Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5741 No. 3: 7 Jan. 2026
Type: Article
Published: 2026-01-07
DOI: 10.11646/zootaxa.5741.3.2
Page range: 441-481
Abstract views: 150
PDF downloaded: 6

Systematic revision of the tribe Ennadini (Lepidoptera: Geometridae) in southern South America, with descriptions of new genera and species

Departamento de Zoología; Facultad de Ciencias Naturales y Oceanográficas; Universidad de Concepción; Casilla 160-C; Concepción; Chile; Programa de Doctorado en Sistemática y Biodiversidad; Facultad de Ciencias Naturales y Oceanográficas; Universidad de Concepción; Chile; Fundación Mongen Ecología y Biodiversidad; Osorno; Chile; “Lepidópteros de Chile” Citizen Science Program; Concepción; Chile
Departamento de Ecología; Facultad de Ciencias; Universidad Católica de la Santísima Concepción; Alonso de Ribera 2850; Concepción; Chile
“Lepidópteros de Chile” Citizen Science Program; Concepción; Chile; Instituto de Humanidades Alfredo Silva Santiago; Colo Colo 24; Concepción; Chile
Departamento de Zoología; Facultad de Ciencias Naturales y Oceanográficas; Universidad de Concepción; Casilla 160-C; Concepción; Chile
Lepidoptera Andean region calibrated phylogeny integrative taxonomy Larentiinae Neotropical region new classification

Abstract

An integrative taxonomic revision of the tribe Ennadini is presented for southern South America, based on adult morphological characters and a time-calibrated molecular phylogeny inferred from 11 genetic regions. The tribe is recovered as monophyletic, with an estimated crown age of 29.8 million years (Oligocene), according to a relaxed lognormal molecular clock model. The genera Ennada Blanchard and Spargania Guenée are redefined, and Praeapodroma Heimlich is transferred to Ennadini. Three new genera are described: Antuluciella gen. nov., Incantia gen. nov., and Kuyenusia gen. nov. Five new combinations are proposed: Antuluciella vittata (Philippi) comb. nov., Incantia edmondsii (Butler) comb. nov., Kuyenusia ceres (Butler) comb. nov., Praeapodroma macerata (Felder & Rogenhofer) comb. nov., and Praeapodroma scarata (Felder & Rogenhofer) comb. nov. Two new synonymies are established: Fidonia incandescens Thierry-Mieg = Incantia edmondsii (Butler), syn. nov.; and Heterusia mesenata Felder & Rogenhofer = Antuluciella vittata (Philippi), syn. nov. The reinstatement of Spargania pastoralis (Butler) comb. rev. is also proposed, and Kuyenusia tenten Ramos-González & Parra sp. nov. is described. Diagnostic characters, redescriptions, and distributional data are provided for all treated species. Lectotypes are designated where necessary to stabilize nomenclature. This revision provides a robust and updated systematic and evolutionary framework for Ennadini moths in the temperate regions of South America.

 

References

  1. Angulo, A.O. & Casanueva, M.E. (1981) Catálogo de los lepidópteros geométridos de Chile (Lepidoptera: Geometridae). Boletín de la Sociedad de Biología de Concepción, Chile, 51, 7–34.
  2. Antonelli, A., Nylander, J.A.A., Persson, C. & Sanmartín, I. (2009) Tracing the impact of the Andean uplift on Neotropical plant evolution. Proceedings of the National Academy of Sciences, 106 (24), 9749–9754. https://doi.org/10.1073/pnas.0811421106
  3. Blanchard, E. (1852) Fauna Chilena, Insectos, Orden VI, Lepidópteros. In: Gay, C. (Ed.), Historia física y política de Chile. Maulde et Renou, Paris, pp. 1–112.
  4. Bouckaert, R. & Drummond, A.J. (2017) bModelTest: Bayesian phylogenetic site model averaging and model comparison. BMC Evolutionary Biology, 17, 42. https://doi.org/10.1186/s12862-017-0890-6
  5. Bouckaert, R., Vaughan, T.G., Barido-Sottani, J., Duchêne, S., Fourment, M., Gavryushkina, A., Heled, J., Jones, G., Kühnert, D., De Maio, N., Matschiner, M., Mendes, F.K., Müller, N.F., Ogilvie, H.A., du Plessis, L., Popinga, A., Rambaut, A., Rasmussen, D., Siveroni, I., Suchard, M.A., Wu, C.H., Xie, D., Zhang, C., Stadler, T. & Drummond, A.J. (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology, 15 (4), e1006650. https://doi.org/10.1371/journal.pcbi.1006650
  6. Brehm, G., Murillo-Ramos, L., Sihvonen, P., Hausmann, A., Schmidt, B., Õunap, E., Moser, A., Mörtter, R., Bolt, D., Bodner, F., Lindt, A., Parra, L.E. & Wahlberg, N. (2019) New World geometrid moths (Lepidoptera: Geometridae): Molecular phylogeny, biogeography, taxonomic updates and description of 11 new tribes. Arthropod Systematics & Phylogeny, 77 (3), 457–486. https://doi.org/10.26049/ASP77-3-2019-5
  7. Bartlett-Calvert, W. (1886) Catálogo de los lepidópteros Rhopalóceros i Heteróceros de Chile. Anales de la Universidad de Chile, 69, 314–352.
  8. Brehm, G. (2015) Three new species of Hagnagora Druce, 1885 (Lepidoptera, Geometridae, Larentiinae) from Ecuador and Costa Rica and a concise revision of the genus. Zookeys, 537, 131–156. https://doi.org/10.3897/zookeys.537.6090
  9. Butler, A.G. (1882) Heterocerous Lepidoptera collected in Chili by Thomas Edmonds, Esq. Part III—Geometrites. Transactions of the Entomological Society of London, 30, 339–427. https://doi.org/10.1111/j.1365-2311.1882.tb01580.x
  10. Butler, A.G. (1893) On a small collection of Lepidoptera from Chili. The Annals and Magazine of Natural History, including Zoology Botany, and Geology, 12 (6), 457–466. https://doi.org/10.1080/00222939308677653
  11. Chalup, A. (2020) The biodiversity of the family Geometridae for Argentina. In: Juñent, S.A., Claps, L.E., Morrone, J.J. (Eds.), Biodiversidad de artrópodos argentinos. Vol. 4. INSUE-UNT Ediciones, San Miguel de Tucumán, pp. 47–62.
  12. Chazot, N., Willmott, K.R., Condamine, F.L., De-Silva, D.L., Freitas, A.V.L., Lamas, G., Morlon, H., Giraldo, C.E., Jiggins, C.D., Joron, M., Mallet, J., Uribe, S. & Elias, M. (2016) Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina. Molecular Ecology, 25, 5765–5784. https://doi.org/10.1111/mec.13773
  13. Cockerell, T.D.A. (1922) A fossil moth from Florissant, Colorado. American Museum Novitates, 34, 1–2.
  14. Donoghue, P.C.J. & Benton, M.J. (2007) Rocks and clocks: calibrating the Tree of Life using fossils and molecules. Trends in Ecology and Evolution, 22 (8), 424–431. https://doi.org/10.1016/j.tree.2007.05.005
  15. Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797. https://doi.org/10.1093/nar/gkh340
  16. Espeland, M., Breinholt, J., Willmott, K.R., Warren, A.D., Vila, R., Toussaint, E., Maunsell, S.C., Aduse-Poku, K., Talavera, G., Eastwood, R., Jarzyna, M.A., Guralnick, R., Lohman, D.J., Pierce, N.E. & Kawahara, A.Y. (2018) A Comprehensive and Dated Phylogenomic Analysis of Butterflies. Current Biology, 28 (5), 770–778. https://doi.org/10.1016/j.cub.2018.01.061
  17. Felder, C. & Rogenhofer, A.F. (1875) Lepidoptera: Atlas. In: Felder, C., Felder, R. & Rogenhofer, A.F. (Eds.), 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. Band 2 (Abtheilung 2). Aus der kaiserlich königlichen Hofund Staatsdruckerel in Kommission bei Karl Gerold’s Sohn, Wien, pp. 1–140.
  18. Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
  19. Forest, F. (2009) Calibrating the Tree of Life: fossils, molecules and evolutionary timescales. Annals of Botany, 104 (5), 789–794. https://doi.org/10.1093/aob/mcp192
  20. Garzione, C., Hoke, G., Libarkin, J., Withers, S. MacFadden, B., Eiler, J., Ghosh, P. & Mulch, A. (2008) Rise of the Andes. Science, 320 (5881), 1304–1307. https://doi.org/10.1126/science.1148615
  21. Gaston, K.J., Scoble, M.J. & Crook, A. (1995) Patterns in species description: a case study using the Geometridae (Lepidoptera). Biological Journal of the Linnean Society, 55, 225–237. https://doi.org/10.1111/j.1095-8312.1995.tb01061.x
  22. Ghanavi, H.R., Chazot, N., Sanmartín, I., Murillo-Ramos, L., Duchêne, S., Sihvonen, P., Brehm, G. & Wahlberg, N. (2025) Region-specific diversification dynamics and biogeographic history of one of the most diverse families of insects. Systematic Entomology, 50 (1), 206–220. https://doi.org/10.1111/syen.12651
  23. Guenée, M.A. (1858) Histoire naturelle des insectes; spécies généraldes lépidoptères. Vol. 10. Uranides et Phalénites. Librairie Encyclopédique de Roret, Paris, 584 pp.
  24. Hall, T.D. (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acid Symposium Series, 41, 95–98.
  25. Hausmann, A. & Parra, L.E. (2009) An unexpected hotspot of moth biodiversity in Chilean northern Patagonia (Lepidoptera, Geometridae). Zootaxa, 1989 (1), 23–38. https://doi.org/10.11646/zootaxa.1989.1
  26. Heimlich, W. (1958) Eine neue Geometride aus Chile (Geom. Larentiinae). Entomologische Zeitschrift, 68, 41–43.
  27. Ho, S.Y.W. & Phillips, M.J. (2009) Accounting for calibration uncertainty in phylogenetic estimation of evolutionary divergence times. Systematic Biology, 58 (3), 367–380. https://doi.org/10.1093/sysbio/syp035
  28. Holloway, J.D. (1997) The Moths of Borneo, pt. 10, Geometridae: Sterrhinae, Larentiinae. Malayan Nature Journal, 51, 1–242.
  29. King, G.E. & Parra, L.E. (2011) Contribution to an understanding of the biology and the morphology of the early stages of a Neotropical larentiine: Hagnagora vittata Philippi, 1859 in Chile (Insecta: Lepidoptera: Geometridae). Acta Zoologica Cracoviensia, 54B (1–2), 5–15. https://doi.org/10.3409/azc.54b_1-2.05-15
  30. Klots, A.B. (1970) Lepidoptera. In: Tuxen, S.L. (Ed.), Taxonomist’s glossary of genitalia in insects. Second Enlarged Edition. Munksgaard, Copenhagen, pp. 115–130.
  31. Lin, C.-P. & Danforth, B.N. (2004) How do insect nuclear and mitochondrial gene substitution patterns differ? Molecular Phylogenetics and Evolution, 30 (3), 686–702. https://doi.org/10.1016/S1055-7903(03)00241-0
  32. Maturana-Russel, P., Brewer, B.J., Klaere, S. & Bouckaert, R. (2019) Model Selection and Parameter Inference in Phylogenetics Using Nested Sampling. Systematic Biology, 68 (2), 219–233. https://doi.org/10.1093/sysbio/syy050
  33. Morrone, J.J. (2015) Biogeographical regionalisation of the Andean region. Zootaxa, 3936 (2), 207–236. https://doi.org/10.11646/zootaxa.3936.2.3
  34. Müller, N.F. & Bouckaert, R. (2020) Adaptive Metropolis-coupled MCMC for BEAST 2. PeerJ, 8, e9473. https://doi.org/10.7717/peerj.9473
  35. Mundaca, E.A. & Parra, L.E. (2004) Geométridos (Lepidoptera) presentes en formaciones vegetales de la Reserva Nacional Isla Mocha (VIII Región Chile). Gayana, 68 (1), 1–8. https://doi.org/10.4067/S0717-65382004000100001
  36. Murillo-Ramos, L., Chazot, N., Sihvonen, P., Ounap, E., Jiang, N., Han, H., Clarke, J.T., Davis, R.B., Tammaru, T. & Wahlberg, N. (2021) Molecular phylogeny, classification, biogeography and diversification patterns of a diverse group of moths (Geometridae: Boarmiini). Molecular Phylogenetics and Evolution, 162, 107198. https://doi.org/10.1016/j.ympev.2021.107198
  37. Murillo-Ramos, L., Twort, V., Wahlberg, N. & Sivhonen, P. (2023) A phylogenomic perspective on the relationships of subfamilies in the family Geometridae (Lepidoptera). Systematic Entomology, 48 (4), 618–632. https://doi.org/10.1111/syen.12594
  38. Ortiz-Jaureguizar, E. & Cladera, G.A. (2006) Paleoenvironmental evolution of southern South America during the Cenozoic. Journal of Arid Environments, 66 (3), 498–532. https://doi.org/10.1016/j.jaridenv.2006.01.007
  39. Õunap, E., Viidalepp, J. & Truuverk, A. (2016) Phylogeny of the subfamily Larentiinae (Lepidoptera: Geometridae): integrating molecular data and traditional classifications. Systematic Entomology, 41 (4), 824–843. https://doi.org/10.1111/syen.12195
  40. Parham, J.F., Donoghue, P.C.J., Bell, C.J., Calway, T.D., Head, J.J., Holroyd, P.A., Inoue, J.G., Irmis, R.B., Joyce, W.G., Ksepka, D.T., Patané, J.S., Smith, N.D., Tarver, J.E., van Tuinen, M., Yang, Z., Angielczyk, K.D., Greenwood, J.M., Hipsley, C.A., Jacobs, L., Makovicky, P.J., Müller, J., Smith, K.T., Theodor, J.M., Warnock, R.C.M. & Benton, M.J. (2012) Best Practices for Justifying Fossil Calibrations, Systematic Biology, 61 (2), 346–359. https://doi.org/10.1093/sysbio/syr107
  41. Parra, L.E. (1991) Revisión y filogenia del género Pachrophylla Blanchard, 1852 (sensu auctorum) (Geometridae: Larentiinae: Trichopterygini). Gayana Zoología (Chile), 55 (2), 145–199.
  42. Parra, L.E. & Alvear, C.A. (2009) Revision of the genus Ennada Blanchard (Lepidoptera: Geometridae). Zootaxa, 2062, 46–56. https://doi.org/10.11646/zootaxa.2062.1.4
  43. Parra, L.E., Vargas, H.A., Sanzana, M.J. & Hernández, C.E. (2017) Phylogenetic study of the genera of Trichopterygini from austral South America (Lepidoptera: Geometridae): a new classification. Gayana, 81 (2), 64–99. https://doi.org/10.4067/S0717-65382017000200064
  44. Philippi, R.A. (1859) Descripción de algunas nuevas especies de mariposas chilenas, principalmente de la provincia de Valdivia. Anales de la Universidad de Chile, 16 (11), 1088–1114.
  45. Pitkin, L.M. (2002) Neotropical ennomine moths: A review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society, 135, 121–401. https://doi.org/10.1046/j.1096-3642.2002.00012.x
  46. Quintero, I. & Jetz, W. (2018) Global elevational diversity and diversification of birds. Nature, 555, 246–250. https://doi.org/10.1038/nature25794
  47. Rajaei, H., Hausmann, A., Scoble, M.J., Wanke, D., Plotkin, D., Brehm, G., Murillo-Ramos, L. & Sihvonen, P. (2022) An online taxonomic facility of Geometridae (Lepidoptera), with an overview of global species richness and systematics. Integrative Systematics, 5 (2), 145–192. https://doi.org/10.18476/2022.577933
  48. Rambaut, A. (2018) Figtree ver 1.4.4. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh.
  49. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
  50. Ramos-González, M.I. (2015) Revisión y Filogenia basada en caracteres morfológicos y moleculares de las especies del género Nebula Bruand, 1846 (Lepidoptera: Geometridae) presentes en Chile. Undergraduate Thesis, Universidad de Concepción, Concepción, 132 pp.
  51. Ramos-González, M.I., Zamora-Manzur, C., Saladrigas Menés, D. & Parra, L.E. (2019) The Trichopterygini (Lepidoptera, Geometridae) of Austral South America: description of new species from Chile. ZooKeys, 832, 91–111. https://doi.org/10.3897/zookeys.832.30851
  52. Ramos-González, M.I., Zamora-Manzur, C. & Parra, L.E. (2024) An overview of geometrid moths (Lepidoptera: Geometridae) in the Andean Region s.l.: A systematic review. Gayana, 88 (1), 111–133. https://doi.org/10.4067/S0717-65382024000100111
  53. Ratnasingham, S. & Hebert, P. (2007) BOLD: The Barcode of Life Data System (www.barcodinglife.org). Molecular Ecology Notes, 7, 355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x
  54. Ratnasingham, S., Wei, C., Chan, D., Agda, J., Agda, J., Ballesteros-Mejia, L., Boutou, H.A., El Bastami, Z.M., Ma, E., Manjunath, R., Rea, D., Ho, C., Telfer, A., McKeowan, J., Rahulan, M., Steinke, C., Dosheimer, J., Milton, M. & Hebert, P.D.N. (2024) BOLD v4: A Centralized Bioinformatics Platform for DNA-Based Biodiversity Data. In: DeSalle, R. (Ed.), DNA Barcoding. Methods in Molecular Biology. Vol. 2744. Humana, New York, New York, pp. 403–441. https://doi.org/10.1007/978-1-0716-3581-0_26
  55. Regier, J.C., Zwick, A., Cummings, M.P., Kawahara, A.Y., Cho, S., Weller, S., Roe, A., Baixeras, J., Brown, J.W., Parr, C., Davis, D.R., Epstein, M., Hallwachs, W., Hausmann, A., Janzen, D.H., Kitching, I.J., Alma Solis, M., Yen, S.-H., Bazinet, A.L. & Mitter, C. (2009) Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology, 9, 280. https://doi.org/10.1186/1471-2148-9-280
  56. Rindge, F.H. (1987) The Euphitecia (Lepidoptera: Geometridae) of Chile. Bulletin of the American Museum of Natural History, 186 (3), 269–363.
  57. Rindge, F.H. (1991) The Eupithecia (Lepidoptera: Geometridae) of Chile II. American Museum Novitates, 3020, 1–14.
  58. Robinson, G.S., Ackery, P.R., Kitching, I., Beccaloni, G.W. & Hernández, L.M. (2023) HOSTS: a Database of the World’s Lepidopteran Hostplants. Natural History Museum. Available from: https://data.nhm.ac.uk/dataset/hosts (accessed 31 August 2025)
  59. Rose-Garrido, C.A., Ramos-González, M.I. & Parra, L.E. (2018) The Larentia of Chile and Argentina, a concise revision. In: Hausmann, A. (Ed.), Proceedings of the ninth Forum Herbulot 2017. The geometrids of southern South America: state of the art—conservation, phylogeny, taxonomy and biodiversity (Concepción, Chile, 9–14 January 2017). Spixiana, 41 (1), pp. 121.
  60. Sauquet, H., Ho, S.Y.W., Gandolfo, M.A., Jordan, G.J., Wilf, P., Cantrill, D.J., Bayly, M.J., Bromham, L., Brown, G.K., Carpenter, R.J., Lee, D.M., Murphy, D.J., Sniderman, J.M.K. & Udovicic, F. (2012) Testing the Impact of Calibration on Molecular Divergence Times Using a Fossil-Rich Group: The Case of Nothofagus (Fagales). Systematic Biology, 61 (2), 289–313. https://doi.org/10.1093/sysbio/syr116
  61. Scoble, M.J. (1995) The Lepidoptera: Form, Function and Diversity. The Natural History Museum & Oxford University Press, London, 416 pp.
  62. Scoble, M.J. (1999) Geometrid moths of the world: a catalogue (Lepidoptera, Geometridae). CSIRO Publishing, Natural History Museum, London, 1016 pp. https://doi.org/10.1071/9780643101050
  63. Skilling, J. (2006) Nested sampling for general Bayesian computation. Bayesian Analysis, 1 (4), 833–859. https://doi.org/10.1214/06-BA127
  64. Thierry-Mieg, P. (1892) Description de lépidoptères nouveaux. Le Naturaliste: Revue Illustrée: Des Sciences Naturelles, 14, 235–237.
  65. Van Nieukerken, E.J., Kaila, L., Kitching, I.J., Kristensen, N.P., Lees, D.C., Minet, J., Mitter, C., Mutanen, M., Regier, J.C., Simonsen, T.J., Wahlberg, N., Yen, S.-H., Zahiri, R., Adamski, D., Baixeras, J., Bartsch, D., Bengtsson, B.A., Brown, J.W., Bucheli, S.R., Davis, D.R., Prins, J.D., Prins, W.D., Epstein, M.E., Gentili-Poole, P., Gielis, C., Hättenschwiler, P., Hausmann, A., Holloway, J.D., Kallies, A., Karsholt, O., Kawahara, A.Y., Koster, S., Kozlov, M.V., Lafontaine, D., Lamas, G., Landry, J.F., Lee, S., Nuss, M., Park, K.T., Penz, C., Rota, J., Schintlmeister, A., Schmidt, B.C., Sohn, J.C., Solis, M.A., Tarmann, G.M., Warren, A.D., Weller, S., Yakovlev, R.V., Zolotuhin, V.V. & Zwick, A. (2011) Order Lepidoptera Linnaeus. In: Zhang, Z.-Q. (Ed.), Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148 (1), 212–221. https://doi.org/10.11646/zootaxa.3148.1.41
  66. Vargas, H.A. & Parra, L.E. (2004) Una nueva especie de Eupithecia Curtis (Lepidoptera: Geometridae) del extremo norte de Chile. Revista Chilena de Historia Natural, 77, 485–490. https://doi.org/10.4067/S0716-078X2004000300007
  67. Vargas, H.A. (2021) A new species of Eupithecia Curtis (Lepidoptera, Geometridae) from the Andes of northern Chile. Nota Lepidopterologica, 44, 239–247. https://doi.org/10.3897/nl.44.73247
  68. Viidalepp, J. (2011) A Morphological review of tribes in Larentiinae (Lepidoptera: Geometridae). Zootaxa, 3136 (1), 1–44. https://doi.org/10.11646/zootaxa.3136.1.1
  69. Vojnits, A.M. (1985) New South American genera and species of Eupithicini (Lepidoptera: Geometridae). Acta Zoologica Hungarica, 31, 405–418.
  70. Vojnits, A.M. (1992) New data to the Eupithecia fauna of Chile and Peru (Lepidoptera: Geometridae). Acta Zoologica Hungarica, 38 (1–2), 125–144.
  71. Vojnits, A.M. (1994) New data on the Eupithecia fauna of Chile (Lepidoptera, Geometridae). Entomofauna, 15 (32), 361–376.
  72. Wahlberg, N., Peña, C., Ahola, M., Wheat, C.W. & Rota, J. (2016) PCR primers for 30 novel gene regions in the nuclear genomes of Lepidoptera. ZooKeys, 596, 129–141. https://doi.org/10.3897/zookeys.596.8399
  73. Warren, W. (1895) New species and genera of Geometridae in the Tring Museum. Novitates Zoologicae, 2, 82–159.
  74. Xia, X. (2017) DAMBE6: New Tools for Microbial Genomics, Phylogenetics, and Molecular Evolution. Journal of Heredity, 108 (4), 431–437. https://doi.org/10.1093/jhered/esx033

How to Cite

Ramos-González, M.I., Zamora-Manzur, C., Garrido, C.R. & Parra, L.E. (2026) Systematic revision of the tribe Ennadini (Lepidoptera: Geometridae) in southern South America, with descriptions of new genera and species. Zootaxa, 5741 (3), 441–481. https://doi.org/10.11646/zootaxa.5741.3.2