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Type: Article
Published: 2023-04-21
Page range: 537-560
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First cladistic analysis of Toxotarsinae (Diptera: Calliphoridae), with insights on the evolution of the group and on the transformation series of some historically controversial characters

Universidade Federal de São Carlos; Campus Sorocaba; CCHB; Departamento de Biologia; Laboratório de Sistemática de Diptera; Rodovia João Leme dos Santos; km 110; Bairro Itinga; Sorocaba—SP; Brazil
Universidade Federal de Pelotas (UFPel); Campus Capão do Leão; IB; Departamento de Ecologia; Zoologia e Genética; Laboratório de Evolução e Genética de Insetos; Travessa André Dreyfus; s/n; prédio 23; Cidade Universitária; Capão do Leão—RS; Brazil
Universidade Federal de São Carlos; Campus Sorocaba; CCHB; Departamento de Biologia; Laboratório de Sistemática de Diptera; Rodovia João Leme dos Santos; km 110; Bairro Itinga; Sorocaba—SP; Brazil
Diptera Neotropical Region Neta Sarconesia Toxotarsus taxonomy

Abstract

Toxotarsinae is a subfamily of Calliphoridae (Diptera: Calyptratae: Oestroidea) with 11 species currently distributed in three genera: Neta Shannon, 1926, Sarconesia Bigot, 1857, and Toxotarsus Macquart, 1851. All known species are endemic to South America, mostly restricted to areas with cooler temperatures—like high elevation areas of the Andean Cordillera—and lowlands and coastal regions in the subtropical and temperate parts of the continent. The classification of the subfamily has been somewhat controversial, with eleven, mostly monotypic nominal genera established to accommodate its species, mostly due to distinct interpretations of certain controversial characters by different authors. To provide a sound basis for a generic classification, we performed the first study aimed to understand the transformation series of those controversial characters and character states under a phylogenetic framework. Our maximum parsimony analyses, under both equal and implied weighting of characters and including all currently known species in the subfamily, retrieved two main clades of Toxotarsinae: the roraima group, composed of Sarconesia roraima (Townsend, 1935), Neta chilensis (Walker, 1836), and S. magellanica (Le Guillou, 1842), and the splendida group, composed of the remaining species of the subfamily. The position of S. magellanica varied, but the species was mostly recovered within the roraima group except in the analyses under implied weighting with k values between 1 and 6. Our results, which include an explicit interpretation of the transformation series associated with some characters and character states commonly used in Toxotarsinae systematics, suggest that both Sarconesia and Neta, along with other genera proposed in the past for species of the subfamily such as Sarconesiomima Lopes & Albuquerque, 1955, Chlorobrachycoma Townsend, 1918, Sarconesiopsis Townsend, 1918, and Roraimomusca Townsend, 1935, previously suggested to be synonyms of Sarconesia, do not properly reflect the evolution of lineages in the subfamily and should not be considered as valid nominal genera. Toxotarsus was the only genus consistently recovered as monophyletic and, based on the phylogenetic relationships recovered herein, we suggest including all known Toxotarsinae within this genus, with the roraima and splendida clades treated as species-groups. Finally, we discuss the characters commonly used to delimit the taxa subordinated to Toxotarsinae, such as the length of the first flagellomere, the position of the rays of the arista relative to the pedicel, the extension of the plumosity of the arista, and the number of postsutural acrostichal setae.

 

References

  1. Aldrich, J.M. & Shannon, R.C. (1923) Boreellus aristatus. In: Shannon, R.C. (Ed.), Genera of Nearctic Calliphoridae, blow-flies, with revision of the Calliphorini (Diptera). Insecutor Inscitiae Menstruus, 11, 101–118.
  2. Amorim, D.S. (1982) Classificação por sequenciação: uma proposta para a denominação dos ramos retardados. Revista Brasileira de Zoologia, 1 (1), 1–9. https://doi.org/10.1590/S0101-81751982000100001 DOI: https://doi.org/10.1590/S0101-81751982000100001
  3. Bigot, J.M.F. (1857) Diptères nouveaux provenant du Chili. Annales de la Société Entomologique de France, 5, 277–308.
  4. Bigot, J.M.F. (1877) Diptères nouveaux ou peu connus. 7e partie. IX. Genre Somomyia (Rondani) Lucilia (Rob-Desv.) Calliphora, Phormia, Chrysomyia (id.). Annales de la Société Entomologique de France, Series 5, 7, 35–48.
  5. Boyes, J.W. & Shewell, G.E. (1975) Cytotaxonomy of Calliphoridae (Diptera). Genetica, 45 (4), 435–488. https://doi.org/10.1007/BF01772870 DOI: https://doi.org/10.1007/BF01772870
  6. Bremer, K. (1994) Branch support and tree stability. Cladistics, 10, 295–304. https://doi.org/10.1111/j.1096-0031.1994.tb00179.x DOI: https://doi.org/10.1111/j.1096-0031.1994.tb00179.x
  7. Buenaventura, E., Lloyd, M.W., Perilla López, J.M., González, V.L., Thomas-Cabianca, A. & Dikow, T. (2021) Protein-encoding ultraconserved elements provide a new phylogenomic perspective of Oestroidea flies (Diptera: Calyptratae). Systematic Entomology, 46, 5–27. https://doi.org/10.1111/syen.12443 DOI: https://doi.org/10.1111/syen.12443
  8. Cerretti, P., Stireman, J.O. III, Pape, T., O’Hara, J.E., Marinho, M.A.T., Rognes, K. & Grimaldi, D.A. (2017) First fossil of an oestroid fly (Diptera: Calyptratae: Oestroidea) and the dating of oestroid divergences. PLoS One, 12, 1–24. https://doi.org/10.1371/journal.pone.0182101 DOI: https://doi.org/10.1371/journal.pone.0182101
  9. Cumming, J.M. & Wood, D.M. (2009) Chapter 2. Adult morphology and terminology. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera. Vol. 1. National Research Council of Canada Monograph Publishing Program, Ottawa, pp. 9–50.
  10. Cumming, J.M. & Wood, D.M. (2017) Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.), Manual of Afrotropical Diptera. Vol. 1. Introductory chapters and keys to Diptera families. Suricata 4. South African National Biodiversity Institute Graphics & Editing, Pretoria, pp. 89–133.
  11. Dear, J.P. (1979) A revision of the Toxotarsinae (Diptera: Calliphoridae). Papéis Avulsos de Zoologia, 32 (13), 145–182.
  12. Dear, J.P. (1985) A revision of the New World Chrysomyini (Diptera: Calliphoridae). Revista Brasileira de Zoologia, 3 (3), 109–169. https://doi.org/10.1590/S0101-81751985000300001 DOI: https://doi.org/10.1590/S0101-81751985000300001
  13. Enderlein, G. (1938) Die Dipterenfauna der Juan Fernández-Inseln und der Oster-Inseln. In: Skottsberg, C. (Ed.), The Natural History of Juan Fernandez and Easter Island. Zoology. Part V. Vol. 3. Almqvist & Wiksells Boktryckeri, Uppsala, pp. 643–688.
  14. Fabricius, J.C. (1794) Entomologia systematica emendata et aucta. Tome IV. C.G. Proft, Hafniae [Copenhagen], pp. 472 + [5] pp.
  15. Farris, J.S. (1989) The retention index and the rescaled consistency index. Cladistics, 5, 417–419. https://doi.org/10.1111/j.1096-0031.1989.tb00573.x DOI: https://doi.org/10.1111/j.1096-0031.1989.tb00573.x
  16. Goloboff, P.A. (1993) Estimating character weights during tree search. Cladistics, 9, 83–91. https://doi.org/10.1111/j.1096-0031.1993.tb00209.x DOI: https://doi.org/10.1111/j.1096-0031.1993.tb00209.x
  17. Goloboff, P.A., Carpenter, J.M., Arias, J.S. & Esquivel, D.R.M. (2008) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 758–773. https://doi.org/10.1111/j.1096-0031.2008.00209.x DOI: https://doi.org/10.1111/j.1096-0031.2008.00209.x
  18. Goloboff, P.A. & Catalano, S.A. (2016) TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics, 32 (3), 221–238. https://doi.org/10.1111/cla.12160 DOI: https://doi.org/10.1111/cla.12160
  19. Goloboff, P.A. & Farris, J.S. (2001) Methods for quick consensus estimation. Cladistics, 17, S26–S34. https://doi.org/10.1111/j.1096-0031.2001.tb00102.x DOI: https://doi.org/10.1111/j.1096-0031.2001.tb00102.x
  20. Hough, G.N. (1899) Synopsis of the Calliphorinae of the United States. Zoological Bulletin, 2, 283–290. https://doi.org/10.2307/1535440 DOI: https://doi.org/10.2307/1535440
  21. James, M.T. (1970) Family Calliphoridae. In: Papavero, N. (Ed.), A catalogue of the Diptera from America South of United States. Vol. 102. Museu de Zoologia, Universidade de São Paulo, São Paulo, pp. 1–28.
  22. Kluge, A.G. & Farris, J.S. (1969) Quantitative phyletics and the evolution of anurans. Systematic Zoology, 18, 1–32. https://doi.org/10.2307/2412407 DOI: https://doi.org/10.1093/sysbio/18.1.1
  23. Kosmann, C., Mello, R.P., Harterreiten-Souza, E.S. & Pujol-Luz, J.R. (2013) A list of current valid blow fly names (Diptera: Calliphoridae) in the Americas South of Mexico with key to the Brazilian species. EntomoBrasilis, 6 (1), 74–85. https://doi.org/10.12741/ebrasilis.v6i1.266 DOI: https://doi.org/10.12741/ebrasilis.v6i1.266
  24. Kutty, S.N., Pape, T., Wiegmann, B.M. & Meier, R. (2010) Molecular phylogeny of the Calyptratae (Diptera: Cyclorrhapha) with an emphasis on the superfamily Oestroidea and the position of Mystacinobiidae and McAlpine’s fly. Systematic Entomology, 35, 614–635. https://doi.org/10.1111/j.1365-3113.2010.00536.x DOI: https://doi.org/10.1111/j.1365-3113.2010.00536.x
  25. Le Guillou, E.J.F. (1842) Description de sept diptères nouveaux, recueillis pendant le voyage antour du monde de l’Astrolabe et la Zélée. Revue Zoologique, 5, 314–316.
  26. Lopes, H.S. (1961) Sobre um novo gênero de Calliphoridae de Ilha de Santo Ambrósio, Chile (Diptera). Revista Brasileira de Biologia, 21, 455–459.
  27. Lopes, H.S. (1962) Sobre dois Calliphoridae do Chile (Diptera). Revista Brasileira de Biologia, 22, 421–426.
  28. Lopes, H.S. & Albuquerque, D.O. (1955) Los insectos de las islas Juan Fernandez. 22. Calliphoridae et Sarcophagidae (Diptera). Revista Chilena de Entomologia, 4, 95–119.
  29. Lopes, H.S. & Albuquerque, D.O. (1982) Notes on Neotropical Calliphoridae (Diptera). Revista Brasileira de Biologia, 42, 63–69.
  30. Macquart, P.J.M. (1834) Insectes diptères du nord de la France. Athericeres: Creophiles, Oestrides, Myopaires, Conopsaires, Scenopiniens, Cephalopsides, L. Danel, Lille, 232 pp., 6 pls.
  31. Macquart, J. (1844) Diptères exotiques nouveaux ou peu connus. Tome deuxieme.—3.e partie. Mémoires de la Société Royale des Sciences, de l’Agriculture et des Arts, de Lille, [1843], 162–460.
  32. Macquart, J. (1851) Diptères exotiques nouveaux ou peu connus. Suite du 4.e supplément publié dans les Mémoires de 1849. Mémoires de la Société Royale des Sciences, de l’Agriculture et des Arts, de Lille, [1850], 134–294.
  33. Macquart, P.J.M. (1855) s.n. In: Diptères exotiques nouveaux ou peu connus. 5e Supplément. Roret, Paris, pp. 5–136.
  34. Mariluis, J.C. (1978a) Contribución al conocimiento del genero Chlorobrachycoma Townsend, 1918. (Diptera, Calliphoridae, Toxotarsinae). Revista de la Sociedad Entomológica Argentina, 37, 107–111.
  35. Mariluis, J.C. (1978b) Contribución al conocimiento de las Calliphoridae de la Argentina (Insecta, Diptera). Ph.D. dissertation, Tesis de doctorado en Ciencias Naturales, Universidad Nacional de La Plata, La Plata. [unknown pagination]
  36. Mariluis, J.C. (1981a) Nuevos Calliphoridae para la Argentina, Bolivia y Equador (Diptera). Revista de la Sociedad Entomologica Argentina, 40 (1–4), 103–105.
  37. Mariluis, J.C. (1981b) Clave para la identificación de los Calliphoridae de la Republica Argentina (Diptera). Revista de la Sociedad Entomológica Argentina, 40 (1–4), 27–30.
  38. Mariluis, J.C. (1982) Contribución al conocimiento de las Calliphoridae de la Argentina (Insecta, Diptera). Opera Lilloana, 33, 1–59.
  39. Mariluis, J.C. & Peris, S.V. (1984) Datos para una sinopsis de los Calliphoridae neotropicales. Eos, 60, 67–86.
  40. Marinho, M.A.T., Junqueira, A.C.M., Paulo, D.F., Esposito, M.C., Villet, M.H. & Azeredo-Espin, A.M.L. (2012) Molecular phylogenetics of Oestroidea (Diptera: Calyptratae) with emphasis on Calliphoridae: insights into the inter-familial relationships and additional evidence for paraphyly among blowflies. Molecular Phylogenetics and Evolution, 65, 840–854. https://doi.org/10.1016/j.ympev.2012.08.007 DOI: https://doi.org/10.1016/j.ympev.2012.08.007
  41. McAlpine, J.F. (1989) Phylogeny and classification of the Muscomorpha. In: McAlpine, J.F. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Vol. 3. Research Branch, Agriculture Canada. Monograph 32. Biosystematics Research Centre, Ottawa, pp. 1397–1518.
  42. Mello, R.P. (1978) Contribuição ao conhecimento do gênero Roraimomusca Townsend, 1935 (Diptera, Calliphoridae). Revista Brasileira de Biologia, 38 (4), 895–898.
  43. Mello, R.P. (2003) Chave para a identificação das formas adultas das espécies da família Calliphoridae (Diptera, Brachycera, Cyclorrhapha) encontradas no Brasil. Entomología y Vectores, 10, 255–268.
  44. Pape, T. (1992) Phylogeny of the Tachinidae family-group (Diptera: Calyptratae). Tijdschrift voor Entomologie, 135, 43–86.
  45. Peris, S.V. (1992) A preliminary key to the World genera of the subfamilies Toxotarsinae, Chrysomyinae and Rhiniinae (Diptera, Calliphoridae). Boletín de la Real Sociedad Española de Historia Natural. Sección Biológica, 88 (1–4), 79–88.
  46. Robineau-Desvoidy, J.B. (1830) Essai sur les Myodaires. Mémoires présentés par divers savans à l’Académie des Sciences de l’Institut de France et imprimés par son ordre. Sciences Mathématiques et Physiques, Series 2, 2, 1–813. https://doi.org/10.5962/bhl.title.8552 DOI: https://doi.org/10.5962/bhl.title.8552
  47. Rognes, K. (1997) The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group. Cladistics, 13, 27–68. https://doi.org/10.1111/j.1096-0031.1997.tb00240.x DOI: https://doi.org/10.1111/j.1096-0031.1997.tb00240.x
  48. Rondani, C. (1851) Dipterorum species aliquae in America aequatoriali collectae a Cajetano Osculati observatae et distinctae novis breviter descriptis. Nuovi Annali delle Scienze Naturali e Rendiconto delle Sessioni della Società Agraria e dell’Accademia delle Scienze dell’Istituto di Bologna, 3 (2), 357–372.[1850]
  49. Schiner, I.R. (1868) Diptera. In: Reise der österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Zweiter Band. 1. Abtheilung. s.n., Wien, vi + 388 pp., 4 pls.
  50. Schnack, J.A. & Mariluis, J.C. (2004) Calliphoridae (Diptera) from Southeastern Argentinean Patagonia: species composition and abundance. Revista de la Sociedad Entomológica Argentina, 63 (1–2), 85–91.
  51. Shannon, R.C. (1926) Synopsis of the American Calliphoridae (Diptera). Proceedings of the Entomological Society of Washington, 28, 115–139.
  52. Shewell, G.E. (1987) Calliphoridae. In: McAlpine, J.F., Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Eds.), Manual of Nearctic Diptera. Vol. 2. Research Branch, Agriculture Canada. Monograph 28. Biosystematics Research Centre, Ottawa, pp. 1133–1145.
  53. Singh, B. & Wells, J.D. (2013) Molecular systematics of the Calliphoridae (Diptera: Oestroidea): evidence from one mitochondrial and three nuclear genes. Journal of Medical Entomology, 50, 15–23. https://doi.org/10.1603/ME11288 DOI: https://doi.org/10.1603/ME11288
  54. Townsend, C.H.T. (1892) Catalogue of the described South American species of calyptrate Muscidae. Annals of the New York Academy of Sciences, 7, 1–44. https://doi.org/10.1111/j.1749-6632.1893.tb55409.x DOI: https://doi.org/10.1111/j.1749-6632.1893.tb55409.x
  55. Townsend, C.H.T. (1908) The taxonomy of the muscoidean flies, including descriptions of new genera and species. Smithsonian Miscellaneous Collections, 51, 1–138.
  56. Townsend, C.H.T. (1918) New muscoid genera, species and synonymy. Insecutor Inscitiae Menstruus, 6, 151–156.
  57. Townsend, C.H.T. (1935) New muscoid genera, mainly from the Neotropical Region. Revista de Entomologia, 5, 68–74.
  58. Walker, F. (1836) Descriptions of the insects collected by Captain P. P. King, R. N., F. R. S., in the survey of the Straits of Magellan. Transactions of the Linnean Society of London, 17, 331–359. https://doi.org/10.1111/j.1095-8339.1834.tb00026.x DOI: https://doi.org/10.1111/j.1095-8339.1834.tb00026.x
  59. Walker, F. (1849) List of the specimens of dipterous insects in the collection of the British Museum. Vol. IV. British Museum, London, pp. 689–1172.
  60. Walker, F. (1853) Diptera. Part IV. In: Saunders, W.W. (Ed.), Insecta Saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders. Vol. 1. Van Voorst, London, pp. 253–474, pls. 7–8.
  61. Whitworth, T.L. (2010) Keys to the genera and species of blow flies (Diptera: Calliphoridae) of the West Indies and description of a new species of Lucilia Robineau-Desvoidy. Zootaxa, 2663 (1), 1–35. https://doi.org/10.11646/zootaxa.2663.1.1 DOI: https://doi.org/10.11646/zootaxa.2663.1.1
  62. Whitworth, T.L. (2012) Identification of Neotropical blow flies of the genus Calliphora Robineau-Desvoidy (Diptera: Calliphoridae) with the description of a new species. Zootaxa, 3209 (1), 1–27. https://doi.org/10.11646/zootaxa.3209.1.1 DOI: https://doi.org/10.11646/zootaxa.3209.1.1
  63. Wiedemann, C.R.W. (1830) Aussereuropäische zweiflügelige Insekten. Als Fortsetzung des Meigenschen Werkes. Zweiter Theil. Schulz, Hamm, xii + 684 pp., 5 pls.
  64. Wiedemann, C.R.W. (1831) Monographia generis Midarun. Abh. K. Leopoldinisch. Carolinisch Deutsche Akademie der Naturforscher, 15, 19–56.
  65. Yan, L., Pape, T., Meusemann, K, Kutty, S.N., Meier, R., Bayless, K.M. & Zhang, D. (2021) Monophyletic blowflies revealed by phylogenomics. BMC Biology, 19, 230. https://doi.org/10.1186/s12915-021-01156-4 DOI: https://doi.org/10.1186/s12915-021-01156-4