Abstract

A list of species of Rutelinae from Bahia state, Northeastern Brazil, is presented. The list is based on specimens deposited in Brazilian collections. The list includes 4 tribes, 23 genera, 101 species and 17 subspecies. The genera Byrsopolis Burmeister, 1844, Pseudodorysthetus Soula, 2008 and Trizogeniates Ohaus, 1917 are recorded for the first time in Bahia and Northeastern Brazil. Thirty species are newly recorded in Bahia: Areoda espiritosantensis Ohaus, 1905, B. laticollis Burmeister, 1855, Bolax flavolineata (Mannerheim, 1829), Chlorota abdominalis Ohaus, 1926, C. espiritosantensis Ohaus, 1912, Dorysthetus espiritosantensis Ohaus, 1905, D. fulgidus (Waterhouse, 1881), Leucothyreus acanthurus Ohaus, 1917, L. albopilosus Ohaus, 1917, L. campestris Burmeister, 1855, L. cayapo Ohaus, 1931, L. duplopunctatus Frey, 1976, L. eligius Ohaus, 1918, L. fluminensis Ohaus, 1918, L. iridipennis Ohaus, 1917, L. lucipetens Ohaus, 1931, L. occipitalis Ohaus, 1931, L. pallefactus Ohaus, 1924, L. paulista Ohaus, 1917, L. punctulatus Blanchard, 1851, L. suturalis Laporte, 1840, L. trochantericus Ohaus, 1917, L. verticalis Ohaus, 1924, Macraspis cincta (Drury, 1872), Paranomala tricostulata (Ohaus, 1897), P. violacea (Burmeister, 1844), Pseudodorysthetus calcaratus (Spinola, 1835), and Trizogeniates planipennis Ohaus, 1917. Pelidnota unicolor unicolor (Drury, 1778) is recorded for the first time in Bahia. Fourteen species are identified and will be described in subsequent papers: 10 of Leucothyreus MacLeay, 1819, 2 of Lobogeniates Ohaus, 1917 and 1 species of Byrsopolis Burmeister, 1844 and Pelidnota MacLeay, 1819. Rutelini is the richest tribe with 16 genera and 49 species. The information presented in the list generates an important set of knowledge regarding the diversity of Rutelinae of Bahia and Brazil and provides the basis for conducting future research within the group.

Keywords:
new records; Northeastern Brazil; species list

Resumo

É apresentada uma lista de espécies de Rutelinae do estado da Bahia, Nordeste do Brasil. A lista é baseada em espécimes depositados em coleções brasileiras. A lista inclui quatro tribos, 23 gêneros, 101 espécies e 17 subespécies. É feito o primeiro registro para a Bahia e região Nordeste dos gêneros Byrsopolis Burmeister, 1844, Pseudodorysthetus Soula, 2008 e Trizogeniates Ohaus, 1917. Trinta espécies de Rutelinae foram registradas pela primeira vez para a Bahia e região Nordeste: Areoda espiritosantensis Ohaus, 1905, B. laticollis Burmeister, 1855, Bolax flavolineata (Mannerheim, 1829), Chlorota abdominalis Ohaus, 1926, C. espiritosantensis Ohaus, 1912, Dorysthetus espiritosantensis Ohaus, 1905, D. fulgidus (Waterhouse, 1881), Leucothyreus acanthurus Ohaus, 1917, L. albopilosus Ohaus, 1917, L. campestris Burmeister, 1855, L. cayapo Ohaus, 1931, L. duplopunctatus Frey, 1976, L. eligius Ohaus, 1918, L. fluminensis Ohaus, 1918, L. iridipennis Ohaus, 1917, L. lucipetens Ohaus, 1931, L. occipitalis Ohaus, 1931, L. pallefactus Ohaus, 1924, L. paulista Ohaus, 1917, L. punctulatus Blanchard, 1851, L. suturalis Laporte, 1840, L. trochantericus Ohaus, 1917, L. verticalis Ohaus, 1924, Macraspis cincta (Drury, 1872), Paranomala tricostulata (Ohaus, 1897), P. violacea (Burmeister, 1844), Pseudodorysthetus calcaratus (Spinola, 1835) e Trizogeniates planipennis Ohaus, 1917. Foi feito o primeiro registro de Pelidnota unicolor unicolor (Drury, 1778) para a Bahia. Foram identificadas 14 espécies, que serão descritas em trabalhos posteriores: dez de Leucothyreus MacLeay, 1819, duas de Lobogeniates Ohaus, 1917 e uma de Byrsopolis Burmeister, 1844 e Pelidnota MacLeay, 1819. A tribo Rutelini apresenta a maior riqueza com 16 gêneros e 49 espécies. Os resultados aqui apresentados têm importantes informações para o conhecimento de Rutelinae, no estado da Bahia e no Brasil, e servirão de base para a realização de futuras pesquisas com o grupo.

Palavras-chave:
lista de espécies; novos registros; Região Nordeste do Brasil

Introduction

According to Kohlmann & Morón (2003)KOHLMANN, B. & MORÓN, M. A. 2003. Análisis histórico de la clasificación de los Coleoptera Scarabaeoidea o Lamellicornia. Acta Zoológica Mexicana (n.s.) 90: 175-280. Scarabaeoidea is represented by three families: Lucanidae, Passalidae and Scarabaeidae, with this last family divided into two informal categories, “Laparosticti” (with spiracles positioned on the pleural membrane) and “Pleurosticti” (with spiracles positioned on the superior part of the abdominal ventrite). The scarabaeids include two feeding groups: coprophagous consisting of subfamilies Aphodiinae and Scarabaeinae (“Laparosticti”) and the phytophagous consisting of subfamilies Cetoniinae, Dynastinae, Melolonthinae and Rutelinae (“Pleurosticti”) (Lawrence & Newton 1995LAWRENCE, J.F. & NEWTON, A.F. 1995. Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names). In Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. (J. Pakaluk & S.A. Slipinski. eds.). Crowson, Warsaw: Muzeum i Instytut Zoologii PAN, p.779-1006.). Cherman & Morón (2014)CHERMAN, M.A. & MORÓN, M.A. 2014. Validación de la Família Melolonthidae Leach, 1819 (Coleoptera: Scarabaeoidea). Acta Zoológica Mexicana (n.s.) 30(1): 201-220. recently considered the six phytophagous subfamilies in Melolonthidae (Euchirinae, Dynastinae, Hopliinae, Melolonthinae, Rutelinae and Sericinae). This paper follows the classification of Cherman & Morón (2014)CHERMAN, M.A. & MORÓN, M.A. 2014. Validación de la Família Melolonthidae Leach, 1819 (Coleoptera: Scarabaeoidea). Acta Zoológica Mexicana (n.s.) 30(1): 201-220..

Rutelinae is the second largest subfamily of Melolonthidae according to the number of species, with approximately 4,197 species described in the world but with the greatest richness in the tropics (Hardy 1991HARDY, A. 1991. A Catalog of the Coleoptera of America North of Mexico - Family: Scarabaeidae, Subfamilies: Rutelinae and Dynastinae. California, United States - Department of Agriculture, Agriculture Handbook., Jameson 2002JAMESON, M. L. 2002. Rutelinae MacLeay 1819. In American Beetles Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. (R.H. Arnett, M.C. Thomas, P.E. Skelley & J.H. Frank eds.). CRC Press, Boca Raton, Florida, p.60-64., Ratcliffe & Jameson 2005, Krajcik 2008KRAJCIK, M. 2008. Checklist of Scarabaeoidea of the World: Rutelinae (Coleoptera: Scarabaeidae: Rutelinae), Vol. 2. Animma. X Supplement 4, Plzen Milan Krajcik., Jameson & Ratcliffe 2011JAMESON, M.L. & RATCLIFFE, B.C. 2011. The Neotropical Scarab Beetle Tribe Anatistini (Coleoptera: Scarabaeidae: Rutelinae). Bulletin of the University of Nebraska State Museum 26: 1-100., Morón & Ramírez-Ponce 2012MORÓN, M.A. & RAMÍREZ-PONCE, A. 2012. Mesoamerican genera of Anomalini (Coleoptera: Melolonthidae: Rutelinae): A brief review. Trends in Entomology 8: 97-114.). Melolonthinae is the richest subfamily of Melolonthidae, with about 11,000 known species (Evans 2002EVANS, A.V. 2002. Melolonthinae. In American Beetles Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. (R.H. Arnett, M.C. Thomas, P.E. Skelley & J.H. Frank. eds.). CRC Press, Boca Raton, Florida, p.51-60.). Seven tribes are currently recognized in Rutelinae: Adoretini, Alvarengiini, Anatistini, Anomalini, Anoplognathini, Geniatini and Rutelini (Bouchard et al. 2011BOUCHARD, P., BOUSQUET, Y., DAVIES, A. E., ALONSO-ZARAZAGA, M. A., LAWRENCE, J. F., LYAL, C. H., NEWTON, A. F., REID, C. A., SCHMITT, M., SLIPIŃSKI, S. A. & SMITH, A. B. 2011. Family-group names in Coleoptera (Insecta). ZooKeys 88: 1-972.).

Adult ruteline are phytophagous, playing an important ecological role in pollination of some plant species. Larvae are saprophytophagus and contribute directly to the decomposition of dead organic matter deposited in forests and in the nutrient cycle (Hardy 1991HARDY, A. 1991. A Catalog of the Coleoptera of America North of Mexico - Family: Scarabaeidae, Subfamilies: Rutelinae and Dynastinae. California, United States - Department of Agriculture, Agriculture Handbook., Morón et al., 1997MORÓN, M.A., RATCLIFFE, B.C. & DELOYA, C. 1997. Atlas de los Escarabajos de México. Coleoptera: Lamellicornia. Vol. 1: Família Melolonthidae. Subfamílias Rutelinae, Dynastinae, Cetoniinae, Trichiinae, Valginae y Melolonthinae. Sociedad Mexicana de Entomologia, A.C., Mexico., Paucar-Cabrera 2003PAUCAR-CABRERA, A. 2003. Systematics and phylogeny of the genus Epectinaspis Blanchard (Coleoptera: Scarabaeidae: Rutelinae) and description of a new genus of Anomalini from Mexico. Coleopterists Society Monographs 2: 1-60.), while some species feed on roots and sometimes become pests (Ritcher 1958, Jameson et al. 2003, Jameson & Howkins 2005). In Brazil, Paranomala testaceipennis (Blanchard, 1851) was recorded in agricultural crops and pasture areas in Mato Grosso do Sul (Rodriguez-del-Bosque 1996RODRIGUEZ-DEL-BOSQUE, L.A. 1996. Population and adult longevity the Phyllophaga crinita, Anomala flavipennis and A. forminosa Coleoptera (Scarabaeidae). Southwestern Entomologist 21 (1): 55-58., 1998RODRIGUEZ-DEL-BOSQUE, L.A. 1998. A sixteen-year study on the bivoltinism of Anomala flavipennis (Coleoptera: Scarabaeidae) in Mexico. Environmental Entomology 27: 248-252., Rodrigues et al. 2008RODRIGUES, S.R., PUKER, A., ABOT, A.R.; BARBOSA, C.L., IDE, S. & COUTINHO, G.V. 2008. Ocorrência e aspectos biológicos de Anomala testaceipennis Blanchard (Coleoptera, Scarabaeidae). Revista Brasileira de Entomologia 52 (1): 68-71.); Leucothyreus albopilosus Ohaus, 1917 was recorded on Eucalyptus citriodora (Hook, 1848); L. dorsalisBlanchard, 1850BLANCHARD, C.E. 1850 (1851). Ordre des Coleoptera. In Muséum d'Histoire Naturelle de Paris. Catalogue de la Collection Entomologique. Classe des Insectes, Vol. 1, Part 2. (H. Milne-Edwards, C.E. Blanchard & H. Lucas. eds.). Gide and Baudry, Paris, p.129-240. was associated to the roots of Acrocomia aculeata (Jacq) Lodd. ex Mart. (Arecaceae) from Mato Grosso do Sul (Puker et al. 2009PUKER, A., RODRIGUES, S.R., TIAGO, E.F. & SANTOS, W.T. 2009. Phytophagous species of Scarabaeidae (Insecta: Coleoptera) associated with the root system of Acrocomia aculeata (Jacq.) Loddi. ex Mart. (Arecaceae). Biota Neotropica 9 (3): 105-109., Pereira et al. 2013PEREIRA, A.F., RODRIGUES, S.R. & MORÓN, M.A. 2013. Biological aspects of Leucothyreus alvarengai Frey and Leucothyreus aff. semipruinosus Ohaus (Coleoptera, Melolonthidae, Rutelinae) in crop succession at central Brazil. Revista Brasileira de Entomologia 57 (3): 323-228.).

In the Neotropical Region, approximately 1,337 species of Rutelinae were recorded (Morón 1990MORÓN, M.A. 1990. The Beetle of the World: Rutelini. Sciences Nat. 10: 1-206., 2004MORÓN, M. A. 2004. Melolontídeos edafícolas. In Pragas de solo no Brasil. (J.R. Salvadori, C.J. Ávila & M.T.B. Silva. eds.). Passo Fundo: Embrapa Trigo; Dourados: Embrapa Agropecuária Oeste; Cruz Alta: Fundacep Fecotrigo, p.41-68., Jameson 2008JAMESON, M.L. 2008. Review of the genus Microchilus Blanchard (Coleoptera: Scarabaeidae: Rutelinae: Geniatini). Insecta Mundi 0025: 1-14., Soula 2011SOULA, M. 2011. Les Coléoptères du Nouveau Monde. Rutelini 5. Geniatini 1, Révision du genre Bolax, Besoiro 5: 1-85., Filippini et al. 2016FILIPPINI, V., MICÓ, E. & GALANTE, E. 2016. Checklist and identification key of Anomalini (Coleoptera, Scarabaeidae, Rutelinae) of Costa Rica. ZooKeys 621: 63-136. http://doi:10.3897/zookeys.621.7565.
https://doi.org/http://doi:10.3897/zooke... , Ferreira et al. 2017FERREIRA, A.S., ALMEIDA, L.M. & BRAVO, F. 2017. Three new species of Pelidnota MacLeay (Coleoptera, Scarabaeidae, Rutelinae) and new distributional records from northeast Brazil. Revista Brasileira de Entomologia 61: 208-223. http://doi.org/10.1016/j.rbe.2017.04.004.
http://doi.org/10.1016/j.rbe.2017.04.004... , Howkins 2017HAWKS, D.C. 2017. Five new species of Chrysina Kirby (Coleoptera: Scarabaeidae: Rutelinae). Insecta Mundi 0544: 1-9., Moore et al., 2017MOORE, M.R., JAMESON, M.L., GARNER, B.H., AUDIBERT, C., SMITH, A.B.T. & SEIDEL, M. 2017. Synopsis of the pelidnotine scarabs (Coleoptera, Scarabaeidae, Rutelinae, Rutelini) and annotated catalog of the species and subspecies. ZooKeys 666: 1-349., Seidel et al. 2017SEIDEL, M., JAMESON, M.L. & STONE, R.L. 2017. A new cryptic species and review of the east-Andean leaf chafer genus Mesomerodon Ohaus, 1905 (Coleoptera, Scarabaeidae, Rutelinae). ZooKeys 671: 61-85. http://doi.org/10.3897/zookeys.671.11815.
http://doi.org/10.3897/zookeys.671.11815... , Sierra 2017SIERRA, J.M. 2017. Four new species of Chrysina Kirby (Coleoptera: Scarabaeidae: Rutelinae) from Guatemala and Honduras. Insecta Mundi 0543: 1-12.), while 436 are from Brazil (Grossi & Vaz-de-Mello 2016GROSSI, P.C. & VAZ-DE-MELLO, F.Z. 2016. Rutelinae in Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/126897 (last access at 10/09/2016).
http://fauna.jbrj.gov.br/fauna/faunadobr... , Ferreira et al. 2017FERREIRA, A.S., ALMEIDA, L.M. & BRAVO, F. 2017. Three new species of Pelidnota MacLeay (Coleoptera, Scarabaeidae, Rutelinae) and new distributional records from northeast Brazil. Revista Brasileira de Entomologia 61: 208-223. http://doi.org/10.1016/j.rbe.2017.04.004.
http://doi.org/10.1016/j.rbe.2017.04.004... ). Grossi & Vaz-de-Mello (2016)GROSSI, P.C. & VAZ-DE-MELLO, F.Z. 2016. Rutelinae in Catálogo Taxonômico da Fauna do Brasil. PNUD. http://fauna.jbrj.gov.br/fauna/faunadobrasil/126897 (last access at 10/09/2016).
http://fauna.jbrj.gov.br/fauna/faunadobr... also included 104 subspecies and 58 genera in their list from Brazil. Only one inventory of Rutelinae is known from Bahia State (Viana et al. 2001VIANA, C.H.P., MARQUES, O.M. & CARVALHO, C.A.L. 2001. Rutelinae (Coleoptera: Scarabaeidae) Coletados em Armadilha Luminosa em Cruz das Almas, Bahia. Magistra, Cruz Das Almas-Bahia 13 (1): 1-7.), which was performed with light traps in Cruz das Almas municipality (approximately 12°40’S–39°06’W). The authors recorded species of four genera: Paranomala Casey, 1915 (Anomalini), Geniates Kirby, 1819 Leucothyreus MacLeay, 1819 (Geniatini) and Pelidnota MacLeay (Rutelini), however, only one species was named and identified, Pelidnota fulva Blanchard, which is very probably a misidentified species, as this taxon occurs in another geographical region, according to the last revision (Soula 2009SOULA, M. 2009. Les Coléoptères du Nouveau Monde. Rutelini 3. Révision des Pelidnotina 3, Besoiro 3: 1-139.).

The most recent information on the diversity of the species from Bahia is scattered among 17 papers (Jameson 1996JAMESON, M.L. 1996. Revision and Phylogeny of the Neotropical genus Cnemida (Coleoptera: Scarabaeidae: Rutelinae). Insecta Mundi 10(1-4): 285-315., Jameson & Hawkins 2005JAMESON, M.L. & HAWKINS, S.J. 2005. Synopsis of the genera of Geniatini (Coleoptera: Scarabaeidae: Rutelinae) with an annotated catalog of species. Zootaxa 874: 1-76., Jameson & Ratcliffe 2011JAMESON, M.L. & RATCLIFFE, B.C. 2011. The Neotropical Scarab Beetle Tribe Anatistini (Coleoptera: Scarabaeidae: Rutelinae). Bulletin of the University of Nebraska State Museum 26: 1-100., Krajcik 2008KRAJCIK, M. 2008. Checklist of Scarabaeoidea of the World: Rutelinae (Coleoptera: Scarabaeidae: Rutelinae), Vol. 2. Animma. X Supplement 4, Plzen Milan Krajcik., Machatschke 1972MACHATSCHKE, J. 1972. Scarabaeidae: Melolonthidae, Rutelinae. Coleopterum Catalogus Supplementa 66(1-2): 1-429., Soula 2002aSOULA, M. 2002a. Les Coléoptères du Nouveau Monde. Rutelini 2. Hillside Books, Canterbury 26: 1-98., bSOULA, M. 2002b. Les Coléoptères du Nouveau Monde. Rutelini 2. Hillside Books, Canterbury 26 (2): 117-296., 2003SOULA, M. 2003. Les Coléoptères du Nouveau Monde. Rutelini 3. Hillside Books, Canterbury 29: 1-76., 2005SOULA, M. 2005. Les Coléoptères du Nouveau Monde. Rutelini 3. Révision Des Anticheirina 3. Hillside Books, Canterbury 26 (3): 294-409., 2006SOULA, M. 2006. Les Coléoptères du Nouveau Monde. Rutelini 1. Révision des Pelidnotina 1 Et des Lasiocalina. Besoiro Suplement 1: 1-176., 2008SOULA, M. 2008. Les Coléoptères du Nouveau Monde. Rutelini 2. Révision des Pelidnotina 2, Besoiro, 2: 1-40., 2009SOULA, M. 2009. Les Coléoptères du Nouveau Monde. Rutelini 3. Révision des Pelidnotina 3, Besoiro 3: 1-139., 2010SOULA, M. 2010. Les Coléoptères du Nouveau Monde. Rutelini 4. Révision des Pelidnotina 4, Besoiro 4: 1-66., 2011SOULA, M. 2011. Les Coléoptères du Nouveau Monde. Rutelini 5. Geniatini 1, Révision du genre Bolax, Besoiro 5: 1-85., Ratcliffe & Jameson 1989RATCLIFFE, B.C. & JAMESON, M.L. 1989. A Synopsis of the Genus Areoda (Coleoptera: Scarabaeidae: Rutelinae). The Coleopterists Bulletin 43(2): 135-144., Viana et al. 2001VIANA, C.H.P., MARQUES, O.M. & CARVALHO, C.A.L. 2001. Rutelinae (Coleoptera: Scarabaeidae) Coletados em Armadilha Luminosa em Cruz das Almas, Bahia. Magistra, Cruz Das Almas-Bahia 13 (1): 1-7., Ferreira et al. 2017FERREIRA, A.S., ALMEIDA, L.M. & BRAVO, F. 2017. Three new species of Pelidnota MacLeay (Coleoptera, Scarabaeidae, Rutelinae) and new distributional records from northeast Brazil. Revista Brasileira de Entomologia 61: 208-223. http://doi.org/10.1016/j.rbe.2017.04.004.
http://doi.org/10.1016/j.rbe.2017.04.004... ) and includes 61 species and nine subspecies in 20 genera of the tribes Anatistini, Anomalini, Geniatini and Rutelini.

In this paper, we present an updated species list of Rutelinae from Bahia that are deposited in seven collections in South, Southeast and Northeast of Brazil. Information about the geographic distribution of the species listed is also provided. The data presented here gather important information that adds new knowledge about the diversity of Rutelinae in Brazil and the Neotropical region and serves as a basis for conducting future research with the group.

Material and Methods

Bahia is one of the biggest states of Brazil (Figure 1) with an area of 564.733,177 km² (Instituto Brasileiro de Geografia e Estatística–IBGE 2013IBGE–INSTITUTO BRASILEIRO DE GEOGRAFIA E ESTATÍSTICA. 2013. Área Territorial Brasileira: Consulta por Unidade de Federação. http://www.ibge.gov.br/home/geociencias/areaterritorial/principal.shtm (las access at 14/09/2015).
http://www.ibge.gov.br/home/geociencias/... ), which represents 37.7% of all the Northeastern territory (Bahia 2013BAHIA. 2013. Meio Ambiente na Bahia. Secretaria de Indústria, Comércio e Mineração/Companhia Baiana de Pesquisa Mineral. http://www.cbpm.com.br/paginas/meio_bahia.php (last access at 14/09/2015).
http://www.cbpm.com.br/paginas/meio_bahi... ). The vegetation, climate, and altitudinal range are heterogeneous, with the presence of three Brazilian Biomes (Caatinga, Atlantic Forest and Cerrado), ecosystems of restingas, mangroves, campos rupestres, and ecotones between the biomes, rainforests, semi-deciduous forest and high montane vegetation, among others (Bahia 2013BAHIA. 2013. Meio Ambiente na Bahia. Secretaria de Indústria, Comércio e Mineração/Companhia Baiana de Pesquisa Mineral. http://www.cbpm.com.br/paginas/meio_bahia.php (last access at 14/09/2015).
http://www.cbpm.com.br/paginas/meio_bahi... ).

Figure 1
Location of the study area and geographic distribution records Rutelinae in the Bahia state.

The collections visited for this study were (acronyms according to Evenhuis (2009) have been used when available): CEIOC – Entomological Collection of Instituto Oswaldo Cruz, Rio de Janeiro, Brazil; CERPE – Entomological Collection of Universidade Federal Rural de Pernambuco, Recife, Pernambuco, Brazil; DZUP – Entomological Collection Pe. J. S. Moure, Department of Zoology, Universidade Federal do Paraná, Curitiba, Paraná, Brazil; EPGC – Everardo and Paschoal Grossi Collection, Nova Friburgo, Rio de Janeiro, Brazil; MNRJ – National Museum, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil; MZFS – Collection Prof. Johann Becker, Museum of Zoology of Universidade Estadual de Feira de Santana, Bahia, Brazil; and MHNBA/MZUFBA – Entomological Collection of Museum of Natural History/ Zoology of Bahia State/Universidade Federal da Bahia, Salvador, Bahia, Brazil.

The species identification was based on Frey (1976)FREY, G. 1976. Neue Sudamerikanischen Ruteliden. Entomologische Arbeiten aus dem Museum 27: 344-356., Ohaus (1905OHAUS, F. 1905. Revision der Amerikanischen Anoplognathiden (Coleoptera Lamellicornia). Stettiner Entomologische Zeitung 1905: 120-167., 1912OHAUS, F. 1912. Beiträge zur Kenntnis der Ruteliden. X. Stettiner Entomologische Zeitung 73: 273-319., 1913OHAUS, F. 1913. XI. Beitrag zur Kenntnis der Ruteliden. (Col.). Deutsche Entomologische Zeitschrift 1913: 487-511., 1917OHAUS, F. 1917. Neue Geniatinen (Col. lamell. Rutelin.). Stettiner Entomologische Zeitung 78: 3-53., 1918aOHAUS, F. 1918a. Scarabaeidae: Euchirinae, Phaenomerinae, Rutelinae. Coleopterum Catalogus 20: 1-124., 1918bOHAUS, F. 1918b. Neue Geniatinen II (Col. lamell. Rutelin.). Stettiner Entomologische Zeitung 79: 350-365., 1924OHAUS, F. 1924. Neue Geniatinen III. Stettiner Entomologische Zeitung 84: 179-186., 1926OHAUS, F. 1926. Three new species of Rutelinae (Coleoptera lamellicornia) in the Carnegie Museum. Annals of the Carnegie Museum 17: 87-89., 1928, 1930OHAUS, F. 1930. XXVI. Beitrag zur Kenntnis der Rutelinen (Col. amell.). Deutsche Entomologische Zeitschrift 1930: 138-158., 1931OHAUS, F. 1931. Neue Geniatinen IV. Stettiner Entomologische Zeitung. Stettin 92: 227-258.), Soula (1998SOULA, M. 1998. Les Coléoptères du Nouveau Monde. Rutelini 2. Revision Des Anticheirina 1. Hillside Books, Canterbury 26 (1): 1-116., 2006SOULA, M. 2006. Les Coléoptères du Nouveau Monde. Rutelini 1. Révision des Pelidnotina 1 Et des Lasiocalina. Besoiro Suplement 1: 1-176., 2009SOULA, M. 2009. Les Coléoptères du Nouveau Monde. Rutelini 3. Révision des Pelidnotina 3, Besoiro 3: 1-139., 2010SOULA, M. 2010. Les Coléoptères du Nouveau Monde. Rutelini 4. Révision des Pelidnotina 4, Besoiro 4: 1-66., 2011SOULA, M. 2011. Les Coléoptères du Nouveau Monde. Rutelini 5. Geniatini 1, Révision du genre Bolax, Besoiro 5: 1-85.), and by comparison with the studied collections.

For the study of male genitalia, each specimen was immersed in boiling water for approximately two to four minutes and the genitalia removed, then studied under a stereomicroscope, glued in a triangle, and pinned just below the specimen.

The geographical coordinates, when not available on the label of the specimen, were obtained by the geoLoc tool on the speciesLink online data platform (http://splink.cria.org.br/geoloc), using the IBGE as a source for the data recovery. The geographical records were plotted on the map of Bahia using QGIS version 2.10.1. The final artwork for the map of geographical records and richness of genera and species by tribe was executed using Abobe Photoshop CS6^®.

The information transcribed from the labels of the examined material usually adhered to the following pattern: COUNTRY, State: (“Locality”; “additional information”); (Geographic Coordinates), Data, Collector. (ACRONYM OF THE MUSEUM# register number).

Results and Discussion

A total of 1,495 specimens, of 20 genera, 79 species and 12 subspecies of the tribes Anomalini, Anatistini, Geniatini and Rutelini were examined. Before the current study, 23 genera, 101 species and 17 subspecies of Rutelinae from Bahia (Table 1) had been reported. With this study, 32 species, 11 subspecies and the genera Anticheiroides Soula, 1998, Oplognathus MacLeay, 1819 and Parhoplognathus Ohaus, 1930, cited from Bahia in the literature by Machatschke (1972)MACHATSCHKE, J. 1972. Scarabaeidae: Melolonthidae, Rutelinae. Coleopterum Catalogus Supplementa 66(1-2): 1-429., Soula (1998SOULA, M. 1998. Les Coléoptères du Nouveau Monde. Rutelini 2. Revision Des Anticheirina 1. Hillside Books, Canterbury 26 (1): 1-116., 2008SOULA, M. 2008. Les Coléoptères du Nouveau Monde. Rutelini 2. Révision des Pelidnotina 2, Besoiro, 2: 1-40.), and Moore et al. (2017)MOORE, M.R., JAMESON, M.L., GARNER, B.H., AUDIBERT, C., SMITH, A.B.T. & SEIDEL, M. 2017. Synopsis of the pelidnotine scarabs (Coleoptera, Scarabaeidae, Rutelinae, Rutelini) and annotated catalog of the species and subspecies. ZooKeys 666: 1-349. respectively were not found in the collections studied (Table 1).

The genera Byrsopolis Burmeister, 1844, Pseudodorysthetus Soula, 2008 and Trizogeniates Ohaus, 1917 were first recorded for Bahia and the Northeast region. Thirty species of Rutelinae were recorded for the first time in Bahia and Northeastern Brazil: Areoda espiritosantensis Ohaus, 1905, Byrsopolis aff. castanea Burmeister, 1844, Bolax flavolineata (Mannerheim, 1829), Chlorota abdominalis Ohaus, 1926, C. espiritosantensis Ohaus, 1912, Dorysthetus espiritosantensis Ohaus, 1905, D. fulgidus (Waterhouse, 1881), Leucothyreus acanthurus Ohaus, 1917, L. albopilosus Ohaus, 1917, L. campestris Burmeister, 1855, L. cayapo Ohaus, 1931, L. duplopunctatus Frey, 1976, L. eligius Ohaus, 1918, L. fluminensis Ohaus, 1918, L. iridipennis Ohaus, 1917, L. lucipetens Ohaus, 1931, L. occipitalis Ohaus, 1931, L. pallefactus Ohaus, 1924, L. paulista Ohaus, 1917, L. punctulatus Blanchard, 1851, L. suturalisLaporte, 1840LAPORTE [=CASTELNAU] FLNCde. 1840. Histoire naturelle des insectes Coléoptères; avec une introduction renfermant l’anatomie et la physiologie des animaux articulés, par M. Brullé. Tome premier. Histoire naturelle des animaux articulés, annélides, crustacés, arachnides, myriapodes et insectes Tome troisième. P. Duménil, Paris, 324p. [+ 19 plates]., L. trochantericus Ohaus, 1917, L. verticalis Ohaus, 1924, Macraspis cincta (Drury, 1872), Paranomala tricostulata (Ohaus, 1897OHAUS, F. 1897. Stettiner Entomologische Zeitung, LVIII.), P. violacea (Burmeister, 1844BURMEISTER, H. 1844. Handbuch der Entomologie. (Coleoptera Lamellicornia Anthobia et Phyllophaga Systellochela), Vol. 4, Pt. 1. Berlin.), Pseudodorysthetus calcaratus (Spinola, 1835), and Trizogeniates planipennis Ohaus, 1917. The subspecies Pelidnota unicolor unicolor (Drury, 1778) was also a new record for Bahia state (Table 1). Gathering this information with the already existing, makes a total of 101 species, and 17 subspecies in 4 tribes and 23 genera (Table 1) only for Bahia. Rutelini presented the greatest richness, with 16 genera, 49 species and 16 subspecies. Geniatini is the second tribe in richness with 5 genera and 46 species (Table 1). Anomalini and Anatistini presented only 1 genera each – Paranomala Casey, 1915, with 5 species and 2 subspecies and Spodochlamys Burmeister, 1855, with only one species (Table 1).

The richest genera were Leucothyreus (Geniatini), with 25 species, and Pelidnota (Rutelini) with 19 species and 9 subspecies (Figure 2). Leucothyreus, with 164 species described (Jameson & Hawkins 2005JAMESON, M.L. & HAWKINS, S.J. 2005. Synopsis of the genera of Geniatini (Coleoptera: Scarabaeidae: Rutelinae) with an annotated catalog of species. Zootaxa 874: 1-76.), and Pelidnota, with 136 species and 60 valid subspecies (Moore et al. 2017MOORE, M.R., JAMESON, M.L., GARNER, B.H., AUDIBERT, C., SMITH, A.B.T. & SEIDEL, M. 2017. Synopsis of the pelidnotine scarabs (Coleoptera, Scarabaeidae, Rutelinae, Rutelini) and annotated catalog of the species and subspecies. ZooKeys 666: 1-349., Ferreira et al. 2017FERREIRA, A.S., ALMEIDA, L.M. & BRAVO, F. 2017. Three new species of Pelidnota MacLeay (Coleoptera, Scarabaeidae, Rutelinae) and new distributional records from northeast Brazil. Revista Brasileira de Entomologia 61: 208-223. http://doi.org/10.1016/j.rbe.2017.04.004.
http://doi.org/10.1016/j.rbe.2017.04.004... ), are the richest genera within Rutelinae.

Figure 2
Species richness of Rutelinae from Bahia state, gathering the literature data with the results presented in this study.

There were recorded Ruteliane species in fifty-seven, taking into account the literature and the examined material, out of a total of 417 Bahian municipalities. Rutelini were found to occur in 41 localities, and Geniatini in 37 (Figure 1), which represents only 13.67% of the total amount of municipalities. Anomalini was recorded in only eight localities in the state and Anatistini only in the municipality of Cachoeira (Figure 1). Rutelini and Geniatini were recorded in all the Biomes, however Rutelini presented a concentration of records in areas of Atlantic Rain Forest near the coast. Geniatini had similar numbers of records in the Caatinga and in the Atlantic Forest (Figure 1). These findings could be attributed to the natural history of these tribes which immatures of Rutelini species occurs in dead wood (Albertoni et al. 2014ALBERTONI, F.F., FUHRMANN, J. & IDE, S. 2014. Lagochile emarginata (Gyllenhal): morphology of immature and imago, and biological records (Coleoptera, Scarabaeidae, Rutelinae). Revista Brasileira de Entomologia 58(1): 32-46.), while Geniatini species develop mainly in the ground (Furhmann 2013FUHRMANN, J. 2013. Description of the third larval instar and pupa of Geniates barbatus Kirby (Coleoptera, Scarabaeidae, Rutelinae). Revista Brasileira de Entomologia 57(1): 40-46.).

The distribution of the Rutelinae species in Bahia has had, in part, direct correlation with the collection trips of the Projeto de Pesquisa em Biodiversidade do Semiárido (PPBio/Ministério da Ciência, Tecnologia, Inovações e Comunicações, MCTIC) since 2005. The ruteline specimens of PPBio Semiarido were deposited in the MZFS and these represent 72.8% (1,064 specimens) of all specimens examined in this study, with 68 species and 5 subspecies. However, although the PPBio has contributed significantly to expand the records of the Rutelinae from Bahia, there are still significant gaps to be explored in the state. The west, southwest and north regions of Bahia present the largest gaps (Figure 1). Certainly, there are many species to be discovered in these regions of the state, which demonstrates the need of a greater collection effort in the regions mentioned above (Oliveira et al. 2016OLIVEIRA, U., PAGLIA, A.P., BRESCOVIT, A.D., CARVALHO, C.J.B., SILVA, D.P., REZENDE, D.T., LEITE, F.S.F., BATISTA, J.A.N., BARBOSA, J.P.P.P., STEHMANN, J.R., ASCHER, J.S., VASCONCELOS, M.R., MARCO JR., P., LÖWENBERG-NETO, PP., DIAS, P.G., FERRO, V.G. & SANTOS, A.J. 2016. The strong influence of collection bias on biodiversity knowledge shortfalls of Brazilian terrestrial biodiversity. Diversity and Distributions 2016: 1-13.). In addition to the mentioned issues, many of the subfamily distribution records in the state are located in areas near the highways that cross Bahia (Figure 1). Oliveira et al. (2016)OLIVEIRA, U., PAGLIA, A.P., BRESCOVIT, A.D., CARVALHO, C.J.B., SILVA, D.P., REZENDE, D.T., LEITE, F.S.F., BATISTA, J.A.N., BARBOSA, J.P.P.P., STEHMANN, J.R., ASCHER, J.S., VASCONCELOS, M.R., MARCO JR., P., LÖWENBERG-NETO, PP., DIAS, P.G., FERRO, V.G. & SANTOS, A.J. 2016. The strong influence of collection bias on biodiversity knowledge shortfalls of Brazilian terrestrial biodiversity. Diversity and Distributions 2016: 1-13. report that species composition decreases as distance from access routes increases, and they suggest that collection localities distant from access routes could increase the possibility of new geographic records as well as records of new species. Most of the records for the south region of Bahia, in Belmonte, Barrolândia, Camacan, Itamaraju, Mucuri, Porto Seguro, Prado and Una, and two localities in the southern region: Encruzilhada and Cândido Sales, are the result of material deposited at MNRJ, CERPE and EPGC (Suplementary Material). Similarly, the collection points located in the north of the state are also from material deposited in MNRJ (examined material). The records for the southern region, Aracatu and Vitória da Conquista (examined material), are deposited in the MFS and were collected by the author.

Among the biomes present in Bahia, the Atlantic Forest has the highest representativity of Rutelinae records, with 32 localities of occurrence: Caatinga with 20 (with the largest collects gap in the southwest, and especially in the north of the state (Figure 1)) and the Cerrado, with only 5 points of records, representing the largest collects gap of all biomes (Figure 1). Atlantic Forest originally accounted for 1,300,000 km² throughout 17 Brazilian states, with 120 million people living in this biome. However, only 22% of the original cover remains, and only 8.5% is in well preserved protection areas, on fragments of over 100 hectares (MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). Caatinga covers 844,453 km², representing 11% of the national territory, and is present in nine states of Northeastern Brazil and in the north of Minas Gerais state (MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). The Caatinga and the Cerrado has been deforested in an accelerated way, especially in the last years, and 46% of the Caatinga total original cover has been deforested (Flores et al. 2012FLORES, M.P., GUIMARÃES, R.F., CARVALHO JÚNIOR, O.A. & GOMES, R.A.T. 2012. Multitemporal Analysis of Agricultural Expansion in the Barreiras Municipality - Bahia. Campo-Território: revista de geografia agrária 7(14): 1-19., MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). Cerrado has been historically transformed in extensive crop plantations in Brazil, causing an increasing reduction of the vegetal cover due to the progressive increase in agricultural economic activity in areas in its domain (Flores et al. 2012FLORES, M.P., GUIMARÃES, R.F., CARVALHO JÚNIOR, O.A. & GOMES, R.A.T. 2012. Multitemporal Analysis of Agricultural Expansion in the Barreiras Municipality - Bahia. Campo-Território: revista de geografia agrária 7(14): 1-19.). This reduction in vegetal cover has, in turn, generated direct consequences for biodiversity loss (Queiroz 2009QUEIROZ, F.A. 2009. Impacts of the international trade of soybeans on the biodiversity of Cerrado Savannah. Sociedade & Natureza, Uberlândia 21 (2): 193-209., MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). However, only 7.6% of the total area of the Caatinga is in the protection area, and only 1% of this area is designated as Units of Integral Protection (MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). In addition, approximately 30% of the Caatinga has already been altered due to human action, and especially due to agriculture. In view of these factors, there is a risk of the rapid loss of unique species in this biome (Araújo et al. 2005ARAÚJO, F.S.; RODAL, M.J.N. & BARBOSA, M.R.V. 2005. Análise das variações da biodiversidade do bioma Caatinga: suporte a estratégias regionais de conservação. Ministério do Meio Ambiente, DF. Brasília.), many of which are yet unknown to science. The Cerrado biome presents the largest collects gap of Rutelinae in Bahia state. Cerrado covers 2,036,448 km² (representing 22% of the national territory), and is present in 12 Brazilian states. It is considered a hotspot of world biodiversity, with an extreme abundance of endemic species (MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). However, the Cerrado is the biome with the lowest percentage of areas of integral protection. Only 8.21% of the total territory is legally protected by Units of Conservation; of this total, 2.85% are Units of Conservation of Integral Protection and 5.36% are Units of Conservation of Sustainable Use, including Particular Reserve of Natural Heritage – RPPNs (0.07%) (MMA 2017MMA–MINISTÉRIO DO MEIO AMBIENTE. 2017. Mata Atlântica. http://www.mma.gov.br/biomas/mata-atlantica (last access at 06/06/2017).
http://www.mma.gov.br/biomas/mata-atlant... ). The findings presented above demonstrate the need to make more collects in the state, especially in Cerrado in the west region of the state, to enable a better understanding of the distribution, expanding the knowledge of the group.

With the results presented here there was an expressive increase, both in the richness of genera and species. After this study the number of genera increased to 23, with the new records of Byrsopolis Burmeister, Pseudodorysthetus Soula and Trizogeniates Ohaus; new distributional records of 30 species and 1 subspecies from Bahia state, and with 14 species, that will be described in subsequent papers, 10 of Leucothyreus (according to the consultation with a researcher, Seidel M., who is conducting the review of the genus), 2 of Lobogeniates (a genus in process of review by ASF) and 1 of Byrsopolis and Pelidnota. With respect to the number of registered species of Rutelinae in Bahia, was increased in more than a half, reaching 101 species and 17 subspecies.

This increase in the number of registered Rutelinae species demonstrates the importance of conducting inventories for group knowledge. In addition, the distribution of the Rutelinae locality records in the Bahia state in only 57 registered municipalities, which is equivalent to 13.67% of the total municipalities in the state, plus the gaps seen in the most richest and poorly known biomes, emphasizes the imminent need for new studies involving the Rutelinae for a better knowledge of fauna, not only in the biomes of this state, but in Brazil as a whole.

Supplementary material

The following online material is available for this article:

Supplementary material

Acknowledgments

The authors would like to thank to the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the master’s fellowship to the first author (ASF), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the research fellowship awarded to LMA (306772/2006-0) and FB (305055/2012-7), Dra. Marcela Monné and Dr. Miguel A. Monné (MNRJ), Dra. Jane Costa and Dr. Márcio Félix (CEIOC–RJ), Dr. Everardo J. Grossi (EPGC), and Dra. Favízia F. Oliveira (MHNBA/MZUFBA) for allowing the study of material, the PPBio Semiarido project (Programa de Pesquisa em Biodiversidade do Semiárido) (agreement CNPq/MCTI – Ministério da Ciência, Tecnologia e Inovação n.º: 457471/2012-3) for financial support for collections and for supporting to visit of Paschoal Grossi visit to the collections at MZFS.

References

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