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Coleoptera of Brazil: what we knew then and what we know now. Insights from the Catálogo Taxonômico da Fauna do Brasil

ABSTRACT

In 2000, Cleide Costa published a paper presenting the state of knowledge of the Neotropical Coleopte ra, with a focus on the Brazilian fauna. Twenty-four years later, thanks to the development of the Coleoptera section of the Taxonomic Catalog of the Brazilian Fauna (CTFB - Catálogo Taxonômico da Fauna do Brasil) through the collaboration of 100 coleopterists from all over the globe, we can build on Costa’s work and present an updated overview of the state of knowledge of the beetles from Brazil. There are currently 35,699 species in 4,958 genera and 116 families known to occur in the country, including representatives of all extant suborders and superfamilies. Our data show that the Brazilian beetle fauna is the richest on the planet, concentrating 9% of the world species diversity, with some estimates accounting to up to 15% of the global total. The most diverse family in numbers of genera is Cerambycidae (1,056 genera), while in number of species it is Chrysomelidae (6,079 species). Conotrachelus Dejean, 1835 (Curculionidae) is the most species-rich genus, with 570 species. The French entomologist Maurice Pic is the author who has contributed the most to the naming of species recorded from Brazil, with 1,794 valid names in 36 families, whereas the Brazilians Ubirajara R. Martins and Maria Helena M. Galileo are the only ones among the top-ten authors to have named species in the 21st century. Currently, approximately 144 new species of Brazilian beetles are described each year, and this average is projected to increase in the next decade to 180 species per year, or about one new Brazilian beetle every two days.

KEY WORDS:
Beetles; biodiversity; list; Neotropical; South America; CTFB

INTRODUCTION

Beetles make up one of the most successful and species-rich clades of animals, with approximately 30,000 genera and 387,000 species described worldwide (Ślipiński et al. 2011Ślipiński SA, Leschen RAB, Lawrence JF (2011) Order Coleoptera Linnaeus, 1758. In: Zhang Z-Q (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 203-208. https://doi.org/10.11646/zootaxa.3148.1.39
https://doi.org/10.11646/zootaxa.3148.1....
, Cai et al. 2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
). One of the most widely accepted hypotheses explaining their evolutionary radiation concerns the development of protective elytra, which allow these insects to occupy spaces not commonly used by other land arthropods (McKenna et al. 2015Mckenna DD, Wild AL, Kanda K, Bellamy CL, Beutel RG, et al. (2015) The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40: 835-880. https://doi.org/10.1111/syen.12132
https://doi.org/10.1111/syen.12132...
, Boudinot et al. 2022Boudinot BE, Fikácek M, Lieberman ZE, Kusy D, Bocak L, McKenna DD, Beutel RG (2022) Systematic bias and the phylogeny of Coleoptera - A response to Cai et al. (2022) following responses to Cai et al. (2020). Systematic Entomology 48(2): 223-232. https://doi.org/10.1111/syen.12570
https://doi.org/10.1111/syen.12570...
, Goczał and Beutel 2023Goczał J, Beutel RG (2023) Beetle elytra: evolution, modifications and biological functions. Biology Letters 19(3): 1-11. http://doi.org/10.1098/rsbl.2022.0559
http://doi.org/10.1098/rsbl.2022.0559...
, Ferreira et al. 2023Ferreira VS, Barbosa FF, Bocakova M, Solodovnikov A (2023) An extraordinary case of elytra loss in Coleoptera (Elateroidea: Lycidae): Discovery and placement of the first anelytrous adult male beetle. Zoological Journal of the Linnean Society 199: 1-14. https://doi.org/10.1093/zoolinnean/zlad026
https://doi.org/10.1093/zoolinnean/zlad0...
).

The body length of adults varies widely, ranging from 0.3 mm for a species of Ptiliidae to 200 mm for a species of Cerambycidae (Casari et al. 2024Casari SA, Biffi G, Ide S (2024) Coleoptera Linnaeus, 1758. In: Rafael JA, Melo GAR, Carvalho CJB, Casari SA, Constantino R (Eds) Insetos do Brasil: diversidade e taxonomia . Instituto Nacional de Pesquisas da Amazônia, Manaus, 2nd ed., 575-698.), and several morphological and physiological adaptations allow them to live in virtually all terrestrial and freshwater ecosystems (Cai et al. 2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
). This wide variation is, among other factors, a result of the ancient age of the group, about 306 to 322 million years old (Cai et al. 2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
), which gives Coleoptera the status of one of the oldest holometabolous orders.

Brazil is the world’s fifth largest country, with an area of 8.5 million km2, occupying almost half of South America, and harbors a megadiverse biota (MMA 2023MMA (2023) Biodiversidade. Ministério do Meio Ambiente, Brasília, Ministério do Meio Ambiente, Brasília, https://antigo.mma.gov.br/biodiversidade.html [Accessed: 07/07/2023]
https://antigo.mma.gov.br/biodiversidade...
). The country encompasses six major biogeographic provinces: the Amazon rainforest, the Caatinga dry forests, the Pampa grasslands, the Pantanal wetlands, the Cerrado savanna, and the Atlantic rainforest. The latter two are known as biodiversity hotspots due to their high concentration of endemic vertebrate and plant species, and their threatened conservation status (Myers et al. 2000Myers N, Mittermeier RA, Mittermeier CG, Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403: 853-858. https://doi.org/10.1038/3500250
https://doi.org/10.1038/3500250...
, MMA 2023MMA (2023) Biodiversidade. Ministério do Meio Ambiente, Brasília, Ministério do Meio Ambiente, Brasília, https://antigo.mma.gov.br/biodiversidade.html [Accessed: 07/07/2023]
https://antigo.mma.gov.br/biodiversidade...
). The Amazon, in turn, comprises about 50% of Brazil’s territory, occupying approximately 4.2 million km2 in the northern part of the country, and congregates the richest biota on Earth (Lewinsohn and Prado 2005Lewinsohn TM, Prado PI (2005) Biodiversidade brasileira: síntese do estado atual do conhecimento. In: Lewinsohn TM, Prado PI (Eds) Síntese do conhecimento atual da biodiversidade brasileira. Contexto, São Paulo, 21-112., MMA 2023MMA (2023) Biodiversidade. Ministério do Meio Ambiente, Brasília, Ministério do Meio Ambiente, Brasília, https://antigo.mma.gov.br/biodiversidade.html [Accessed: 07/07/2023]
https://antigo.mma.gov.br/biodiversidade...
).

Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.) produced the first list of Brazilian Coleoptera, comparing the Brazilian fauna with that of the rest of the Neotropical region and the world. For that work, the author listed the number of genera and species recorded from Brazil for each known family at that time. Species numbers were subsequently updated for the species by Casari and Ide (2012Casari SA, Ide S (2012) Coleoptera Linnaeus, 1758. In: Rafael JA, Melo GAR, Carvalho CJB, Casari SA, Constantino R (Eds) Insetos do Brasil: diversidade e taxonomia. Holos Editora, Ribeirão Preto, 453-535.) and Casari et al. (2024Casari SA, Biffi G, Ide S (2024) Coleoptera Linnaeus, 1758. In: Rafael JA, Melo GAR, Carvalho CJB, Casari SA, Constantino R (Eds) Insetos do Brasil: diversidade e taxonomia . Instituto Nacional de Pesquisas da Amazônia, Manaus, 2nd ed., 575-698.). In 2015, an initiative was started by Brazilian zoologists and overseas partners to build a website named Taxonomic Catalog of the Brazilian Fauna (or Catálogo Taxonômico da Fauna do Brasil, hereafter shorten to its Portuguese abbreviation CTFB), aiming at cataloging the entire fauna of the country. The Coleoptera section is captained by Marcela L. Monné and Cleide Costa, and since then, the CTFB has been constantly updated and a growing number of coleopterists have been involved.

The present survey brings together the most up-to-date data on the Brazilian beetle fauna, the result of the collaboration of 100 Brazilian and foreign researchers, the latter based in as varied countries as Argentina, Australia, Canada, Colombia, the Czech Republic, France, Germany, Italy, Latvia, New Zealand, Poland, Peru, Spain, and the USA (Supplementary Material 1 Supplementary material 1 Supplementary S1. Beetle groups recorded in the CTFB and the authors responsible for them. Classification and taxon names follow Table 1. Authors: E. Caron, M.L. Monné, V.S. Ferreira, et al. Data type: Authors of beetle groups. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zoologia.41.e23072 ). We also provide an updated classification for the beetle families found in Brazil, a brief historical overview of beetle studies in Brazil, and a projection of the expected advancements in discovering new species for the fauna in the coming years.

MATERIAL AND METHODS

Data was retrieved from the Coleoptera section of the CTFB website (http://fauna.jbrj.gov.br/) (Monné and Costa 2023Monné ML, Costa C (2023) Coleoptera. In: Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/223 [Accessed: 10/05/2023]
http://fauna.jbrj.gov.br/fauna/faunadobr...
). An Excel spreadsheet containing all Coleoptera data was extracted from the website on March 7, 2023 (Supplementary Material 2 Supplementary material 2 Supplementary S2. Excel spreadsheet containing all Coleoptera data extracted from the website on March 7, 2023. Data obtained from the Coleoptera section of the CTFB website (http://fauna.jbrj.gov.br/ - Monné and Costa 2023). Authors: E. Caron, M.L. Monné, V.S. Ferreira, et al. Data type: Species data. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zoologia.41.e23072 ), which served as the basis for the analyses performed in this study. The higher-level classification and family-group names of Coleoptera follow Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
), with the changes indicated in Table 1 and discussed in the following section.

Table 1
Coleoptera of Brazil. Data from Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.) and CTFB, the latter as of March 7, 2023. All world beetle families are included on the list following the Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
) classification. Full reference of each taxa name see Bouchard et al. (2011Bouchard P, Bousquet Y, Davies AE, Alonso-Zarazaga MA, Lawrence JF, Lyal CHC, Newton AF, Reid CAM, Schmitt M, Ślipiński SA, Smith ABT (2011) Family-group names in Coleoptera (Insecta). ZooKeys 88: 1-972. https://doi: 10.3897/zookeys.88.807
https://doi: 10.3897/zookeys.88.807...
) or Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
). The world’s genera and species percentages were calculated using Ślipiński et al. (2011Ślipiński SA, Leschen RAB, Lawrence JF (2011) Order Coleoptera Linnaeus, 1758. In: Zhang Z-Q (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 203-208. https://doi.org/10.11646/zootaxa.3148.1.39
https://doi.org/10.11646/zootaxa.3148.1....
). Additional comments are indicated by superscripts.

RESULTS AND DISCUSSION

Our catalog differs from Costa’s (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.) list in that we follow the updated classification recently established by Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
). The following changes have accordingly been made for the CTFB family-level classification: 1) all three families in Archostemata are included in Cupedoidea; 2) In Adephaga, Rhysodinae is now treated as a subfamily of Carabidae; 3) Scirtoidea now includes only Scirtidae, with Clambidae and Eucinetidae transferred to Clamboidea; 4) Nosodendridae is moved from Derodontoidea (now a junior synonym of Clamboidea) to Nosodendroidea; 5) Histeridae is moved from Hydrophiloidea to Histeroidea; 6) Rhadalidae is raised from subfamily status under Melyridae to full family; 7) Byrrhoidea includes just Byrrhidae, with 10 other families previously included in it now transferred to Dryopoidea (Elmidae, Dryopidae, Lutrochidae, Limnichidae, Heteroceridae, Psephenidae, Cneoglossidae, Ptilodactylidae, Chelonariidae and Callirhipidae); and 8) Cucujoidea now consists of seven families (Cryptophagidae, Silvanidae, Cucujidae, Cavognathidae, Passandridae, Phalacridae and Laemophloeidae), having Erotyloidea (with only Erotylidae) and Nitiduloidea (Sphindidae, Monotomidae, Kateretidae, Nitidulidae) removed from it. The three points which we differ from Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
), however, relate to the following taxa: 1) Cicindelinae remains a subfamily of Carabidae, following Gough et al. (2019Gough HM, Duran DP, Kawahara AY, Toussaint EFA (2019) A comprehensive molecular phylogeny of tiger beetles (Coleoptera, Carabidae, Cicindelinae). Systematic Entomology 44: 305-321. https://doi.org/10.1111/syen.12324
https://doi.org/10.1111/syen.12324...
), and not a distinct family; 2) Melolonthidae and Cetoniidae are treated as separate families from Scarabaeidae following Cherman and Morón (2014Cherman MA, Morón MA (2014) Validación de la família Melolonthidae Leach, 1819 (Coleoptera: Scarabaeoidea). Acta Zoológica Mexicana 30: 201-220.) instead of subfamilies of the latter; and 3) Lagrioididae is classified as a family following Lawrence et al. (2023Lawrence JF, Leschen RAB, Elgueta M, Porch N, Ślipiński A (2023) The family Lagrioididae Abdullah & Abdullah, stat. nov. (Coleoptera: Tenebrionoidea), with description of two new Australian species. Annales Zoologici 73: 261-292. https://doi.org/10.3161/00034541ANZ2023.73.2.009
https://doi.org/10.3161/00034541ANZ2023....
) instead of an incertae sedis subfamily of Tenebrionoidea as interpreted by Cai et al. (2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
). These different treatments adhere to more detailed and, sometimes, recent classificatory studies.

Of the 235 families of Coleoptera (Cai et al. 2022Cai C, Tihelka E, Giacomelli M, Lawrence JF, Ślipiński SA, Kundrata R, et al. (2022) Integrated phylogenomics and fossil data illuminate the evolution of beetles. Royal Society Open Science 9: 211771. https://doi.org/10.1098/rsos.211771
https://doi.org/10.1098/rsos.211771...
, with Cicindelinae in Carabidae, and considering Cetoniidae, Melolonthidae, and Lagrioididae as families), 116 have so far been recorded from Brazil (Table 1). They belong to all the living suborders and superfamilies, but no family known exclusively from the fossil record has been found in the country. All fossil species so far described from Brazil belong to extant families. For example, in Staphylinidae, there are three fossil species recorded from Brazil: Caririderma pilosa Martins-Neto, 1990, Apticax solidus Schomann and Solodovnikov, 2012, and A. volans Schomann and Solodovnikov, 2012 (Supplementary Material 2 Supplementary material 2 Supplementary S2. Excel spreadsheet containing all Coleoptera data extracted from the website on March 7, 2023. Data obtained from the Coleoptera section of the CTFB website (http://fauna.jbrj.gov.br/ - Monné and Costa 2023). Authors: E. Caron, M.L. Monné, V.S. Ferreira, et al. Data type: Species data. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zoologia.41.e23072 ). Passalidae has Protopassalus araripensis Santos et al., 2020, the oldest known passalid fossil (Santos et al. 2021Santos MFA, Mattos I, Mermudes JRM, Scheffler SM, Reyes-Castillo P (2021) A new passalid fossil (Insecta: Coleoptera) from the Santana Formation (Crato member, Lower Cretaceous), Araripe Basin, NE Brazil: Paleoecological and paleobiogeographic implications. Cretaceous Research 118: 104664. https://doi.org/10.1016/j.cretres.2020.104664
https://doi.org/10.1016/j.cretres.2020.1...
), whereas Curculionoidea encompasses two fossil species from the Santana Formation, Preclarusbelus vanini Santos et al., 2007, in Nemonychidae, and Arariperhinus monnei Santos et al., 2011, in Curculionidae (Santos et al. 2007Santos MFA, Mermudes JRM, Fonseca VMM (2007) Description of a new genus and species of Belinae (Belidae, Curculionoidea, Coleoptera) from the Santana Formation (Crato member, Lower Cretaceous) of the Araripe basin, northeastern Brazil. In: Carvalho IS, Cassab RCT, Schwanke C (Eds) Paleontology: life scenarios. Interscience, Rio de Janeiro, vol. 1, 449-455., 2011Santos MFA, Mermudes JRM, Fonseca VMM (2011) A specimen of Curculioninae (Curculionidae, Coleoptera) from the Lower Cretaceous, Araripe Basin, north-eastern Brazil. Palaeontology 54: 807-814. https://doi.org/10.1111/j.1475-4983.2011.01057.x
https://doi.org/10.1111/j.1475-4983.2011...
, Anderson et al. 2014Anderson RS, Oberprieler RG, Marvaldi AE (2014) 3.1 Nemonychidae Bedel, 1882. In: Leschen RAB, Beutel RG (Eds) Handbook of Zoology. Walter de Gruyter, Berlin, vol. 3.4, 301-309., Caldara et al. 2014Caldara R, Franz NM, Oberprieler RG (2014) 3.7.19 Curculionidae Latreille, 1802. In: Leschen RAB, Beutel RG (Eds) Handbook of Zoology. Walter de Gruyter, Berlin, vol. 3.4, 589-628.).

Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.) listed 104 families for Brazil, 97 if based on the current classification (Table 1, notes 1, 4, 5, 10 and 14). Our numbers, therefore, record 12 (or 19) more families. This in part reflects a more “splitter” classification adopted here: Melolonthidae, Cetoniidae, Vesperidae and Disteniidae, for example, were all present on Costa’s list not as families, but as subfamilies. Others, however, are genuine additions: Clambidae was first recorded from the country by Casari and Ide (2012Casari SA, Ide S (2012) Coleoptera Linnaeus, 1758. In: Rafael JA, Melo GAR, Carvalho CJB, Casari SA, Constantino R (Eds) Insetos do Brasil: diversidade e taxonomia. Holos Editora, Ribeirão Preto, 453-535.), whereas Jurasaidae was described as a new family endemic to Brazil only four years ago (Rosa et al. 2020Rosa SP, Costa C, Kramp K, Kundrata R (2020) Hidden diversity in the Brazilian Atlantic rainforest: the discovery of Jurasaidae, a new beetle family (Coleoptera, Elateroidea) with neotenic females. Scientific Reports 10: 1544. https://doi.org/10.1038/s41598-020-58416-6
https://doi.org/10.1038/s41598-020-58416...
).

A number of families are known to occur in the Neotropical region, but have so far failed to be collected in Brazil: Lepiceridae*, Sphaeriusidae* (= Microsporidae), Trachypachidae, Meruidae*, Synteliidae, Glaphyridae, Dascillidae*, Derodontidae, Protocucujidae, Hobartiidae, Smicripidae*, Ulodidae, Promecheilidae (= Perimylopidae), Pyrochroidae and Caridae (Costa 2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114., Spangler and Steiner 2005Spangler PJ, Steiner WE Jr (2005) A new aquatic family, Meruidae, from Venezuela (Coleoptera: Adephaga). Systematic Entomology 30: 339-357. https://doi.org/10.1111/j.1365-3113.2005.00288.x
https://doi.org/10.1111/j.1365-3113.2005...
). We expect, however, that the ones indicated by an asterisk (*) will eventually prove to be present in the country as they are known from neighboring areas (e.g., Meruidae) or their species inhabit warm tropical climates in other parts of the globe. The remaining families have rather restricted distributions and tend to live in temperate regions and may be true absentees (e.g., Caridae, which has been recorded in South America solely from Chile and southern Argentina).

Concerning the number of genera, Brazil currently counts 4,958 of them. In comparison with Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.), there was an increase of nearly 30% (Table 1, note 17). The five most diverse families in the number of genera are Cerambycidae (1,056 genera), Curculionidae (737), Chrysomelidae (562), Staphylinidae (474) and Carabidae (226). Together, these five families represent 61% of the genera so far recorded from Brazil.

As for the species, Brazil currently records 35,699 of them. Compared with Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.), the increase was in the same proportion as that observed for the genera, approximately 30% (Table 1). The list of the five most diverse families in number of species shares the same members as the one for the genera: Chrysomelidae (6,079 species), Curculionidae (5,904), Cerambycidae (4,366), Staphylinidae (2,829) and Carabidae (1,841). Following the latter closely, Melolonthidae (1,592) and Buprestidae (1,562 species) join the list of the seven families with over 1,500 species recorded from Brazil. These seven families encompass 67% of the Brazilian beetle fauna.

Only a small number of countries have had their beetle fauna recently counted and cataloged and our data show that the Brazilian fauna is the richest among them, both in terms of genera and species. For example, the Australian fauna accounted for 22,901 species in 3,265 genera, while New Zealand had 5,525 species in 1,094 genera, Chile 3,947 species in 1,196 genera and Peru with about 10,000 species (Klimaszewski and Watt 1997Klimaszewski J, Watt JC (1997) Coleoptera: family-group review and keys to identification. Fauna of New Zealand 37: 1-199., Elgueta 2000Elgueta M (2000) Coleoptera de Chile. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica. SEA, Zaragoza , vol. 1, 145-154., Yeates et al. 2003Yeates DK, Harvey MS, Austin AD (2003) New estimates for terrestrial arthropod species-richness in Australia. Records of the South Australian Museum, Monograph Series 7: 231-241., Chaboo 2015Chaboo CS (2015) Beetles (Coleoptera) of Peru: a survey of the Families. Part I. Overview. Journal of the Kansas Entomological Society 88(2): 135-139. https://doi.org/10.2317/0022-8567-88.2.135
https://doi.org/10.2317/0022-8567-88.2.1...
, Bouchard et al. 2017Bouchard P, Smith ABT, Douglas H, Gimmel M, Brunke A, Kanda Kojun (2017) Biodiversity of Coleoptera. In: Foottit RG, Adler PH (Eds) Insect Biodiversity: Science and Society. John Wiley & Sons, Hoboken, 2nd ed., vol. 1, 337-417.). Our beetle fauna confirms Brazil’s status as a megadiverse country. Brazil harbors a beetle fauna that is actually roughly 40% higher than that listed for the entire North American continent, including Canada, the USA, and northern Mexico combined (Marske and Ivie 2003Marske KA, Ivie MA (2003) Beetle fauna of the United States and Canada. Coleopterists Bulletin 57: 495-503. https://doi.org/10.1649/663
https://doi.org/10.1649/663...
).

And what about the world’s fauna? Brazil boasts approximately 9% of the described species and 17% of the genera (see Table 1). While estimating the entire beetle biodiversity of the country poses challenges, we can use a conservative approach by selecting a large and relatively well-inventoried group, Cerambycidae, as a proxy. This method suggests that our fauna may actually contribute up to 15% of the world’s beetle species. This estimation prompts the identification of families requiring taxonomic advancement to align with this potential global representation. Notably, families such as Carabidae (5% of the world’s species), Staphylinidae (5%), and Tenebrionidae (7%) emerge as candidates for further taxonomic attention.

The knowledge of the Brazilian fauna for certain groups has greatly improved since Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.). For example, Hydraenidae shows a staggering 1,550% increase in the number of species, something also observed for Attelabidae (500%), Heteroceridae (300%), Ciidae (275%), Gyrinidae (250%), Limnichidae (217%) and Coccinellidae (139%). In stark contrast, none of the families of Archostemata and Myxophaga has received taxonomic attention, the same situation of a handful of Polyphaga families such as Glaresidae, Cneoglossidae, Chelonariidae (see note 7 in Table 1), Byrrhidae, Artematopodidae, Throscidae, Biphyllidae, Sphindidae, Kateretidae, Cucujidae, Discolomatidae, Corylophidae, Salpingidae and Scraptiidae. They may be promising groups for study.

Another challenge would be to explore genera with a large number of species (Table 2). Among the top-ten genera, Conotrachelus Dejean, 1835 (Curculionidae) is the richest, with 570 species in the country (Table 2, Fig. 1), followed by Agrilus Curtis, 1825 (Buprestidae), with 450 species, the latter representing about 30% of the diversity of its family in Brazil. Similarly, Chauliognathus Hentz, 1830 (Cantharidae), with 169 species, represents 37% of the Brazilian fauna of the family. The family Tenebrionidae has two genera on the top-ten list, Statira Saint-Fargeau and Audinet-Serville, 1828 and Strongylium Kirby, 1819, which together account for 26% of the family in Brazil. Chrysomelidae is also represented on the top-ten list by two genera, Chlamisus Rafinesque, 1815 and Platyphora Gistel, 1857, but their combined share in the family’s richness is not as significant, merely 6.5% of its species diversity. Other families making the list, each with one representative, are Carabidae (Agra Fabricius, 1801), Melolonthidae (Plectris Saint-Fargeau and Audinet-Serville, 1828), and Staphylinidae (Euconnus Thomson, 1859). Chelonariidae has a single genus of 112 species, Chelonarium Fabricius, 1801 (Table 1). Unfortunately, none of these genera has a taxonomic revision that clarifies their phylogenetic status, species limits, and distribution in Brazil.

Figure 1
Habitus of species of the top-ten richest beetle genera in Brazil. (A, B) Conotrachelus camelus Fiedler, 1940, dorsal and lateral view, courtesy of DZUP; (C, D) Plectris variipennis (Moser, 1921), lateral and dorsal view, DZUP; (E, F) Chlamisus chapadensis Bokermann, 1962, dorsal and lateral view, DZUP, paratype; (G, J) Agrilus oceanicus Cobos, 1959, dorsal and lateral view, DUZP, holotype; (H, K) Chauliognathus flavipes (Fabricius, 1781). Hentz, 1830 dorsal and lateral view, DZUP; (I, L) Strongylium militare Mäklin, 1864, dorsal and lateral view, DZUP; (M) Agra sp. Fabricius, 1801, dorsal view, CELC; (N) Statira sp. Saint-Fargeau and Audinet-Serville, 1828, dorsal view, CELC; (O) Platyphora fulvovittata (Bechyné, 1948), dorsal view, MNRJ; (P) Euconnus sp. Thomson, 1859, dorsal view, CESP. Scale-bars: E, F, G, J, P = 1 mm, A, C, D, I, L, M = 2 mm; H, K, N = 5 mm; O = 10 mm. See Acknowledgements for full photo credits.

Table 2
Ten most speciose genera in the Brazilian beetle fauna.

Concerning the authors who have named species living in Brazil (Fig. 2), the French entomologist Maurice Pic (1866-1957) is the most productive, with 1,794 valid names in 36 families of beetles (Table 3). The list of the ten most productive authors is completed by another Frenchman, a Swede, a German, an American, two British, a Czech, and two Brazilians. Remarkably, two of these authors have over 1,000 valid names in a single family, the Czech Jan Karel Bechyné (1920-1973) in Chrysomelidae and the German Carl Fiedler (1864-1955) in Curculionidae. The two sole Brazilian authors on this list, Ubirajara Ribeiro Martins (1932-2015) and Maria Helena Mainieri Galileo (b. 1950), the only ones to provide names in the 21st century and the latter the only still living, were responsible for the description of hundreds of new species of Cerambycidae. A consequence of the fact that most of the prolific authors were foreigners is that the type specimens they established are usually held in overseas institutions, far from where taxonomic studies on the fauna are now mostly performed, in Brazil. The study of this material is becoming easier through image and data sharing by curators and travel funding. Despite being competitive, this funding enables taxonomists to travel globally.

Figure 2
Top-ten authors of beetle species occurring in Brazil. (A) Maurice Pic (Groll 2016aGroll EK (2016a) Pic, Maurice. Biographies of the Entomologists of the World, details. Senckenberg World Biodiversity. https://sdei.senckenberg.de/biographies/index.php [Accessed: 05/07/2023]
https://sdei.senckenberg.de/biographies/...
); (B) Carl Henrich Boheman (Bauer 2021Bauer C (2021) Boheman, Carl Heinrich. Biographies of the Entomologists of the World, details. Senckenberg World Biodiversity. https://sdei.senckenberg.de/biographies/index.php [Accessed: 05/07/2023]
https://sdei.senckenberg.de/biographies/...
); (C) Jan Karel Bechyné (Schubert 2022Schubert E (2022) Bechyné, Jan Karel. Biographies of the Entomologists of the World, details. Senckenberg World Biodiversity. https://sdei.senckenberg.de/biographies/index.php [Acessed: 05/07/2023]
https://sdei.senckenberg.de/biographies/...
); (D) Carl Fiedler, courtesy of Editha Schubert, from the archives of the SDEI; (E) Ubirajara Ribeiro Martins (Galileo and Santos-Silva 2015Galileo MHM, Santos-Silva A (2015) Necrológio Ubirajara Ribeiro Martins de Souza (1932-2015). Arquivos de Zoologia, Museu de Zoologia da Universidade de São Paulo 46: 41-64. https://doi.org/10.11606/issn.2176-7793.v46i2-11p41-64
https://doi.org/10.11606/issn.2176-7793....
); (F) Henry Walter Bates (Ulbrich 2022Ulbrich S (2022) Bates, Henry Walter. Biographies of the Entomologists of the World, details. Senckenberg World Biodiversity. https://sdei.senckenberg.de/biographies/index.php [Accessed: 05/07/2023]
https://sdei.senckenberg.de/biographies/...
); (G) Thomas Lincoln Casey (Smetana and Herman 2001Smetana A, Herman L (2001) Brief history of taxonomic studies of the Staphylinidae including biographical sketches of the investigators. Catalog of the Staphylinidae (Insecta: Coleoptera) 1758 to the end of the second millennium. Bulletin of the American Museum of Natural History 265: 17-159.); (H) David Sharp (Smetana and Herman 2001Smetana A, Herman L (2001) Brief history of taxonomic studies of the Staphylinidae including biographical sketches of the investigators. Catalog of the Staphylinidae (Insecta: Coleoptera) 1758 to the end of the second millennium. Bulletin of the American Museum of Natural History 265: 17-159.); (I) Alphonse-Adrien Hustache (Maldes and Péricart 1979Maldes J-M, Péricart J (1979) Pour un Trentenaire Rétrospective sur la Biographie et l’Œuvre Entomologique d’Alphonse Hustache (1872-1949). Annales de la Société entomologique de France 15: 194-208. https://doi.org/10.1080/21686351.1979.12278203
https://doi.org/10.1080/21686351.1979.12...
); (J) Maria Helena Mainieri Galileo (Souza et al. 2022Souza D, Jorge I, Marinoni L (2022) Maria Helena Mainieri Galileo. Biografia. Repositório R. Marinoni. https://www.types-rmarinoni.com.br/biografia.php?pesquisador=Maria%20Helena%20Mainieri%20Galileo [Accessed: 05/07/2023]
https://www.types-rmarinoni.com.br/biogr...
).

Table 3
Top ten authors with the most valid names for species recorded from Brazil.

In the inaugural work of modern zoological nomenclature, the Swedish naturalist Carl Linnaeus (1758Linnaeus C (1758) Systema naturae, per regna tria naturae, secundum Classes, Ordines, Genera, Species, cum characteribus, differentiis, synonymis, locis. Laurentii Salvii, Holmiae, vol. 1,10th ed., 824 pp.) described 69 species that we now know to occur in the Brazilian territory, these belonging to 23 families. Around the 1820s, this number had grown to 1,000. A hundred years later, approximately half of the Brazilian beetle fauna had been described, totaling 18,000 species (Fig. 3). Over the last 30 years, the description of new species present in Brazil has increased by more than 25% per decade (Table 4). Currently, approximately 144 new species are described each year. If this trend continues, the next decade, 2022-2031, is expected to see an average of about 180 new species described per year, equating to a new Brazilian beetle discovery about every two days. These results show how much we still have to learn about the Brazilian beetle fauna, most evidently demonstrated by the description from material exclusively collected in the country of the most recently discovered living Coleoptera family, Jurasaidae (Rosa et al. 2020Rosa SP, Costa C, Kramp K, Kundrata R (2020) Hidden diversity in the Brazilian Atlantic rainforest: the discovery of Jurasaidae, a new beetle family (Coleoptera, Elateroidea) with neotenic females. Scientific Reports 10: 1544. https://doi.org/10.1038/s41598-020-58416-6
https://doi.org/10.1038/s41598-020-58416...
, Nunes et al. 2023Nunes JP, Nascimento M, Pereira-Colavite A (2023) A new species of Jurasai Rosa et al., 2020 and the first record of the family Jurasaidae (Coleoptera: Elateroidea) in the northern Brazilian Atlantic Forest. Zootaxa 5323: 524-534. https://doi.org/10.11646/zootaxa.5323.4.4
https://doi.org/10.11646/zootaxa.5323.4....
).

Figure 3
Accumulation curve of Brazilian Coleoptera species by decade from 1758 to 2022.

Table 4
Annual average and percentage increase in the description of new Coleoptera species occurring in Brazil over the past three decades, 1992 to 2021.

To continue improving our knowledge about the Brazilian beetle fauna, we must advance in taxonomic studies of all families. Particularly promising are families with substantial species numbers but whose described Brazilian fauna represents less than 10% of the world’s diversity (refer to Table 1), for this situation likely betrays an understudied taxonomy. Also worthy of close attention are those neglected groups showing low growth rates since Costa (2000Costa C (2000) Estado de conocimiento de los Coleoptera neotropicales. In: Martin-Piera F, Morrone JJ, Melic A (Eds) Hacia un Proyecto CYTED para el Inventario y Estimación de la Diversidad Entomológica en Iberoamérica: PrIBES-2000. SEA, Zaragoza, vol. 1, 99-114.), typically involving families with small-sized species (also noted by Navarrete-Heredia et al. 2022Navarrete-Heredia JL, Arriaga-Varela E, Contreras-Félix G (2022) Mexican beetle species described between 2000-2020: Analysis from Zoological Record of patterns and trends. Southwestern Entomologist 47: 399-409. for Mexico), as well as highly speciose genera (Table 2). Finally, it would likely pay to be alert to the potential presence of families that have not yet been recorded from Brazil, such as Meruidae, Sphaeriusidae, and others.

ACKNOWLEDGEMENTS

We are grateful to the CTFB managing team for technical support. Many of the pictures illustrating this text were kindly provided by colleagues: Editha Schubert (SDEI, Senckenberg Deutsche Entomologische Institut, Müncheberg, Germany; the Carl Fiedler portrait), Artur Orsetti (CELC, Coleção Entomológica do Laboratório de Sistemática e Biologia de Coleoptera, Universidade Federal de Viçosa, Viçosa, Brazil; pictures of Agra and Statira species), Norma Ganho, Paulo Alvarenga and Keli Morais (Taxonline project and DZUP, Coleção Pe. Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil; pictures of Chauliognathus, Chlamisus, Conotrachelus, Plectris and Strongylium species) and Nicolly Barbosa (Taxonline project and CESP, Coleção Entomológica do Setor Palotina, Palotina, Brazil; the Euconnus picture). EC and JRMM thank Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the research fellowship (311390/2021-8 and 312786/2022-0, respectively). MC was supported by a technician fellowship provided by Paraná state’s Fundação Araucária. VSF is grateful to the CNPq for his PhD scholarship and novation process (202559/2015-7). GB is grateful to Instituto Tecnológico Vale and Fundação Guamá (Belém, PA). Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the United States Department of Agriculture. The United States Department of Agriculture is an equal-opportunity employer and provider.

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ADDITIONAL NOTES

Supplementary material 1

Supplementary S1. Beetle groups recorded in the CTFB and the authors responsible for them. Classification and taxon names follow Table 1.

Authors: E. Caron, M.L. Monné, V.S. Ferreira, et al.

Data type: Authors of beetle groups.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/zoologia.41.e23072

Supplementary material 2

Supplementary S2. Excel spreadsheet containing all Coleoptera data extracted from the website on March 7, 2023. Data obtained from the Coleoptera section of the CTFB website (http://fauna.jbrj.gov.br/ - Monné and Costa 2023).

Authors: E. Caron, M.L. Monné, V.S. Ferreira, et al.

Data type: Species data.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/zoologia.41.e23072

Edited by

Editorial responsibility

Sionei R. Bonatto

Data availability

Data citations

Monné ML, Costa C (2023) Coleoptera. In: Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/223 [Accessed: 10/05/2023]

Publication Dates

  • Publication in this collection
    06 Sept 2024
  • Date of issue
    2024

History

  • Received
    10 Oct 2023
  • Accepted
    01 July 2024
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