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Type: Monograph
Published: 2020-08-31
Page range: 1–98
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Systematic revision and morphological phylogenetic analysis of Anchylorhynchus Schoenherr, 1836 (Coleoptera, Curculionidae: Derelomini)

Smithsonian Tropical Research Institute, Panama City, Panama Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA
Instituto de Biociências, Universidade de São Paulo, São Paulo—SP, Brazil
Coleoptera systematics morphology taxonomic revision Curculionidae Arecaceae Syagrus Butia Oenocarpus Euterpe


Anchylorhynchus Schoenherr is a genus of palm-associated weevils currently including 22 described species in the Neotropics. These weevils engage in brood pollination interactions with species in at least four genera of palms (Arecaceae), representing an emerging system for the study of mutualisms. Here we revise the taxonomy of Anchylorhynchus and propose the first phylogenetic hypothesis for the group, based on morphology. Anchylorhynchus chrysomeloides sp. nov., Anchylorhynchus goiano sp. nov., Anchylorhynchus imitator sp. nov., Anchylorhynchus latipes sp. nov., Anchylorhynchus multisquamis sp. nov. and Anchylorhynchus rectus sp. nov. are described, Anchylorhynchus gottsbergerorum Vanin is a new junior subjective synonym of Anchylorhynchus bicarinatus O’Brien, and Anchylorhynchus eriospathae Bondar and Anchylorhynchus pictipennis Hustache are new junior subjective synonyms of Anchylorhynchus tremolerasi Hustache, resulting in 25 valid species for the genus. We provide genus and species descriptions with a new dichotomous key to the species and updated information on geography and host associations based on museum records and extensive new collections. We also produce a new morphological matrix with 113 characters, 11 of them based on measurements while accounting for allometry, correlation and power to delimit groups. The software pipeline to produce these characters was encoded in a graphical user interface named DiscretzR, made available here. Analysis of this matrix under parsimony and Bayesian inference resulted in strong support for the monophyly of Anchylorhynchus, but weaker support for most clades within the genus. Reconstruction of the host plant associations indicates that the common ancestor of Anchylorhynchus visited flowers of Syagrus Mart., with later transitions to other genera restricted to one of the major clades of Anchylorhynchus. This taxonomic revision with the first phylogeny for the group provides a foundation for future evolutionary studies in the genus.



  1. Agassiz, L. (1846) Nomenclator Zoologicus. Nomina Systematica generum animalium. (Nomina Systematica generum coleopterorum). Jent et Gassmann, Soloduri, 170 pp.

    Agassiz, L. (1848) Nomenclatoris zoologici index universalis. Jent et Gassmann, Soloduri, 1097 pp.

    Baker, W.J., Savolainen, V., Asmussen-Lange, C.B., Chase, M.W., Dransfield, J., Forest, F., Harley, M.M., Uhl, N.W. & Wilkinson, M. (2009) Complete Generic-Level Phylogenetic Analyses of Palms (Arecaceae) with Comparisons of Supertree and Supermatrix Approaches. Systematic Biology, 58, 240–256.

    Barrett, C.F., Bacon, C.D., Antonelli, A., Cano, Á. & Hofmann, T. (2016) An introduction to plant phylogenomics with a focus on palms. Botanical Journal of the Linnean Society, 182, 234–255.

    Beaulieu, J.M., Oliver, J.C. & O’Meara, B. (2017) corHMM: Analysis of Binary Character Evolution. Available from: (accessed 8 July 2020)

    Blackwelder, R.E. (1947) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part. 5. United States National Museum Bulletin, 185, 765–925.

    Bockmann, F.A., Rodrigues, M.T., Kohsldorf, T., Straker, L.C., Grant, T., de Pinna, M.C.C., Mantelatto, F.L.M., Datovo, A., Pombal, J.P., McNamara, J.C., de Almeida, E.A.B., Klein, W., Hsiou, A.S., Groppo, M., e Castro, R.M.C. & de Souza Amorim, D. (2018) Brazil’s government attacks biodiversity. Science, 360, 865.1–865.

    Bondar, G.G. (1940a) Notas Entomológicas da Baía. V. Revista de Entomologia, 11, 199–214.

    Bondar, G.G. (1940b) Notas Entomológicas da Baía. VI. Revista de Entomologia, 11, 842–861.

    Bondar, G.G. (1941a) Notas Entomológicas da Baía. VII. Revista de Entomologia, 12, 268–303.

    Bondar, G.G. (1941b) Notas Entomológicas da Baía. VIII. Revista de Entomologia, 12, 427–470.

    Bondar, G.G. (1943a) Notas Entomológicas da Baía. XIII. Revista de Entomologia, 14, 337–388.

    Bondar, G.G. (1943b) Notas Entomológicas da Baía XII. Revista de Entomologia, 14, 85–134.

    Bondar, G.G. (1950) Notas Entomológicas da Baía. XXII. Revista de Entomologia, 21, 449–480.

    Bruch, C. (1915) Catálogo sistemático de los Coleópteros de la República Argentina. Pars VII. Revista del Museo de La Plata, 19, 401–441.

    Caldara, R., Franz, N.M. & Oberprieler, R.G. (2014) Curculioninae Latreille, 1802. In: Leschen, R.A.B. & Beutel, R.G. (Eds.), Handbook of Zoology. Arthropoda: Insecta: Coleoptera, Beetles. Vol. 3. Morphology and Systematics (Phytophaga). de Gruyter, Berlin, pp. 589–628.

    Cardona-Duque, J. & Franz, N.M. (2012) Description and phylogeny of a new Neotropical genus of Acalyptini (Coleoptera: Curculionidae: Curculioninae) associated with the staminodes of Cyclanthaceae. Zoological Journal of the Linnean Society, 166, 559–623.

    Castelnau, F. comte de (1850) Histoire Naturelle des insectes, coléoptères. Vol. 2. P. Dumménil Éditeur, Paris, 563 pp.

    Chang, W., Cheng, J., Allaire, J.J., Xie, Y. & McPherson, J. (2019) shiny: Web Application Framework for R. Available from: (accessed 8 July 2020)

    Church, S.H., Donoughe, S., de Medeiros, B.A.S. & Extavour, C.G. (2019a) A dataset of egg size and shape from more than 6,700 insect species. Scientific Data, 6, 104.

    Church, S.H., Donoughe, S., de Medeiros, B.A.S. & Extavour, C.G. (2019b) Insect egg size and shape evolve with ecology but not developmental rate. Nature, 571, 58–62.

    Coleman, C.O. (2003) “Digital inking”: how to make perfect line drawings on computers. Organisms Diversity & Evolution, 3, 303–304.

    Coleman, C.O. (2006) Substituting time-consuming pencil drawings in arthropod taxonomy using stacks of digital photographs. Zootaxa, 1360 (1), 61–68.

    Coleman, C.O. (2009) Drawing setae the digital way. Zoosystematics and Evolution, 85, 305–310.

    Dallwitz, M.J. (2000) CSIRO Delta for Windows. Available from: (accessed 8 July 2020)

    Davis, S.R. (2009) Morphology of Baridinae and related groups (Coleoptera, Curculionidae). ZooKeys, 10, 1–136.

    Davis, S.R. (2011) Delimiting baridine weevil evolution (Coleoptera: Curculionidae: Baridinae). Zoological Journal of the Linnean Society, 161, 88–156.

    Dejean, P.F.M.A. comte (1833) Catalogue des coléoptères de la collection de M. le comte Dejean. Chez Méquignon-Marvis Père et Fils, Paris, 443 pp.

    Faldermann, M.F. (1831) Some account of a Russian natural history expedition in Brazil, during the seven years preceding April, 1831. Magazine of natural history and journal of zoology, botany, mineralogy, geology and meteorology, 4, 394–403.

    Faurby, S., Eiserhardt, W.L., Baker, W.J. & Svenning, J.C. (2016) An all-evidence species-level supertree for the palms (Arecaceae). Molecular Phylogenetics and Evolution, 100, 57–69.

    Faust, J. (1894) Russelkafer der alten und neuen Welt. Stettiner Entomologische Zeitung, 55, 140–159.

    Ferreira, V.S., Prado, L.P. & Seripierri, D. (2016) The Entomological Collection of Ricardo von Diringshofen (1900–1986) and its incorporation to the Museu de Zoologia da Universidade de São Paulo. Revista Brasileira de Entomologia, 60, 117–122.

    Franz, N.M. (2006) Towards a phylogenetic system of derelomine flower weevils (Coleoptera: Curculionidae). Systematic Entomology, 31, 220–287.

    Franz, N.M. (2012) Phylogenetic reassessment of the Exophthalmus genus complex (Curculionidae: Entiminae: Eustylini, Geonemini). Zoological Journal of the Linnean Society, 164, 510–557.

    Franz, N.M. & Valente, R.M. (2005) Evolutionary trends in derelomine flower weevils (Coleoptera : Curculionidae): from associations to homology. Invertebrate Systematics, 19, 499–530.

    Gemminger, M. & Harold, B. (1871) Tom. VIII: Curculionidae. In: Gemminger, M. & Harold, B. (Eds.), Catalogus Coleopterorum hucusque descriptorum synonymicus et systematics. Tom VIII. Sumptu E. H. Gummi, Monachii, pp. 2181–2668.

    Goloboff, P.A., Farris, J.S., Källersjö, M., Oxelman, B., Ramírez, M.J. & Szumik, C.A. (2003) Improvements to resampling measures of group support. Cladistics, 19, 324–332.

    Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.

    Goloboff, P.A., Mattoni, C.I. & Quinteros, A.S. (2006) Continuous characters analyzed as such. Cladistics, 22, 589–601.

    Gottsberger, G. & Silberbauer-Gottsberger, I. (2006) Life in the Cerrado. Reta Verlag, Ulm, 383 pp.

    Guerrero-Olaya, N.Y. & Núñez-Avellaneda, L.A. (2017) Ecología de la polinización de Syagrus smithii (Arecaceae), una palma cantarofila de la Amazonia Colombiana. Revista Peruana de Biologia, 24, 43–54.

    Hadley, A. (2008) CombineZM. Avalable from: (accessed 31 March 2016)

    Hembry, D.H. & Althoff, D.M. (2016) Diversification and coevolution in brood pollination mutualisms: windows into the role of biotic interactions in generating biological diversity. American Journal of Botany, 103, 1783–1792.

    Herre, E.A., Jandér, K.C. & Machado, C.A. (2008) Evolutionary ecology of figs and their associates: Recent progress and outstanding puzzles. Annual Review of Ecology, Evolution, and Systematics, 39, 439–458.

    Hustache, A. (1937) Curculionides Nouveaux de L’Uruguay. Revista de la Sociedad Entomológica Argentina, 9, 7–11.

    Hustache, A. (1940) Curculionides nouveaux du Brèsil. Revista de Entomologia, 11, 690–713.

    Kawakita, A. & Kato, M. (2009) Repeated independent evolution of obligate pollination mutualism in the Phyllantheae-Epicephala association. Proceedings of the Royal Society B: Biological Sciences, 276, 417–426.

    Kuschel, G. (1952) Los Curculionidae de la Cordillera chilenoargenliina (Ia. parte) (Aporte 13 de Coleoptera Curculionidae). Revista Chilena de Entomologia, 2, 229–279.

    Kuschel, G. (1995) A phylogenetic classification of Curculionoidea to families and subfamilies. Memoirs of the Entomological Society of Washington, 14, 5–33.

    Lacordaire, T. (1863) Histoire naturelle des insectes. Vol. 6. Librairie Encyclopedique de Roret, Paris, 637 pp.

    Lev-Yadun, S., Dafni, A., Flaishman, M.A., Inbar, M., Izhaki, I., Katzir, G. & Ne’eman, G. (2004) Plant coloration undermines herbivorous insect camouflage. BioEssays, 26, 1126–1130.

    Lorenzi, H., Noblick, L.R., Kahn, F. & Ferreira, E. (2010) Flora brasileira: Arecaceae - Palmeiras. Editora Plantarum, Nova Odessa, São Paulo, 368 pp.

    Lyal, C.H.C. & King, T. (1996) Elytro-tergal stridulation in weevils (Insecta: Coleoptera: Curculionoidea). Journal of Natural History, 30, 703–773.

    Maddison, W.P. & Maddison, D.R. (2010) Mesquite: A modular system for evolutionary analysis. Available at: (accessed 8 July 2020)

    McKenna, D.D., Clarke, D., Anderson, R.S., Astrin, J., Brown, S., Chamorro, L., Davis, S.R., de Medeiros, B., del Rio, M., Haran, J., Kuschel, G., Franz, N.M., Jordal, B., Lanteri, A., Leschen, R., Letsch, H., Lyal, C., Marvaldi, A., Mermudes, J., Oberprieler, R., Schütte, A., Sequeira, A., Shin, S., Van Dam, M. & Zhang, G. (2018) Morphological and Molecular Perspectives on the Phylogeny, Evolution, and Classification of Weevils (Coleoptera: Curculionoidea): Proceedings from the 2016 International Weevil Meeting. Diversity, 10, 64.

    McKenna, D.D., Sequeira, A.S., Marvaldi, A.E. & Farrell, B.D. (2009) Temporal lags and overlap in the diversification of weevils and flowering plants. Proceedings of the National Academy of Sciences, 106, 7083–7088.

    McKenna, D.D., Shin, S., Ahrens, D., Balke, M., Beza-Beza, C., Clarke, D.J., Donath, A., Escalona, H.E., Friedrich, F., Letsch, H., Liu, S., Maddison, D., Mayer, C., Misof, B., Murin, P.J., Niehuis, O., Peters, R.S., Podsiadlowski, L., Pohl, H., Scully, E.D., Yan, E.V., Zhou, X., Ślipiński, A. & Beutel, R.G. (2019) The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences, 16, 24729–24737.

    de Medeiros, B.A.S. (2011) Análise filogenética e revisão sistemática do gênero Anchylorhynchus Schoenherr, 1836 (Curculionidae: Derelomini), com o uso de caracteres morfológicos contínuos e discretos. M. Sc. Thesis Universidade de São Paulo, São Paulo, 328 pp.

    de Medeiros, B.A.S., Bená, D. de C. & Vanin, S.A. (2014) Curculio Curculis lupus: biology, behavior and morphology of immatures of the cannibal weevil Anchylorhynchus eriospathae G. G. Bondar, 1943. PeerJ, 2, e502.

    de Medeiros, B.A.S. & Farrell, B.D. (2019) Evaluating species interactions as a driver of phytophagous insect divergence. bioRxiv, 842153.

    de Medeiros, B.A.S. & Núñez-Avellaneda, L.A. (2013) Three new species of Anchylorhynchus Schoenherr, 1836 from Colombia (Coleoptera: Curculionidae; Curculioninae; Acalyptini). Zootaxa, 3636 (2), 394–400.

    de Medeiros, B.A.S., Núñez-Avellaneda, L.A., Hernandez, A.M. & Farrell, B.D. (2019) Flower visitors of the licuri palm (Syagrus coronata): brood pollinators coexist with a diverse community of antagonists and mutualists. Biological Journal of the Linnean Society, 126, 666–687.

    Meerow, A.W., Noblick, L.R., Salas-leiva, D.E., Sanchez, V., Francisco-ortega, J., Jestrow, B. & Nakamura, K. (2015) Phylogeny and historical biogeography of the cocosoid palms (Areaceae, Arecoideae, Cocoseae) inferred from sequences of six WRKY gene family loci. Cladistics, 31, 509–534.

    Melo, D., Porto, A., Cheverud, J.M. & Marroig, G. (2016) Modularity: Genes, Development, and Evolution. Annual Review of Ecology, Evolution, and Systematics, 47, 463–486.

    Mirande, J. (2009) Weighted parsimony phylogeny of the family Characidae (Teleostei: Characiformes). Cladistics, 25, 574–613.

    Nixon, K.C. (2002) Winclada. Available from: (accessed 8 July 2020)

    Noblick, L.R. (2017) A revision of the genus Syagrus (Arecaceae). Phytotaxa, 294, 001–262.

    Nunes, C.E.P., Maruyama, P.K., Azevedo-Silva, M. & Sazima, M. (2018) Parasitoids turn herbivores into mutualists in a nursery system involving active pollination. Current Biology, 28 (e3), 980–986.

    Núñez-Avellaneda, L.A. (2014) Patrones de asociación entre insectos polinizadores y palmas silvestres en Colombia con énfasis en palmas de importancia económica. PhD Thesis. Universidad Nacional de Colombia, Bogotá, 348 pp.

    Núñez-Avellaneda, L.A. & Carreño, J.I. (2017) Polinización por abejas en Syagrus orinocensis (Arecaceae) en la Orinoquia colombiana. Acta Biológica Colombiana, 22, 221–233.

    Núñez-Avellaneda, L.A., Isaza, C. & Galeano, G. (2015) Ecología de la polinización de tres especies de Oenocarpus (Arecaceae) simpátricas en la Amazonia colombiana. Revista de Biologia Tropical, 63, 35–55.

    Núñez-Avellaneda, L.A. & Rojas-Robles, R. (2008) Biología reproductiva y ecología de la polinización de la palma milpesos Oenocarpus bataua en los Andes colombianos. Caldasia, 30, 101–125.

    O’Brien, C.W. (1981) A New Anchylorhynchus from Panama (Petalochilinae: Curculionidae: Coleoptera). Annals of Entomological Society of America, 74, 501–503.

    O’Leary, M.A. & Kaufman, S. (2011) MorphoBank: Phylophenomics in the “cloud.” Cladistics, 27, 529–537.

    Pascoe, F.P. (1870) I. Contributions towards a Knowledge of the Curculionidae. Zoological Journal of the Linnean Society, 10, 459–503.

    Pellmyr, O. (2003) Yuccas, Yucca moths, and coevolution: a review. Annals of the Missouri Botanical Garden, 90, 35.

    Pichardo-Marcano, F.J., Nieto-Blázquez, M.E., MacDonald, A.N., Galeano, G. & Roncal, J. (2019) Phylogeny, historical biogeography and diversification rates in an economically important group of Neotropical palms: Tribe Euterpeae. Molecular Phylogenetics and Evolution, 133, 67–81.

    Price, N., Green, S., Troscianko, J., Tregenza, T. & Stevens, M. (2019) Background matching and disruptive coloration as habitat-specific strategies for camouflage. Scientific Reports, 9, 1–10.

    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904.

    Revell, L.J. (2012) phytools: an R package for phylogenetic comparative biology (and other things). Methods in Ecology and Evolution, 3, 217–223.

    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mrbayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.

    Schoenherr, C.J. (1833) Genera et species curculionidum, cum synonymia hujus familiæ. T. 1. Librairie Encyclopedique de Roret, Paris, 316 pp.

    Schoenherr, C.J. (1836) Genera et species curculionidum, cum synonymia hujus familiæ. T. 3. Librairie Encyclopedique de Roret, Paris, 858 pp.

    Schoenherr, C.J. (1843) Genera et species curculionidum, cum synonymia hujus familiæ. T. 7. Librairie Encyclopedique de Roret, Paris, 461 pp.

    Scrucca, L., Fop, M., Murphy, T.B. & Raftery, A.E. (2016) McLust 5: clustering, classification and density estimation using Gaussian finite mixture models. The R Journal, 8, 205–233.

    Sereno, P.C. (2007) Logical basis for morphological characters in phylogenetics. Cladistics, 23, 565–587.

    Sherborn, C.D. (1922) Index Animalium sive Index Nominum Quae ab A.D. MDCCLVIII generibus et speciebus animalium imposita sunt. Sectio Segunda. Part I. Oxford University Press, London, 128 pp.

    Sherborn, C.D. (1923) Index Animalium sive Index Nominum Quae ab A.D. MDCCLVIII generibus et speciebus animalium imposita sunt. Sectio Segunda. Part II. Oxford University Press, London, 384 pp.

    Silberbauer-Gottsberger, I. (1973) Blüten- und Fruchtbiologie von Butia leiospatha (Arecaceae). Österreichische Botanische Zeitschrift, 121, 171–185.

    Silberbauer-Gottsberger, I., Vanin, S.A. & Gottsberger, G. (2013) Interactions of the Cerrado palms Butia paraguayensis and Syagrus petraea with parasitic and pollinating insects. Sociobiology, 60, 306–316.

    Silva, F.R., Begnini, R.M., Lopes, B.C., Castellani, T.T., da Silva, F.R., Begnini, R.M., Lopes, B.C. & Castellani, T.T. (2012) Temporal variation in seed predation by insects in a population of Syagrus romanzoffiana (Arecaceae) in Santa Catarina Island, SC, Brazil. Neotropical Entomology, 41, 17–21.

    STRI (2015) Tree Abundance in BCI. Available from: (accessed 11 October 2018)

    Sukumaran, J. & Holder, M.T. (2010) DendroPy: A Python library for phylogenetic computing. Bioinformatics, 26, 1569–1571.

    Valente, R.M. (2005) O gênero Celetes Schoenherr, 1836 (Curculionidae: Erirhininae): filogenia, sistemática e evolução com palmeiras hospedeiras. PhD Thesis, Universidade de São Paulo, São Paulo, 258 pp.

    Valente, R.M. & de Medeiros, B.A.S. (2013) A new species of Anchylorhynchus Schoenherr (Coleoptera: Curculionidae) from the Amazon, with a record of a new host palm for the genus. Zootaxa, 3709 (4), 394–400.

    Vanin, S.A. (1973) Nova espécie do gênero Anchylorhynchus Schoenherr (Coleoptera, Curculionidae, Petalochilinae). Revista Brasileira de Entomologia, 17, 37–40.

    Vanin, S.A. (1995) Three new species of palm weevils from the Amazonian region (Coleoptera, Curculionidae). Revista Brasileira de Entomologia, 39, 871–881.

    Vaurie, P. (1954) Revision of the genera Anchylorhynchus and Petalochilus of the Petalochilinae (Coleoptera, Curculionidae). American Museum Novitates, 1651, 1–58.

    Viana, M.J. (1975) Revision de “Petalochilinae” argentinos. Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, 5, 1–57.

    Voss, E. (1943) Ein Uberbliek über die bisher bekannt gewordenen Arten der Gattung Ancylorrhynchus aus der Unterfamilie Petalochilinae (Col., Curc.). Entomologische Blätter, 39, 60–64.

    Wanat, M. (2007) Alignment and homology of male terminalia in Curculionoidea and other Coleoptera. Invertebrate Systematics, 21, 147.

    Wibmer, G.J. & O’Brien, C.W. (1986) Annotated checklist of the weevils (Curculionidae sensu lato) of South America (Coleoptera: Curculionoidea). Memoirs of the American Entomological Institute, 39, 1–563.

    Wilcox, D., Dove, B., McDavid, D. & Greer, D. (1995) UTHSCSA ImageTool for Windows. Available from: (accessed 20 March 2012)

    Wright, A.M., Lloyd, G.T. & Hillis, D.M. (2016) Modeling Character Change Heterogeneity in Phylogenetic Analyses of Morphology through the Use of Priors. Systematic Biology, 65, 602–611.

    Yu, G., Lam, T.T.Y., Zhu, H. & Guan, Y. (2018) Two methods for mapping and visualizing associated data on phylogeny using GGTree. Molecular Biology and Evolution, 35, 3041–3043.

    Yu, G., Smith, D.K., Zhu, H., Guan, Y. & Lam, T.T.Y. (2017) Ggtree: an R package for visualization and annotation of phylogenetic trees with their covariates and other associated data. Methods in Ecology and Evolution, 8, 28–36.

    Zapata, F. & Jiménez, I. (2012) Species delimitation: Inferring gaps in morphology across geography. Systematic Biology, 61, 179–194.