Abstract
The genus Metaphotina Piza, 1964 is revised based on comparative morphology. Three species are recognized: Metaphotina bimaculata (Saussure, 1870), Metaphotina austri Ferraz, Souza-Dias & Rivera, 2023, and a newly described species from Bahia, Brazil, Metaphotina akaraje sp. nov. Three new synonyms of M. bimaculata are established after a detailed analysis of primary type specimens and their taxonomic and nomenclatural history: Acontista brevipennis Saussure, 1872 syn. nov., Acontista rehni Giglio-Tos, 1927 syn. nov., and Metaphotina piracicabensis Piza, 1964 syn. nov. The analysis of distributional records confirm that Metaphotina is associated with semiarid environments across the eastern South American Dry Diagonal, mainly encompassing the Cerrado, Caatinga, and Chaco biomes. Field observations enabled the identification of the egg cases, discovering the ability of females to produce parthenogenetic offspring, and the recognition and formal description of the myrmecomorphic protonymph of M. bimaculata. Notably, we report on nymphs of M. bimaculata feeding on extrafloral nectaries (EFNs), marking the first known instance of praying mantises interacting with these plant structures. A dichotomous key, extensive graphic documentation of intra- and interspecific variation, and distribution maps are provided to facilitate the identification of Metaphotina spp..
References
- Agudelo, A.A. & Ferraz, B.R. (2024) Mantodea. Catálogo Taxonômico da Fauna do Brasil. Available from: http://fauna.jbrj.gov.br/fauna/faunadobrasil/348 (accessed 5 March 2024)
- Agudelo Rondón, A.A. & Chica, M.L. (2002) Mántidos: Introducción al conocimiento del orden Mantodea. Universidad Distrital Francisco José de Caldas, Bogotá, 90 pp.
- Agudelo Rondón, A.A., Lombardo, F. & Jantsch, L.J. (2007) Checklist of the Neotropical mantids (Insecta, Dictyoptera, Mantodea). Biota Colombiana, 8 (2), 105–158.
- Agudelo, A.A. & Rafael, J.A. (2014) Genus Mantillica Westwood, 1889: Rediscovery and review of the Amazonian “ant-mantis” (Mantodea: Thespidae: Oligonicinae). Entomological Science, 17, 400–408. https://doi.org/10.1111/ens.12072
- Anderson, K. (2018) Acontista cordillerae Saussure, 1869 in Florida. Soothsayer: Journal of Mantodea Research, 2 (1), 97–106.
- Beier, W. von M. (1934) Mantodea. Fam. Mantidae. Subfam. Hymenopodinae. In: Genera Insectorum. L. Desmet-Verteneuil, Brussels, pp. 37.
- Beier, M. (1964) Blattopteroidea-Mantodea. Bronn’s Klassen und Ordnungen des Tierreichs. Vol. 5. In: Bronn, H.G. (Ed.), Insecta-Arthropoda. Part III. Akademische Verlagsgesellschaft Geest & Portig, Leipzig, pp. 850–970.
- Beier, M. (1968) Mantodea (Fangheuschrecken). In: Helmcke, J.-G., Starck, D. & Wermuth, H. (Eds.), Handbuch der Zoologie. Walter de Gruyter & Co, Berlin, pp. 1–47.
- Belchior, C., Sendoya, S.F. & Del-Claro, K. (2016) Temporal variation in the abundance and richness of foliage-dwelling ants mediated by extrafloral nectar. PLoS One, 11 (7), e0158283. https://doi.org/10.1371/journal.pone.0158283
- Blanckenhorn, W.U. & Demont, M. (2004) Bergmann and Converse Bergmann latitudinal clines in arthropods: Two ends of a continuum? Integrative and Comparative Biology, 44, 413–424. https://doi.org/10.1093/icb/44.6.413
- Boudouris, J. & Queenborough, S.A. (2013) Diversity and distribution of extra-floral nectaries in the cerrado savanna vegetation of Brazil. PeerJ, 1, e219. https://doi.org/10.7717/peerj.219
- Brannoch, S.K., Wieland, F., Rivera, J., Klass, K.D., Béthoux, O. & Svenson, G.J. (2017) Manual of praying mantis morphology, nomenclature, and practices (Insecta, Mantodea). ZooKeys, 696, 1–100. https://doi.org/10.3897/zookeys.696.12542
- Caudell, A.N. (1904) On a collection of non-saltatorial Orthoptera from Paraguay. Journal of the New York Entomological Society, 12, 179–188.
- Cerdá, F.J. (1993) Valor taxonomico del complejo falico en mantidos neotropicales (Dictyoptera: Mantodea). Boletín de Entomología Venezolana, New Series, 8, 33–52.
- Chown, S.L., Jaco, C., Chown, K., Klok, S.L., Chown, S.L. & Klok, C.J. (2003) Altitudinal body size clines: latitudinal effects associated with changing seasonality. Ecography, 26, 445–455. https://doi.org/10.1034/j.1600-0587.2003.03479.x
- Dussé, K. & Hurd, L.E. (1997) Food limitation reduces body length in mantid nymphs, Tenodera sinensis Saussure (Mantodea: Mantidae): Implications for fitness. Proceedings of the Entomological Society of Washington, 99, 490–493.
- Ehrmann, R. (2002) Mantodea Gottesanbeterinnen der Welt. 1st Edition. Natur und Tier-Verlag, Münster, 519 pp.
- Ferraz, B.R., Souza-Dias, P.G.B. & Rivera, J. (2023) A hidden gem from northeastern Brazil: A new species of Metaphotina Piza, 1964 (Mantodea, Acontistidae) ecologically linked to the Caatinga. Zootaxa, 5343 (1), 31–54. https://doi.org/10.11646/zootaxa.5343.1.2
- Giglio-Tos, E. (1894) Viaggio del dott. Alfredo Borelli nella Republica Argentina e nel Paraguay. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Universita di Torino, 9, 1–46. https://doi.org/10.5962/bhl.part.8048
- Giglio-Tos, E. (1897) Viaggio del Dott. A. Borelli nel Chaco Boliviano e nella Republica Argentina. Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universita di Torino, 12, 1–47. https://doi.org/10.5962/bhl.part.4564
- Giglio-Tos, E. (1915) Mantidi esotici. Bollettino della Società Entomologica Italiana, 46, 31–108.
- Giglio-Tos, E. (1927) Das Tierreich. Orthoptera-Mantidae. Walter de Gruyter & Co., Berlin, 707 pp. https://doi.org/10.1515/9783111430669
- Hebard, M. (1924) Studies in the Dermaptera and Orthoptera of Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia, 76, 109–248.
- Heitzmann-Fontenelle, T. (1965) Acontiothespis concinna (Perty, 1832) (Mantodea, Acontiothespinae): Descrição morfológica. Papéis Avulsos do Departamento de Zoologia, 17, 277–289. https://doi.org/10.11606/0031-1049.1965.17p277-289
- Heitzmann-Fontenelle, T.J. (1968) Acontiothespis brevipennis (Saussure, 1871) (Mantodea, Acontiothespinae): Descrição morfológica. Papéis Avulsos de Zoologia, 21, 99–110. https://doi.org/10.11606/0031-1049.1968.21.p99-110
- International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
- Kirby, W.F. (1904) A synonymic catalogue of Orthoptera. British Museum of Natural History, London, 501 pp.
- Lombardo, F. (1996) Neotropical mantids. IV. The Genera Tithrone Stål and Paratithrone n. gen. (Mantoidea: Mantidae: Acontiothespini). Journal of Orthoptera Research, 5, 45–50. https://doi.org/10.2307/3503578
- Luebert, F. (2021) The two South American dry diagonals. Frontiers of Biogeography, 13 (4), e51267. https://doi.org/10.21425/F5FBG51267
- Morrone, J.J., Escalante, T., Rodríguez-Tapia, G., Carmona, A., Arana, M. & Mercado-Gómez, J.D. (2022) Biogrographic regionalization of the Neotropical region: New map and shapefile. Anais da Academia Brasileira de Ciências, 94 (1), e20211167. https://doi.org/10.1590/0001-3765202220211167
- O’Hanlon, J.C. (2014) The roles of colour and shape in pollinator deception in the orchid mantis Hymenopus coronatus. Ethology, 120, 652–661. https://doi.org/10.1111/eth.12238
- O’Hanlon, J.C., Holwell, G.I. & Herberstein, M.E. (2014a) Pollinator deception in the Orchid Mantis. The American Naturalist, 183, 126–132. https://doi.org/10.1086/673858
- O’Hanlon, J.C., Holwell, G.I. & Herberstein, M.E. (2014b) Predatory pollinator deception: Does the orchid mantis resemble a model species? Current Zoology, 60, 90–103. https://doi.org/10.1093/czoolo/60.1.90
- Oliveira, P.S. (1988) Ant-mimicry in some Brazilian salticid and clubionid spiders (Araneae: Salticidae, Glubionidae). Biological Journal of the Linnean Society, 33 (1), 1–15. https://doi.org/10.1111/j.1095-8312.1988.tb00443.x
- Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: A new map of life on Earth. BioScience, 51, 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
- Otte, D. & Spearman, L.A. (2005) Mantida species file: catalog of the mantids of the world. Insect Diversity Association, Philadelphia, 489 pp.
- Pfäuti, P. & Hollier, J. (2012) Additions to the catalogue of mantid type material deposited in the Muséum d’histoire naturelle de Genève (Insecta: Mantodea). Revue Suisse de Zoologie, 119, 261–267. https://doi.org/10.5962/bhl.part.151211
- Pie, M.R. & Del-Claro, K. (2002) Male-male agonistic behavior and ant-mimicry in a Neotropical richardiid (Diptera: Richardiidae). Studies on Neotropical Fauna and Environment, 37 (1), 19–22. https://doi.org/10.1076/snfe.37.1.19.2114
- Piza, S.de T. (1964) Um novo gênero de Mantidae do Brasil. Anais do Segundo Congresso Latino-Americano de Zoologia, 1962, 125.
- Rehn, J.A.G. (1907) Non-saltatorial and acridoid Orthoptera from Sapucay, Paraguay. Proceedings of the Academy of Natural Sciences of Philadelphia, 59, 151–192.
- Rehn, J.A.G. (1916) The Stanford Expedition to Brazil, 1911. Dermaptera and Orthoptera I. Transactions of the American Entomological Society, 42, 215–308.
- Rehn, J.A.G. (1920) Records and descriptions of Brazilian Orthoptera. Proceedings of the Academy of Natural Sciences of Philadelphia, 72, 214–293.
- Rehn, J. (1935) The Orthoptera of Costa Rica, Part I: Mantidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 87, 167–272.
- Rivera, J. & Svenson, G.J. (2014) Editorial - A revived focus on the praying mantises (Insecta: Mantodea). Zootaxa, 3797 (1), 5–6. https://doi.org/10.11646/zootaxa.3797.1.3
- Rivera, J. & Svenson, G.J. (2016) The Neotropical “polymorphic earless praying mantises” - Part I: Molecular phylogeny and revised higher-level systematics (Insecta: Mantodea, Acanthopoidea). Systematic Entomology, 41, 607–649. https://doi.org/10.1111/syen.12178
- Rivera, J. & Svenson, G.J. (2020) The Neotropical ‘polymorphic earless praying mantises’ – Part II: Taxonomic review of the genera and checklist of species (Insecta: Mantodea, Acanthopoidea). Thomas Say Publications in Entomology, Monograph 37, Entomological Society of America, Annapolis, Maryland, 236 pp. https://doi.org/10.4182/QQHU9127
- Rivera, J. & Vergara-Cobián, C. (2017) A checklist of the praying mantises of Peru: new records, one new genus (Piscomantis gen. n.) and biogeographic remarks (Insecta, Mantodea). Zootaxa, 4337 (3), 361–389. https://doi.org/10.11646/zootaxa.4337.3.3
- Roy, R. (2004) “Acontista” Saussure & Zehntner, 1894, genre valid (Dict. Mantodea). Bulletin de la Société Entomologique de France, 109, 231–235. https://doi.org/10.3406/bsef.2004.16119
- Roy, R. (2006) Vue d’ensemble sur les Acontistinae Giglio-Tos, 1919 (Dictyoptera, Acanthopidae). Bulletin de la Société Entomologique de France, 111, 327–338. https://doi.org/10.3406/bsef.2006.16331
- Salazar, J.E. (2003) On some mimetic resemblaces of Acontista multicolor in Colonbia (Mantodea). Lambillionea, 103, 649–654.
- Santos, J.C. & Del-Claro, K. (2001) Interação entre formigas, herbívoros e nectários extraflorais em Tocoyena formosa (Cham. & Schlechtd.) K. Schum. (Rubiaceae) na vegetação do cerrado. Revista Brasileira de Zoociências, 3 (1), 77–92.
- Saussure, H. (1870) Additions au Système des Mantides. Mitteilungen der Schweuzerischen entomologischen Gesellschaft, 3, 221–244.
- Saussure, H. (1871) Synopsis des Mantides Américains. In: Memories pour servir a L’Histoire Naturelle du Mexique, des Antilles et États-Unids. H. George, Genéve et Bale, pp. 186.
- Saussure, H. (1872a) Mélanges Orthoptérologiques. Tome 2me. Fasc. 4–6. Ramboz et Schuchard, Genéve, 836 pp.
- Saussure, H. (1872b) Recherches Zoologiques pour servir à l’histoire de la Faune de l’Amerique Centrale et du Mexique. Études sur les Myriapodes et les insectes, famille des Mantides. Mission Mexico, 6, 202–295.
- Saussure, H. & Zehntner, L. (1894) Fam. Mantidae. In: Biologia Centrali-Americana. Société Entomologique, pp. 123–197.
- Scherrer, M.V. & Aguiar, A.P. (2021) Myrmecomorphy in early nymphs of Acontista concinna (Perty) (Mantodea: Acanthopidae: Acontistini), with notes on a new chromatic form. Annales de la Societe Entomologique de France, 57, 379–390. https://doi.org/10.1080/00379271.2021.1974314
- Scherrer, M.V. & Aguiar, A.P. (2022) Comparative morphology of the post-embryonic development reveals the existence of distinct nymphal semaphoronts for Mantodea (Dictyoptera). Annales de la Societe Entomologique de France, 58, 469–498. https://doi.org/10.1080/00379271.2022.2142158
- Schwarz, C.J. & Roy, R. (2019) The systematics of Mantodea revisited: an usdpted classification incorporating multiple data sources (Insecta: Dictyoptera). Annales de la Societe Entomologique de France, 55, 101–196. https://doi.org/10.1080/00379271.2018.1556567
- Schwarz, C.J., Ehrmann, R., Stiewe, M.B.D., Mörtter, R. & Falkenberg, M. (2020) Mantodea of Panguana (Insecta: Dictyoptera). Zootaxa, 4824 (1), 1–66. https://doi.org/10.11646/zootaxa.4824.1.1
- Sebastiani, R., Fujihara, R.T. & Garcia, M. (2017) Occurrence of ants (Hymenoptera: Formicidae) in Banisteriopsis laevifolia (A. Juss.) B. Gates (Malpighiaceae) in São Paulo State, Brazil. Brazilian Journal of Biology, 77, 662–663. https://doi.org/10.1590/1519-6984.03816
- Terra, P.S. (1995) Revisão sistemática dos gêneros de louva-a-deus da região neotropical (Mantodea). Revista Brasileira de Entomologia, 39, 13–94.
- Trillo, M.C., Agudelo, A.A., Guerrero, J.C., Miguel, L. & Lorier, E. (2021) Mantodea (Insecta) of Uruguay: Diversity and distribution. Zootaxa, 4963 (3), 423–456. https://doi.org/10.11646/zootaxa.4963.3.3
- Weber, M.G. & Keeler, K.H. (2013) The phylogenetic distribution of extrafloral nectaries in plants. Annals of Botany, 111 (6), 1251–1261. https://doi.org/10.1093/aob/mcs225
- Westwood, J.O. (1889) Revision Insectorum Familiae Mantidarim, speciebus novis aut minus cognitis descriptis et delineatis. Gurney & Jackson, London, 55 pp.
- Wieland, F. & Svenson, G.J. (2018) Biodiversity of Mantodea. In: Foottit, R.G. & Adler, P.H. (Eds.), Insect Biodiversity: Science and Society. Wiley-Blackwell, Hoboken, New Jersey, pp. 389–416. https://doi.org/10.1002/9781118945582.ch15
- Zanella, F.C.V. (2011) Evolução da biota da diagonal de formações abertas secas da América do Sul. In: Carvalho, C.B. & Alemida, E. (Eds.), Biogeografia da América do Sul: padrões e processos. Roca, São Paulo, pp. 198–220.