Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Molecular phylogenetics of the Muscidae (Diptera : Calyptratae): new ideas in a congruence context

Guilherme Schnell e Schuehli A C , Claudio José Barros de Carvalho A and Brian M. Wiegmann B

A Departamento de Zoologia, Universidade Federal do Paraná, CP 19020, Curitiba 81.531-980, Brazil.

B North Carolina State University, Department of Entomology, College of Agriculture and Life Sciences, Gardner Hall, Campus Box 7613, Raleigh, NC 27695-7613, USA.

C Corresponding author. Email: schnell@onda.com.br

Invertebrate Systematics 21(3) 263-278 https://doi.org/10.1071/IS06026
Submitted: 12 July 2006  Accepted: 20 March 2007   Published: 27 June 2007

Abstract

Hypotheses about the evolution of Muscidae have long been the subject of continuous re-evaluation and reinterpretation. Current understandings of the relationships among these flies are based mainly on a single set of characters and are therefore questionable. Our understanding of muscid phylogeny thus needs greater support and further corroboration from additional suites of characters. In the current study, we analysed phylogenetic relationships among 24 species of muscid flies (18 genera and six subfamilies) using 2989 characters derived from sequences of mitochondrial (COI and COII) and nuclear genes (CAD and EF-1α). Data from each gene partition were analysed both in combined and separate phylogenetic analyses using maximum parsimony, maximum likelihood, and Bayesian inference. Support was found for the monophyly of the Muscidae in all analyses and for a sister-group relationship between Coenosiini and Phaoniinae. The latter group was placed in a clade with sampled species of Reinwardtiini and Cyrtoneurininae. The genera Ophyra and Hydrotaea were placed in the Muscinae and a sister-group relationship for Musca and Stomoxys was supported. Sampled species of Polietina form a monophyletic lineage, while Morellia was found to be paraphyletic. Combined analysis of gene partitions improved support and resolution for resulting topologies despite significant incongruence between data partitions found through application of the Incongruence Length Difference test.

Additional keywords: cytochrome oxidase, CAD, EF-1α, molecular systematics, Muscoidea, nuclear genes.


Acknowledgements

We gratefully acknowledge Walter Boeger, Marcia Souto Couri, Ana Maria Azeredo-Espin, Gabriel A. R. Melo and especially Marcio R. Pie for discussing the manuscript and their valuable suggestions. We also would like to express our thanks to Adrian Charles Pont, who provided the European species, and Lisete M. Lorini for the loan of several of the South American specimens included in the study. We appreciate the relevant comments made by the anonymous reviewers regarding this publication.


This project was supported by a fellowship from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES (Programa de Doutorado com Estágio no Exterior – PDEE/BEX0143/02–2), and grants from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), process number 140700/2001–3 (GSS) and 304.148/2002–4 (CJBC), and the US National Science Foundation (0098745, BMW and J. Thorne; and 0334948, BMW). This paper is the contribution number 01707 of the Departamento de Zoologia da Universidade Federal do Paraná (DZUP).


References


Adams M. D., Celniker S. E., Holt R. A., Evans C. A., Gocayne J. D. , et al.. (2000) The genome sequence of Drosophila melanogaster. Science 287, 2185–2195.
CrossRef | PubMed | open url image1

Akaike H. (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control 19, 716–723.
CrossRef | open url image1

Amorim D. S., and Pires M. R. S. (1996). Neotropical biogeography and a method for maximum biodiversity estimation. In ‘Biodiversity in Brazil, A First Approach’. (Eds C. E. M. Bicudo and N. A. Menezes.) pp. 183–219. (CNPq: São Paulo, Brazil.)

Baker R. H., DeSalle R. (1997) Multiple sources of character information and the phylogeny of the Hawaiian drosophilids. Systematic Biology 46, 654–673.
CrossRef | PubMed | open url image1

Baker R. H., Wilkinson G. S., DeSalle R. (2001) Phylogenetic utility of different types of molecular data used to infer evolutionary relationships among Stalk-Eyed flies (Diopsidae). Systematic Biology 50, 87–105.
CrossRef | PubMed | open url image1

Beckenbach A. T., Borkent A. (2003) Molecular analysis of the biting midges (Diptera: Ceratopogonidae), based on mitochondrial cytocrome oxidase subunit 2. Molecular Phylogenetics and Evolution 27, 21–35.
CrossRef | PubMed | open url image1

Beckenbach A. T., Wei Y. W., Liu H. (1993) Relationships in the Drosophila obscura species group, inferred from mitochondrial cytochrome oxidase II sequences. Molecular Biology and Evolution 10, 619–634.
PubMed |
open url image1

Bernasconi M. V., Pawlowski J., Valsangiacomo C., Piffaretti J.-C., Ward P. I. (2000a) Phylogeny of the Scathophagidae (Diptera, Calyptratae) based on mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 16, 308–315.
CrossRef | PubMed | open url image1

Bernasconi M. V., Valsangiacomo C., Piffaretti J.-C., Ward P. I. (2000b) Phylogenetic relationships among Muscoidea (Diptera: Calyptratae) based on mitochondrial DNA sequences. Insect Molecular Biology 9, 67–74.
CrossRef | PubMed | open url image1

Bernasconi M. V., Pawlowski J., Valsangiacomo C., Piffaretti J.-C., Ward P. I. (2001) Phylogeny of the genus Scathophaga (Diptera: Scathophagidae) inferred from mitochondrial DNA sequences. Canadian Journal of Zoology 79, 517–524.
CrossRef | open url image1

Bremer K. (1988) The Limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42, 795–803.
CrossRef | open url image1

Bremer K. (1994) Branch support and tree stability. Cladistics 10, 295–304.
CrossRef | open url image1

Brower A. (1996) Combining data in phylogenetic analysis. Trends in Ecology & Evolution 11, 334–335.
CrossRef | open url image1

Brower A. V. Z., DeSalle R. (1994) Practical and theoretical considerations for choice of a DNA sequence region in insect molecular systematics, with a short review of published studies using nuclear gene regions. Annals of the Entomological Society of America 87, 702–716. open url image1

Brown J. M., Pellmyr O., Thompson J. N., Harrison R. G. (1994) Phylogeny of Greya (Lepidoptera: Proxidae) based on nucleotide sequence variation in mitochondrial cytochrome oxidase I and II: congruence with morphological data. Molecular Biology and Evolution 11, 128–141.
PubMed |
open url image1

Bull J. J., Huelsenbeck J. P., Cunningham C. W., Swofford D. L., Waddell P. J. (1993) Partitioning and combining data in phylogenetic analysis. Systematic Biology 42, 384–397.
CrossRef | open url image1

Capaldi R. A., Malatesta F., Darley-Usmar V. M. (1983) Structure of cytochrome oxidase. Biochimica et Biophysica Acta 726, 135–148.
PubMed |
open url image1

Carvalho C. J. B. de (1989) Classificação de Muscidae (Diptera): uma proposta através da análise cladística. Revista Brasileira de Zoologia 6, 627–648. open url image1

Carvalho C. J. B. de (1999) Revision, cladistics and biogeography of the neotropical genus Souzalopesmyia Albuquerque (Diptera: Muscidae). Proceedings of the Entomological Society of Washington 101, 123–137. open url image1

Carvalho C. J. B. de (2002). ‘Muscidae (Diptera) of the Neotropical Region.’ (Editora Universidade Federal do Paraná: Curitiba, Brazil.)

Carvalho C. J. B. de, Couri M. S. (2002) A cladistic and biogeographic analysis of Apsil Malloch and Reynoldsia Malloch (Diptera, Muscidae) of southern South America. Proceedings of the Entomological Society of Washington 104, 309–317. open url image1

Carvalho C. J. B. de, Nihei S. S. (2002) A new genus of Phaoniinae (Diptera: Muscidae) from Central America. Annales Zoologici 52, 327–331. open url image1

Carvalho C. J. B. de, Pont A. C. (2006) Taxonomy, cladistics and biogeography of the South American genus Brachygasterina Macquart (Diptera: Muscidae). Zootaxa 1151, 1–26. open url image1

Carvalho C. J. B. de, Pont A. C., Couri M. S., Pamplona D. (2003) A catalogue of the Fannidae (Diptera) of the Neotropical Region. Zootaxa 210, 1–32. open url image1

Carvalho C. J. B. de, Couri M. S., Pont A. C., Pamplona D., Lopes S. M. (2005) A Catalogue of the Muscidae (Diptera) of the Neotropical Region. Zootaxa 860, 1–282. open url image1

Caterino M. S., Sperling F. A. H. (1999) Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes. Molecular Phylogenetics and Evolution 11, 122–137.
CrossRef | PubMed | open url image1

Cho S., Mitchell A., Regier J. C., Mitter C., Poole C., Friedlander T. P., Zhao S. (1995) A highly conserved nuclear gene for low-level phylogenetics: elongation factor 1-alpha recovers morphology-based tree for Heliothine moths. Molecular Biology and Evolution 12, 650–656.
PubMed |
open url image1

Collins K. P., Wiegmann B. M. (2002) Phylogenetic relationships of the lower Cyclorrhapha (Diptera: Brachycera) based on 28s rDNA sequences. Insect Systematics and Evolution 33, 445–456. open url image1

Couri M. S., Carvalho C. J. B. (1997) Revision of Polietina Schnabl & Dziedzicki (Diptera, Muscidae) and considerations on its new systematic position. Revista. Revista Brasileira de Zoologia 14, 255–281. open url image1

Couri M. S., Carvalho C. J. B. de (2003) Systematic relations among Philornis Meinert, Passeromyia Rodhain & Villeneuve and allied genera (Diptera, Muscidae). Brazilian Journal of Biology 63, 223–232.
CrossRef | open url image1

Couri M. S., Pont A. C. (2000) A cladistic analysis of Coenosiinae (Diptera, Muscidae). Systematic Entomology 25, 373–392.
CrossRef | open url image1

Cunningham C. W., Zhu H., Hillis D. M. (1998) Best-fit maximum-likelihood models for phylogenetic inference: empirical tests with known phylogenies. Evolution 52, 978–987.
CrossRef | open url image1

Cuny R. (1980) Revision of the genus Eudasyphora Townsend (Diptera: Muscidae), and reflections on its evolution. Canadian Entomologist 112, 345–373. open url image1

Danforth B. N., Ji S. (1998) Elongation factor-1α occurs as two copies in bees: implications for phylogenetics analysis of EF-1α in insects. Molecular Biology and Evolution 15, 225–235.
PubMed |
open url image1

Eastwood R., Hughes J. M. (2003) Molecular phylogeny and evolutionary biology of Acrodipsas (Lepidoptera: Lycaenidae). Molecular Phylogenetics and Evolution 27, 93–102.
CrossRef | PubMed | open url image1

Evenhuis N. L. (1994). ‘Catalogue of the Fossil Flies of the World (Insecta: Diptera).’ (Leiden: Backhuys, The Netherlands.)

Farris J. S., Källersjö M., Kluge A. G., Bult C. (1994) Testing significance of incongruence. Cladistics 10, 315–319.
CrossRef | open url image1

Felsenstein J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39, 783–791.
CrossRef | open url image1

Felsenstein J. (1993). PHYLIP (phylogeny inference package), Version3.5c. (Department of Genetics, University of Washington: Seattle, USA.)

Frailey C. D. (2002). Neogene paleogeography of the Amazon basin. In ‘TER-QUA Symposium Series, 3’. (Ed. W. Dort Jr.) pp. 71–97. (Institute for Tertiary-Quaternary Studies: Lincoln, USA.)

Frati F., Simon C., Sullivan J., Swofford D. L. (1997) Evolution of the mitochondrial cytochrome oxidase II gene in Collembola. Journal of Molecular Evolution 44, 145–158.
CrossRef | PubMed | open url image1

Hecker K. H., Roux K. H. (1996) High and low annealing temperatures increase both specificity and yield in “Touchdown” and “StepDown” PCR. BioTechniques 20, 478–485.
PubMed |
open url image1

Hennig W. (1955–1964). Muscidae. In ‘Die Fliegen der Paläarktischen Region, Teil 63b’. (Ed. E. Lindner.) pp. 97–1056. (Schweizerbartische Verlagsbuchhandlung: Stuttgart, Denmark.)

Hennig W. (1965) Vorarbeiten zu einem phylogenetischen System der Muscidae (Diptera: Cyclorrhapha). Stuttgarter Beitrage zur Naturkunde 141, 000–000. open url image1

Hennig W. (1973). Diptera. In ‘Handbuch der Zoologie, IV: Arthropoda’. (Ed W. Kukenthal). pp. 1–337. (De Gruyter: New York, USA.)

Holt R. A., Subramanian G. M., Halpern A., Sutton G. G., Charlab R., Nusskern D. R., Wincker P. , et al.. (2002) The genome sequence of the malaria mosquito Anopheles gambiae.  , 129–149.
CrossRef | open url image1

Howland D. E., Hewitt G. M. (1995) Phylogeny of the Coleoptera based on mitochondrial cytochrome oxidase I sequence data. Insect Molecular Biology 4, 203–215.
PubMed |
open url image1

Huelsenbeck J. P., Rannala B. (1997) Phylogenetic methods come of age: testing hypothesis in an evolutionary context. Science 276, 227–232.
CrossRef | PubMed | open url image1

Huelsenbeck J. P., Ronquist F. (2001) MRBAYES: Bayesian inference of phylogeny. Bioinformatics 17, 754–755.
CrossRef | PubMed | open url image1

Johnson K. P., Moylet R. G., Witt C. C., Faucett R. C., Weckstein J. D. (2001) Phylogenetic relationships in the louse genus Penenirmus based on nuclear (EF-1α) and mitochondrial (COI) DNA sequences. Systematic Entomology 26, 491–497.
CrossRef | open url image1

Johnson L. A., and Soltis D. E. (1998). Assessing congruence: empirical examples from molecular data. In ‘Molecular Systematics of Plants.’ Vol. II. (Eds P. S. Soltis, D. E. Soltis and J. J. Doyle.) pp. 297–343. (Chapman and Hall: New York, USA.)

Kawasaki E. S. (1990). Amplification of RNA. In ‘PCR Protocols’. (Eds M. A. Innis, D. H. Gelfand, J. J. Sninsky and T. J. White.) pp. 21–27. (Academic Press: San Diego, USA.)

Kluge A. G. (1989) A concern for evidence and a phylogenetic hypothesis for relationships among Epicrates (Boidae, Serpentes). Systematic Zoology 38, 7–25.
CrossRef | open url image1

Kluge A. G., Wolf A. J. (1993) Cladistics: what’s in a word? Cladistics 9, 183–199.
CrossRef | open url image1

Lemmon A. R., Moriarty E. C. (2004) The importance of proper model assumption in bayesian phylogenetics. Systematic Biology 53, 265–277.
CrossRef | PubMed | open url image1

Lessinger A. C., Martins Junqueira A. C., Lemos T. A., Kemper E. L., da Silva F. R., Vettore A. L., Arruda P., Azeredo-Espin A. M. L. (2000) The mitochondrial genome of the primary screwworm fly Cochliomyia homnivorax (Diptera: Calliphoridae). Insect Molecular Biology 9, 521–529.
CrossRef | PubMed | open url image1

Liu H., Beckenbach A. T. (1992) Evolution of the mitochondrial cytochrome oxidase II gene among 10 orders of insects. Molecular Phylogenetics and Evolution 1, 41–52.
CrossRef | PubMed | open url image1

Lundberg J. G., Marshall L. G., Guerrero J., Horton B., Malabarba M. C. S. L., and Wesselingh F. (1998). The stage for Neotropical fish diversification: a history of tropical South American rivers. In ‘Phylogeny and Classification of Neotropical Fishes’. (Eds L. R. Malabarba, R. E. Reis, R. P Vari, Z. M. S. Lucena and C. A. S. Lucena.) pp. 13–48. (EDIPUCRS: Porto Alegre, Brazil.)

Malloch J. R. (1934). Muscidae. In ‘Diptera of Patagonia and South Chile 7’. pp. 171–346. (British Museum of Natural History: London.)

McAlpine J. F. (1989). Phylogeny and classification of the Muscomorpha. In ‘Manual of Nearctic Diptera, Vol 3’. (Eds. J. F. McAlpine and D. M. Wood.) pp. 1397–1518. (Research Branch of Agriculture and Agri-Food Canada: Ottawa, Canada.)

Méier R., Wiegmann B. M. (2002) A phylogenetic analysis of Coelopidae (Diptera) based on morphological and DNA sequence data. Molecular Phylogenetics and Evolution 25, 393–407.
CrossRef | PubMed | open url image1

Michelsen V. (1991) Revision of the aberrant New World genus Coenosopsia (Diptera: Anthomyiidae), with a discussion of anthomyiid relationships. Systematic Entomology 16, 85–104. open url image1

Mitchell A., Cho S., Regier J. C., Mitter C., Poole R. W., Matthews M. (1997) Phylogenetic utility of elongation factor-1α in Noctuoidea (Insecta: Lepidoptera): the limits of synonymous substitution. Molecular Biology and Evolution 14, 381–390.
PubMed |
open url image1

Miyamoto M. M., Fitch W. M. (1995) Testing species phylogenies and phylogenetic methods with congruence. Systematic Biology 44, 64–76.
CrossRef | open url image1

Motta H. C. G., Couri M. S. (1999) Revisão das espécies de Bithoracochaeta Stein (Diptera, Muscidae). Revista Brasileira de Zoologia 16, 665–689. open url image1

Moulton J. K., Wiegmann B. M. (2004) Evolution and phylogenetic utility of CAD (rudimentary) among Mesozoic-aged Eremoneuran Diptera (Insecta). Molecular Phylogenetics and Evolution 31, 363–378.
CrossRef | PubMed | open url image1

Nihei S. S. (2004). Sistemática e biogeografia de Muscini (Diptera, Muscidae). Ph.D. Thesis, Curso de Pós-Graduação em Entomologia, Universidade Federal do Paraná, Curitiba, Brasil.

Nihei S. S., Carvalho C. J. B. de (2004) Taxonomy, cladistics and biogeography of Coenosopsia Malloch (Diptera, Anthomyiidae) and its significance to the evolution of anthomyiids in the Neotropics. Systematic Entomology 29, 260–275.
CrossRef | open url image1

Page D. M., and Holmes E. C. (1998). ‘Molecular Evolution: A Phylogenetic Approach.’ (Blackwell: Malden, MA, USA.)

Pamplona D. M. (1986) Revalidação de Biopyrellia Townsend, 1932 com redescrição de B. bipuncta (Wiedemann, 1830) (Diptera, Muscidae, Muscinae). Revista Brasileira de Biologia 46, 49–54. open url image1

Pamplona D. M. (1999) Nova caracterização de Cyrtoneurina Giglio-Tos, 1893 e descrição de Paracyrtoneurina g.n. (Diptera, Muscidae). Revista Brasileira de Entomologia 43, 9–24. open url image1

Petri S., and Fúlfaro V. J. (1983). ‘Geologia do Brasil (Fanerozóico).’ (T. A. Queiroz and EDUSP: São Paulo, Brazil.)

Pitman W. C., Cande S., LaBrecque J., and Pindell J. (1993). Fragmentation of Gondwana: the separation of Africa and South America. In ‘Biological Relationships between Africa and South America’. (Ed. P. Goldblatt.) pp. 15–34. (Yale University Press: New Haven, CT, USA.)

Pont A. C. (1972). Family Muscidae. In ‘A Catalogue of the Diptera of the Americas South of the United States, 97’. (Ed. N. Papavero) pp. 1–111. (Museu de Zoologia, Universidade de São Paulo: São Paulo, Brazil.)

Pont A. C. (1986). Family Muscidae. In:‘Catalogue of the Palaearctic Diptera, Vol. 11’. (Eds A. S. Sóos and L. Papp.) pp. 57–215. (Hungarian Natural History Museum: Budapeste, Hungary.)

Pont A. C., Carvalho C. J. B. (1997) Three new species of Muscidae (Diptera) from Dominican amber. Studia Dipterologica 4, 173–181. open url image1

Posada D., Crandall K. A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics 14, 817–818.
CrossRef | PubMed | open url image1

Roback S. S. (1951) A classification of the muscoid calyptrate Diptera. Annals of the Entomological Society of America 44, 327–361. open url image1

Rokas A., Nylander J. A. A., Ronquist F., Stone G. N. (2002) A maximum-likelihood analysis of eight phylogenetic markers in gallwasps (Hymenoptera: Cynipidae): implications for insect phylogenetic studies. Molecular Phylogenetics and Evolution 22, 206–219.
CrossRef | PubMed | open url image1

Savage J., Wheeler T. A. (2004) Phylogeny of the Azeliini (Diptera: Muscidae). Studia Dipterologica 11, 259–299. open url image1

Savage J., Wheeler T. A., Wiegmann B. M. (2004) Phylogenetic analysis of the genus Thricops Rondani (Diptera: Muscidae) based on molecular and morphological characters. Systematic Entomology 29, 395–414.
CrossRef | open url image1

Scheffer S., Wiegmann B. M. (2000) Molecular phylogenetics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): species limits, speciation, and dietary specialization. Molecular Phylogenetics and Evolution 17, 244–255.
CrossRef | PubMed | open url image1

Schmidt H. A., Strimmer K., Vingron M., von Haeseler A. (2002) TREE-PUZZLE: maximum likelihood phylogenetic analysis using quartets and parallel computing. Bioinformatics 18, 502–504.
CrossRef | PubMed | open url image1

Schuehli G. S., Carvalho C. J. B. de (2005) Revision and cladistics of the Neotropical genus Pseudoptilolepis Snyder, (Diptera: Muscidae). Revista Brasileira de Zoologia 22, 23–34.
CrossRef | open url image1

Schuehli G. S., Carvalho C. J. B. de, Wiegmann B. M. (2004) Regarding the taxonomic status of Ophyra Robineau-Desvoidy (Diptera: Muscidae): a molecular approach. Zootaxa 712, 1–12. open url image1

Séguy E. (1937). Diptera, Family Muscidae. In ‘Genera Insectorum. Fasc. 205’. (Ed. P. Wytsman.) pp. 1–604. (P. Wystman: Brussels, Belgium.)

Sharpe R. G., Harbach R. E., Butlin R. K. (2000) Molecular variation and phylogeny of members of the Mimus group of Anopheles subgenus Cellia (Diptera: Culicidae). Systematic Entomology 25, 263–272.
CrossRef | open url image1

Simon C., Frati F., Beckenbach A., Crespi B., Liu H., Flook P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America 87, 651–701. open url image1

Skidmore P. (1985). ‘The Biology of the Muscidae of the World.’ (Springer: Dordrecht, Germany.)

Smith S. W., Overbeek R., Woese C. R., Gilbert W., Gillevet P. M. (1994) The genetic data enviroment and expandible GUI for multiple sequence analysis. Computer Applications in the Biosciences 10, 671–675.
PubMed |
open url image1

Snyder F. M. (1954) A revision of Cyrtoneurina Giglio-Tos, with notes on related genera (Diptera, Muscidae). Bulletin of the American Museum of Natural History 103, 417–464. open url image1

Soares E. D. G., Carvalho C. J. B. de (2005) Taxonomia, análise cladística e descrição de uma espécie nova de Palpibracus Rondani (Diptera, Muscidae). Revista Brasileira de Entomologia 49, 171–180.
CrossRef | open url image1

Sorenson M. D. (1999). ‘TreeRot, Version 2.’ (Boston University: Boston, MA, USA.)

Sperling F. A. H., Hickey D. A. (1994) Mitochondrial DNA sequence variation in the spruce budworm species complex (Choristoneura: Lepidoptera). Molecular Biology and Evolution 11, 656–665.
PubMed |
open url image1

Sperling F. A. H., Anderson G. S., Hickey D. A. (1994) A DNA-based approach to the identification of insect species used for postmortem interval estimation. Journal of Forensic Sciences 39, 418–427.
PubMed |
open url image1

Strimmer K., von Haeseler A. (1996) Quartet puzzling: a quartet maximum likelihood method for reconstructing tree topologies. Molecular Biology and Evolution 13, 964–969. open url image1

Swofford D. L. (2002). ‘PAUP*. Phylogenetic analysis using parsimony (* and other methods) Version 4’. (Sinauer Associates: Sunderland, MA, USA.)

Swofford D. L., Waddell P. J., Huelsenbeck J. P., Foster P. G., Lewis P. O., Rogers J. S. (2001) Bias in phylogenetic estimation and its relevance to the choice between parsimony and likelihood methods. Systematic Biology 50, 525–539.
CrossRef | PubMed | open url image1

Yang L., Wiegmann B. M., Yeates D. K., Irwin M. E. (2000) Higher-level phylogeny of the Therevidae (Diptera: Insecta) based on 28s ribosomal and Elongation Factor-1α gene sequences. Molecular Phylogenetics and Evolution 15, 440–451.
CrossRef | PubMed | open url image1

Yeates D. K., Wiegmann B. M. (1999) Congruence and controversy: toward a higher-level phylogeny of Diptera. Annual Review of Entomology 44, 397–428.
CrossRef | PubMed | open url image1










Appendix 1.  Species sampled and GenBank accession number for each gene fragment
Note that the sequences for COI (2792/C2N3389), tRNA-Leu and COII (2792/C2N3389) are continuous and so are listed with identical GenBank accession numbers. An asterisk indicates sequences originally published in Schuehli et al. (2004)
Click to zoom



Appendix 2.  Species included in the analysis, their respective subfamilies and tribes (based on Carvalho 2002; Couri and Carvalho 2003), and geographical origin of specimen
Click to zoom



Rent Article (via Deepdyve) Export Citation Cited By (13)