Hennig's orphans revisited: Testing morphological hypotheses in the "Opomyzoidea" (Diptera: Schizophora)

Molecular Phylogenetics and Evolution

Volumn 54, Issue 3, 2010, Pages 746-762

  Winkler, Isaac S ^a,d   Rung, Alessandra ^b,c,d   Scheffer, Sonja J ^c,d  

^a NORTH CAROLINA STATE UNIVERSITY   (United States)

^b CALIFORNIA DEPARTMENT OF FOOD AND AGRICULTURE   (United States)

^c AGRICULTURAL RESEARCH SERVICE   (United States)

^d Bldg 142   (United States)

Author keywords

Acalyptrate; Higher classification; Phylogeny; Rapid radiation

Indexed keywords

RNA 28S;

ANIMAL; ARTICLE; BIOLOGICAL MODEL; CLASSIFICATION; DNA SEQUENCE; FLY; GENE; GENETICS; MOLECULAR EVOLUTION; PHYLOGENY; STATISTICAL MODEL;

ANIMALS; DIPTERA; EVOLUTION, MOLECULAR; GENES, INSECT; LIKELIHOOD FUNCTIONS; MODELS, GENETIC; PHYLOGENY; RNA, RIBOSOMAL, 28S; SEQUENCE ANALYSIS, DNA;

AULACIGASTRIDAE; AUSTRALIMYZIDAE; CYAMOPS; DIPTERA; OPOMYZOIDEA; PERISCELIDIDAE; SCHIZOPHORA; STENOMICRA; XENASTEIIDAE;

EID: 76449085509     PISSN: 10557903     EISSN: 10959513     Source Type: Journal    
DOI: 10.1016/j.ympev.2009.12.016     Document Type: Article

References (113)

1. Baptista A.R., and Mathis W.N. A revision of New World Cyamops Melander (Diptera, Periscelididae). Smithson. Contrib. Zool. 563 (1994) 1-28
2. Barraclough D.A., and McAlpine D.K. Natalimyzidae, a new African family of acalyptrate flies (Diptera: Schizophora: Sciomyzoidea). Afr. Invertebr. 47 (2006) 117-134
3. Baurain D., Brinkmann H., and Philippe H. Lack of resolution in the animal phylogeny: closely spaced cladogeneses or undetected systematic errors?. Mol. Biol. Evol. 24 (2007) 6-9
4. Bergsten J. A review of long branch attraction. Cladistics 21 (2005) 163-193
5. Bernasconi M.V., Valsangiacomo C., Piffaretti J.C., and Ward P.I. Phylogenetic relationships among Muscoidea (Diptera: Calyptratae) based on mitochondrial DNA sequences. Insect Mol. Biol. 9 (2000) 67-74
6. Bertone M.A., Courtney G.W., and Wiegmann B.M. Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes. Syst. Entomol. 33 (2008) 668-687
7. Blagoderov V.A., Lukashevich E.D., and Mostovski M.B. Order Diptera Linné, 1758. The true flies (= Muscida Laicharting, 1781). In: Rasnitsyn A.P., and Quicke D.L.J. (Eds). History of Insects (2002), Kluwer Academic Publishers, Dordrecht 227-240
8. Blanquart S., and Lartillot N. A Bayesian compound stochastic process for modeling non-stationary and nonhomogeneous sequence evolution. Mol. Biol. Evol. 23 (2006) 2058-2071
9. Brake I. Phylogenetic systematics of the Milichiidae (Diptera, Schizophora). Entomol. Scand. 57 (2000) 1-58
10. Brake I., and Mathis W.N. Revision of the genus Australimyza Harrison (Diptera: Australimyzidae). Syst. Entomol. 32 (2007) 252-275
11. Britton, D., 2009. Rare - or just hidden? Australian Museum Online. Available from: (accessed Jan. 2010).
12. Brunel E. Family Opomyzidae. In: Soós A., and Papp L. (Eds). Catalogue of Palearctic Diptera (1998), Elsevier Science Publishers and Akadémiai Kiadó, Budapest 259-266
13. Buck M. A new family and genus of acalypterate flies from the Neotropical region, with a phylogenetic analysis of Carnoidea family relationships (Diptera, Schizophora). Syst. Entomol. 31 (2006) 377-404
14. Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 17 (2000) 540-552
15. Castresana, J., 2002. Gblocks v.0.91b. Available from: .
16. Colless D.H., and McAlpine D.K. Diptera (flies). In: Naumann I.D. (Ed). The Insects of Australia. A Textbook for Students and Research Workers. second ed. vol. II (1991), Melbourne University Press, Carlton, Victoria, Australia 717-786
17. Currie G.A. Galls on Eucalyptus trees: a new type of association between flies and nematodes. Proc. Linn. Soc. N. S. W. 62 (1937) 147-174
18. Edgar R.C. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res. 32 (2004) 1792-1797. http://www.drive5.com/muscle Available from:
19. Evenhuis, N.L., 1994. Catalogue of the Fossil Flies of the World (Insecta: Diptera). Available from: (accessed Oct. 2008).
20. Evenhuis, N.L., Pape, T., Pont, A.C., Thompson, F.C. (Eds.), 2008. Biosystematic Database of World Diptera, Working Version. Available from: (accessed Dec. 2008).
21. Fish D. Phytotelmata: Flora and Fauna. In: Frank J.H., and Lounibos L.P. (Eds). Phytotelmata: Terrestrial Plants as Hosts for Aquatic Insect Communities (1983), Plexus, Medford, New Jersey 1-27
22. Freidberg A. Nemula, a new genus of Neminidae (Diptera) from Madagascar. Proc. Entomol. Soc. Wash. 96 (1994) 471-482
23. Freidberg A. Recent advances in the study of Asteiidae. Abstracts Volume, Fifth International Congress of Dipterology, Brisbane, Australia (2002), University of Queensland, Brisbane p. 73
24. Gaimari, S.D., Mathis, W.N., in press. World catalog and conspectus of the family Odiniidae (Diptera: Schizophora). Myia.
25. Goloboff P.A. Self-weighted optimization: tree searches and character state reconstructions under implied transformation costs. Cladistics 13 (1997) 225-245
26. Gowri-Shankar V., and Rattray M. A reversible jump method for Bayesian phylogenetic inference with a nonhomogeneous substitution model. Mol. Biol. Evol. 24 (2007) 1286-1299
27. Griffiths G.C.D. The phylogenetic classification of Diptera Cyclorrhapha with special reference to the structure of the male postabdomen. Ser. Entomol. 8 (1972) 1-340 The Hague
28. Grimaldi D.A., and Engel M.S. Evolution of the Insects (2005), Cambridge University Press, Cambridge, UK
29. Grimaldi D.A., and Mathis W.N. Fossil Periscelididae (Diptera). Proc. Entomol. Soc. Wash. 95 (1993) 383-403
30. Grimaldi D.A., Beck C.W., and Boon J.J. Occurrence, chemical characteristics, and paleontology of the fossil resins from New Jersey. Am. Mus. Novit. 2948 (1989) 1-28
31. Han H.Y., and McPheron B.A. Molecular phylogenetic study of Tephritidae (Insecta: Diptera: Tephritidae) using partial sequences of the mitochondrial 16S ribosomal DNA. Mol. Phylogenet. Evol. 7 (1997) 17-32
32. Han H.Y., and Ro K.E. Molecular phylogeny of the superfamily Tephritoidea (Insecta: Diptera): new evidence from the mitochondrial 12S, 16S, and COII genes. Mol. Phylogenet. Evol. 34 (2005) 416-430
33. Hardy D.E. Xenasteiidae, a new family of Schizophora from the Pacific and Indian oceans. Proc. Hawaii Entomol. Soc. 23 (1980) 205-225
34. Hennig W. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen. Beitr. Entomol. 8 (1958) 505-688
35. Hennig W. Die Acalyptraten des Baltischer Bernsteins. Stuttgarter Beitr. Naturkd. 145 (1965) 1-215
36. Hennig W. Neue Gattungen und Arten der Acalypteratae. Can. Entomol. 101 (1969) 589-633
37. Hennig W. Neue Untersuchungen über die Familien der Diptera Schizophora (Diptera: Cyclorrhapha). Stuttgarter Beitr. Naturkd. 226 (1971) 1-76
38. Hennig W. Diptera. In: Kukenthal W. (Ed). Handbuch der Zoologie, IV: Arthropoda, Section 2, Part 2 vol. 31 (1973), De Gruyter, New York 1-337
39. Hoffeins C., and Rung A. Procyamops succini, a new genus and new species of Periscelididae (Diptera, Brachycera) from Baltic amber (Tertiary, Eocene). Stud. Dipterol. 12 (2005) 23-27
40. Jian S., Soltis P.S., Gitzendanner M.A., Moore M.J., Li R., Hendry T.A., Qiu Y.-L., Dhingra A., Bell C.D., and Soltis D.E. Resolving an ancient rapid radiation in Saxifragales. Syst. Biol. 57 (2008) 38-57
41. Khoo K.C., and Sabrosky C.W. Family Stenomicridae. In: Evenhuis N.L. (Ed). Catalog of the Diptera of the Australasian and Oceanian Regions. B.P. Bishop Museum Special Publication No. 86. (1989), E.J. Brill and Bishop Museum Press, Honolulu p. 551
42. Kotrba M., and Baptista A.R. The internal reproductive tract of the Opomyzidae (Diptera, Schizophora). Stud. Dipterol. 9 (2002) 57-71
43. Kutty S.N., Pape T., Pont A., Wiegmann B.M., and Meier R. The Muscoidea (Diptera: Calyptratae) is paraphyletic: evidence from four mitochondrial and four nuclear genes. Mol. Phylogenet. Evol. 49 (2008) 639-652
44. Lartillot N., and Philippe H. A Bayesian mixture model for across-site heterogeneities in the amino-acid replacement process. Mol. Biol. Evol. 21 (2004) 1095-1109
45. Lartillot, N., Blanquart, S., Lepage, T., 2004. PhyloBayes 2.3: a Bayesian software for phylogenetic reconstruction using mixture models. Available from: .
46. Lartillot N., Brinkmann H., and Philippe H. Suppression of long-branch attraction artefacts in the animal phylogeny using a site-heterogeneous model. BMC Evol. Biol. 7 Suppl. 1 (2007) S4
47. Lonsdale O., and Marshall S.A. Redefinition of the Clusiinae and Clusiodinae, description of the new subfamily Sobarocephalinae, revision of the genus Chaetoclusia and a description of Procerosoma gen. n. (Diptera: Clusiidae). Eur. J. Entomol. 103 (2006) 163-182
48. Lonsdale O., and Marshall S.A. Redefinition of the genera Clusiodes and Hendelia (Diptera: Clusiidae: Clusiodinae) with a review of Clusiodes. Stud. Dipterol. 14 (2007) 117-159
49. Lonsdale O., and Marshall S.A. Synonymy within Clusia and description of the new genus Melanoclusia (Diptera: Clusiidae: Clusiinae). Ann. Entomol. Soc. Am. 101 (2008) 327-330
50. Lutzoni F., Wagner P., Reeb V., and Zoller S. Integrating ambiguously aligned regions of DNA sequences in phylogenetic analyses without violating positional homology. Syst. Biol. 49 (2000) 628-651
51. Mathis, W.N., in press. World Catalog and Conspectus on the Family Periscelididae (Diptera: Schizophora). Myia.
52. Mathis W.N., and Papp L. A new genus of Periscelididae (Diptera) from the Neotropics. Proc. Biol. Soc. Wash. 105 (1992) 366-372
53. Mathis W.N., and Papp L. Family Periscelididae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 285-294
54. Mathis W.N., and Rung A. Redescription of the genus Diopsosoma Malloch (Diptera, Periscelididae). Rev. Bras. Entomol. 48 (2004) 303-309
55. McAlpine J.F. First record of calypterate flies in the Mesozoic era (Diptera: Caliphoridae). Can. Entomol. 102 (1970) 342-346
56. McAlpine D.K. Description and biology of a new genus of flies related to Anthoclusia and representing a new family (Diptera, Schizophora, Neurochaetidae). Ann. Natal Mus. 23 (1978) 273-295
57. McAlpine D.K. A new subfamily of Aulacigastridae (Diptera: Schizophora), with a discussion of aulacigastrid classification. Aust. J. Zool. 31 (1983) 55-78
58. McAlpine D.K. Studies in upside-down flies (Diptera: Neurochaetidae). Part I. Systematics and phylogeny. Proc. Linn. Soc. N. S. W 110 (1988) 31-58
59. McAlpine D.K. Studies in upside-down flies (Diptera: Neurochaetidae). Part II. Biology, adaptations and specific mating mechanisms. Proc. Linn. Soc. N. S. W 110 (1988) 59-82
60. McAlpine, J.F., 1989. Phylogeny and classification of the Muscomorpha. In: McAlpine, J.F., Wood, D.M. (Eds.), Manual of Nearctic Diptera, Volume 3: Research Branch, Agriculture Canada, Monograph 32, pp. 1397-1518.
61. McAlpine D.K. Marginidae, a new Afrotropical family of Diptera (Schizophora: Opomyzoidea). Ann. Natal Mus. 32 (1991) 167-177
62. McAlpine D.K. Review of the upside-down flies (Diptera: Neurochaetidae) of Madagascar and Africa, and evolution of neurochaetid host plant associations. Rec. Aust. Mus. 45 (1993) 221-239
63. McAlpine D.K. Gobryidae, a new family of acalyptrate flies (Diptera: Diopsoidea), and a discussion of relationships of the diopsoid families. Rec. Aust. Mus. 49 (1997) 167-194
64. McAlpine D.K. Neminidae (Nobody flies). In: Oosterbroek P. (Ed). The Families of Diptera of the Malay Archipelago. Fauna Malesiana Handbook 1 (1998), Brill, Leiden 123-124
65. McAlpine D.K. Review of the Borboroidini or wombat flies (Diptera: Heteromyzidae), with reconsideration of the status of families Heleomyzidae and Sphaeroceridae, and descriptions of femoral gland-baskets. Rec. Aust. Mus. 59 (2007) 143-219
66. McAlpine D.K., and de Keyzer R.G. Generic classification of the fern flies (Diptera: Teratomyzidae) with a larval description. Syst. Entomol. 19 (1994) 305-326
67. Meier R. Role of dipterology in phylogenetic systematics: the insight of Willi Hennig. In: Yeates D.K., and Wiegmann B.M. (Eds). The Evoutionary Biology of Flies (2005), Columbia University Press, New York 45-62
68. Meier R., and Baker R. A cladistic analysis of Diopsidae (Diptera) based on morphological and DNA sequence data. Insect Syst. Evol. 33 (2002) 325-336
69. Meier R., and Wiegmann B.M. A phylogenetic analysis of Coelopidae (Diptera) based on morphological and DNA sequence data. Mol. Phylogenet. Evol. 25 (2002) 393-407
70. Moulton J.K., and Wiegmann B.M. Evolution and phylogenetic utility of CAD (rudimentary) among Mesozoic-aged eremoneuran Diptera (Insecta). Mol. Phylogenet. Evol. 31 (2004) 363-378
71. Moulton J.K., and Wiegmann B.M. The phylogenetic relationships of flies in the superfamily Empidoidea (Insecta: Diptera). Mol. Phylogenet. Evol. 43 (2007) 701-713
72. Murphy D.H. The natural history of insect herbivory on mangrove trees in and near Singapore. Raffles Bull. Zool. 38 (1990) 119-203
73. Nirmala X., Hypsa V., and Zurovec M. Molecular phylogeny of Calyptratae (Diptera: Brachycera): the evolution of 18S and 16S ribosomal rDNAs in higher dipterans and their use in phylogenetic inference. Insect Mol. Biol. 10 (2001) 475-485
74. Ortiz-Rivas B., Moya A., and Martínez-Torres D. Molecular systematics of aphids (Homoptera: Aphididae): new insights from the long-wavelength opsin gene. Mol. Phylogenet. Evol. 30 (2004) 24-37
75. Papp L. New taxa of acalyptrate flies (Diptera: Tunisimyiidae fam. n., Risidae, Ephydridae: Nannodastiinae subfam. n.). Acta Zool. Acad. Sci. Hung. 26 (1980) 415-431
76. Papp L. Family Aulacigastridae. In: Soós A., and Papp L. (Eds). Catalogue of Palearctic Diptera (1984), Elsevier Science Publishers and Akadémiai Kiadó, Budapest 60-61
77. Papp L. Family Aulacigastridae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 279-284
78. Papp L. Family Asteiidae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 295-303
79. Papp L. Family Xenasteiidae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 305-308
80. Papp L., and Ozerov A.L. Family Acartophthalmidae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 227-232
81. Papp L., Merz B., and Földvári M. Diptera of Thailand: a summary of the families and genera with references to the species representations. Acta Zool. Acad. Sci. Hung. 52 (2006) 97-268
82. Petersen F.T., Meier R., Kutty S.N., and Wiegmann B.M. The phylogeny and evolution of host choice in the Hippoboscoidea (Diptera) as reconstructed using four molecular markers. Mol. Phylogenet. Evol. 45 (2007) 111-122
83. Remsen J., and O'Grady P.M. Phylogeny of Drosophilidae (Diptera), with comments on combined analysis and character support. Mol. Phylogenet. Evol. 24 (2002) 248-263
84. Roháček J. Taxonomic limits, phylogeny and higher classification of Anthomyzidae (Diptera), with special regard to fossil record. Eur. J. Entomol. 95 (1998) 141-177
85. Roháček J. Family Anthomyzidae. In: Papp L., and Darvas B. (Eds). Contributions to a Manual of Palaearctic Diptera, Volume 3: Higher Brachycera (1998), Science Herald, Budapest 267-278
86. Roháček J. A monograph of Palaearctic Anthomyzidae (Diptera) part 1. Čas. Slez. Muz. Opava (A) 55 suppl. 1 (2006) 1-328
87. Ronquist F., and Huelsenbeck J.P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19 (2003) 1572-1574
88. Rung A., Mathis W.N., and Papp L. Curiosimusca, gen. nov., and three new species in the family Aulacigastridae from the Oriental Region (Diptera: Opomyzoidea). Zootaxa 1009 (2005) 21-36
89. Scheffer S.J., Giblin-Davis R.M., Davies K.A., Taylor G.S., Purcell M., Lewis M.L., Goolsby J., and Center T.D. Phylogenetic relationships, species limits, and host specificity of gall-forming Fergusonina flies (Diptera: Fergusoninidae) feeding on Melaleuca (Myrtaceae). Ann. Entomol. Soc. Am. 97 (2004) 1216-1221
90. Scheffer S.J., Winkler I.S., and Wiegmann B.M. Phylogenetic relationships within the leaf-mining flies (Diptera: Agromyzidae) inferred from sequence data from multiple genes. Mol. Phylogenet. Evol. 42 (2007) 756-775
91. Shatalkin A.I. On the taxonomy of the flies of the family Strongylophthalmyiidae (Diptera). Zool. Zh. 72 (1993) 124-131
92. Shimodaira H. An approximately unbiased test of phylogenetic tree selection. Syst. Biol. 51 (2002) 492-508
93. Shimodaira H., and Hasegawa M. Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Mol. Biol. Evol. 16 (1999) 1114-1116
94. Shimodaira H., and Hasegawa M. CONSEL: for assessing the confidence of phylogenetic tree selection. Bioinformatics 17 (2001) 1246-1247. http://www.is.titech.ac.jp/~shimo/prog/consel/ Available from:
95. Speight M.C.D. The prothoracic morphology of Acalyptrates (Diptera) and its use in systematics. Trans. R. Entomol. Soc. Lond. 121 (1969) 325-421
96. Spencer, K.A., 1990. Host specialization in the world Agromyzidae (Diptera). Ser. Entomol. 45, 1-444. Kluwer Academic Publishers, Dordrecht.
97. Su F.-Y.K., Kutty S., and Meier R. Morphology versus molecules: the phylogenetic relationships of Sepsidae (Diptera: Cyclorrhapha) based on morphology and DNA sequence data from ten genes. Cladistics 24 (2008) 902-916
98. Suzuki Y., Glazko G.V., and Nei M. Overcredibility of molecular phylogenies obtained by Bayesian phylogenetics. Proc. Natl. Acad. Sci. USA 99 (2002) 16138-16143
99. Swofford D.L. PAUP* 4.0 beta 10. Phylogenetic Analysis Using Parsimony (*and Other Methods) (2001), Sinauer Associates, Massachussets
100. Tautz D., Hancock J.M., Webb D.A., Tautz C., and Dover G.A. Complete sequences of the rRNA genes of Drosophila melanogaster. Mol. Biol. Evol. 5 (1988) 366-376
101. Taylor G.S., Davies K.A., and Giblin-Davis R.M. Species-richness in gall-flies (Diptera: Fergusoninidae), nematodes (Nematoda: Sphaerulariidae) and associated parasitoids and inquilines on Myrtaceae. Rec. S. Aust. Mus. Monogr. Ser. 7 (2004) 249-256
102. Taylor G.S., Head E.R., and Davies K.A. Gall flies (Diptera: Fergusoninidae) on Myrtaceae: a mutualistic association between flies and nematodes. In: Raman A., Schaefer C.W., and Withers T.M. (Eds). Biology, Ecology, and Evolution of Gall-inducing Arthropods (2005), Science Publishers, Enfield, New Hampshire 643-671
103. Thompson J.D., Higgins D.G., and Gibson T.J. Clustal W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Res. 22 (1994) 4673-4680
104. Vockeroth, J.R., 1987. Family Opomyzidae. In: McAlpine, J.F., Wood, D.M. (Eds), Manual of Nearctic Diptera, Volume 2: Research Branch. Agriculture Canada, Monograph 32, pp. 881-885.
105. von Tschirnhaus M., and Hoffeins C. Fossil flies in Baltic amber-insights in the diversity of Tertiary Acalyptratae (Diptera, Schizophora), with new morphological characters and a key based on 1000 collected inclusions. Denisia 26 (2009) 171-212
106. Wheeler T.A. Systematics of Mallochianamyia Santos-Neto (= Gayomyia Malloch), an unplaced genus of Neotropical acalyptrate flies (Diptera: Schizophora). Insect Syst. Evol. 31 (2000) 209-224
107. Wheeler W.C., Aagesen L., Arango C.P., Faivovich J., Grant T., D'Haese C., Janies D., Smith W.L., Varón A., and Giribet G. Dynamic Homology and Phylogenetic Systematics: A Unified Approach Using POY (2006), American Museum of Natural History, New York pp 365
108. Whitfield J.B., and Kjer K.M. Ancient rapid radiations of insects: challenges for phylogenetic analysis. Annu. Rev. Entomol. 53 (2008) 449-472
109. Wiegmann B.M., Tsaur S.C., Webb D.W., Yeates D.K., and Cassel B.K. Monophyly and relationships of the Tabanomorpha (Diptera: Brachycera) based on 28S ribosomal gene sequences. Ann. Entomol. Soc. Am. 93 (2000) 1031-1038
110. Winterton S.L., Wiegmann B.M., and Schlinger E.I. Phylogeny and Bayesian divergence time estimations of small-headed flies (Diptera: Acroceridae) using multiple molecular markers. Mol. Phylogenet. Evol. 43 (2006) 808-832
111. Yeates D.K., and Wiegmann B.M. Congruence and controversy: Toward a higher-level phylogeny of Diptera. Annu. Rev. Entomol. 44 (1999) 397-428
112. Yeates D.K., Wiegmann B.M., Courtney G.W., Meier R., Lambkin C., and Pape T. Phylogeny and systematics of Diptera: two decades of progress and prospects. Zootaxa 1668 (2007) 565-590
113. Zwickl, D.J., 2006. Genetic algorithm approaches for the phylogneetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. Dissertation, the University of Texas at Austin. Available from: .