Meinert (mon.). Diagnosis. Male holoptic (Fig. 100) or intermediate (Fig. 102); palpus a little enlarged apically; arista with long plumae; mesonotum usually with four brown vittae; anepimeron hairy; postalar wall with a tuft of cilia; posterior spiracle with cilia; dorsocentral setae 2:4; calcar absent; anal vein long; veins R 4+5 and M divergent, the latter a weakly bent forward apically (Fig. 99). References. COURI, 1989, 1999b. Monophyly. Genus certainly monophyletic having as sister-group Passeromyia with occurrence on the Afrotropical, Oriental and Australian Regions (COURI & CARVALHO, 2000). Several authors (e.g. PONT, 1974; SKIDMORE, 1985) had already called attention to the similarities among the two genera. Geographical record. Mexico, Costa Rica, Panama, Cuba, Dominican Republic, Trinidad, Puerto Rico, Venezuela, Ecuador, Peru, Brazil, Chile, Uruguay, Argentina; United States of America. Comments. P. cinnamomina (Stein), P. convexa (Stein), P. mima (Townsend), P. nielseni Dodge, P. obscura (Wulp), P. umanani Garcia, P. steini (Pont) and P. molesta Meinert were not included in the key, since we had access only to the data from the literature, which did not provide decisive species characters. Also, the last mentioned species is known only from one larval phase. Philornis species are largely distributed in the Neotropical Region, with three species known from the U.S.A., P. angustifrons (Loew, 1861), P. mimicola Dodge, 1968, and P. porteri Dodge, 1955, the latter known only from Florida (COURI, 1999b).

Key to species (modified from CO URI, 1999b) 1.

Vein R4+5 and/or CuA 1 with cilia on ventral surface and/or dorsal surface (Fig. 99); male: eyes approximated (Fig. 100); female: inner margin eye parallel (Fig. 10 1).......... 2 Veins bare; male: distance between eye greater than diameter of anterior ocellus (Fig. 102); female: inner eye margin converging above (Fig. 103) ....................................... 10

88



2.

Gena yellow-haired ............................................ ,................................................. 3 Gena black-haired ................................................................................................ 4

3.

Wing slightly brown, calypteres uniforrnlybrown (Costa Rica) ........P schildiDodge, 1963 Wing hyaline, calypteres light yellow, except for brown margin of upper calypter (Brazil)

.................................................................................. ...... P setinervis Dodge, 1963 4.

Intra-alar setae 1:3 (Panama) ...... ................................ P rettenmeyeri Dodge, 1963 Intra-alar setae 1: 1 or 1:2 ........................................................................................ 5

5.

Mid femur on posterior surface with three preapical setae (Brazil) ........................ ..

...................................................................................... P amazonensis Couri, 1983 Mid femur on posterior surface with two preapical setae ......................................... 6 6.

Vein R4+5 with cilia only on ventral surface ................................................................ 7 Vein R 4+5 with cilia both on dorsal and ventral surfaces ........................................... 9

7.

Supra-alar setae 1:5; intrapostalar seta as long as posterior intra-alar one (Brazil)

..................................... .......................................................... P lopesi Couri, 1983 Supra-alar setae 1:3; intrapostalar seta shorter than posterior intra-alar one .............. 8 8.

Vein CuA! bare; vein R 4+5 on ventral surface with 2-3 cilia (Trinidad, Peru, Brazil)

.............................................................................................P aitkeni Dodge, 1963 Vein CuA! on ventral surface with 2 cilia; vein R 4+5 on ventral surface with 7 cilia (Brazil) .................................................. ..................... P rufoscutellaris Couri, 1983 9.

Vein R4+5 with 2 cilia on dorsal surface and 5 on ventral surface; vein CuA! on ventral surface bare (Mexico) ................................................. Pfasciventris (Wulp, 1896) Vein R 4+5 with 1-3 cilia on dorsal surface and 3-4 on ventral surface; vein CuA l on ventral surface with at least one cilia (Panama) ...................... .P zeketi Dodge, 1963

10. Gena black-haired ................................................................................................. 11 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

89


Gena yellow-haired .............................................................................................. 15 11. Mid femur on posterior surface with 3 preapical setae (Trinidad, Brazil)

......................................................................... .... P downsi Dodge & Aitken, 1968 Mid femur on posterior surface with 2 preapical setae ......................................... 12 12. Hind tibia on anterodorsal surface with 2 setae on middle third; flagellomere dark brown (Argentina) ....................................................... ... P blanchardi Garcia, 1952 Hind tibia on anterodorsal surface with 1 setae on middle third; flagellomere yellow.... 13 13. Legs brown; tergite 5 reddish (United States of America.) ......P porteriDodge, 1955 Legs yellow with brown tarsi; tergite 5 brown with gray pruinescence....................... 14 14. Intrapostalar seta long, approximately 0.80 as long as posterior intra-alar one (Fig. 104); basicosta and epaulet brown (Guatemala, Panama, Peru, Brazil; United States of America) .................................................................... P mimicola Dodge, 1968 Intrapostalar seta ciliform; basicosta and epaulet yellow (Panama, Trinidad)

............................................................................... P nigra Dodge & Aitken, 1968 15. Hind tibia on anterodorsal surface no median setae or at most one seta.................. .16 Hind tibia on anterodorsal surface with more than one seta .................................. .19 16. Costal margin of wing with a brown elongated spot extending from middle of wing to apex (Fig. 105) (Peru, Brazil) ........................................... Pfomicosta Dodge, 1968 Wing hyaline ........................................................................................................ 17 17. Thorax light brown with one median brown vitae (Brazil) .......P univittata Dodge, 1968 Thorax brown with gray pruinescence and with four brown vittae .......................... .18 18. Mid tibia on posterior surface with 3 setae; intrapostalar seta ciliform; calypter light brown (Costa Rica, Panama, Trinidad, Ecuador, Brazil) .......P. falsifica Dodge & Aitken, 1968 Mid tibia on posterior surface with 2 setae; intrapostalar seta developed; calypter 90



white (Brazil) .................................................... ................. P. petersoni Couri, 1984 19. Wing with dark brown spots on crossveins r-m and dm-cu (well marked in P. masoni, Fig. 106) .............................................................................................................. 20 Wing without brown spots on crossveins r-m and dm-cu ..................................... .21 20. Scutellum dark brown on sides, with extreme apex yellow; upper calypter with brown margin; fore tibia on dorsal surface with setae only at apex (Uruguay) ................ ..

...............................................................................................P. lnasoni Couri, 1986 Scutellum entirely brown; upper calypter yellowish; fore tibia on dorsal surface with 3-4 short setae on middle third (Argentina) .............................. P. seguyi Garcia, 1952 21. Wing light brown with brown clouds around longitudinal veins (Fig. 107) ............. 22 Wing without clouds, sometimes crossvein r-m surrounded by a brown spot.. ........ 25 22. Mid femur on posterior surface with 3 preapical setae; mid tibia on posterior surface with more than 3 setae on middle third (Mexico, Ecuador, Brazil) ............................. ..

............ ...............................................................................P. grandis Couri, 1984 Mid femur on posterior surface with 2 preapical setae; mid tibia on posterior surface with 1-3 setae on middle third .............................................................................. 23 23. Distance between eyes, at level of anterior ocellus, 0.20-0.22 of head width in male and 0.32-0.34 in female (Brazil) ................................ P. sabroskyi Albuquerque, 1957 Distance between eyes, at level of anterior ocellus, 0.06-0.09 of head width in male and 0.19-0.23 in female ....................................................................................... 24 24. Hind tibia on anterodorsal surface with 3 setae; calypteres white, except for brown margin of upper calypter (Brazil) ................................... P. obscurinervis Couri, 1984 Hind tibia on anterodorsal surface with 4 setae; calypteres light brown, margin of upper calypter dark brown and of lower one yellow (Costa Rica)

........................................................................................P. carinata Dodge, 1968 25. Hind tibia on anterodorsal surface with more than 5 setae .................................... .26 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

91


Hind tibia on anterodorsal surface with at most 4 setae ............................................ 27 26. Legs yellow with brown tarsi in both sexes; calypteres light brown, margin of upper calypter brown; male: hind tibia not bowed (Mexico) .... ............... P bella Couri, 1984 Legs yellow in male and brown in female; calypteres white, margin of upper calypter brown; male: hind tibia distinctly bowed (Fig. 108) (Brazil, Argentina) .................. .

....... .............. ........... ........ ..... ...... ........ .............. ............... P torquans Nielsen, 1913 27. Distance between eyes, at level of anterior ocellus, of 0.03-0.05 of head width in males and of 0.09-0.14 in female ......................................................................... 28 Distance between eyes, at level of anterior ocellus, more than 0.05 if head width ........ 30 28. Basicosta brown; tergites 4 and 5 paler than thorax (Trinidad, Panama) ................ ..

...................................................... .............. ......... P querula Dodge & Aitken, 1968 Basicosta yellow; tergites 4 and 5 concolour with thorax ....................................... 29 29. Legs yellow in both sexes; mid femur on posterior surface with 3 preapical setae (Cuba, Costa Rica, Trinidad, Guyana, Brazil; United States of America) ..........

.................................................................................... P angustifrons Loew, 1861 Legs yellow in male and brown in female; mid femur on posterior surface with 2 pre apical setae (Trinidad, Brazil) ............................ .P. glaucinis Dodge & Aitken, 1968 30. Basicosta brown .................................................................................................... 31 Basicosta yellow.................................................................................................. 35 31. Wing without spots ............................................................................................. 32 Wing with brown spot at crossvein r-m ............................................................... 33 32. Abdomen dark brown with grey pruinescence; frons red at lunule; male: distance between eyes, at level of anterior ocellus, 0.10-0.12 of head width (Puerto Rico, Trinidad, Venezuela, Brazil) .............................. .P. deceptiva Dodge & Aitken, 1968 Abdomen brown with grey pruinescence, entire tergite 1+2 and basal half oftergite 3 92



yellow; frons brown; male: distance between eyes, at level of anterior ocellus, 0.060.07 of head width (Costa Rica, Panama, Cuba, Dominican Republic, Puerto Rico, Trinidad, Venezuela, Brazil, Argentina) ............................... .P pici (Macquart, 1854) 33. Thorax light brown with yellow pruinescence; mid tibia on posterior surface with 3 setae (Trinidad, Brazil)....................................... P sanguinis Dodge & Aitken, 1968 Thorax brown with grey pruinescence; mid tibia on posterior surface with 2 setae ...... 34 34. Katepistemum with black cilia; hind tibia on anterodorsal surface with 2 setae (Brazil)

........................... ................................................................. P vulgaris Couri, 1984 Katepistemum with yellow cilia; hind tibia on anterodorsal surface with 3 setae (Brazil) ........................................................................................ p mediana Couri, 1984

35. Wing with black spot at crossvein r-m ................................................................... 36 Wing without a spot at crossvein r-m .................................................................. 38 36. Supra-alar setae 1:5; frons of both sexes with margins strongly diverging toward lunule (Brazil) .............................................. ................................. Pfrontalis Couri, 1984 Supra-alar setae 1:3; frons of both sexes with margins slightly diverging toward lunule .................................................................................................................. 37 37. Calypteres light brown, margin of upper one dark brown (Peru) .................. .

.................................................................................. .. P vespidicola Dodge, 1968 Calypteres white, margin of upper one brown (Peru) ...... .P albuquerquei Couri, 1983 38. Mid tibia on posterior surface with 3 setae; legs of both sexes yellow...................... 39 Mid tibia on posterior surface with 2 setae; legs of male yellow and of female brown (male of P spermophila unknown) ...................................................................... .40 39. Calypteres white, margin of upper one brown, greater ampulla brown, darker in male (Brazil) .............................................. ................................. P diminuta Couri, 1984 Calypteres white, greater ampulla yellow (Brazil) ...................P insularis Couri, 1983 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

93


40. Flagellomere brown (Trinidad, Brazil) .................P. trinitensis Dodge & Aitken, 1968 Flagellomere yellow............................................................................................. 41 41. Hind femur on anteroventral surface with 3 strong setae on apical third; hind tibia on anterodorsal surface with 3-4 setae (Trinidad) ......................... P. gagnei Couri, 1983 Hind femur on anteroventral surface with 4 strong setae on apical third; hind tibia on anterodorsal surface with 2 setae (Jamaica, Trinidad) .................. . ..........................................................................P. spermophila (Townsend, 1895)

1 mrn

1mm

Figs. 99-103. characters of Philornis (modified from co URI. 1999b): 99. P. aitkeni. wing. dorsal view; 100. P. aitkeni. male. head. frontal view; 101. P. aitkeni. female. head. frontal view; 102. P. downsi. male. head. frontal view; 103. P. dOlVnsi. female, head, frontal view.

94



1 mm

1 mm

1 mm

r-m

1 mm

Figs. 104-108, characters of Phi/omis (modified from COURI, 1999b): 104, P. mimicola, mesonotum, dorsal view; 105, P. fumicosta, wing, dorsal view; 106, P. masoni, wing, dorsal view; 107, P. grandis, wing, dorsal view; 108, P. torquans, hind leg, lateral view. Abbreviations: ipal, intrapostalar seta; r-m, crossvein; dm-cu, crossvein.


95


Psilochaeta Stein, 1911 (Plate 2) Type-species. Limnophora chlorogaster Bigot (orig. des.) = chlorogaster Wiedemann. Diagnosis. General coloration metallic violet blue, sometimes with greyish abdomen (P. chlorogaster); male holoptic; eyes with cilia; arista pubescent, with cilia smaller than the basal width of the arista; flagellomere slightly dilated apically; pre sutural acrostichal developed; prealar seta strong; katepimeron with few setulae; mesonotum with 3 dark vittae on dorsal surface; dorsocentral setae 2:3-4; acrostichals 2-3:3-6 but not forming lines; humerals 3; katepisternals 1-2:3-4; wing veins bare; calcar present; sternite 1 bare; female: ovipositor long; tergite 6 anchor-type; tergite 7 narrow; sternites 6 and 7 divided posteriorly. Reference. CARVALHO, 1989b. Monophyly. Psilochaeta was always associated to Brachygasterina Macquart, probably because of the same metallic coloration and, in part for their partly overlapping geographical distribution. CARVALHO (1989c) did not find autapomorphic characters for the genus. Geographical record. Colombia, Peru, Ecuador, Bolivia, Brazil, Uruguay, Argentina, Chile, Juan Fernandez Is. Comments. Among the endemic genera of Reinwardtiini in the southern South America, Psilochaeta has the largest known geographical distribution, with some important species for the antropobiocenosis (CARVALHO, 1989c). Psilochaetafulvolateralis Stein was not included in the key, since we could not exam its type. The holotype is probably deposited atSMT (PAPAVERO, 1971; CARVALHOetal., 1993). Psilochaeta carnifex Stein, 1911 (Chile) was recently transferred to Correntosia by PONT (2001).

Key to species (modified from CARVALHO, 1989c) 1.

Thorax dark brown with greyish pruinescence, with 4 brown vittae; abdomen metallic greenish, prealar seta shorter than the anterior notopleural seta; female: ovipositor longer than abdomen length (peru, Ecuador, Bolivia, Brazil, Uruguay, Chile, Argentina)

.........................................................................P. chlorogaster (Wiedemann, 1830) 96


•


Thorax metallic violet blue, with 2 or 4 dark brown vitta:e; abdomen metallic blue to greenish blue; prealar seta similar to the anterior notoplemal in length; female: ovipositor shorter than abdomen length ................................................................................... 2 2.

Thorax with 4 dark brown vittae; notopleural setulae sparse; veins R4+5 and M parallel apically; male: distance between eyes shorter than flagellomere diameter; female: parafacial narrow, about half as wide as height of gena at lower margin of eyes (Brazil, Chile, Argentina) ................................... P. pampiana (Shannon & Del Ponte, 1926) Thorax with 2-4 dark brown vittae, more conspicuous presuturally; notoplemal setulae concentrated around posterior notoplemal seta; veins R4+5 and M divergent apically; male: distance between eyes similar to flagellomere diameter; female: parafacial wide, as wide as height of gena at lower margin of eyes .................................................. 3

3.

General coloration uniform; male: antennal axis longer than oral axis; antennal insertion below mid-level of eye; cercal plate with a large triangular, membranous area anteriorly and lower margin convex; paramere acute apically (Colombia, Brazil, Uruguay, . Argentina, Chile, Juan Fernandez Is.) .........................P. chalybea (Wiedemann, 1830) General coloration not uniform; greenish blue abdomen contrasting with mesonotum coloration; male: antennal axis shorter than oral axis; antennal insertion above midlevel of eye; cercal plate with a narrow triangular membranous area and lower margin almost linear; paramere obtuse apically (Chile) ................ P. violescens (Dodge, 1967)

Reinwardtia Brauer & Bergenstamm, 1889 Type-species. Reinwardtia tachinina Brauer & Bergenstamm (mon.) . Diagnosis. General coloration metallic blue; eye setulose; gena high; arista with very short cilia; prealar strong; katepimeron and anepimeron bare; katepisternals 1:2; calcar strong; wing veins bare; vein M strongly cmved forward to R4+5; lower calypter not very enlarged, about 2.5 times as long as upper calypter; abdominal setae strong; sternite 1 bare.


97


References. BRAUER&BERGENSTAMM, 1889; HENNIG, 1965. Monophyly. The monophyly of the genus is not defined yet. HENNIG (1965) related this genus to Calliphoroides Malloch, which has only one described species from New Zealand. Geographical record. Venezuela, Ecuador. Comments. Monobasic genus: Reinwardtia tachinina Brauer & Bergenstamm, 1889.

Synthesiomyia Brauer & Bergenstamm, 1893 Type-species. Synthesiomyia brasiliana Brauer & Bergenstamm (orig. des.) =nudiseta Wulp. Diagnosis. Eye bare; arista with very short cilia; acrostichal presutural not differentiated; prealar strong; katepimeron and anepimeron bare; dorsocentrals 2:4; katepistemals 1:2; wing veins bare; vein M strongly curved forward apically; lower calypter very enlarged; stemite 1 setulose. Reference. HENNIG 1963a. Monophyly. The monophyly of the genus is not defined yet. S. nudiseta is closely related to Muscina based both on characters of the larvae and adults (SKIDMORE, 1985). Geographical record. Mexico, Nicaragua, Costa Rica, Panama, Virgin Is., Jamaica, Cuba, St. Domingo, Puerto Rico, Trinidad, Guyana, Venezuela, Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; pantropical (SKIDMORE, 1985). Comments. Monobasic genus: Synthesiomyia nudiseta (Wulp, 1883). This species has been amply distributed throughout the world by human transport (HENNIG, 1965).

Subfamily Phaoniinae Malloch, 1917 Diagnosis. Calcar present or absent; anepimeron bare; female: ovipositor elongated; cerci free, surpassing the hypoproct; sternite 8 reduced. Monophy ly. HENNIG (1965) pointed out that the Phaoniinae did not constitute 98



a monophyletic group. In CARVALHO's (1989d) cladistic analysis for the Muscidae, the subfamily showed no derived character. Helina Robineau-Desvoidy came out in a trichotomy at the base of the subfamily cladogram. Comments. The subfamily has 27 genera in the world, four of them in the Neotropical Region; 158 species are ascribed to this region.

Dolichophaonia Carvalho, 1993, stat. rev. (Figs. 109-120, Plate 2) Type-species. Phaonia brasiliensis Albuquerque (orig. des.). Diagnosis. Male holoptic; eye with short cilia; arista plumose; pre sutural acrostichals often differentiated; dorsocentral setae 2:3-4; prealar present, except in D. vockerothi, shorter than notopleural anterior seta; katepisternals 1:2; meron haired or not; wing veins bare; vein M parallel or very slightly forward-curved apically; calcar present, about twice as long as the basal width of hind tibia; female: clypeus, in lateral view, with a strong, hook-shaped anterior tip, posteriorly with a prominent sclerotization (Figs. 109118); ovipositor with large tergites and sternites (119-120). Reference. CARVALHO, 1993a. Monophyly. The monophyly of the genus was indicated by CARVALHO (1993a) based on the cladistic analysis of the family (CARVALHO, 1989d), where 2 characters of the female proboscis supported its monophyly. CARVALHO (1993a) also indicated the restrict geographical distribution, with only 2 species occurring in the United States of America, as an evidence for the monophyly of the genus. Geographical record. Mexico, Jamaica, Panama, Nicaragua, St. Vincent, Brazil, Argentina, Chile, Ecuador, Peru; south of United States of America. Comments. The synonymy between Dolichophaonia with Phaonia proposed by VOCKEROTH (1996) was not considered here, as he probably did not analyse the characters supporting the monophyly of Dolichophaonia indicated by CARVALHO (1993a). This is essentially a Neotropical genus with 27 species, two of them occurring in the southern of United States of America.


99


Key to species (modified from CARVALHO, 1993a) 1.

Two proepimeral setae developed; mid tibia with three median posterior setae; male and female palpus filiform ...................................................................................... 2 One proepimeral setae developed; mid tibia often with 2 median posterior setae; female palpus more dilated than in male; if female palpus filiform, then tergite 5 bright yellow.................................................................................................................. 5

2.

Wing with or without a brown cloud on erossvein dm-eu; meron bare ..................... 3 Wing with two dark brown clouds on erossvein dm-eu; meron haired .................... ..4

3.

Dorsoeentral setae 2:4; mid and hind femora yellow apically; wing without clouds on erossvein dm-eu; male: eye densely haired (Chile )............ .D.femorata (Stein, 1911) Dorsoeentral setae 2: 3; mid and hind femora dark brown apically; wing with a cloud on erossvein dm-eu; male: eye with sparse hairs (Chile) ...... .D. regina Carvalho, 1993

4.

Palpus yellow; Dagellomere dark brown, except for yellow base (Mexico; United States of Ameriea) ........................................................ D. texensis (Malloch, 1923) Palpus dark brown apically; flagellomere entirely dark brown (United States of America) .......................................................... ........... D. limbinervis (Stein, 1918)

5. Sternite 1 setulose ................................................................................................ 6 Stemite 1 bare ..................................................................................................... 11 6.

Palpus yellow; hind femur with 2 preapical dorsal setae (Ecuador, Peru)

............................................................................ D. compressipalpis (Stein, 1911) Palpus light to dark brown; hind femur with 1 preapieal dorsal seta.......................... 7 7.

Dorsoeentral setae 2:3; both calypteres entirely whitish .......................................... 8 Dorsocentral setae 2:4; both calypteres or only the upper one with dark brown margins .................................................................................................................. 9

8. 100

Prealar strong, a little shorter than the anterior notopleural seta; wing yellowish, more Muscidae (Diptera) of the Neotropical Region: TAXONOMY


evident on costal margin (Peru) ............................... .D. sensitarsis (Carvalho, 1983) Prealar weak:, shorter than the anterior notopleural and similar to posterior notopleural; wing hyaline, with costal margin, from R j to apex, strongly dark brown (Brazil)

.......................................................................... ..... D. paranaensis Carvalho, 1993 9.

Upper calypter entirely yellowish; wing hyaline; cercal plate with a deep posterior incision (Brazil) ......................................... .D. santoamarensis (Albuquerque, 1958) Upper calypter yellowish with dark brown margins; wing hyaline with the costal margin yellowish; cercal plate with only a shallow posterior incision .................................... 10

10. Pa1pus yellow basally; tergite 5 dark brown, yellow apically; one prescutelar seta (Brazil) ............................................................. .D. catharinensis (Carvalho, 1983) Palpus entirely dark brown; tergite 5 light brown; 2 prescutelar setae (Brazil)

................................................................... ...... D. machadoi (Albuquerque, 1958) 11. Prealar absent (St. Vincent) ................................... .D. vockerothi (Carvalho, 1983) Prealar present, shorter than notopleural anterior seta .......................................... 12 12. One intra-alar postsutural seta; tergite 5 bright yellow (Jamaica) ....................... .

...................................................... ..................... D.jamaicensis (Carvalho, 1983) Two intra-alars postsutural setae; tergite 5 not as above ........................................ 13 13. Wing with inconspicuous clouds on crossveins; calypteres whitish; dorsocentral setae 2:3 ........................................................................................................................ 14 Wing with conspicuous clouds on crossveins; calypteres whitish to yellowish with dark margins or not; dorsocentral setae 2:3-4 ....................................................... 15 14. Palpus yellowish; flagellomere partly yellow, especially its base; lateral thoracic vittae interrupted at suture; calcar inserted on apical fifth of hind tibia (Nicaragua)

......................................................................... ............. D. trigonata (Wulp, 1896) Palpus dark brown; flagellomere dark brown; lateral thoracic vittae extending up to Muscidae (Diptera) of the Neotropical Region: TAXONOMY

101


middle of mesonotum; calcar inserted on apical fourth of hind tibia (Mexico)

......................................................................................... .D. unica Carvalho, 1993 15. Wing with a cloud on basal crossvein dm-cu and another on apex of vein R 1••••••••• 16 Wing without clouds on basal crossvein dm-cu or vein R1 ...................................... 17 16. Palpus light brown; calypteres whitish; dorsocentral setae 2:4 (Brazil) ............... .

............................................................................... D. anoctiluca (Carvalho, 1983) Palpus dark brown, except for yellow base; calypteres yellowish with dark brown margins; dorsocentral setae 2:3 (Brazil) ................. D. noctiluca (Albuquerque, 1958) 17. Wing with two clouds on crossvein dm-cu ........................................................... 18 Wing with one cloud on crossvein dm-cu ............................................................. 20 18. Upper calypter yellowish with dark brown margin; wing with costal margin yellowish (Brazil, Uruguay, Argentina) ...................... .D. trigona (Shannon & Del Ponte, 1926) Upper calypter whitish; wing without yellowish costal margin ................................. 19 19. Mid and hind femora yellowish; prealar seta a little shorter than anterior notopleural seta (Argentina) ......................................................... ... D. cacheuta (Snyder, 1957) Mid and hind femora dark brown; prealar seta strongly shorter than anterior notopleural seta (Argentina) ....................................................... .. D. catamacla (Snyder, 1957) 20. Meron haired below posterior spiracle; male: tergite 3 with discal setae (Brazil)

...................................................... .................. D. tachnoides (Albuquerque, 1958) Meron bare; male: tergite 3 without discal setae ..................................................... 21 21. Frontal vitta light brown (Brazil) ............................ D. gallicola (Albuquerque, 1958) Frontal vitta dark brown ...................................................................................... 22 22. Thorax light brown (Brazil) ................................ D. brasiliensis (Albuquerque, 1958) Thorax dark brown .............................................................................................. 23 102



23. DorsocentraI setae 2:3; wing with only a inconspicuous cloud on crossvein dmcu ....................................................................................................................... 24 Dorsocentral setae 2:4; wing with a conspicuous cloud on crossvein dm-cu ............. .25 24. Palpus yellow; wing yellowish; acrostichal setae not differentiated; tergite 5 yellow at apex (Panama, Ecuador, Peru, Argentina) .................. .D. spontanea Carvalho, 1993 Palpus dark brown; wing hyaline with dorsal margin yellowish; presutural acrostichal setae differentiated; tergite 5 concolour with other abdominal tergites (Brazil) ........................................................................... .D. simplex (Albuquerque, 1958) 25. Wing hyaline without yellowish or dark brown costal margin; upper calypter whitish; post-humeral seta as long as anterior notopleura (Brazil) ................... . ....................................................... .......................... D. giacomeli (Carvalho, 1981) Wing hyaline with costal margin yellowish to dark brown; upper calypter yellowish, sometimes with dark brown margin; post-humeral seta shorter than anterior notopleural. .......................................................................................................... 26 26. Upper calypter uniformly yellowish; one prescutelar seta; mid femur yellowish (Brazil) ..............................................................................D. elongata (Albuquerque, 1958) Upper calypter yellowish with dark brown costal margin; two prescutelar setae; mid femur dark brown (Brazil) ...................................... .D. plaumanni (Carvalho, 1983)


103


.>. :' ..... ):~:+

~ ;,

:,0';'

"

109

O,5mm

stn6

O,5mm

1,Omm

stn8

hyp

Figs. 109-120, female proboscis of Dolichophaonia, lateral view (modified from CARVALHO, 1993a): 109, D. anoctiluca; 110, D. brasiliensis; III, D. cacheuta; 112, D. catharinensis; 113, D. jemorata; 114, plaumanni; 115, D. santoamarensis; 116, D. simplex; 117, D. spontanea; 118, D. trigona; Figs. 119-120, D. regina, ovipositor; 119, ventral view; 120, dorsal view. Abbreviations: c1p, clypeus; hyp, hypoproct; pre, prementum; stn, sternite; spt, spermatheca.

104



Helina Robineau-Desvoidy, 1830 (Plate 2) Type-species. Helina euphemioidea Robineau-Desvoidy (COQUILLET, 1901: 137) =pertusa Meigen. Diagnosis. Male hoi optic or dichoptic; eyes slightly pubescent; arista plumose; presutural acrostichal setae differentiated or not; dorsocentral setae 2:3; fore tibia without median setae; veins R 4+5 and M conspicuously divergent apically; pro sternum and anepimeron bare; wing veins bare; prealarpresent in male, fIliform; calcar absent; mesonotum with four dark vittae; abdomen usually without dark dorsal spots on tergites. Geographical record. Mexico, Jamaica, Guatemala, Costa Rica, Panama, Venezuela, Colombia, Peru, Ecuador, Bolivia, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, northern parts of Oriental and Afrotropical Regions. References. MALLOCH, 1934; SNYDER, 1941. Monophyly. The genus is probably not monophyletic. Two subgenera are considered-Helina s.s. and Quadrularia Huckett, 1965 (see SKIDMORE, 1985). The genus is considered a repository of poorly studied species. See Monophy ly item under Phaonia. Geographical record. Mexico, Panama, Guatemala, Costa Rica, Jamaica, Colombia, Venezuela, Peru, Ecuador, Bolivia, Brazil, Argentina, Chile, Falklands Is.; Nearctic, Palaearctic, Afrotropical, northern Oriental and Australian Regions. Comments. Helina needs a thorough taxonomic revision. The keys for identification available in the literature (STEIN, 1911; MALLOCH, 1934; SNYDER, 1941) were all included here, with modifications (some species appear in more than one key). At the end of this section, there is a list of species not included in the keys, with their known geographical distribution. Helina cruciata Snyder, 1941 was included by SNYDER (1941) in his key (see below) because he thought this species might occur also in the Neotropical Region.

Key to species (modified from STEIN, 1911) 1.

Postsutural dorsocentral setae 3 ............................................................................ 2 Postsutural dorsocentral setae 4 ........................................................................... 10


105


2.

Fore tibia with setae .............................................................................................. 3 Fore tibia without setae .......................................................................................... 5

3.

Wing with anterior margin dark brown (Bolivia) ..........H. nigromarginata (Stein, 1911) Wing with anterior margin not dark brown ............................................................. .4

4.

Pre sutural acrostichal setae differentiated (Peru) ...................H. biseta (Stein, 1904) Presutural acrostichal setae not differentiated (Peru) ..... H. echinogaster (Stein, 1911)

5.

General polinosity yellowish; antennae yellowish-white; legs light yellow (Panama, Peru, Bolivia, Brazil) ............................................... ..H. angustipennis (Stein, 1911) General polinosity grey; antennae dark brown or yellowish; legs never light yellow ................................................................................................................... 6

6.

Wing with anterior margin light brown (peru; Brazil) .....H. pedella (Wiedemann, 1830) Wing with anterior margin hyaline or, if margin pigmented, then prealar seta absent... .. 7

7.

Male dichoptic (Peru) .............................................. ....H. eurycephaZa (Stein, 1911) Male holoptic ....................................................................................................... 8

8.

Prealar seta present (Peru) .....................................H. consanguinea (Stein, 1911) Prealar seta absent. ............................................................................................... 9

9.

Dorsal dark vittae on mesonotum not well marked; palpus black (Peru, Brazil) .................................................................................H. nigrina (Wiedemann, 1830)

Dorsal dark vittae on mesonotum well marked; palpus yellow (Bolivia) ........................................................................................H. piliceps (Stein, 1911)

10. Fore tibia with setae (Peru, Brazil) ....................................H. tesselata (Stein, 1904) Fore tibia without setae (Peru) ............................................... H.fallax (Stein, 1911)

106



Key to species (modified from MALLOCH, 1934) 1.

Metallic violet-blue species; calypteres and halter black; eyes conspicuously longhaired; arista almost bare; palpus black, strap-like; prealar seta over half as long as the one behind it; legs entirely black (Argentina) ....................H. viola Malloch, 1934 Black species, with more or less distinct grey or brownish-yellow dusting; calypteres and halter pale ........................................................................................................ 2

2.

Mid and hind femora entirely orange-yellow............................................................. 3 Mid and hind femora partly or entirely black ............................................................ 5

3.

All femora and tibiae entirely orange-yellow; male: hind femur with about 6 strong setae on apical half or more of anteroventral surface, unarmed on basal half of posteroventral surface (Chile)....... ...................... H. australis Carvalho & Pont, 1993 Fore femur almost entirely black; fore tibia usually with darkened base; mid and hind femora and tibiae orange yellow; male: hind femur with a greater portion of the anteroventral surface bristled ................................................................................. 4

4.

Hind femur in both sexes with long curved setae on apical half of anteroventral surface that decreases in length and strength toward the middle of femur; posteroventral surface without long setae; arista with its longest hairs about three times as long as its basal diameter (Chile, Argentina) ..............................................H. connexa Malloch, 1934 Hind femur in male with a complete series of anteroventral setae, weaker near base, and a series of long, slender setae on basal half of posteroventral surface; in female with a less extensive anteroventral series and fewer and shorter posteroventral setae; arista with its longest hairs hardly longer than its basal diameter (Chile, Argentina)

......................................................... ..................... H. nigrimana (Macquart, 1851) 5.

Postsutural dorsocentral setae 3 ............................................................................ 6 Postsutural dorsocentral setae 4 .............................................................................. 8

6.

Mid and hind femora broadly orange yellow at bases, blackened at apices; wing with


107


crossveins distinctly with brown clouds, the inner one the more conspicuous, almost spot-like; abdomen densely grey dusted with a pair of small dark brown spots on tergite 3 and tergite 4 (Chile, Argentina) ............................... H. xena Malloch, 1934 Mid and hind femora blackened at bases; wing with crossveins not noticeably clouded; abdomen without evident dark paired spots on dorsum ......................................... 7 7.

Frons quite densely pale grey dusted, at narrowest point about 1.5 times as wide as flagellomere; orbits not bristled immediately in front of anterior ocellus; arista short pubescent; mid femur with apical third and hind femur with apical three-fourths brownish-yellow (Peru, Argentina) ....................................H. discolor (Stein, 1911) Frons entirely black, at narrowest point about three times as wide as flagellomere; orbits with setae on their entire extent, upper seta on each outcurved; arista short-haired, the longest hairs above, fully twice as long as basal diameter of arista; mid and hind femur with apical third reddish-yellow (Argentina) .............. ... .H. rufoapicata Malloch, 1934

8.

Mid tibia with a strong submedian seta on posteroventral surface; frons at narrowest point wider than flagellomere; prealar seta much less than one-third as long as the one behind it. ............................................................................................................... 9 Mid tibia with posterior but no posterodorsal submedian setae; other characters not as above in toto .................................................................................................. 11

9.

Arista with its longest hairs not longer than its basal diameter; mid tibia with 1-2 submedian anterodorsal setae (Peru, Argentina) .............. ....H. discolor (Stein, 1911) Arista with much longer hairs ............................................................................... 10

10. Mid tibia without submedian anterodorsal setae (Chile) ......H. chilensis Malloch, 1934 Mid tibia with 1-2 submedian anterodorsal setae (Chile, Argentina) ................ ..

......................... .............................................................. .H. simplex Malloch, 1934 11. Prealar seta less than one-third as long as the one behind it; hind femur without long setae on basal half of posteroventral surface; mid and hind femora orange-yellow to 108



reddish yellow on at least the apical third ............................................................. 12 Prealar seta at least one-third as long as the one behind it, frequently much longer; hind femur entirely black or with fine long setae on at least basal half of posteroventral surface ................................................................................................................ 14 12. Mid and hind femora with less than their basal half blackened; male: frons at narrowest point about twice as wide as flagellomere (Chile, Argentina) ... .H. simplex Malloch, 1934 Mid and hind femora with more than their basal half blackened............................. 13 13. Male: frons at narrowest point hardly wider than flagellomere; orbits unarmed on upper third in front of ocelli (Chile, Argentina) ......H.juscocalyptrata Malloch, 1934 Male: frons at narrowest point about three times as wide as flagellomere; orbits bristled on entire extent (Argentina) ........................................H. ruJoapicata Malloch, 1934 14. Arista short haired, its longest hairs about five times as long as its basal diameter; hind femur in male black, with a faint yellowish-brown tinge at extreme apex, in female much more broadly fulvous-yellow, posteroventral surface in both sexes with no well developed erect setae on basal half; prealar seta over half as long as the one behind it; fore tibia rarely with a submedian posterior seta; calypter and bases of wing-veins fulvous-yellow (Chile )........................................................ H. bigoti Malloch, 1934 Arista pubescent, its longest hairs not or very little longer than its basal diameter; hind femur distinctly fulvous or reddish-yellow at apex and in male at least with a series of long fine erect setae at least on basal half of posteroventral surface; prealar seta less than half as long as the one behind it; calypter fulvous-yellow, but bases of wing veins fuscous (Chile, Argentina) ....................................... H. nigrimana (Macquart, 1851)

Key to species (modified from SNYDER, 1941)

Male 1. Strong postsutural dorsocentral setae 3............................................................ .2 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

109


Strong postsutural dorsocentral setae 4 ................................................................ 12 2.

Costal margin, crossveins and apex of veins broadly infuscated; stem of halter at base of knob darkened; hind femur with slender, almost hair-like setae on at least basal half of posteroventral surface ........................................................................................ 3 Wings not so marked or exactly not as above ......................................................... .4

3.

Frontal vitta at narrowest part at most 1.5 times as wide as the diameter of anterior ocellus; when viewed in profile, antenna inserted opposite middle of eyes; flagellomere narrow not more than two and 2.5 times as long as pedicel (Colombia) ............ ..

............. ................................................................. .......... .. H. vierecki Snyder, 1941 Frontal vitta at narrowest part almost 2.5 times as wide as the distance across posterior ocelli, inclusive; when viewed in profile, antenna inserted opposite dorsal third of eye; flagellomere broad almost 4 times as long as pedicel (Colombia, Peru, Bolivia)

...................................................................... .......... H. marginipennis (Stein, 1904) 4.

One or two pairs of strongly developed pre sutural acrostichal setae ........................... 5 With at most a pair of well developed acrostichal setae which are never more than half as long and strong as the adjacent dorsocentral setae, or eyes not separated by a distance greater than diameter ocelli, inclusive ......................................................... 6

5.

Deep blackish pruinescent species with only the apices of mid and hind femora orange; hind tibia with three anterodorsal setae and three or four shorter posterior ones on middle (Argentina) ..................................................... H. ruJoapicata Malloch, 1934 Light yellowish-gray pruinescent species with at most only the fore femur indistinctly darkened; hind tibia with only one anterodorsal and no posterior setae (Ecuador)

............................................................... ...................... H. acrosticalis Snyder, 1941 6.

Longest hairs on arista shorter than or only as long as greatest diameter of arista........ 7 Longest hairs on arista at least half as long as the width of flagellomere ...................... 9

7. 110

Palpus, at least on basal two-thirds, pedicel and fore femur yellowish to fulvous Muscidae (Diptera) of the Neotropical Region: TAXONOMY

a


(Ecuador, Peru) .......................................................... ;..... ..H. acrinis Snyder, 1941 Palpus, pedicel and fore femur black ..................................................................... 8 8.

Hind tibia with only one anteroventral seta which is situated on the apical fourth (Chile, Argentina) ................................................................. .........H. xena Malloch, 1934 Hind tibia with two or three anteroventral setae which are situated on the apical two thirds (?Mexico, Colombia, Peru) ...................................H. adelpha (Schiner, 1868)

9.

Hind tibia with one or more posterodorsal setae on at least basal half; crossveins very narrowly infuscated; abdominal stemites with a marginal fringe oflong setae (Mexico, North America) .............................................. ...............H. lasiosterna Snyder, 1941 Hind tibia without a basal posterodorsal seta; crossveins very broadly infuscated; abdominal stemites without long marginal setae ................................................... 10

10. Hind femur with at most only the anteroventral row of setae (Mexico, Costa Rica)

..................................................................................... H. refusa (Giglio-Tos, 1893) Hind femur with a complete row of anteroventral and posteroventral setae or long bristly hairs ............................................................................................................ 11 11. Presence of a conspicuous dark spot at base of discal cell below juncture of veins R 2+3 and R 4+5; crossveins clear (Brazil) ....................H. poeciloptera (Schiner, 1968) Absence of a conspicuous dark spot at base of discal cell below juncture of veins R4+5 and M; crossveins infuscated (Mexico ).....................H. signatipennis (Wulp, 1896) 12. All legs black or very dark brown ........................................................................ 13 At least some parts of the legs much lighter coloured............................................. 15 13. Margins of upper calypter darkened; hind femur without posteroventral hair-like setulae or setae (Ecuador) ............................................. .....................H. browni Snyder, 1941 Margins of upper calypter not darkened; hind femur with an almost complete row of posteroventral hair-like setae or setulae................................................................... 14


111


14. Hind tibia with a series of very long slender posteroventral and ventral setae on basal three-quarters which are strongest at the base and become shorter toward apex (Peru) ................................................. ...............................H. longipila (Stein, 1918) Hind tibia without long hair-like posteroventral and ventral setulae on basal threequarters (Ecuador) .................................................... ........H. equator Snyder, 1941 15. With only one pre sutural acrostichal seta (Mexico )...............H. copiosa (Wulp, 1896) With one or more pairs of distinct presutural acrostichal setae which are always stronger than the adj acent acrostichal setae ....................................................................... 16 16. Vein M strongly bent posteriorly at apex; crossvein dm-cu strongly curved; both crossveins broadly infuscated (Peru) .....................................H. inepta (Stein, 1911) Vein M not strongly bent posteriorly at apex; other characters not exactly as above .................................................................................................................. 17 17. Mid tibia with a strong median posteroventral seta; frons distinctly wider than width of flagellomere; prealar less than one third as long as the seta behind it... ..... '" ............ 18 Mid tibia with no posteroventral setae or if present the other characters not exactly as above.,. .. " ......................................................................................................... 20 18. Longest hairs on arista not longer than its greatest diameter; mid tibia with one or two submedian anterodorsal setae (Peru, Argentina) ................... H. discolor (Stein, 1911) Longest hairs on arista distinctly longer than its greatest diameter............................ 19 19. Mid tibia with no anterodorsal setae; scutellar setulae descending below level of marginals (Chile) ........................... ...................................H. chilensis Malloch, 1934 Mid tibia with one or two anterodorsal setae; scutellar setulae not descending below level of marginals (Chile or Argentina) ................................H. simplex Malloch, 1934 20. All femora and tibia entirely fulvous (Chile )..........H. australis Carvalho & Pont, 1993 At least saIne parts offemora or tibia infuscated .................................................... 21

112



21. Prealar less than one-third as long as first postsutural dorsocentral seta ................... 22 Prealar as long and strong as the first postsutural dorsocentral seta ....................... .25 22. Hind femur and tibia with long, slender posteroventral setae at base (Peru)

...................................................... ................................. H.longipila (Stein, 1918) Hind femur and tibia not so armed ........................................................................ 23 23. Frons at narrowest part equal to length of flagellomere; parafacial setae extending all the way to vertex (Argentina) .............................. .......H. ruJoapicata Malloch, 1934 Frons at narrowest part less wide as length of flagellomere; at most only slightly wider than distance across posterior ocelli, inclusive ........................................................ 24 24. Mid tibia without median anterodorsal setae; hind femur and tibia darkened only on basal fourth (Peru) ................................................. ...........H. townsendi Snyder, 1941 Mid tibia with one or two median anterodorsal setae; hind femur darkened on basal half and the hind tibiae darkened on basal two-thirds (Argentina) ............. .

................ ............................... ....................... ...... H. Julvocalyptrata Malloch, 1934 25. Longest hairs on arista fully one-half as long as the width of flagellomere; hind femur with anteroventral hair-like setae not as long as the diameter offemur at the portion in which they are situated; legs mostly blackish (Chile ) ............H. bigoti Malloch, 1934 Longest hairs on arista not longer than its greatest diameter; hind femur with anteroventral hair-like seta at least 1.5 as long as diameter offemur at the portion in which they are situated; legs mostly yellow........................................................... 26 26. Mid tibia with one or more short but distinct submedian anterodorsal setae; hind femur with a number oflong, hair-like posteroventral setae which are distinctly longer than diameter of femur at the portion in which they are situated (Chile, Argentina)

..................................................................................... .H. connexa Malloch, 1934 Mid tibia with no anterodorsal setae; hind femur without long, hair-like posteroventral setae (Chile, Argentina) ...........................................H. nigrimana (Macquart, 1851) Muscidae (Diptera) of the Neotropical Region: TAXONOMY

113


Female 1. Strong postsutural dorsocentral setae 3................................................................. 2 Strong postsutural dorsocentral setae 4 ................................................................ 13 2.

Crossveins, costal margins and apex of vein R4+S and M broadly infuscated; no prescutellar acrostichal seta; prealar absent; halter black (Colombia, Peru, Bolivia) ...............................................................................H. marginipennis (Stein, 1904)

At most only the crossveins infuscated or not exactly as above ................................. 3 3. Hind tibia with one or more distinct posterodorsal setae on basal half or less .......... ..4 Hind tibia with no posterodorsal setae .....................................................................7 4. Frontal vitta with a pair of interfrontal setae; gena almost half as high as eyes; fore tibia without median posterior and anterodorsal setae; prealar almost as long and strong as posterior notopleural (North America; ?Neotropical Region) ....H. cruciata Snyder, 1941 Frontal vitta without interfrontal setae; gena less than one-fourth of eye height; fore tibia with median posterior or anterodorsal setae or with both .................................. 5 5. Pedicel mostly or entirely light yellow to orange (Mexico) ..... .H. parvula (Wulp, 1986) Pedicel entirely black or dark brown ...................................................................... 6 6. Hind tibiae with two or three posterodorsal setae; longest hairs on arista not quite as long as greatest width of flagellomere (Mexico )..................... .H. copiosa (Wulp, 1896) Hind tibia with only one posterodorsal seta on basal third or less; longest hairs on arista not over half as long as greatest width of flagellomere (Mexico) .............. .. ............................................................................................H. caneD Snyder, 1941

7.

Arista with hairs at most slightly longer than the greatest diameter of arista............... 8 Arista with hairs distinctly longer than greatest diameter of arista, usually at least half as long as greatest width of flagellomere ................................................................ 12

8. Thorax, abdomen and palpus largely or entirely yellow in ground colour; fore tibia 114



with one median anterodorsal seta (Ecuador, Peru) ................H. acrinis Snyder, 1941 Thorax entirely and abdomen and palpus usually black; fore tibia without median anterodorsal seta ........................................................... 9.

0 .......................................

9

Mid tibia with two median anterodorsal setae (Ecuador) ...H. brevivena Snyder, 1941 Mid tibia without median anterodorsal setae ......................................................... 10

10. Without presutural acrostichal setae; crossveins at most only moderately infuscated (Mexico) ..................................................................... .......... H. socia (Wulp, 1896) With one or more pairs of short but distinct pre sutural acrostichal setae; crossveins very broadly infuscated ........................................................................................... 11 11. Mid and hind femora infuscated on apical third, the basal two-thirds orange-yellow; hind tibia with only one anteroventral seta which is situated on apical fourth (Chile, Argentina) ......................................................... ..................... H. xena Malloch, 1934 All femora almost entirely infuscated; hind tibia with one anteroventral seta situated on its middle (?Mexico, Colombia, Peru) .............................H. adelpha (Schiner, 1868) 12. Veins R J and R 2, crossveins and M at base of discal cell broadly infuscated; crossveins most distinctly so infuscated; katepisternals 1:2 (Brazil) ... H. poeciloptera (Schiner, 1968) Only the crossveins infuscated; katepisternals 2:2 (Mexico; Costa Rica)

.................. H. refusa (Giglio-Tos, 1893) and (Mexico) H. signatipennis (Wulp, 1896) 13. Metallic violet-blue non pruinescent species with entirely black calypter and halter (Argentina) ..................................................... .....................H. viola Malloch, 1934 Species not so coloured ....................................................................................... 14 14. Legs entirely black to dark brown ......................................................................... 15 Legs partly or entirely yellow to reddish .............................................................. 17 15. Margins of both calypteres and entire halter black; scutellar setulae descending well below level of marginals, especially at base of scutellum; abdomen distinctly bluish, Muscidae (Diptera) of the Neotropical Region: TAXONOMY

115


with sparse greyish pruinescence (Ecuador) ............................ H. notha Snyder, 1941 Margins of calypteres and entire halter not black; at most with one or two setulae descending below level of marginals on middle of scutellum. .................................. 16 16. Crossvein dm-cu joining the vein M at about the middle of the portion beyond the crossvein r-m; costal cell basad the vein R1 light brownish hyaline, scarcely darker than the lightest portion of wing membrane (Ecuador) ........ ....H. anubes Snyder, 1941 Crossvein dm-cu joining vein M well beyond the middle of the portion beyond cross vein r-m; costal cell before juncture basad vein R1 almost opaque, black, very much darker than the lightest portion of wing membrane (Ecuador) ........ H. equator Snyder, 1941 17. Mid tibia with a median posteroventral seta (Chile or Argentina) ........................... .

...................................................... ................................ H. simplex Malloch, 1934 Mid tibia without a submedian posteroventral seta............................................... 18 18. Mid tibia with two or more strong median anterodorsal setae ............................... 19 Mid tibia bare or with a single very weak median anterodorsal seta ........................ 21 19. Prealar as long as or longer than posterior notopleural seta; mid and hind femora entirely yellow (Chile, Argentina) ...................................H. connexa Malloch, 1934 Prealar at most half as long as the posterior notopleural seta ................................. 20 20. Thorax with dense, golden yellow pruinescence; all tibiae fulvous; without a strong pair of prescutellar acrostichal seta (Ecuador) ................. H. brevivena Snyder, 1941 Thorax with greyish pruinescence; only the hind tibia fulvous, others of legs black; with a pair of strong prescutellar acrostichal seta (Argentina) ............... ..

........................................................................... H.Julvocalyptrata Malloch, 1934 21. Vein M very strongly bent posteriorly at apex; crossvein broadly infuscated, dm-cu most broadly so, at its juncture with veins M and CuA 1; eye with short but distinct hairs (Peru) ........................................................................... H. inepta (Stein, 1911)

116



Vein M subparallel with R4+5 at apex; cross veins not broadly infuscated ................ 22 22. Mid and hind femora entirely yellow (Chile, Argentina) ... .H. nigrimana (Macquart, 1851) Mid and hind femora mostly infuscated ................................................................. 23 23. Prealar distinctly longer than the posterior notopleural seta; mid tibia with four or five median posterior setae; fore and mid tibiae entirely fulvous (Chile) ............ ..

.......................................... ................................................. .H. bigoti Malloch, 1934 Prealar at most half as long as the posterior notopleural seta; mid tibia with two median posterior setae; fore and mid tibiae entirely infuscated (Ecuador) .............. .

.......................................................................................... .. H. anubes Snyder, 1941

Species of Helina not included in the key and their respective geographical distribution Species


H. albuquerquei Pont, 1972

Brazil

H. argentina Snyder, 1957

Argentina

H. auricolis Albuquerque, 1980

Brazil

H. circulatrix (Walker, 1861)

Mexico, Guatemala

H. crepedoseta Snyder, 1940

Mexico, Guatemala, Costa Rica, Panama

H. crocea Snyder, 1940

Brazil, Argentina

H. cyanicolor (Stein, 1911)

Bolivia

H. dasyophthalma Malloch, 1928

Peru, Bolivia

H. dasyops (Macquart, 1843)

Falkland Is.

H. dichroma (Wiedemann, 1830)

Brazil

H. discreta (Wulp, 1896)

Mexico, Jamaica


117


H. etesia (Giglio-Tos, 1893)

Mexico

H. evecta (Harris, 1780)

Mexico, Venezuela; Nearctic, Palaearctic, northern parts of Oriental and Afrotropical Regions

lIS

H. Julvapoda Snyder, 1940

Mexico, Guatemala

H. gigantea Albuquerque, 1956

Brazil

H. grandis (Schiner, 1868)

South America

H. latipennis (Stein, 1904)

Colombia, Bolivia

H.leucocephala (Wulp, 1896)

Mexico, Panama

H. limbata (Wiedemann, 1830)

Brazil

H. luteola Albuquerque, 1956

Brazil

H. lyncii (Wulp, 1883)

Argentina

H. maculipes (Stein, 1918)

Mexico

H. meraca (Wulp, 1896)

Mexico; U. S. A.

H. monacha (Schiner, 1868)

South America

H. mulcata (Giglio-Tos, 1893)

Mexico

H. neosimplex Snyder, 1957

Argentina

H. neotropica Snyder, 1951

Peru

H. nivaloides Albuquerque, 1956

Brazil

H. nobilis Albuquerque, 1956

Brazil

H. parsura (Giglio-Tos, 1893)

Mexico

H. praecipua (Walker, 1853)

Brazil



H. procedens (Walker, 1861)

Mexico,U.S.A.

H. prolatifrons Snyder, 1940

Panama

H. regobarrosi Albuquerque, 1958

Brazil

H. rubripalpis (Wulp, 1896)

Mexico, Costa Rica, U.S.A.

H. sera (Giglio-Tos, 1893)

Mexico

H. tarsalis (Stein, 1918)

Mexico

H. trichops (Stein, 1918)

Peru

H. umbrosa (Wulp, 1896)

Costa Rica, Venezuela

H. walkeri Carvalho & Pont, 1993

Brazil, Paraguay

Phaonia Robineau-Desvoidy, 1830 Type-species. P haonia viarum Robineau -Desvoidy (des. COQUILLET, 1901 a: 140) = valida Harris. Diagnosis. Male dichoptic; eyes ciliated; arista plumose; dorsocentral setae 12:3-4; notopleuron with covering setulae and with 2 setae, the posterior one weaker; prealar seta present (absent in P. lentiginosa Snyder); lower calypter glossiform, Phaoniatype; Rs node bare or ciliated; vein M usually curved forward apically; hind tibia on posterodorsal surface with the calcar about as long as the width of the tibia at calcar insertion; female: ovipositor elongated, tubular, tergites narrow; stemite 8 reduced to 2 sclerites; microtrichia usually well developed only on the membrane; cerci free. Reference. COELHO, 2000. Monophy ly. Phaonia is a very heterogeneous genus with more than 300 species described in the world and it is certainly not monophyletic (HENNIG, 1965; ZINOVJEY, 1981; SKIDMORE, 1985; HUCKETT & VOCKEROTH, 1987; CARVALHO, 1989d). Except for the Neotropical Region, the species from other biogeographical regions have been segregated into several groups (seeZINOVJEV, 1981), howeverthe monophyly of none of these "groups" has been established. SKIDMORE (1985) considered Phaonia


119


s.s. more closely related to subgenus Quadrularia of Helina than to the other species placed in Phaonia. Geographical record. Mexico, Guatemala, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Uruguay, Argentina; cosmopolitan. Comments: The species P costipennis (Snyder, 1957); P marginata Stein, 1918; P nervicincta Stein, 1918; P pteropsila Stein, 1918 andP tetragona Gaminara, 1930 were not included in this key. According to COELHO (2000), their original descriptions are not sufficient to recognise them. Besides this, their types have not been located and are probably destroyed.

Key to species (modified from COELHO, 2000) 1.

Wing on radial node with cilia on dorsal and/or ventral surface; dorsocentral setae 12:3-4 .................................................................................................................... 2 Wing on radial node with no cilia; dorsocentral setae 2:4 .......................................... 31

2.

Presutural dorsocentral seta 1............................................................................... 3 Presutural dorsocentral setae 2 .............................................................................. 5

3.

Prescutellar acrostichal setulae present; posterior spiracle with cilia on posteroventral margin; wing with dark clouds on apical portion of veins Sc and R}, apical half of~+3 and on crossveins r-m and dm-cu; female: faint cloud on apical portions ofR4+5 and M (Brazil, Peru, Bolivia) ............................................. P praesuturalis (Stein, 1904) Prescutellar acrostichal setulae absent; posterior spiracle bare on posteroventral margin; wing not as above ..................................................................................... 4

4.

General coloration brown to dark brown; fore tibia bare on anterior surface; male: tarsi with claws and pulvilli short, shorter than the length of tarsomere 5 (Brazil)

........................................................................... ....... P bigoti (Albuquerque, 1957) General coloration dark brown with tergite 1 and base oftergite 2 light brown; fore tibia with an anterior median seta; male: tarsi with claws and pulvilli similar to tarsomere 5 in length (Argentina) ................................................ P monochaeta Snyder, 1957 120



5.

Postsutural dorsocentral setae 3 ............................................................................ 6 Postsutural dorsocentral setae 4 ........................................................................... .28

6.

Prealar seta absent (Brazil) ........................................ ...P. lengitinosa Snyder, 1957 Prealar seta present, similar to notopleural anterior seta in length ............................. 7

7.

General coloration light brown, at most, abdomen darker...................................... 8 General coloration dark brown to black, at most the scutellum yellowish brown; abdomen entirely dark brown or with tergites 1-3 light brown ............................... 14

8.

Thorax and abdomen light brown (Brazil) ..P. aurantica (Albuquerque & Medeiros, 1980) Abdomen dark brown, in contrast to thorax, or at most tergites 1+2 light brown ..... 9

9. Abdomen dark brown, with tergites 1+2 light brown (Brazil, Argentina) ...........................................................................P. shannoni Carvalho & Pont, 1993

Abdomen entirely dark brown ............................................................................. 10 10. Prescutelar acrostichal setulae present. .................................................................. 11 Prescutelar acrostichal setulae absent.. ................................................................. 13 11. Posterior spiracle with cilia on posteroventral margin (Brazil, Argentina) ....................................................................... P. nigriventris (Albuquerque, 1954)

Posterior spiracle bare on posteroventral margin .................................................. 12 12. Hind tibia with 4 median setae (Brazil) ...P. benjamini (Albuquerque & Medeiros, 1980) Hind tibia with 1-2- median setae (Brazil) ................ P. similata (Albuquerque, 1957) 13. Dark brown vitta on mesonotum reaching the row of intra-alar setae; male: ocellar setae strong (Brazil) ............................................... P. annulata (Albuquerque, 1957) Dark brown vitta on mesonotum not reaching the row of intra-alar setae; male: ocellar setae weak (Brazil) ........................................... P. grajauensis (Albuquerque, 1957) 14. Scutellum yellowish brown apically (Venezuela, Brazil, Argentina, Uruguay) Muscidae (Diptera) of the Neotropical Region: TAXONOMY

121


......................................................................................... P. trispila (Bigot, 1885) Scutellum concolour with thorax .......................................................................... 15 15. Posterior spiracle with cilia on posteroventral margin ............................................. 16 Posterior spiracle bare on posteroventral margin .................................................. 18 16. Male: tarsi with claws and pulvilli short, shorter than the length of tarsomere 5; female: ovipositor with microtrichia, except on tergite 6 (Peru, Brazil) .................................. ..

.................. ..... ............ ...... .............. ..... ................. ........ .... P. quercus Coelho, 1998 Male: tarsi with claws and pulvilli similar to tarsomere 5 in length; female: ovipositor not as above ........................................................................................................ 17 17. Wing with dark brown clouds only on crossveins r-m and dm-cu; male: paramere with a concavity on ventral surface; gonopod with the anterior region short and shorter than the width of paramere (peru, Bolivia, Brazil, Argentina) ................ P.latinervis (Stein, 1904) Wing with dark clouds on apices of veins R4+5 and M, another cloud extending from apices of Sc and R1 to apex of R2+3, and one cloud on crossveins r-m and dm-cu; male: paramere and gonopod not as above (Ecuador, Peru, Bolivia) ........................ .

.................................. ........................... ........................... .... P. maculosa Stein, 1911 18. Abdomen dark brown with tergites 1+2 and base oftergite 3light brown; male: tarsi with claws and pulvilli short, shorter than length of tars omere 5; female: ovipositor with microtrichia, except on tergite 6 (Panama, Brazil, Ecuador, Peru) .................. .

....... .......... ..... ........... ................. ....... ............... .............. P. hugonis Carvalho, 1989 Abdomen entirely dark brown; male: tarsi with claws and pulvilli long, longer than or similar in length to tarsomere 5; female: ovipositor not as above .............................. 19 19. Prescutelar acrostichal setulae present.. ................................................................ 20 Prescutelar acrostichal setulae absent. .................................................................. 27 20. Fore tibia with 1-2 anterior median setae ............................................................. 21

122



Fore tibia bare on anterior surface ........................................................................ 24 21. Mid tibia bare on anterior surface ........................................................................ 22 Mid tibia with 1-2 anterior median setae ............................................................... 23 22. Wing with dark brown clouds only on crossveins r-m and dm-cu; male: distance between eyes, at level of anterior ocelli, about 0.26 of head width (Venezuela, Ecuador) ............................................................ ....... P. bipunctata (Schiner, 1868) Wing with dark brown clouds on apices of veins Sc, R] and M and on crossveins r-m and dm-cu; male: distance between eyes, at level of anterior ocellus, about 0.24 of head width (Colombia, Ecuador, Peru, Bolivia) ...................P. curvata (Stein, 1904) Hind tibia with two anterior and one posterodorsal sub-basal setae; dorsal surface with one strong apical seta (Peru, Boli via, Argentina) .......P. punctinervis Stein, 1911 23 Hind tibia with no anterior or posterodorsal sub-basal setae; dorsal surface without apical seta (Peru) .......................................................... .P. punoensis Coelho, 1998 24. Hind tibia with 1-2 sub-basal anterior setae ............................................................ 25 Hind tibia with no sub-basal anterior setae ........................................................... 26 25. Hind tibia with one submedian posterodorsal seta (Bolivia) .....P. soratiensis Coelho, 1998 Hind tibia with no submedian posterodorsal seta (Bolivia) .....P. nigripuncta Stein, 1911 26. Female: ovipositor with microtrichia only on membrane (Brazil, Ecuador, Peru)

..........................................................................................P. triseta (Curran, 1931) Female: ovipositor with microtrichia on membrane, tergites and sternites (Brazil)

......................................................................................... P. grandis (Couri, 1982) 27. Fore tibia with a median anterior seta (Mexico, Panama, Colombia, Ecuador, Bolivia)

...................................................................................P. abdita (Giglio-Tos, 1893) Fore tibia bare on anterior surface (Brazil; Bolivia; Argentina) ...P. advena Snyder, 1957 28. Fore tibia with 1-2 anterior median setae .............................................................. 29 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

123


Fore tibia without anterior median seta .................................................................. 30 29. Hind femur on anterior surface with a preapical seta; hind tibia without an anterior sub-basal seta (Colombia, Ecuador) ............................. P. equatoralis Coelho, 1998 Hind femur on anterior surface bare; hind tibia with one anterior sub-basal seta (Bolivia)

......................................................................................P. boliviana Coelho, 1998 30. Apical portion of vein M strongly bent forward; male: gonopod with a long and curved anterior region, surpassing the paramere basally (Mexico, Ecuador, Peru, Bolivia) ............................................... ............................. P. venicurva (Stein, 1904) Apical portion of vein M slightly bent forward; male: gonopod not as above (Mexico) .................................................................. ...... P. mexala (Snyder, 1957) 31. Stemite 1 setulose ................................................................................................ 32 Stemite 1 bare ...................................................................................................... 34 32. Flagellomere entirely dark brown; calypteres yellow with dark brown margins; hind femur on anteroventral surface with setae only on apical half; meron with many setulae; male: paramere on ventral surface with a weak concavity; gonopod with the anterior region long, straight and not surpassing the width of the paramere (Mexico)

.....................................................................................P. nigerrima Carvalho, 1984 Flagellomere dark brown with a yellow base; calypteres whitish; hind femur on anteroventral surface with a complete series of strong setae; meron with few setulae ................................................................................................................ 33 33. Pedicel dark brown, yellow apically; wing hyaline; hind femur dark brown (Mexico) ............................................. ........................... P. robusta Carvalho, 1984 Pedicel entirely dark brown; wing yellowish; hind femur yellowish; male: paramere with the ventral surface almost straight, with a small cavity near its apex; gonopod with the anterior region long, straight and not surpassing the width of the paramere (Mexico) ................................................................ ....P. mexicana Carvalho, 1984 34. Arista with long plumae; cerci with a deep posterior incision .................................. 35 124



Arista with few plumae; cerci not as above .......................................................... 36 35. Palpus dark brown, with a yellow base; humerals 3; fore tibia with 1-2 anteroventral submedian setae; male: paramere small; gonopod with the anterior region short (Mexico) .......................................................................... P. major Carvalho, 1984 Palpus entirely dark brown; humerals 2; fore tibia without an anteroventral submedian seta; male: paramere large; gonopod with the anterior region curved (Mexico)

........................................................................................ P. minuta Carvalho, 1984 36. Lower-posterior katepistemal seta weak; fore tibia without one anterior median seta; hind tibia on posterodorsal surface with a strong median seta, in addition the calcar (Ecuador) ..............................................................................P. ponti Coelho, 1998 Lower-posterior katepistemal seta strong; fore tibia with one anterior median seta; hind tibia on posterodorsal surface only with the calcar.. ....................................... 37 37. Hind tibia with one anterior submedian seta; male: paramere with a concavity on ventral surface (Ecuador) ...............................................P. carvalhoi Coelho, 1998 Mid tibia without an anterior submedian seta; male: paramere not as above (Ecuador) ......................................................................... P. hucketti Coelho, 1998

Species of Phaonia not included in the key and their respective geographical distribution Species


P. costipennis (Snyder, 1957)

Brazil

P. marginata Stein, 1918

Peru

P. nervicincta Stein, 1918

Paraguay

P. pteropsila Stein, 1918

Ecuador

P. tetragona Gaminara, 1930

Uruguay


125


Souzalopesmyia Albuquerque, 1951 (Figs. 121-132) Type-species. Souzalopesmyia carioca Albuquerque (orig. des.). Diagnosis. Male dichoptic, frons narrower than in female, with reclinate orbital seta, and without interfrontal setae; ocellar setae absent; antenna long, reaching epistoma (Fig. 121); arista plumose, longest hairs as long as maximum antennal diameter; parafacial setulose on upper half; dorsocentral setae 1-2: 3; acrostichal setulae not distinct from ground setulae, except in female of S. amazonica; humerals 2; intra-alars 1:2; prealar absent; anepisternum with a short seta on upper anterior comer, meron bare; katepisternals 1:2 (not 1: 1: 1 as stated by ALBUQUERQUE, 1951); posterior spiracle small, triangular, with yellow setulae on its margin; prostemum bare; fore tibia with one posterodorsal submedian seta; fore tarsomere 1 with one ventral seta; mid femur with 1 anterodorsal, 1 dorsal, 1 posterodorsal and 1 posterior preapical seta; mid tibia with 2-4 posterior median setae, 1 strong ventral seta apically; hind tibia without calcar and with 1 anterodorsal median seta, and dorsal, anterodorsal and anteroventral surfaces each with one apical seta (Fig. 122); veins bare, except costa; vein M slightly curved forward apically (Figs. 123, 124); lower calypter of Phaonia-type; sternite 1 bare; ovipositor with long tergites; sternites and membranes covered with rnicrotrichia (Figs. 125, 126). Reference. CARVALHO, 1999. Geographical record. Peru, Bolivia, Brazil. Monophyly. Souzalopesmyia is a monophyletic genus based on the following synapomorphies: head elongated in lateral view; ocellar setae absent and general coloration yellow (CARVALHO, 1999). Comments. Souzalopesmyia species are rare and very similar to each other. The species, based on the known collected material, are found in forests and are probably nocturnal, since they have been very rarely collected during the day (CARVALHO, 1999).

Key to species (modified from CARVALHO, 1999) 1.

Inner vertical setae divergent (Fig. 121); crossvein dm-cu oblique, weakly curved (Fig. 124); female: 2 frontals (Brazil) ............................S. carioca Albuquerque, 1951 Inner vertical setae parallel; crossvein dm-cu oblique, almost straight (Fig. 123); female:

126



3 frontals ................................................................................................................ 2 2.

Dorsocentrals setae 2:3; male: postocular row of setulae incomplete, not reaching epistoma; setulae black; female: some presutural acrostichal setulae stronger than the ground setulae (Brazil) ......................................... S. amazonica Albuquerque, 1951 Dorsocentrals setae 1: 3; male: postocular row of setae complete, setulae black and yellow, the latter beginning beyond basal half of eye; female: pre sutural acrostichal setulae undifferentiated from the ground setulae ........................................................ 3

3.

One proepistemal seta weak, about three-quarters as long as upper anepistemal setae; male: stemite 5 without sharp depression on posterior margin (Fig. 127); cercal plate heart-shaped (Fig. 128) (Brazil) ................................... S. paraensis Carvalho, 1999 One proepistemal seta strong, similar to the upper anepistemal setae; male: stemite 5 with sharp depression on posterior margin (Figs. 129, 130); cercal plate rounded (Figs. 131, 132) ................................................................................................... 4

4.

Species ranging from 8.0-9.0 mm in length; posterior katepistemal seta strong, about twice the length of the anterior one; male: both frontal setae cruciate; stemite 5 as in Fig. 129; cercal plate as in Fig. 131 (Peru, Bolivia) ........... S. singularis (Stein, 1919) Species ranging from 6.5 to 8.0 mm in length; posterior katepistemal seta very strong, about three times as long as the anterior one; male: lower frontal setae cruciate, upper reclinate; stemite 5 as in Fig. 130; cereal plate as in Fig. 132 (Brazil) .............. ..

................. ........... ..... ................ ................. .................... .S. sulina Carvalho, 1999


t

127


~3 1 mm

1 mm

't', ••':.:::."

.•••

O,5mm

126

Figs. 121-126, characters of Souzalopesmyia (modified from ALBUQUERQUE. 1951): 121. S. carioca. head. frontal view; 122. S. carioca. hind leg. anterior view; 123. S. amazonica. male wing; 124. S. carioca. female; 125. S. carioca, ovipositor; 126, S. amazonica, ovipositor.

128



129 130

Figs. 127-132, characters of Souzalopesmyia (modified from CARVALHO, 1999): 127, S. paraensis, sternite 5; 128, S. paraensis, cereal plate; 129, S. singularis, sternite 5; 130, S. sulina, sternite 5; 131, S. singularis, cereal plate; 132, S. sulina, cereal plate.


129


Table 1. Subfamilies, tribes and genera of basal Muscidae, including the number of species in Brazil, South America and the Neotropical Region. Subfamily

Genus

Tribe

Brazil

South

Neotropical

America Atherigoninae Muscinae

Muscini

Stomoxyini

Azeliinae

Azeliini

Reinwardtiini

Phaoniinae

Atherigona Biopyrellia Cyacyrtoneura Dasymorellia Morellia Musca Neomyia Neorypellia Parapyrellia Polietina Sarcopromusca Xenomorellia Haematobia Neivamyia Stomoxys Azelia Drymeia Hydrotaea Micropotamia Ophyra Potamia Thricops Brachygasterina Chaetagenia Correniosia Dalcyella ltatingamyia Muscina Palpibracus

I

I

I

I

I

I

I

2

7

9

14

17

1

I

19 2

I

I

I

I

I

I

2 13

2 15

2 18

2 2

2 2

2 2

I

I

I

3

4

4

1

1

I

0

1

I

0 3 4

0 7 3 6

8 4 7

0

2

2

0

0

I

0 I

3 I

3 I

I

I

2

I

I

I

I

I

I

I 4

17

I

I

0

2 17

Phi/om is

29

38

49

Psilochaeta Reinwardtia Synthesiomyia Dolichophaonia stat.rev. Helina

3

5

5

I

I

I

I

I

I

15

22

27

17

61

84

16

36

42

Phaonia Souwiopesmyia Total

I

1

37 genera

3

5

5

144

271

332

References (see end of Part II) 130


,

Claudio Jose Barros de Carvalho & Marcia Souto Couri ,,

Plate 1

•

•

•

•

Morellia humeralis (Stein, 1918)

Polietina nigra Couri & Carvalho, 1996

(6mm)

(7 mm)

•

Xenomorellia montanhesa Albuquerque, 1952

Palpibracus spicatus (Malloch, 1934)

(7 mm)

(5 mm) •

Muscidae (Diptera) ofthe Neotropical Region: TAXONOMY

131

•


Plate 2

•

•

I

..

..

Philornis trinitensis Dodge & Aitken, 1968 . (7 mm)

L Psilochaeta chlorogaster (Wiedemann, 1830) (6mm)

•

I I

Dolichophaonia gallicola (Albuquerque, 1958) (8mm)

Helina longipila (Stein, 1918) (8 mm) •

132


•

PART II

APICAL GROUPS By Marcia Souto Couri* & Claudio Jose Barros de Carvalho**

INTRODUCTION Part II of this book deals with the remaining three subfamilies of Muscidae Cyrtoneurininae, Mydacinae and Coenosiinae - considered as apical groups within Muscidae, in particular the two last subfamilies (CARVALHO, 1989d). As in Part I, the main scope of this section is to provide means of identifying the species included in these subfamilies. However, one of the difficulties to reach this goal is that some species could not be included in the identification keys, especially in the genera for which a thorough revision is needed. On the other hand, an indirect goal of this chapter, and for extension of this book as a whole, is to identify weak points in our knowledge of muscid taxonomic and diversity for the neotropics (e.g. taxonomic problems, need of generic revisions, need of more collecting in some groups with few known species, among others). CARVALHO et al. (1993) considered Cyrtoneurininae as a Muscidae subfamily, although recognising that' 'this is not a natural group, and several genera have already been transferred

* **

Museu Nacional do Rio de Janeiro, Departamento de Entomologia, Quinta da Boa Vista, Rio de Janeiro, 20.940040, Brazil; e-mail: [email protected] Universidade Federal do Parana, Departamento de Zoologia, C.P. 19020, Curitiba, 81.531-980, Parana, Brazil; email: [email protected]


133

Part II - Apical Groups

to other subfamilies". Following this and considering the taxonomic changes and the new taxa described, from the last catalogue until now, the subfamily includes 11 genera and about one hundred species in the Neotropical Region. Cyrtoneuropsis Malloch and Neomuscina Townsend are, by far, the largest genera of Cyrtoneurininae, housing together more than 60% of the species of the subfamily. Three genera remain monobasic - Arthurella Albuquerque, Chortinus Aldrich and Neurotricha Shannon & Del Ponte. Some Cyrtoneurininae show a very interesting biology - e.g., Charadrella malacophaga Lopes breeds in dead gastropods (CARVALHO, 1980, 1985a). Viviparity has been recorded in some genera. The other two subfamilies seem to be close related, although this has not been clearly established. Mydaeinae has been traditionally treated as a separate subfamily, especially based on adult characters (CARVALHO, 1989d). SKIDMORE (1985), however, did not find characters from the larva morphology to "readily separate" them from the Reinwardtinae. Mydaeinae comprises seven genera, including Mydaea, a very speciose and poorly studied genus in South America. The Coenosiinae is the most speciose subfamily of Muscidae, housing over than 40% of all known muscid species in the Neotropical region. Two tribes are included in this subfamily - Limnophorini and Coenosiini. CARVALHO (1989d) provided evidence for the monophyly of the Coenosiini, although only two genera were included - Bithoracochaeta andNeodexiopsis. COURI & PONT (2000) proposed a worldwide classification for the genera of Coenosiini, using cladistic methodology. In their analysis, the monophyly of Coenosiinae was supported mainly by absence of a prealar seta, while the monophyly of Coenosiini was supported by the number and arrangement of the katepistemal setae (three setae forming an equilateral triangle), the dichoptic male frons, the presence of only one pair of reclinate orbital seta in both sexes, the presence of usually only one anterodorsal setae on hind tibia, and the elongated, tube-like hypandrium (COURI & PONT, 2(00). As a result of their analysis, some taxonomic changes were proposed, including the removal of six genera to tribe Lirnnophorini, four of them with occurrence in the neotropics - Agenamyia Albuquerque, Drepanocnemis Stein, Pachyceramyia Albuquerque and Rhabdotoptera Stein. Talking these changes into consideration, Coenosiini now comprises 16 genera, being Neodexiopsis Malloch the largest one and Limnophorini, 12 genera. About 43% of 28 genera of Coenosiinae found in South America are not represented in Brazil. The material and methods follow the Part I of this book, including the terminology and geographical distribution. 134


Marcia Souto Couri & Claudio Jose Barros de Carvalho

DIAGNOSIS AND KEY TO SPECIES The classification adopted here is the one proposed by CARVALHO et al. (1993), including the taxonomic updates and changes published posteriorly, especially the ones recently proposed by COURI & PONT (2000) on Coenosiini. Table 1 (page 259) summarizes the genera and species of the subfamilies Cyrtoneurininae, Mydaeinae and Coenosiinae occurring in Brazil, South America and the Neotropical Region.

Subfamily Cyrtoneurininae Snyder, 1954 Diagnosis. Male: dichoptic or holoptic; anepimeron setulose in most genera; hind tibia usually with no calcar; female: fronto-orbital seta absent. Monophyly. This is a polyphyletic subfamily (COURI & CARVALHO, 1997b). HENNIG (1965) stated that the Cyrtoneurininae could constitute a monophy letic group and several authors in the Neotropical Region followed his concept. However CARVALHO et al. (1993), despite including this subfamily in their catalogue for the Neotropical Region, did not recognised it as a natural group. Comments. After HENNIG's (1965) consideration about the monophyly of the Cyrtoneurininae, PONT (1972) included 15 genera in this subfamily but posteriorly some of them were excluded. Currently, there are 12 genera and 108 species in the Neotropical Region.

ArthurellaAlbuquerque, 1954 Type-species. Arthurella nudiseta Albuquerque (orig. des.). Diagnosis. Male holoptic; arista bare; pro sternum, anepimeron and me ron setulose; dorsocentrals 2:4; presutural acrostichals 2; intra-alars 2; katepisternals 2:2; vein R j with dorsal setulae on middle third; base of vein R 4+5 setulose on both surfaces; veins R 2+3 and R4+5 divergent apically; mid tibia with 4 setae on posterior surface; hind tibia with calcar; sternite 1 bare. References. ALBUQUERQUE, 1954a; LOPES, 1985a. Monophyly. Not yet established. Geographical record. Chile. Comments. Monobasic genus: Arthurella nudiseta Albuquerque, 1954. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

135

.~


ARAUJO & COURI (1993) redescribed this species, including the description ofthe puparium and suggested that the species might belong in the Azeliinae, Reinwardtiini.

Cariocamyia Snyder, 1951 (Plate 1) Type-species. Cariocamyia maculosa Snyder (orig. des.). Diagnosis. Facial border with developed setulae that continue above the vibrissae up to the basal quarter of the antenna; male: strong pair of reclinate setae opposite to the anterior ocellus; female: interfrontal setae present; prosternum with numerous setulae over a large portion of the lateral area; proepisternum, anepimeron and suprasquamal ridge bare; meron with setulae only on the subtriangular portion above hind coxa; posterior thoracic spiracle with fine black hairs; vein R 2+3 setulose on both surfaces from node to midway to crossvein r-m; vein R4+5 with cilia on both surfaces from node to half of distance to crossvein r-m, strongly curved forward; lower calypter not truncate apically; mid femur with a short, subapical, anterodorsal seta; hind tibia without calcar and hind coxa bare above; female: sixth abdominal spiracle present Reference. SNYDER, 1951. Monophyly. One of the synapomorphies for Muscidae is the abdomen offemale showing less than 7 pairs of spiracles. This reduction in the number of spiracles is a unique apomorphy among the Calyptratae. Spiracles 6 and 7 were lost, but some genera, not related amongst themselves have reacquired the spiracle 6. Cariocamyia (see VOCKEROTH, 1972) has six spiracles, a condition acquired independently and considered apomorphic for the genus (see MICHELSEN, 1991). Only Achanthiptera Rondani, a genus apparently not phylogenetically related to Cariocamyia, has also reacquired the spiracle 6. This is a monobasic Palaearctic genus. Geographical record. Colombia, Brazil. Comments. Only two species occur in the region, both in South America.

Key to species 1.

Hind tibia on anteroventral surface with one seta (Colombia, Brazil) ........... .

...................................................................................... C. maculosaSnyder, 1951 136

Muscidae (Diptera) of the Neotropica1 Region: TAXONOMY


Hind tibia on anteroventral surface with two setae (Brazil) .............. .

...................................................... .............................. C. vibrissata (Stein, 1918)

Charadrella Wulp, 1896 (Figs. 1-9, Plate 1) Type-species. Charadrella macrossoma Wulp (mon.). Diagnosis. Male dichoptic; eyes with sparse, short and fme yellowish cilia; antenna long, reaching epistoma; arista plumose; the 2 upper pairs of frontal setae backwardly directed; dorsocentrals 3:3; acrostichals not differentiated; prealars 1 or 2; anepimeron setulose; proepimeron setulose; lower calypter enlarged reaching the area below scutellum; veins M and R4+5 divergent apically; calcar almost undifferentiated; sternite 1 setulose. References. CARVALHO, 1980, 1985a. Monophyly. Although the monophyly of C haradrella has not yet been established, a series of characters seems to support its monophyly, like the configuration of the head in both sexes and the shape of the lower calypter. Geographical record. Mexico, Panama, Trinidad, Tobago, Ecuador, Brazil. Comments. A very characteristic genus with three species in the Neotropical Region. Charadrella macrossoma, whose type-locality is in Yucatan, Mexico, has the largest geographical distribution within the genus, occurring from southern Mexico to Ecuador.

Key to species 1.

Male: femora yellow; fore and mid tibia light brown on apical third; sternite 5 triangular (Fig. 4); female: hypoproct rectangular (Fig. 5) (Brazil) ................ .

...................................................... ....................... C. albuquerquei Carvalho, 1985 Male: femora dark brown on basal third; fore and mid tibia dark brown on apical third; sternite 5 rectangular (Figs. 1,2); female: hypoproct triangular (Fig. 3) ............ .2 2.

Wmgs with a spot on apex of vein M (Fig. 6); male: cercal plate without median incision and with a prolongation on posterior margin (Fig. 7) (Mexico, Guatemala, Panama, Ecuador


137


Trinidad, Tobago, Venezuela, Guyana)................................ C. macrossoma Wulp, 1896 Wings without a spot on apex of vein M (Fig. 9); male: cereal plate with a deep median incision on posterior margin (Fig. 8) (Brazil) .... C. malacophaga Lopes, 1938

0,1 mm

0,1 mm

M

sur

0,1 mm

1 mm Figs. 1-9, characters of Charadrella (modified from CARVALHO, 1981, 1985a): I, C. malacophaga, male, sternite 5, dorsal view; 2. C. macrossoma, male, sternite 5, dorsal view; 3, C. malacophaga, female, hypoproct and cerci, dorsal view; 4, C. albuquerquei, male, sternite 5; 5, C. albuquerquei, female, hypoproct and cerci; 6, C. macrossoma, wing, dorsal view; 7, C. macrossoma, male, cerci and surstyli; 8, C. malacophaga, male, cerci and surstyli; 9, C. malacophaga, wing, dorsal view. Abbreviations: c pI, cereal plate (cerci); M, vein; sur, surstylus.

138



Chortinus Aldrich, 1932 Type-species. Chortinus bequaerti Aldrich (orig. des.). Diagnosis. Eyes with short and sparse cilia; male with enlarged anterointemal ommatidia; male holoptic; arista with short setulae; acrostichals 0: 1; dorsocentrals 2:4; intra-alars 0:2; prealar strong; scutellum bare on lateroventral margin; prostemum bare; propleuron bare; anepistemun, katepistemum and anepimeron with long cilia; veins R j and R4+5 with dorsal and ventral setulae; veins R4+5 and M almost parallel apically; stem vein on dorsal surface with short setulae on basal third; hind tibia on anteroventral, anterodorsal and posterodorsal surfaces with one short seta on apical third; female: ovipositor short. Reference. PAMPLONA & COURI, 1995a. Monophyly. PAMPLONA & COURI (199Sa) presented a key to segregate Chortinus from Cyrtoneurina, but did not indicate the monophyly of the genus. Geographical record. Guatemala, Costa Rica, Panama. Comment: Monobasic genus: Chortinus bequaerti Aldrich, 1932 (validated by PAMPLONA & COURI, 1995a).

Cyrtoneurina Giglio-Tos, 1893 (Fig. 10) Type-species. Cyrtoneurina uber Giglio-Tos (des. COQUILLET, 1910). Diagnosis. Male eyes with enlarged anterointemal ommatidia; arista with long cilia (12 to 19 cilia); dorsal secondary cilia short (Fig. 10), absent in C. arleriopsis; acrostichals 0: 1; dorsocentrals 2:4; intra-alars 0: 1; intrapostalar seta present; scutellum with setulae on lateroventral margin; prostemum bare or with a very fine lateral pubescence; propleuron bare; katepistemals 1:2; anepimeron setulose; vein R1 on ventral surface bare; vein R 4+5 with cilia on dorsal (except in C. costalis) and ventral surfaces (except in C. costalis and C. crispaseta); ovipositor long, epiproct with 2 setae, tergites slender. Reference. PAMPLONA, 1999. Monophyly. According to PAMPLONA (1999), Cyrtoneurina is monophyletic based on the following synapomorphies: presence of microtrichia on the membrane between the segments 6-7 and 7 -S. Geographical record. Mexico, Belize, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Grenada, Trinidad, Colombia, Venezuela, Guyana, Peru, Brazil. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

139


Comments. COURI & PAMPLONA (2000) reevaluated the position of C alifusca Couri, 1982 and kept it in this genus.


Vein R] with dorsal cilia; entire palpus yellow......................................................... 2 Vein R] bare on dorsal surface; palpus brown, at least on basal half.. ....................... 3

2.

Hind femur on anterodorsal surface with a series of setae; anteroventral surface with 2 setae on apical third; posterodorsal surface with one preapical seta (Brazil) .......................................................................................... C. alifusca Couri, 1982 Setae on hind femur distributed not as above (peru, Brazil) .... C crispaseta Snyder, 1954

3. Vein R 4+5 bare on ventral surface; hind tibia on middle third of posterodorsal surface without setae (Brazil) ....................................................... C costalis (Walker, 1853) Vein R 4+5 with cilia on ventral surface; hind tibia on middle third of posterodorsal surface with one seta ............................................................................................. 4 4.

Prealar seta absent; hind femur on basal third of ventral surface with one strong seta with round apex (?Guatemala or Panama - La Pita) ........ C geminata (Stein, 1904) Prealar seta present; hind femur on basal third of ventral surface without one strong seta with round apex ............................................................................................... 5

5.

Arista with up to 12 cilia; secondary dorsal cilia absent; palpus with apical half yellow and basal half brown (Brazil) ........................................... C arleriopsis Couri, 1982 Arista with more than 12 cilia; secondary dorsal cilia present; palpus entirely brown ....... 6

6.

Body length about 4.0 mm; mid tibia on basal half of posterior surface with a complete series of short setae (Brazil) .................................................. C biseta Snyder, 1954 Body length about 5.0 to 6.0 mm; mid tibia on basal half of posterior surface without a series of short setae ............................................................................................ 7

140



7.

Scutellum on basal half of lateroventral margin with up to 5 cilia; prostemum bare; arista with about 15 cilia (Costa Rica, Panama, Brazil) ......... c. confusa Snyder, 1954 Scutellum on basal half oflateroventral margin with more than 5 cilia; pro sternum with a very fine and inconspicuous pubescence; arista with about 19 cilia (Mexico, Guatemala, El Salvador, Belize, Nicaragua, Panama, Grenada, Trinidad, Colombia, Peru, Venezuela, Guyana) .................................................... c. uber Giglio-Tos, 1893

Species of Cyrtoneurina not included in the key and its respective geographical distribution Species


C. cylindrica (Macquart, 1846)

Brazil

O,5mm

10 Fig. 10, Cyrtoneurina uber (modified from PAMPLONA, 1999), antenna.

Cyrtoneuropsis Malloch, 1925 (Figs. 11-13) Type-species. Spilogasterveniseta Stein (orig. des.). Diagnosis. Male eye with enlarged anterointernal ommatidia; arista with long cilia (18 to 29 cilia, except C. seriata with 9, C. ocasionalis with 10 and C. spiloptera with 14); dorsal secondary cilia short or long (Fig. 11); acrostichals 0: 1; dorsocentrals 2:4 (2:3 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

141


in C. gluta and C. spiloptera); intra-alars 0: 1; intrapostalar seta absent or present; scutellum with many cilia on latera-ventral margin (few cilia in C. gluta, C. maculipennis and C. walkeri); prosternum setulose or bare, with fine lateral pubescence (c. neotrita and C. ocasionalis); propleuron bare or setulose in C. beebei, C. multomaculata and C. neotrita; katepisternals 1:2; anepimeron setulose; veins R] (except in C. brunnea and C. paUipes) and R4+5 with cilia on dorsal and ventral surfaces; ovipositor short or medium sized, tergites slender or large; sternite 8 with or without strong setae with round apex. References. PAMPLONA, 1999; PONT & PAMPLONA, 2000. Monophyly. According to PAMPLONA (1999), the genus is monophyletic and supported by five synapomorphies: presence of ventral cilia on vein R]; medium to short length of the non-delicate ovipositor; cerci short and flattened; tergites 6 and 7 very sclerotized and hypoproct with a lateral incision. Geographical record. Mexico, Belize, EI Salvador, Guatemala, Nicaragua, Costa Rica, Panama, Trinidad, Cuba, Puerto Rico, Colombia, Venezuela, Guyana, Suriname, Ecuador, Bolivia, Brazil, Argentina. Comments. PONT & PAMPLONA (2000) synonymized Paracyrtoneuropsis Pamplona under Cyrtoneuropsis. COURI & PAMPLONA (2000) transferred Cyrtoneurina brunnea and Cyrtoneurina pallipes to this genus and they are included in the key below.


Male: prealar seta absent or, if present, only slightly longer than ground setulae (Fig. 12); female: interfrontal seta absent... ..................................................................... 2 Male: prealar seta present and strong; female: interfrontal seta present... .................. 17

2.

Fore tibia with one seta on middle third; dorsocentrals 2:3 (Mexico, Guatemala, Guyana, Costa Rica, Panama, Canal Zone, Venezuela) ....... c. gluta (Giglio-Tos, 1893) Fore tibia without one seta on middle third; dorsocentrals 2:4 (the posterior one very short in C.fuscicosta) ............................................................................................ 3

3. 142

Pro sternum setulose ................................................................................................ 4



Prosternum bare .................................................................................................. 15 4.

Male ................................................................................................................... 5 Female .............................................................................................................. 11

5.

Hind tibia on ventral surface, at least, with one seta ................................................ 6 Hind tibia on ventral surface without a long submedian seta .................................... 7

6.

Hind tibia on ventral surface with one long submedian seta (Panama, Trinidad, British Guyana, Brazil) ................................................................ c. armipes (Stein, 1911) Hind tibia on ventral surface with three setae on middle third (Paraguay) ........................................................................................ C. paUipes (Stein, 1918)

7.

Wings without a brown cloud between apex of veins Sc and R J ; hind tibia on posterior to posteroventral surface with at most one seta ...................................................... 8 Wings with a brown cloud between apex of vein Sc and R J (sometimes faint); hind tibia on posterior to posteroventral surface with long setae or cilia ........................... 9

8.

Katepisternum with short and sparse ground setulae (Fig. 13); hind tibia on posterior surface with a series of setae (Mexico, Guatemala, Belize, El Salvador, Costa Rica, Panama, Colombia, Peru, Venezuela, Brazil, Paraguay) ............... c. gem ina (Wiedemann, 1830) Katepisternum bare; hind tibia on posterior surface without a series of setae (Guatemala, Belize, Nicaragua, El Salvador, Costa Rica, Panama, Grenada, Trinidad, Colombia, Peru, Venezuela, Guyana, Brazil?) ................................... C. immunda (Stein, 1911)

9.

Katepimeron with one seta and one setula; pal pus yellow; arista with about 9 cilia (Colombia, Peru, Brazil) ..................................................... C. seriata (Stein, 1911) Katepimeron bare; palpus brown, at least basally; arista with more than 20 cilia.......... 10

10. Hind tibia on anterodorsal surface with 2 setae on middle third; dm-cu straight; pro sternum bare (Peru, Guyana, Brazil) ............................. C. conspersa (Stein, 1911) Hind tibia on anterodorsal surface with 1 seta on middle third; dm-cu sinuose; Muscidae (Diptera) of the Neotropical Region: TAXONOMY

143


prostemum setulose (Brazil) ................................... C.jlaviantennata (Couri, 1982) 11. Wings without a brown cloud between apex of veins Sc and R J ................................ 12 Wings with a brown cloud between apex of veins Sc and R J ( sometimes faint) ............ 13 12. Vein R J with cilia (Mexico, Guatemala, Belize, Salvador, Costa Rica, Panama, Colombia, Peru, Venezuela, Brazil, Paraguay) ..............................

c. gemina (Wiedemann, 1830)

Vein R J without cilia (Paraguay, Brazil) ................................... C. paZZipes (Stein, 1918) 13. Mesonotum, when viewed posterolaterally, with an conspicuous golden pruinescence, and without vittae (Peru, Guyana, Brazil) .......................... C. conspersa (Stein, 1911) Mesonotum, when viewed posterolaterally, with grey pruinescence, and with vittae .... 14 14. Arista with 8.bout 9 cilia; prealar seta absent; fore coxa and tibia brown (Colombia, Peru, Brazil) ..........................................................................

c. seriata (Stein, 1911)

Arista with about 20 cilia; prealar seta present; fore coxa yellow and fore tibia yellowish on basal half (Panama, Trinidad, Guyana, Brazil) ................

c. armipes (Stein, 1911)

15. Wings with brown clouds from base to apex of costal vein, on crossveins r-m and dm-cu (Guyana, Brazil) .................................................. C.juscicosta (Curran, 1934) Wings without brown clouds on costal vein, cross veins i·-m and dm-cu .................... 16 16. Presutural dorsocentral setae with similar length; male: hind tibia on ventral to posteroventral surface with one long submedian seta and with a series of shorter setae (Mexico, Guatemala, El Salvador, Belize, Panama, Colombia) ....................................

c. steini (Snyder, 1954)

Anterior presutural dorsocentral seta longer than the second; male: hind tibia on ventral to posteroventral surface without a long submedian seta and without a series of shorter setae (Mexico, Guatemala, Belize, El Salvador, Costa Rica, Panama, Colombia, Venezuela, Peru, Brazil, Paraguay) .............................

c. gemina (Wiedemann, 1830)

17. Prosternum setulose ............................................................................................ 18 Prostemum bare or with a very fine and inconspicuous pubescence .......................... 23 144



18. Postsutural dorsocentrals 3 (Belize, Panama, Suriname, Ecuador, Peru, Bolivia, Brazil)

........................................................................... c. spiloptera (Wiedemann, 1830) Postsutural dorsocentrals 4 ................................................................................. 19 19. Propleuron bare .................................................................................................. 20 Propleuron ciliated ................................................................................................ 22 20. Stem vein bare; anepimeron ciliated only posteriorly (Panama) ...................... .

...................................................... ................................. C. mimica (Snyder, 1954) Stem vein ciliated; anepimeron entirely ciliated...................................................... .21 21. Vein R J with cilia after the humeral crossvein (Guatemala, Belize, Costa Rica, Panama, Peru, Guyana, Brazil) ............................................... ...........

c. dubia (Snyder, 1954)

Vein R J without cilia after the humeral crossvein (Brazil; Argentina) ............... . ..................................................................................... C. brunnea (Hough, 1900) 22. Lateroventral margin of scutellum with many short cilia reaching its apex; stem vein ciliated on ventral surface at level of humeral vein (?Costa Rica, Colombia, Peru, Guyana, Brazil, Bolivia) ............................................ c. multomaculata (Stein, 1904) Lateroventral margin of scutellum with many short cilia restricted to its basal half; stem vein bare on ventral surface at level of humeral vein (Brazil) .................. .

.................................................................................. C. protosetosa (Snyder, 1954) 23. Propleuron ciliated ............................................................................................. 24 Propleuron bare .................................................................................................. 25 24. Hind tibia on posterodorsal surface with one seta (Peru) ..... c. neotrita (Snyder, 1954) Hind tibia on posterodorsal surface without setae (Trinidad, Guyana) .......... .

.................................................................... ..................... C. beebei (Curran, 1934) 25. Postsutural dorsocentrals 3 (Brazil) ..................................... C. walkeri (Pont, 1972) Postsutural dorsocentrals 4 ................................................................................ 26 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

145


26. Hind coxa ciliated on posterior surface (Mexico, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Peru, Bolivia, Brazil, Argentina)

...................................................... .................................. C. veniseta (Stein, 1904) Hind coxa bare on posterior surface ....................................................................... 27 27. Prealar setae 2; arista with about 10 cilia (Brazil) ............. C ocasionalis (Couri, 1982) Prealar setae 1; arista with more than 16 cilia ...................................................... 28 28. Stem vein with cilia on ventral surface (Costa Rica, Panama, Guyana, Bolivia, Brazil)

...................................................... ..................... C. maculipennis (Macquart, 1843) Stem vein bare on ventral surface ........................................................................... 29 29. Stem vein without cilia on dorsal surface ................................................................ 30 Stem vein with cilia on dorsal surface ................................................................... 37 30. Wings without clouds on apex of vein R 2+3 ; hind femur on posteroventral to ventral surface with, at least, 2 strong basal setae ............................................................. 31 Wings with a brown cloud on apex of vein R 2+3; hind femur on posteroventral to ventral surface without strong basal setae ............................................................. 32 31. Pedicel brown; legs dark brown (Brazil) ....................... C pararescita (Couri,1995) Pedicel yellow; legs light brown (Mexico, Guatemala, Belize, Nicaragua Costa Rica, Panama, Cuba, Puerto Rico, Trinidad, Colombia, Ecuador, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay) ............................................. ...... C rescita (Walker, 1861) 32. Halter yellow at least at knob .............................................................................. 33 Halter brown, with knob darker........................................................................... 35 33. Calypteres entirely whitish; arista with about 18 cilia; hind femur, on ventral surface without setae (Brazil) .............................................. .......... C similata (Couri, 1982) Calypteres brown or at least lower one with brown margins; arista with more than 21 146



cilia; hind femur, on ventral surface with 3-4 setae ................................................ 34 34. Hind tibia on anterodorsal surface with one seta on middle third; arista with dorsal secondary cilia short; eyes bare (Brazil) .......................... C incognita (Snyder, 1954) Hind tibia on anterodorsal surface with 2 setae on middle third; arista with long dorsal secondary cilia; eyes with short and sparse cilia (difficult to observe) (Mexico, Guatemala, El Salvador, Costa Rica, Panama, Brazil, Paraguay) ................... .

...................................................... .............................. C. polystigma (Wulp, 1896) 35. Wing without brown clouds on apex of costal vein and on crossvein dm-cu (Panama, Grenada, Trinidad, Peru, Brazil) ................................... C praenubila (Snyder, 1954) Wing with a brown cloud on apex of costal vein and on crossvein dm-cu .............. 36 36. Humeral callus yellowish; calypteres white or yellowish; costal vein with an elongated brown cloud along its entire extension (Mexico, Nicaragua, Costa Rica, Panama, Trinidad) ............................................. .............................. C wulpi (Snyder, 1954) Humeral callus brownish; calypteres light brown with darker margins; costal vein with a brown cloud only on its apical half (Brazil) ............. Cjitscisquama (Snyder, 1954) 37. Pleura and fore leg yellow..................................................................................... 38 Pleura and fore leg light brown .............................................................................. 39 38. Proepimeral cilia yellow; wing with a brown cloud only on apex of vein R) (Brazil, Paraguay) ............................................. .............................. C mellina (Stein, 1918) Proepimeral cilia brown; wing with a brown cloud on upper margin from vein Sc up to wing apex (Brazil) ........................................................... C varicolor(Hough, 1900) 39. Anepimeron with long cilia; vein R 2+3 with a brown cloud; arista with long dorsal secondary cilia (Mexico, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Peru, Bolivia, Brazil, Argentina) ............ .

.......................................................................................... C. veniseta (Stein, 1904)


147


Anepimeron with short cilia; vein R 2+3 without a brown cloud; arista with short dorsal secondary cilia (Mexico, Guatemala, El Salvador, Costa Rica, Panama, Guyana, Argentina) ..................................................... ............

c. inuber(Giglio-Tos, 1983)

Species of Cyrtoneuropsis not included in the key and its respective geographical distribution Species


C. monstrata (Wulp, 1896)

Mexico

1 mm

c Figs, 11-13, characters of Cyrtoneuropsis (modified from PAMPLONA, 1999): II, C. dubia, antenna; 12, C. dubia, thorax, lateral view; 13, C. gemina, katepisternum. Abbreviation: c, lower calypter.

148



Mulfordia Malloch, 1928 Type-species. Mulfordiajerruginea Malloch (orig. des.). Diagnosis. Female: inner vertical setae long and strong, outer pair very short; dorsocentraI2:3; prealar absent; suprasquamal ridge setulose; katepisternal setae 1: 1; pro sternum and center of anepimeron bare; posterior spiracle with a few black hairs on hind margin; meron with fine hairs on center; sternite 1 bare; hind coxa setulose above on posterior surface; hind tibia without a posterodorsal seta; vein R4+5 with few short black setulae on its base, on ventral surface. Reference. MALLOCH, 1928. Monophyly. Not yet established. Geographical record. Bolivia, Peru. Comments. CARVALHO et aI. (1993b) placed M. longipes (Stein, 1911) tentatively in Mulfordia. This species was included in the key.

Key to species

1.

Veins R4+5 and M sub-parallel apically (Peru) .......................M. secunda Snyder, 1951 Vein M slightly curved forward .............................................................................. 2

2.

Pedicel and palpus fuscous; wings yellowish hyaline with yellow veins (Bolivia)

.................................................................................. .M. jerruginea Malloch, 1928 Entire antenna greyish black; palpus black; wings infuscated (Bolivia)

......................................................................................... .M. longipes (Stein, 1911)

Neomuscina Townsend, 1919 (Figs. 14-15) Type-species. Musca scutellaris Fabricius (orig. des.) =tripunctata Wulp. Diagnosis. Eyes bare; arista plumose; anepimeron with discal cilia; prosternum, proepimeron and proepisternum bare; postalar wall bare; supra and infra squamal ridges bare; anterior intra-alar seta absent; female with 2 posterior intra-alar; acrostichals 0: 1 or 1: 1; calcar absent; hind coxa on posterior surface bare; vein R 2+3 bare or with few cilia at Muscidae (Diptera) of the Neotropical Region: TAXONOMY

149


node, on dorsal and ventral surfaces; apical portion of stem vein setulose (Fig. 14) on ventral surface and usually with one or 2 setulae on dorsal surface; vein M curved forward and ending at or close to wing apex; vein R4+5 ending well before wing apex (Fig. 15). Reference. SNYDER, 1954a. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, Costa Rica, Panama, Trinidad, Cuba, Guyana, Colombia, Venezuela, Peru, Brazil, Paraguay, Argentina; United States of America. Comments. Neomuscina needs an extensive revision. Except for SNYDER (1954a), there is no revisional paper and a few new species have been described after his work. At the end of the following key, there is a list of species that were not included in it. COURI & PAMPLONA (2000) transferred Cyrtoneurina scutellata Johnson to this genus.

Key to species (modified from SNYDER, 1954a) 1.

Vein R 4+5entirely bare ............................................................................................. 2 Vein R4+5 with one or more hairs on ventral surface beyond the node .................... 28

2.

With one or more pairs of distinct presutural acrostichal setae .................................. 3 Without distinct presutural acrostichal setae .......................................................... 13

3.

Meron with hairs or setulae below the spiracle (Mexico, Puerto Rico, Venezuela; southern United States of America.) .............................. .N. tripunctata (Wulp, 1896) Meron bare below spiracle .................................................................................... 4

4.

Mid femur with two strong preapical, dorsal to posterior setae ............................ . ...... ...... .......... .. (Brazil, Argentina) N. zosteris (Shannon & Del Ponte, 1926) [in part] ................................................................. (Brazil) N. ponti Lopes & Khouri, 1995 Mid femur with three strong preapical, dorsal to posterior setae ............................. 5

5.

150

Hind tibia with two median anterodorsal setae ....................................................... 6



Hind tibia with one median anterodorsal seta.......................................................... 8 6.

Mid and hind femora fulvous on basal half or more (Venezuela) ................ .

.......................................... .......................................... .N. nudistigma Snyder, 1949 Mid and hind femora fulvous, with the possible exception of extreme base ............... 7 7.

Male: margin of upper calypter broadly infuscated; eyes facets not conspicuously enlarged above and in front; diameter of anterior ocellus subequal to diameter of one of the posterior ocelli; female: palpus conspicuously enlarged near apex, this apical portion 1.7 times the basal diameter (Mexico) .............. .N. vecta (Giglio-Tos, 1893) Male: margin of upper calypter not broadly infuscated; eyes facets conspicuously enlarged above and in front; diameter of anterior ocellus approximately two times that of one ofthe posterior ocelli; female: palpus not conspicuously enlarged apically (Mexico ) ..................................................... .............. .N. paralis (Giglio-Tos, 1893)

8.

Base of femora infuscated, usually one or more pairs of coxae with dark streaks ...... 9 Base offemora and all coxae entirely yellow to fulvous ......................................... 11

9.

Crossveins not surrounded by dark clouds ........................................................... 10 Crossveins surrounded by extensive, infuscated c1ouds ....................................... .23

10. Male: intra-alars 2; eye facets not conspicuously enlarged above and in front (Mexico, Costa Rica, Panama, Venezuela, Brazil) ....................... ..N. dorsipuncta (Stein, 1918) Male: intra-alar 1; eyes with an area of conspicuously enlarged facets above and in front (Venezuela) ........................................... ..................... N. macrops Snyder, 1949 11. Without dark accessory setulae adjacent to either the prothoracic or stigmatal seta (Brazil) .............................................. ............................. N. sanespra Snyder, 1949 With dark accessory setulae adjacent to both the prothoracic and the stigmatal seta..... 12 12. Dorsocentrals 2:3 (Mexico, Guatemala) ................................ .N. triseta Snyder, 1949 Dorsocentrals 2:4 (Mexico, Guatemala) ....................... N. mexicana (Macquart, 1843) Muscidae (Diptera) of the Neotropical Region: TAXONOMY

151


13. Dorsocentrals 2:3 ................................................................................................ 14 Dorsocentrals 2:4 ................................................................................................ 16 14. Mid femur with one posterodorsal and one posterior preapical setae (Barro Colorado Is., Trinidad) ..................................................................... .N. capalta Snyder, 1949 Mid femur with one dorsal, one posterodorsal and one posterior preapical setae ........ 15 15. Calypteres deep brown, the margins infuscated; hind femur with a complete row of anteroventral setae; vein R4+5 bare on ventral surface (Brazil) ... .N. inflexa (Stein, 1918) Calypteres white to pale yellow, the margins not infuscated; hind femur with only two to three long anteroventral setae at apex; vein R4+5 with a single ventral setulae beyond node (Barro Colorado Is.) .....................................................N. currani Snyder, 1949 16. Mid femur with 2 preapical setae ......................................................................... 17 Mid femur with 3 preapical setae .......................................................................... 23 17. Hind tibia with 1 submedian anterodorsal seta; thoracic pleura mostly fulvous ............ 18 Hind tibia with 2 submedian anterodorsal setae; thoracic pleura mostly infuscated ............................................................................................................................ 21 18. Mid and hind femora infuscated on at least basal half (Brazil) ..................... . .................... ..... .................. ..... ........ .......... ..... ........ ...... .. N. mediana Snyder, 1949 .................................... .................................. N. goianensis Lopes & Khoury, 1995

Mid and hind femora entirely fulvous to light brown ............................................... 19 19. Calypteres and their margins darkened (Brazil, Argentina) ............................ .. .................................................. .N. zosteris (Shannon and Del Ponte, 1926) [in part]

Calypteres and their margins not darkened .......................................................... .20

20. Wings with an uninterrupted dark cloud from apex of vein Sc to apex of vein R 2+3 ; fore tibiae with a short submedian anterodorsal seta (Trinidad, Guyana) ................................................................................... .N. nigricosta Snyder, 1949 152



Wing with a distinct dark spot at apex of vein Sc and a faint spot near apex ofR2+3 , the interspace not darkened; fore tibia without a submedian anterodorsal seta

...................................................................... ..N. transporta Snyder, 1949 [in part] 21. Wings without a trace of a dark cloud along costal margin or surrounding crossveins; hind femur with 2-4 posteroventral setae on basal half only (Costa Rica, Panama)

........................................................................................... .N. atincta Snyder, 1949 Wings with a faint to distinct dark cloud along costal margin and surrounding cross veins r-m and dm-cu; hind femur with a complete row of strong posteroventral setae ...... .22 22. Thorax and femora entirely fulvous (South America) ................................ .. .................... ........ ........................ ...... ............N. transporta Snyder, 1949 [in part]

Thorax and at least the mid and hind femora mostly infuscated (Venezuela) .....................................................................N. nudistigma Snyder, 1949 [in part]

23. Posterior thoracic spiracle with several black setulae in at least the posteroventral comer in addition to the flap-like covering of hairs (Mexico, Guatemala, EI Salvador, Costa Rica, Panama, Venezuela, Guyana, Peru, Brazil, Paraguay) [N. pictipennis (Bigot, 1878) sensu lato] .......................................................................................... 24 Posterior thoracic spiracle without any black setulae in addition to the flap-like covering of hairs ................................................................................................................ 25 24. Thoracic pleurae and femora mostly infuscated (Mexico, Guatemala, EI Salvador, Costa Rica, Panama, Venezuela, Guyana, Peru) .........N. pictipennis nudinervis (Stein, 1918) Thoracic pleura and femora fulvous (Venezuela, Brazil, Paraguay) ................. .. .......... ...................... ...... ..... ............ ..........N. pictipennis pictipennis (Bigot, 1878)

25. Wings without infuscated areas ............................................................................ 26 Wings with infuscated areas ................................................................................ 27 26. The basal three-quarters or more of at least hind femur infuscated; male: intra-alar 1 (Panama, Trinidad, Brazil) ................................................N. instabilis Snyder, 1949 Muscidae (Diptera) of the Neotropical Region: TAXONOMY

153


All femora entirely fulvous; male: intra-alars 2 (Paraguay) .. .N. schadei Snyder, 1949 ................................................................................ (Peru) N. stabilis (Stein, 1911) .......................................................... (Brazil) N. vittoriae Lopes & Khoury, 1995 27. Posteriorfourth offemora infuscated (Trinidad, Venezuela, Guyana) .................... .

....................................................................................... .N. similata Snyder, 1949 Posterior fourth of femora mostly fulvous (Brazil, Paraguay) ........................ .

................................................................................. ... .N. neosimilis Snyder, 1949 ............................. (United States of America, Florida) N. rufoscutella Dodge, 1955 28. Dorsocentrals 2:3 (Panama) .................................. N. currani Snyder, 1949 [in part] Dorsocentrals 2:4 ............................................................................................... 29 29. Costal margin darkened ...................................................................................... 30 Costal margin not darkened (Brazil) .............................. N. atincticosta Snyder, 1949 ................................................................. (Jamaica) N. scutellata (Johnson, 1919) 30. Margins of calypteres infuscated (Brazil) ......................N. tinctinervis (Stein, 1918) Margins of calypteres not infuscated (Colombia, Peru) ........... N. apicata (Stein, 1904)

Species of Neomuscina not included in the key and their respective geographical distribution

154

Species

Geographical Record

N. arcuata (Wiedemann, 1830)

Brazil

N. neotropica Curran, 1934

Guyana

N. sparsiplumata (Stein, 1918)

Brazil

N. tauota Lopes, 1984

Cuba Muscidae (Diptera) of the Neotropical Region: TAXONOMY


15

Figs. 14-15, characters of Neomuscina (modified from SNYDER, 1954a): 14, N. tripullctata, wing base, ventral view; IS, N. tripunctata, wing tip. Abbreviations: sv, stem vein; R4+5 , vein; M, vein.

Neomusciniopsis Albuquerque & Lopes, 1982 Type-species. Neomusciniopsis Iatifrontata Albuquerque & Lopes (orig. des.). Diagnosis. Male with wide frons, eyes separated by no more than 1,2 - 1,3 times the width of antenna; arista with moderately long plumae; dorsocentral setae 2:4; anepimeron and katepimeron bare; lower calypter enlarged, reaching scutellum; katepistemals 1:2, wing veins bare; vein M slightly curved forward apically; hind tibia without calcar; stemite 1 setulose. References. ALBUQUERQUE & LOPES, 1982; COURI et aI. (in press). Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: N. latifrontata Albuquerque & Lopes, 1982. The two previously described species were recently synonymized by COURI et al. (in press).

Neurotrixa Shannon & Del Ponte, 1926 Type-species. Neurotrixa mirata Shannon & Del Ponte (orig. des.) = felsina Walker. Diagnosis. Eyes with short and sparse cilia; arista pubescent; anepimeron partly ciliated; proepimeron and meron bare; intra-alar setae developed; vein R J with dorsal cilia; vein R4+5 with cilia from node to apex; dorsocentrals 2:4; katepistemals 2:2; hind tibia Muscidae (Diptera) of the Neotropical Region: TAXONOMY

155


with one median fine and short seta on anterodorsal, posterodorsal and anteroventral surfaces. Reference. LOPES, 1985b. Monophyly. Not yet established. Geographical record. Brazil, Uruguay, Argentina. Comments. Monobasic genus: Neurotrixafelsina (Walker, 1849).

Pseudoptilolepis Snyder, 1949 (Figs. 16-19) Type-species. Pseudoptilolepisfulvapoda Snyder (orig. des.). Diagnosis. Anepimeron, postalar wall, meron above posterior spiracle and proepimeron setulose; posterior thoracic spiracle with many setae on its superior margin; wings with vein M strongly curved forward apically; vein R 4+5 with weak cilia along its whole extension on ventral surface, on dorsal surface with strong and sparse cilia on its basal half; lower calypter with fine discal setae on dorsal surface; hind coxa with a series of small setae on posterior surface apically; calcar absent; male: aedeagus with the distiphallus bifid. Reference. LOPES & CARVALHO, 1985. Monophyly. The presence of cilia on the dorsal surface of the lower calypter, as well as the bifid aedeagus, suggest that the genus is monophyletic. These characters are rarely seen within the Muscidae, although a bifid aedeagus is also found in Chaetagenia Malloch, perhaps a synapomorphy uniting these two genera (SCHUHLr & CARVALHO, in press). Geographical record. Costa Rica, Nicaragua, Panama, Venezuela, Brazil, Paraguay. Comments. All the six described species occur in the Neotropical Region. The type material of P. latipalpis (Stein, 1918) was not examined, but it was included in the key based on characters mentioned in the original description.

156



Key to species (modified from LOPES & CARVALHO, 1985) 1.

Postsutural dorsocentral setae 4 (Brazil) ................ ............ P. latipalpis (Stein, 1918) Postsutural dorsoeentral setae 3 ............................................................................ 2

2.

Proepistemum with diseal yellow hairs .................................................................... 3 Proepisternum bare ............................................................................................... 4

3.

Male with stemite 5 longer than wide (Fig 16); cercal plate with a upper incision that ends before the middle of the plate (Fig. 17) (Brazil) ........................ . ....................................................................... P. jluminensis Albuquerque, 1954

Male with stemite 5 wider than long (Fig. 18); cereal plate with a long upper incision that ends beyond the middle of the plate (Fig. 19) (Nicaragua, Venezuela, Brazil) .................................................................. ...................... P.fulvapoda Snyder, 1949

4.

Posterior femur yellowish brown; lowercalypteryellowish brown with darker margins (Nicaragua, Costa Rica, Panama, Venezuela, Brazil) .......... P. nigripoda Snyder, 1949 Femora yellow; lower calypter yellowish ............................................................... 5

5.

Fore tibia with an apical anterodorsal and dorsal seta subequal or equal to tibial width (Brazil, Paraguay) ...................................................... .P. nudapleura Snyder, 1949 Fore tibia without an apical anterodorsal seta or, if present, short, shorter than the dorsal apical and tibial width (Costa Riea, Venezuela, Brazil).............. P. confusa Snyder, 1949


157


17

O,2mm

O,2mm

O,2mm O,2mm Figs. 16-19, characters of Pseudoptilolepis (modified from LOPES & CARVALHO, 1985): 16, P. jluminensis. sternite 5. male; 17, P. jluminensis, cereal plate and surstyIi; 18, P. julvapoda, sternite 5, male; 19, P. julvapoda, cereal plate and surstyli.

Xenothoracochaeta Malloch, 1921

(Plate 1) Type-species. Xenothoracochaeta prima Malloch (orig. des.). Diagnosis. Male holoptic; arista plumose; fIrst pair of frontal setae strong; scutellum with discal setulae extending up to the margins of the ventral surface; prealar seta present in both sexes (sometimes vestigial in male); meron setulose; anepimeron with cilia below calypter; katepistemals 1:2; metapleuron with a group of cilia; posterior spiracle with setae on posterior margin; veins R4+5 and M divergent apically; stemite 1 bare. 158



References. SNYDER, 1949a; ALBUQUERQUE & LOPES 1979b. Monophyly. Not yet established. Geographical record. Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay, Argentina. Comments. There are six species in the Neotropical region, all them occurring in South America.

Key to species (modified from SNYDER, 1949a) 1.

Scutellar setulae descending onto ventral surface of scutellum; with two equally long and strong stigmatal setae ....................................................................................... 2 Scutellar setulae not invading ventral surface of scutellum; with a single stigmatal seta on each side or with a strong and a very much weaker one ...................................... 5

2.

Antenna and palpus mostly yellow.......................................................................... 3 Antenna and pal pus not yellow.............................................................................. .4

3.

Pro sternum setulose (Peru, Bolivia, Brazil, Paraguay) ......................................... ..

.................................................................... .......... X. ruJiguttata (Macquart, 1851) Prosternum bare (Brazil) ....................... .x. subreptilia Albuquerque & Lopes, 1979 4.

Antenna, palpus and entire scutellum infuscated (Venezuela) .......................... .

...... ................................... ... ..................... ............ .......x. caerulea (Snyder, 1949) Antenna mostly black, brownish at apex of pedicel and base of flagellomere; palpus black; scutellum greyish (Argentina) .................................. .x. golbachi Snyder, 1957 5.

Dorsocentrals 2:3 (Colombia) ................................ X.foscomarginata (Snyder, 1949) Dorsocentrals 1:3 (Colombia, Peru) ....................................X. prima Malloch, 1921


159


Subfamily Mydaeinae Verral, 1888 Diagnosis. Prealar seta usually present; calcar absent; wings with the veins usually ciliated; female: ovipositor reduced; cerci reduced, sclerotized and with cilia only on outer surface; sternite 7 divided; egg Mydaea-type (according to HERTING, 1957). Monophyly. Not well established yet. SKIDMORE (1985) did not find unique characters for the subfamily. CARVALHO (1989d) found the shape of the stemite 7 (divided transversely in two pieces) as a synapomorphy for the genus-type of the subfamily. Comments. The subfamily has 22 genera in the world, eight of them occurring in the Neotropical Region. There are 46 species occurring in this region.

Brontaea Kowarz, 1873 (Figs. 20-32) Type-species. Anthomyia polystigma Meigen (des. COQUILLETT, 1910: 516). Diagnosis. Male holoptic; female: frons without proclinate orbitals and without crossed setae on frontal vitta; arista bare (longest hairs shorter than its basal diameter); intra-alars 2; dorsocentrals 2:4; prealar absent; intrapostalar 1; proepistemal 1; disk of notopleuron bare; postalar declivity, suprasquamal ridge, pro sternum and proepisternal depression bare; anepisternum with a short seta in upper anterior corner; anepimeron, greater ampulla and meron bare; katepistemals 1:2; methatoracic spiracle small, triangular, bare on margins; mid tibia with 1 submedian posterior seta and 1 strong ventral apical seta; hind tibia without calcar, with 1 short submedian anterodorsal seta, dorsal apical seta strong; veins bare; vein M slightly curved towards R 4+5 on apical portion; lower calypter "Phaonia-type", bare; sternite 1 with stiffsetulae laterally (Fig. 20); female ovipositor (Figs. 21, 22) and egg "Mydaea-type". Reference. CARVALHO & PONT, 1998. Monophyly. The genus Brontaea is probably polyphyletic. Some of its species have aPhaonia-type ovipositor (according to HERTING, 1957), while others, have a Mydaea-type ovipositor. All species herein considered have a Mydaea-type ovipositor. This genus is here considered as a Mydaeinae as previously suggested by SKIDMORE (1985) and then by CARVALHO (1989d) and CARVALHO & PONT (1998). 160



Geographical record. Mexico, Cuba, St. VicentIs., Jamaica, Puerto Rico, Haiti, Peru, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. The genus occurs in all biogeographical regions but it is absent from New Zealand and Oceania and was accidentally introduced in Hawaii (CARVALHO & PONT, 1998).

Key to species (modified from CARVALHO & PONT, 1998) 1.

Mesonotum with dark brown vittae, very conspicuous and well-defined (Figs. 25); male: outer rows of presutural acrostichal setulae equidistant between the dorsocentral setae and each other; mesonotum with the dark central vittae continuing onto scutellum; abdomen with the dark markings on tergite 3 L-shaped (Fig. 29); cercal plate quadrate; female: sternite 2 wider than long (Mexico, Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay) ............................................................. .......... .B. nonnata (Bigot, 1885) Mesonotum with dark vittae absent or, when present, not well marked and not sharply defined; male: outer row of presutural acrostichal setulae usually closer to the dorsocentral setae than to each other; scutum with the dark central vitta discontinued at scutellum or continuing as an ill-defined and suffused mark; abdomen with the dark markings on tergite 3 L-shaped or quadrate; cercal plate not as described above .......... .2

2.

Palpus yellow, darkened at tip; larger species, body length 4.6-6.7 mm but only rarely less than 5.0 mm (a few female specimens); trochanters broadly yellow, especially on fore leg; anterior postsutural intra-alar seta reduced or absent, hardly distinct from ground setulae; male: abdomen extensively yellow on basal segments, and the dark markings quadrate especially on tergite 3 (Fig. 27); cercal plate heart-shaped; epiphallus abruptly enlarged apically, in dorsal view expanded mushroom-shaped; female: frontal triangle yellowish-grey dusted, very well marked; sternite 2 longer than wide (Mexico, St. Vincent, Jamaica, Puerto Rico, Haiti, Venezuela, Brazil, Argentina; Southern United States of America) .............................................................. B.debilis (Williston, 1896)


161


Palpus black; smaller specimens, body-length 2.6-5.3 mm; trochanters only narrowly or not at all yellow; male: abdomen wholly black, the dark markings on tergites 3 and 4 as in figures 29 and 30; cereal plate and epiphallus not as described above; female: frontal triangle dull grey to grey, less well marked .................................................... 3 3. Crossvein dm-cu more or less straight; tergite 1+2 narrowly grey dusted along hind margin; sternite 5 wider than long, with 4-6 setulae placed on apical margin; anterior postsutural intra-alar seta very short, at most twice as long as ground setulae; male: ocellar setae very short, hardly longer than length of ocellar tubercle; 3-4 frontal setae; parafacial sunken, in lateral view partly obscured by eye; hind femur with 3-4 strong anteroventral setae preceded by a series of much shorter but distinct setae along basal part of this surface (Fig. 31); cercal plate expanded at base and narrowing suddenly in apieal part; distiphallus without spinules on basal part; female: proepimeral seta with no or at most one adjacent setula; fronto-orbital plate with 1-3 setulae outside lower frontal setae; tergites 4 and 5 with fewer, smaller and sparser brown seta-dots; sternite 7 present as short rod (Mexico, Cuba, Puerto Rico, Peru, Bolivia, Ecuador, Brazil, Chile; United States of America, mainly southern states)...... ........ B. quadristigma (Thomson, 1869) Crossvein dm-cu distinctly curved; sternite 5 longer than wide, with numerous setulae (usually 15-20) on lateral and apical margins; anterior postsutural intra-alar seta longer, at least twice as long as ground setulae, often longer; male: ocellar setae long and strong, at least equal to lowest frontal seta; 5-6 frontal setae; parafacial not sunken, visible in lateral view; hind femur with only 3-4 strong anteroventral setae in apical third (Fig. 32); cercal plate narrowing uniformly from base to apex; distiphallus with spinules on basal part; female: proepimeral seta with 3-6 adjacent setulae; frontoorbital plate with 4-6 setulae outside lower frontal setae; tergites 4 and 5 with numerous large brown seta-dots; sternite 7 present only as 2 seta-dots (Mexico, West Indies, Brazil; United States of America, Canada; Hawaii) ................B. delecta (Wulp, 1896)

162



~20

epi

21

0,5 mm 0,25 mm

0,25 mm

1 mm

1 mm

Figs. 20-32, characters of Brontaea (modified from CARVALHO & PONT, 1998): 20, B. debilis, sternite 1; 21, B. debilis, ovipositor, lateral view; 22, B. normata, ovipositor, lateral view; 23, B. debilis, mesonotum, dorsal view; 24, B. quadristigma, mesonotum, dorsal view; 25, B. normata, mesonotum, dorsal view; 26, B. delecta, mesonotum, dorsal view; 27, B. debilis, abdomen, dorsal view; 28, B. quadristigma, abdomen, dorsal view; 29, B. llormata, abdomen, dorsal view; 30, B. delecta, abdomen, dorsal view; 31, B. quadristigma, hind femur and hind tibia, lateral view; 32, B. delecta, hind femur and hind tibia, lateral view. Abbreviations: epi, epiproct; hyp, hypoproct.

Graphomya Robineau-Desvoidy, 1830 (Figs. 33-37, Plate 1) Type-species. Musca maculata Scopoli (des. WESTWOOD, 1840: 141). Diagnosis. Characteristic marks on mesonotum and abdomen (Figs. 33-34); male holoptic; female without interfrontal setae; arista enlarged on basal third, plumose; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge and pro sternum bare; anepimeron, center of proepimeron and prosternum bare; katepisternals 1: 1, both Muscidae (Diptera) of the Neotropical Region: TAXONOMY

163


on upper posterior angle; meron setulose on the upper margin, both in front of and below the spiracle; methatoracic spiracle bare on margins; mid tibia with 1 submedian posterior seta and 1 strong ventral apical seta; hind tibia without calcar; vein M curved towards R4+5 on apical portion (Fig. 35); vein R4+5 setulose basally base; lower calypter about twice the length of the upper one; stemite 1 setulose. References. MALLOCH, 1934; ARNTFIELD, 1975. Monophyly. The monophyly and affinities of Graphomya have been long regarded as a problem (SKIDMORE, 1985). HENNIG (1965) included it among the Mydaeinae, a concept followed by VOCKEROTH (1972). SKIDMORE (1985) also included the genus among the Mydaeinae, although having the same view as LOBANOV (1977), that Graphomya has more affinities with the Coenosiinae than with the Mydaeinae. Geographical record. Mexico, Cuba, Puerto Rico, Peru, Brazil, Chile, Venezuela, Uruguay, Argentina; Afrotropical, Palaearctic, Nearctic, Oriental. Essentially cosmopolitan in temperate and tropical areas, absent in New Zealand and Oceania, except Micronesia, better developed in Afrotropical and Oriental Regions (VOCKEROTH, 1972). Comments. Some South American records, especially those related to the typespecies, are probably misidentifications.


Male ................................................................................................................... 2 Female ................................................................................................................ 4

2.

Frons at narrowest point as wide as length of flagellomere; parafacial with anterior part of frontal orbits, and most of gena, densely golden yellow dusted; abdomen black, with grey dust and the usual irregular tripunctate dorsum; tergite 5 conspicuously differentiated by dense golden yellow dusting; mid tibia with at least one anterodorsal seta beyond middle; 2 or more posterodorsal and 1 or 2 posteroventral setae (Peru, Brazil, Chile, Argentina) .................................. G. analis (Macquart, 1851) [in part] Frons at narrowest point not, or very little, over half as wide as length of flagellomere; parafacial dusting not as above; tergite 5 not conspicuously differentiated by goldenyellow dusting from the others; mid tibia without anterodorsal setae beyond

164



middle .................................................................................................................. 3 3.

Pre sutural area between dorsocentrals with a broad black vitta that is but inconspicuously if at all divided by a central grey vitta (Puerto Rico, Venezuela, Uruguay, Argentina; Afrotropical, Palaearctic and Oriental Regions; Melanesia) .................. . .................................................................................. G. maculata (Scopoli, 1763) Pre sutural area between dorsocentrals with 2 narrow black vittae that are separated by a central dusted vitta as conspicuous as those on each side of the dorsocentrals (Peru, Argentina, Chile) ...................................................... G. chilensis Bigot, 1888

4.

Mid tibia with an anterodorsal seta beyond middle; tergite 5 with dense golden-yellow dust that contrasts sharply with the grey dusting of the other tergites; orbits, parafacial and gena with the same dense golden-yellow dusting (Peru, Brazil, Chile, Argentina) ......................................................................... G. analis(Macquart, 1851) [in part] Mid tibia without an anterodorsal seta beyond middle; tergite 5 not, or very slightly, more noticeably yellowish dusted than the other tergites; head usually with much less distinct yellow dusting ............................................................................................ 5

5.

Frontal orbits, parafacial and the greater part of gena densely golden-yellow dusted (Argentina) ...................................................................... G. auriceps Malloch, 1934 Frontal orbits, parafacial and most of gena grey dusted (Puerto Rico, Venezuela, Uruguay, Argentina; Afrotropical; Palaearctic and Oriental Regions; Melanesia) .................................................................................... G. maculata (Scopoli, 1763)

Species of Graphomya not included in the key and their respective geographical distribution Species


G. mendozana (Enderlein, 1935)

Argentina

G. meridionalis Townsend, 1892

Brazil


165


G. mexicana Giglio-Tos, 1893

Mexico, Panama, Bolivia, Argentina

G. occidentalis Arntfield, 1975

Mexico, California, Hawaii (*)

G. panamensis Dodge, 1965

Panama

G. podexaurea (Enderlein, 1935)

Chile

G. stipata (Walker, 1853)

Puerto Rico, Venezuela, Guyana

(*) Keyed in ARNTFIELD (1975) with other North American Graphomya

Figs. 33-35, characters of Graphomya sp. (modified from ARNTFIELD, 1975): 33, mesonotum, dorsal view; 34, abdomen. dorsal view; 35. wing. dorsal view. Abbreviation: ntp ant. notopleural anterior seta.

166



Hemichlora Wulp, 1893 Type-species. Curtonevra vittigera Bigot (mon.) =scordalus Walker. Diagnosis. Male hoI optic; female frons without interfrontal setae; arista with short cilia only on dorsal surface; intra-alars 2; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge, pro sternum and proepisternal depression bare; anepimeron setulose; propleuron and prosternum bare; meron setulose below spiracle; katepisternals 1: 1, both setae on upper posterior angle; methatoracic spiracle bare on margins; hind tibia without calcar; veins bare; M slightly curved towards R4+5 on apical portion; lower calypter about 1.5 times the length of the upper one; sternite 1 setulose. Reference. LOPES, 1978. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, Costa Rica, Brazil. Comments. Monobasic genus: Hemichlora scordalus (Walker, 1861).

Mydaea Robineau-Desvoidy, 1830 (Plate 2) Type-species. Mydaea scutellaris Robineau-Desvoidy (des. COQUILLET, 1901: 139) = corni Scopoli. Diagnosis. Male hoI optic; female frons without interfrontal setae; arista plumose, with plumae moderately long on basal half, becoming shorter on apical half; intra-alars 2; dorsocentrals 2:4; prealar present; anepimeron and meron bare; katepisternals 1:2; hind tibia without calcar; veins bare; vein M straight; lower calypter about twice the length of the upper one; sternite 1 bare; female ovipositor and egg "Mydaea-type". Reference. SNYDER, 1949a. Monophyly. Not yet established, but according to VOCKEROTII (1972) Mydaea and Myospila are closely related, especially based on the morphology of the ovipositor. Geographical record. Mexico, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. Mydaea is poorly studied in the Neotropical Region and a great number of new species to be described remain. VOCKEROTH (1996) synonymized Scenetes Malloch, 1936 with Mydaea, specially based on the structure of the ovipositor. However, the Neotropical Mydaea were not yet studied phylogenetic ally together with Muscidae (Diptera) of the Neotropical Region: TAXONOMY

167


the Mydaea of the other biogeographical regions. The genus has monophyletic "groups" that could be raised to a higher taxonomic level. Based on these considerations, we will not recognise this synonymy until a generic revision of Mydaea is done.

Key to species (modified from SNYDER, 1949a) 1. Dorsocentral setae 2:3 ........................................................................................... 2 Dorsocentral setae 2:4 .......................................................................................... 4 2.

Fore tibia without a median posterior seta; margins of calypteres darkened (Venezuela, Ecuador, Bolivia) .......... .................................................. M. nubivena Snyder, 1941 Fore tibia with a median posterior seta; margins of calypteres not darkened ............. 3

3.

Palpus and at least the second antennal segment yellow (Mexico) .......................... .

..... .................. .................. ............. .............. ....... ....... ..... ...M. meridia Snyder, 1941 Palpus and antenna entirely black (Mexico ) ........................... M. cresa Snyder, 1949 4.

Wing with a dark costal cloud; fore tibia with a median posterior seta (Ecuador) ...................................................................................... M. Jumicosta Snyder, 1941 Wing without a dark costal cloud; fore tibia without a median posterior seta ............ ..

S.

s

Margins of upper calypteres infuscated; crossveins r-m and dm-cu with narrow infuscated clouds (Mexico, Paraguay, Brazil) ...... ..............M. plaumanni Snyder, 1941 Margins of upper calypteres and crossveins nn and dm-cu not infuscated .................. 6

6.

Frontal vitta, at narrowest part of front, three to four times as wide as fronto-orbital plate (Venezuela) ........................................... .......................M. caras Snyder, 1949 Frontal vitta, at narrowest part of front, only as wide as width of fronto-orbital plate (Colombia) ....................................................... ..M. compressicornis Snyder, 1949

168



Species of Mydaea not included in the key and their respective geographical distribution Species


M. castanea Albuquerque, 1955

Brazil

M. centralis Stein, 1911

Peru; Chile

M.fumipennis (Wulp, 1896)

Mexico

M. insons (Giglio-Tos, 1896)

Mexico

M. latomensis Snyder, 1957

Argentina

M. longiseta (Wulp, 1896)

Mexico

M. orba Stein, 1911

Bolivia

M. pansa (Giglio-Tos, 1893)

Mexico

M. pauciseta Stein, 1911

Peru

M. sexpunctata (Wulp, 1883)

Argentina

Unplaced species (CARVALH 0 etal. 1993) Species


M. inermis (Stein, 1904)

Bolivia

M. latipennis (Hough, 1900) *

Brazil

M. marginalis (Stein, 1904)**

Bolivia

* probably Cyrtoneurininae; ** probably Helina.


169


Myospila Rondani, 1856 (Figs. 36-37) Type-species. Musca meditabunda Fabricius (orig. des.). Diagnosis. Male holoptic; female with a pair of interfrontal setae; arista enlarged on basal third, plumose; intra-alars 2; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge, pro sternum, anepimeron and meron bare; katepisternals 2:2; methatoracic spiracle bare on margins; vein M slightly curved forward apically; vein R4+5 with few cilia basally; hind tibia without calcar; sternite 1 bare; female: ovipositor and egg Mydaea-type (Figs. 36-37). References: SNYDER, 1940; COURI & LOPES, 1988a, b. Monophyly. See discussion under Mydaea. Geographical record. Mexico, Costa Rica, Panama, Jamaica, Cuba, Puerto Rico, Haiti, S., Domingo, Guatemala, Venezuela, Guyana, Ecuador, Bolivia, Peru, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. Only six species are know in the Neotropical Region.

Key to species (modified from SNYDER, 1940) 1.

Thorax and abdomen entirely shiny blue; halter black (Argentina, Chile, Falkland Is.)

...................................................... ..............................M. cyanea (Macquart, 1843) Thorax and abdomen not shiny blue; halter yellow to orange ................................ '" .. 2 2.

Wing with a distinct darkened area on apices of the veins Sc and R J ; at most the margins of both calypteres distinctly infuscated (Peru, Brazil, Argentina)

............................................................... .M. obscura (Shannon & Del Ponte, 1926) 3.

Wings without a distinct darkened area on apices of the veins Sc and R J ; at most the margins of the upper calypter slightly brownish; the lower margined with yellow............ 3 Pedicel, flagellomere and the fore and mid tibiae black; intra-alars 2 (Mexico, Guatemala, Ecuador, Peru, Bolivia; Nearctic and Palaearctic Regions)

...................................................... .................... M. meditabunda (Fabricius, 1781) 170



Pedicel and flagellomere light brown to orange or yellow; all tibiae light brown to orange; intra-alar 1................................................................................................ 4 4.

Palpus and flagellomere light brown or yellow; wing with a brown spot on cross vein rm (Mexico, Costa Rica, Panama, Puerto Rico, Haiti, Cuba, S. Domingo, Jamaica, Venezuela, Guyana, Peru, Bolivia, Brazil) ......................M. pallidicomis (Bigot, 1887) Palpus and flagellomere light brown; wings hyaline .................................................. 5

5.

Calypteres white with brown margins in female; hind tibia on anterodorsal surface with 2 setae on middle third (Brazil) ............M. matogrossensis Couri & Lopes, 1988 Calypteres entirely white in female; hind tibia on anterodorsal surface with 4 setae on middle third (Brazil) .........................................M.jluminensis Couri & Lopes, 1988

Figs. 36-37, characters of Myospi/a (modified from COURI & LOPES, 1988a): 36, M. matogrossensis, ovipositor, lateral view; 37, egg, dorsal view.

Scenetes Malloch, 1936, stat.rev. Type-species. Scenetes cardini Malloch (orig. des.). Diagnosis. Head in profile almost twice as high as its greatest length, distinctly narrower at vibrissae than at antennal bases, gena about as high as width of flagellomere and twice as high as width of parafacial on middle; facial ridges rather closely short setulose to a little above middle of flagellomere, the latter from 6 to 8 times as long as pedicel; arista plumose on basal half or more, bare apically; eyes almost bare, frons in both sexes about one-fifth of the head width at vertex, widened toward anterior margin; orbits narrow, each Muscidae (Diptera) of the Neotropical Region: TAXONOMY

171


with fourto six incurved anterior and two recurved posterior setae; interfrontal setae absent; ocellar and inner vertical setae long; outer vertical setae undeveloped; prostemum, center of anepimeron and meron bare; lower calypter narrowly rounded at apex and much larger than the upper calypter; vein M not curved forward apically; R 4+5setulose on both surfaces from base to about midway to the crossvein r-m; calcar absent. Reference. CARVALHO, 1985b. Monophyly. Not yet established. See discussion under Mydaea. Geographical record. Cuba and Puerto Rico. Comments. Monobasic genus: Scenetes cardini Malloch, 1936.

Scutellomusca Townsend, 1931 (Figs. 38-42) Type-species. Musca scutellaris Fabricius (orig. des.). Diagnosis. Eyes with short and sparse cilia; male holoptic; arista plumose; 2 orbital setae, the upper ones backward inclinated; prealar seta short; scutellum with lateral cilia on basal half; proepimeron setulose; katepistemals 1:2; lower calypter about twice as long as the upper one; suprasquamal ridge haired; katepimeron with few setulae; vein Rl with dorsal cilia; vein R4+5 with dorsal and ventral cilia; vein M strongly curved forward apically; mid tibia with a posteroventral seta inserted on the limit of basal third and a posterior seta on the limit of apical third; hind tibia with no setae on posterodorsal surface; stemite 1 setulose; male: cercal plate with short lower projections; female: ovipositor and egg "Mydaea-type" (Figs. 38-42). References. ALBUQUERQUE, 1954c; COURI & CARVALHO, 1992. Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: Scutellomusca scutellaris (Fabricius, 1805).

172



38

0,3 mm

41

Figs. 38-42, characters of Scutellomusca scutellaris (modified from COURI & CARVALHO, 1992): 38, ovipositor, dorsal view; 39, ovipositor, ventral view; 40, ovipositor, lateral view; 41, egg, dorsal view; 42, egg, lateral view.

Subfamily Coenosiinae Verral, 1890 Diagnosis. Proboscis adapted to predation, prestomal teeth developed, prementum shiny; prealar seta absent; mid femur on anterior surface with a preapical seta; anepimeron bare; mesonotum with 3 or 5 dorsal vittae; calcar absent; abdomen with lateral spots on tergites. Monophyly. The subfamily is probably monophyletic, the strongest evidence being the absence of the prealar seta (COURI & PONT, 2000). Comments. Two tribes are traditionally included in the subfamily: Limnophorini and Coenosiini, as presented by SKIDMORE (1985), CARVALHO (1989d) and CARVALHO et al. (1993). The cosmopolitan distribution of several genera has been taken as an indication of the antiquity of the group (HENNIG, 1959). Muscidae (Diptera) of the Neotropical Region: TAXONOMY

173


Tribe Limnophorini Villeneuve, 1902 Diagnosis. Prostemum setulose or bare; apical portion of vein M bent forward apically or not; female: stemite 8 usually absent; membrane on segment 8 usually with spicules; hypoproct with modifications. Monophyly. Not yet established, but the tribe is probably paraphyletic. Several Coenosiini genera were recently transferred to Limnophorini (COURI & PONT, 2000). Comments. There are 12 genera and 129 species recorded from the Neotropical Region. Agenamyia Albuquerque, 1953

(Figs. 43-50) Type-species. Agenamyiafumipennis Albuquerque (orig. des.) Diagnosis. Male dichoptic; eyes bare, at level of anterior ocellus separated by 3.43.6 of head width; gena and peristome linear (Fig. 43a, 43b); prestomal teeth developed; arista bare; one orbital reclinate seta; prealar seta absent; lower proepimeral seta downcurved; dorsocentral setae 1:3; anepimeron bare; scutellum bare on lower and lateral margins; prostemum bare; katepistemals 1: 1: 1, not equidistant from each other, the anterior one weaker than the others; lower calypter about twice the length of upper one; wing veins bare; M and R4+5 parallel apically (Fig. 44); calcar absent; stemite 1bare; cercal plate with lateral projections (Fig. 45); ovipositor moderately long with large tergites (Figs. 46-48). References. CARVALHO & COURI, 1992; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. Probably monophyletic, based on the presence of unique muscid characters as the shape of the gena and the linear peristome. Geographical record. Panama, Brazil. Comments. Agenamyia has been recently transferred to Limnophorini (COURI & PONT, 2000).

Key to species (modified from CARVALHO & COURI, 1992) 1. Male: hind femur with three strong anteroventral setae on middle third; cerci, in lateral 174



view, with the apical third enlarged; stemite 5 as in figure 99; female: legs usually dark, sometimes femora and tibiae paler (Panama) ....... A. exotica Carvalho & Couri, 1992 Male: hind femur with two strong anteroventral setae on middle third; cerci, in lateral view with the apical third uniform; stemite 5 as in Fig. 50; female: unknown (Brazil) ............................................................................A.fumipennis Albuquerque, 1953

1 mm

~

U a

1 mm

"~o,'mm gn_~r.;:"5'fI:J'

44

43

O,Smm

Figs. 43-50, characters of Agenamyia (modified from ALBUQUERQUE, 1953; CARVALHO & COURI, 1992): 43, A. jumipennis, male head, (a) view frontal, (b) lateral view; 44, A. jumipennis, wing; 45, A. jumipennis, cereal plate; 46, A. exotica, ovipositor, ventral view; 47, A. exotica, ovipositor, dorsal view; 48, A. exotica, ovipositor, lateral view; 49, A. exotica, sternite 5, dorsal view; 50, A. jumipennis, sternite 5, dorsal view. Abbreviations: gn, gena; tg, tergite.


175


Albertinella Couri & Carvalho, 1997 (Figs. 51-55) Type-species. Hypsomyia nebulicola Couri & Carvalho (orig. des.). Diagnosis. Male holoptic; eyes with cilia, more conspicuously in male; arista pubescent with plumes shorter than basal width of arista (some exemplars with 1-2 cilia more or less long on basal half of arista and easily differentiated from the others); gena slightly prominent (Fig. 51); prestomal teeth developed; labella reduced (Fig. 52); prosternum setulose laterally; mesonotum with two silver pruinose vittae coincident with dorsocentral row, in female the vittae reach the apex of scutellum, in male they are discontinuous and interrupted below suture up to posterior dorsocentral pair of setae and do not reach the apex of scutellum; acrostichal setae ciliform, not differentiated from ground setulae; prealar seta absent; lower proepimeral seta upcurved; dorsocentral setae 0: 1; katepistemals 1:2, the upper posterior seta strong and long, about three times the length of the others; anepimeron bare; lower calypter about twice the length of the upper one; vein R j with cilia on apical half of dorsal surface; vein R4+5 with cilia along its entire extension on both surfaces; hind tibia with a long and fine median anteroventral seta and another shorter submedian seta; no posterodorsal preapical setae; stemite 1 bare; ovipositor (Fig. 53-55) moderately long; segment 8 directed upwards with spicules (Fig. 55); hypoproct elongated, upward curved and with strong setae and spines; cerci short; egg Phaonia-type. References. COURI & CARVALHO, 1995, 1997a. Monophyly. Albertinella is the only Limnophorini genus with dorsocentral seta 0: 1. Geographical record. Brazil. Comments. Monobasic genus: Albertinella nebulicola (Couri & Carvalho, 1995).

176



... _-------- ........

O,5mm

53 O,3mm

O,3mm

,

I

,

I I

I I

I I I

I

Figs. 51-55, characters of Albertinella nebulicola (modified from COURI & CARVALHO, 1995): 51, male head, lateral view; 52, female proboscis, lateral view; 53, ovipositor, dorsal view; 54, ovipositor, ventral view; 55, ovipositor, lateral view. Abbreviation: hyp, hypoproct.

Drepanocnemis Stein, 1911 Type-species. Drepanocnemis dorae Stein (des. MALLOCH, 1928: 471). Diagnosis. Male dichoptic; arista bare; 1 pair of reclinate orbital setae; prestomal teeth developed; labella reduced; prealar seta absent; lower proepimeral seta directed upward; dorsocentral setae 1:3; katepistemum with many ground setulae; lower calypter about twice the upper one; male fore tibia very broad, with a long, posterodorsal subbasal seta; stemite 1 bare. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

177


References. STEIN, 1911; MALLOCH, 1928; COURI & PONT 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) indicated five characters for the genus, three of them reversals: labella not developed; prealar seta present and disk of katepistemum with many setulae. The modification of the male fore tibia can be interpreted as an autapomorphy for the genus. Geographical record. Peru, Bolivia. Comments. Drepanocnemis was recently transferred from Coenosiini to Limnophorini (COURI & PONT, 2000). Only three species are known in the Neotropical Region: D. dorae Stein, 1911 (Peru); D. griseovirens Malloch, 1928 (Peru) and D. hirticeps Stein, 1911 (Peru, Bolivia). The characters available in literature are insufficient to identify and key out the species.

Limnophora Robineau-Desvoidy, 1830 (Plate 2) Type-species. Limnophora palustris Robineau-Desvoidy (des. COQUILLETT, 1910: 561) = maculosa (Meigen). Diagnosis. Male eyes separated by no more than one-fifth ofthe head width (at least in the South American species, see MALLOCH, 1934); eyes almost invariably bare; prestomal teeth developed; prealar seta absent; lower proepimeral seta upcurved; postsutural dorsocentral setae 3 or 4; prostemum setulose; apical portion of vein M slightly bent forward; wing with cilia on both sides of base of radial sector; stemite 1 bare; female: ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. STEIN, 1911; MALLOCH, 1934. Monophyly. Although Limnophora belong to a monophy letic group that includes many genera occurring in all biogeographical regions, HENNIG (1965) understood that the genus is paraphyletic. The Limnophora group is considered the sister-group of Spilogona (HENNIG, 1965; SKIDMORE, 1985). Geographical record. Mexico, Bahamas, Puerto Rico, Cuba, Belize, Guatemala, Costa Rica, Panama, St. Vincent Is., Jamaica, Venezuela, Colombia, Guyana, Bolivia, Peru, Ecuador, Galapagos Is., Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. 178



Comments. The genus needs an extensive taxonomic revision. The keys of STEIN (1911) and MALLOCH (1934) are presented below with modifications. The species not included in the key are listed at the end.


Both crossveins r-m and dm-cu with a brown cloud (Mexico, Guyana) .................. . . .......................... ........ .................... ............. ........... ...... L. elegans Macquart, 1843

Crossveins r-m and dm-cu without brown clouds .................................................... 2 2.

Mesonotum almost entirely dark brown, vittae, if present almost imperceptible (Peru) ................................................................................. .L. marginipennis Stein, 1911

Mesonotum grey with 3 brown very distinct vittae ................................................... 3 3.

Wing completely hyaline or at least yellowish (Mexico, Peru, Brazil) .......................... .. . ............. .............................. ............. ......... ............ ......... ..... L. deleta (Wulp, 1896)

Wing distinctly blackish, specially on anterior margin ................................................ .4 4.

Orbit, face and gena with yellowish pruinescence (female) (Peru, Bolivia, Argentina) ........................................................................... ............... L. aurifacies Stein, 1911

Orbit, face and gena with grey or blackish pruinescence .......................................... 5 5.

Prescutum entirely black (Mexico, Guatemala, Costa Rica, Panama, Venezuela, Guyana, Ecuador, Peru, Bolivia, Brazil, Argentina) .......... .L. saeva (Wiedemann, 1830) Prescutum with spots or vittae ................................................................................ 6

6.

Arista with long plumae, longer than width of flagellomere (Peru, Guyana, Brazil, Argentina) ................................... ......................................... .L. piliseta Stein, 1919 Arista with plumae shorter than width of flagellomere ............................................. 7

7.

Mesonotal vittae forming a large vittae after suture; vein R] bare (Mexico, Bahamas, Puerto Rico, Cuba, Venezuela, Guyana, Peru, Brazil, Paraguay, Argentina, Chile;


179


Nearctic region) ................................................................. .L. narona (Walker, 1849) Mesonotal vittae not forming a unique large vittae after suture; vein R J with short setae near apex ............................................................................................................. 8 8.

Mesothorax with a transversal brown vitta after suture (Mexico, Guatemala, Belize, Costa Rica, Puerto Rico, Venezuela, Peru, Brazil, Chile; southern United States of America) .............................................. ................... .L. corvina (Giglio-Tos, 1893) Mesothorax without a transversal brown vitta after suture ....................................... 9

9.

Arista slightly pubescent; mesonotal vittae almost confluent after suture (Chile)

...................................................... ............................................... L.laeta Stein, 1911 Arista distinctly pubescent or with short plumae; mesonotal vittae not confluent after suture (Bolivia) ....................................................................... L. alacris Stein, 1911


Longest hairs on arista longer than width of flagellomere; eyes of male bare, separated at narrowest part of frons by a mere line, at which point the orbits obliterate the interfrontalia; hind femur with 2-3 preapical anteroventral setae, and a series of widely spaced weak setae on basal half or more of posteroventral surface (Peru, Guyana, Brazil, Argentina) .................... '" ..................................... .... .L. piliseta Stein, 1919 Longest hairs on arista not as long as the width of the flagellomere; other characters different from above .................................................................... '" ....................... 2

2.

Postsutural dorsocentral setae 3, all strong; longest cilia on arista about half as long as width of flagellomere (Peru, Argentina) ........................... .L. brevihirta Malloch, 1934 Postsutural dorsocentral setae 4; the two anterior pairs behind suture much shorter and weaker than the two posterior pairs ................................................................ 3

3. 180

Arista pubescent, longest shorter not longer than its basal diameter; both sexes with Muscidae (Diptera) of the Neotropical Region: TAXONOMY

Mar ;ia Souto Couri & Claudio Jose Barros de Carvalho

some well-developed setae on basal half of posteroventral surface of hind femur (Argentina, Chile, Juan Fernandez Is.) ................ ......... .L. patagonica Malloch, 1934 Arista with longest hairs about twice as long as its basal diameter; female without well-developed setae on basal half of posteroventral surface of hind femur (Argentina, Chile) ........................................ species not named according to MALLOCH, 1934

Species of Limnophora not included in the key and their respective geographical distribution Species


L. aczeli (Snyder, 1957)

Argentina

L. albuquerquei (Lopes & Couri, 1987)

Brazil

L. altaneira (Albuquerque, 1954)

Belize, Venezuela, Brazil

L. cubana Johnson, 1919

Cuba

L. exulWilliston, 1896

Cuba,Jamaica,St. Vincent Is.

L. garrula (Giglio-Tos, 1893)

Mexico, Brazil, Argentina

L. iniqua (Stein, 1911)

Peru, Chile

L. leucotelus (Walker, 1853)

Brazil

L. limbata Bigot, 1885

Mexico, Chile

L. longiseta (Lopes & Khouri, 1989)

Brazil

L. marginata Stein, 1904

Colombia, Ecuador, Brazil, Argentina

L. minuscula (Wulp, 1896)

Mexico

L. nigrargentata (Albuquerque, 1954)

Brazil

L. ovativentris (Macquart, 1851)

Uruguay


181


L. paranaensis (Albuquerque, 1954)

Brazil

L. paulistana (Lopes & Khouri, 1991)

Brazil

L. pica (Macquart, 1851)

Costa Rica, Jamaica, Brazil, Galapagos Is.

L. platystoma (Thomson, 1869)

Ecuador

L. spreta Malloch, 1921

Brazil

L. strigata (Giglio-Tos, 1893)

Mexico

L. vicaria (Walker, 1853)

Brazil

L. vicina Robineau-Desvoidy, 1830

Brazil

L. virgata (Wiedemann, 1830)

Brazil

L vittata (Macquart, 1851)

Brazil, Argentina.

Unplaced species of Limnophora s.lat. (CARVALHO etal., 1993) Species


* L. porteri (Brethes, 1920)

Chile

* Probably not Limnophorini Lispe Latreille, 1797 Type-species. Musca tentaculata De Geer (mon. LATREILLE, 1802: 462). Diagnosis. Male dichoptic; palpus enlarged apically, spatula-like; prestomal teeth developed; without interfrontal setae and proclinate supraorbital, parafacialia setulose; dorsocentral setae 2:3; prealar seta absent; anepimeron setulose in the center; lower proepimeral seta upcurved; katepistemals 1:2; male: gonopod absent; female: ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. SNYDER, 1949b, 1954b. 182



Monophyly. A monophyletic group with many apomorphic characters (HENNIG, 1965; SKIDMORE, 1985). See sister-group relationships under Lispoides. Geographical record. Mexico, Bahamas Is., Virgin Is., Cuba, Puerto Rico, Guadalupe Is., Guatemala, Guyana, Bolivia, Peru, Brazil, Argentina, Chile, Paraguay; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. The genus needs an extensive taxonomic revision on the Neotropical Region. A key of SNYDER's (1949b) is here presented with modifications and the species not included in the key are listed at the end.

Key to species (modified from SNYDER, 1949b) 1.

Mid tibia with only one posterior median seta ........................................................ 2 Mid tibia with one submedian anterior or anterodorsal seta in addition to the posterior median seta ........................................................................................................... 4

2.

Hind femur with a row of short, stout, posteroventral setae on basal half (Argentina, Chile) ............................................................................ .L. setuligera (Stein, 1911) Hind femur with no posteroventral seta .................................................................. 3

3.

Male with a prolongation offore metatarsus (Guadalupe Is., Guatemala, Peru, North America, Palaearctic Region) ................................. .L. tentaculata (De Geer, 1776) Male with a simple fore metatarsus (Bolivia) ........................... .L. vilis (Stein, 1911)

4.

Pre sutural dorsocentral seta weak, the anterior one, if present, never so strong as the outter vertical, and the posterior one smaller then the anterior ocellar seta (Brazil)

......................................................................................... .. L. latana Snyder, 1949 Presutural dorsocentral seta strong; the anterior one as strong as the outer vertical and the posterior one as long as the anterior ocellar seta ................................................ 5 5.

Postsutural dorsocentral setae 4, the two anterior pairs weak (Mexico, Puerto Rico, Guyana, Brazil, Paraguay) .................................................. .L. seratina Wulp, 1896 Postsutural dorsocentral setae 3, all strong ............................................................. 6


183


6.

The anterior postsutural dorsocentral seta strong; pedicel with only a small fulvous spot at apex; with well-defined dark abdominal markings (Mexico, Chile; United States of America) ...................................................... .......... .L. levis (Stein, 1911) The anterior postsutural dorsocentral seta very small, scarcely developed; pedicel entirely fulvous; posterior margins of abdominal tergites broadly fulvous and with illdefined markings (Brazil) .................................................... .L. lisarba Snyder, 1949

Species of Lispe not included in the key and their respective geographical distribution Species


L. albitarsis (Stein, 1898)

Bahamas Is.; Nearctic Region

L. bahama Snyder, 1958

Bahamas Is.

L. cotidiana Snyder, 1954

Mexico; Nearctic Region; Central and East Asia

L. nasoni (Stein, 1898)

Bahamas Is., Virgin Is., Cuba, Puerto Rico; Nearctic Region

L. rufitibialis Macquart, 1843

Puerto Rico, Brazil or Chile

Lispoides Malloch, 1920 Type-species. Limnophora aequifrons Stein (orig. des.). Diagnosis. Frons of male moderately wide, sometimes about as wide as that of the female; prestomal teeth developed; pro sternum bare; prealar seta absent; lower proepimeral seta upcurved; vein R4+5 bare; sternite 1 setulose; ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. STEIN, 1911; MALLOCH, 1934. Monophyly. HENNIG (1965) suggested that Lispoides and Lispe could be sister-groups. SKIDMORE (1985), based on characters of the larvae, indicated that Lispoides is not closely related to Limnophora and that the common characters found in both genera are due to adaptative convergence. 184



Geographical record. Guatemala, Venezuela, Colombia, Bolivia, Peru, Argentina, Chile, Juan Fernandez Is.; Nearctic Region. Comments. Recently, PONT (2001) transferred Limnophora pubiceps Stein, 1911 to this genus.


Eye distinctly haired; crossveins r-m and dm-cu surrounded by a brown cloud (Bolivia)

...................................................................................... .. L. elegantula Pont, 1972 Eye bare; crossveins r-m and dm-cu not surrounded by a brown c1oud ................... 2 2.

Crossvein r-m greyish (Bolivia, Peru) ........................L. abnonninervis (Stein, 1911) Crossvein r-m not greyish ..................................................................................... 3

3.

Halter whitish or greyish with daker margin .......................................................... ..4 Halter whitish without daker margin ....................................................................... 6

4.

Mesonotum bare beyond the strongest seta (Venezuela, Peru) ... .L. laevis (Stein, 1911) Mesonotum setulose beyond the strongest setae .................................................... 5

5.

Mid femur with strong and long setae on posteroventral surface; mid tibia with many setae; hind femur with strong setae on ventral surface, a little before distal apex (Peru)

.....................................................................................L. atrisquama (Stein, 1904) Mid femur with fine and short setae on posteroventral surface; mid tibia with only one median seta; hind femur bare on ventral surface (Peru) .. ....L. propinqua (Stein, 1911) 6.

Median vitta on frons thin in both sexes, or at most as wide as orbits; general coloration grey-whitish (Chile) ................................................................ L. diluta (Stein, 1911) Median vitta on frons wider than the orbits; general coloration dark .......................... 7

7.

Mesonotum almost without vittae; abdomen much longer than thorax (Bolivia, Chile)

............................................................................................ L. gracilis (Stein, 1911) Muscidae (Diptera) of the Neotropical Region: TAXONOMY

185


Mesonotum with vittae faint or distinctly marked; abdomen as long as thorax ............ 8 8.

Mesonotum with faint dorsal vittae, all having the same width (Peru, Bolivia) ...................................................... ................................ L. nigribasis (Stein, 1911)

Mesonotum with distinct dorsal vittae, the median one thinner than the lateral ones (Peru, Bolivia, Chile) ................................................ ........... .L. triplex (Stein, 1911)


Thorax densely pale, grey dusted; mesonotum with three conspicuous brownish-black vittae, the central one narrowest, of uniform width and sharply margined on its entire extent, occupying one-third of the intra-dorsocentral area; pleurae with a dark upper marginal vitta; frons of male about one fourth of the head width and of female about one third ................................................................................................................ 2 Thorax much less conspicuously vittate, central vitta neither uniformly wide nor sharply margined; pleurae without a dark upper marginal vitta; frons of male and of female about equally wide, at most one-fourth of the head width ....................................... 3

2.

Meron with some setulose hairs on upper margin in front of spiracle (Argentina, Chile) .............................................................................. .L. inaequifrons Malloch, 1934

Meron bare (Argentina, Chile?) .................................. .L. argenticeps Malloch, 1934 3.

With some hairs on fronto-orbital plate below level of lower frontal seta; abdomen less densely greyish dusted, the dark markings not sharply defined but more extensive, specially in male (Colombia, North America) .................. .L. aequifrons (Stein, 1898) No hairs on fronto-orbital plate below level oflower frontal setae; abdomen densely brownish-grey dusted, with the dark central line and subtriangular paired spots well defined and conspicuous in both sexes .................................................................. .4

4.

Mid tibia with one posterior seta; hind tibia with one long submedian anterodorsal seta (Peru) ................................................. ....................... .L. uniseta Malloch, 1934

186



Mid tibia with several posterior setae; hind tibia with two or more unequal anterodorsal setae (Argentina) .......................................................... ,L. argentina Malloch, 1934 Species of Lispoides not included in the keys and their respective geographical distribution [including L. puhiceps (Stein, 1911) recently transferred to this genus by PONT,2001] Species

Geographical Distribution

L. guatemala Snyder, 1951

Guatemala

L. insularis Hennig, 1957

Juan Fernandez Is.

L. latifrons Snyder, 1957

Argentina

L. lopesi Albuquerque, 1955

Brazil

L. puhiceps (Stein, 1911)

Peru,Bolivia

Pachyceramyia Albuquerque, 1955 (Figs. 56-61) Type-species. Pachyceramyia enigmatica Albuquerque (orig. des.). Diagnosis. Head with the antennal axis longer than the oral axis (Figs. 56-57); male dichoptic; upper pair of frontal setae directed outward; arista short, wider on basal two-thirds, pubescent and a little longer than flagellomere; gena and facialia large; frontoorbital plate with a series of strong short cilia; prealar seta absent; acrostichal prescutellar seta ciliform; dorsocentral setae 2:4; katepisternals 2:2; wing veins bare; veins R4+5 and M parallel apically (Fig. 58); lower calypter about 1.5 as long as upper one; hind tibia with two anterodorsal (one median to submedian and another supramedian) , one posterodorsal on apical third and one posterodorsal preapical setae (Fig. 59); female: ovipositor moderately long with tergites 6 and 7 large (Figs. 60, 61). References. HUCKETT, 1936; ALBUQUERQUE, 1955b. Monophyly. Not yet established. Geographical record. Bahamas Is., South America, Brazil; United States of America. Muscidae (Diptera) of the Neotropical Region: TAXONOMY

187


Comments. P. pantherina (Wiedemann, 1824) was not recognised after the original description and is not herein included in the key. Key to species 1.

Postsutural dorsocentral setae 4 (Brazil) ............... P. enigmatica Albuquerque, 1955 Postsutural dorsocentral setae 3............................................................................ 2

2.

Arista plumose; hind tibia with the distal posterodorsal seta much longer (about 1.5 times) than proximal posterodorsal seta (United States of America)

................................................................................. P. longispina (Malloch, 1923)

1 mm

O.5mm

61 Figs. 56-61, characters of Pachyceramyia (modified from ALBUQUERQUE, 1955b; COURI & PONT, 2000): 56, P. enigmatica, head, lateral view; 57, P. enigmatica, % view; 58, P. enigmatica, wing; 59, P. enigmatica, hind tibia; 60, P. cordyluroides, ovipositor, dorsal view; 61, P. cordyluroides, ovipositor, ventral view. Abbreviations: tg, tergite; st, sternite.

188



Arista bare or with very short setulae; hind tibia with the distal posterodorsal seta as long as or shorter than proximal posterodorsal seta (Bahamas Is.; United States of America) ............................................... ................... P. cordyluroides (Stein, 1898)

Species of Pachyceramyia not included in the key and its respective geographical distribution Species

Geographical Distribution

P. pantherina (Wiedemann, 1824)

South America

Rhabdotoptera Stein, 1919 (Figs. 62··67) Type-species. Shoenomyza striatipennis Stein (mon.). Diagnosis. Eyes with small and sparse cilia; male dichoptic; 1 pair of reclinate orbital setae; epistoma not produced; antenna short, apex of flagellomere rounded; arista bare; palpus slender; scutellum bare below and laterally; acrostichals present before and after suture; dorsocentrals 2:3, the anterior one about half as long as the posterior setae; prealar absent; prosternum bare; proepisternals 2; propleurals 1, all directed upwards; katepisternals 1: 1: 1, forming an equilateral triangle and with an extra seta laterally to anterior one (Fig. 62); lower calypter short, hardly projecting beyond the upper one; wings with bare veins; veins R 4+s and M parallel apically (Fig. 67); legs with claws and pulvilli very small; hind tibia on anterodorsal surface with one long median seta, anteroventral surface with 2 short setae on middle third, dorsal surface with a preapical seta; sternite 1 with few short setulae; male: sternite 5 sub-triangular (Fig. 63); cercal plate sub-quadrate (Figs. 64, 65); hypandrium curved (Fig. 66). Reference. COURI, 1996a. Monophyly. Not yet established. Geographical record. Peru. Comments. Monobasic genus: Rhabdotoptera striatipennis (Stein, 1898).


189


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