ARTICLE An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhão, Brazil Lívia Pires do Prado¹⁶; Rodrigo Machado Feitosa²⁷; Stefania Pinzón Triana³⁸; Jhonatan Andrés Muñoz Gutiérrez³⁹; Guillaume Xavier Rousseau³¹⁰; Raimunda Alves Silva⁴¹¹; Glécio Machado Siqueira⁴¹²; Ciro Líbio Caldas dos Santos⁵¹³; Francisco Veras Silva⁵¹⁴; Thiago Sanches Ranzani da Silva²¹⁵; Alexandre Casadei-Ferreira²¹⁶; Rogério Rosa da Silva¹¹⁷ & Joudellys Andrade-Silva¹¹⁸ ¹ Museu Paraense Emílio Goeldi (MPEG), Coordenação de Ciências da Terra e Ecologia (COCTE), Laboratório de Morfologia e Ecologia Funcional de Formigas (AntMor), Programa de Pós-Graduação em Zoologia (PPGZOOL). Belém, PA, Brasil. ² Universidade Federal do Paraná (UFPR), Departamento de Zoologia (DZOO), Laboratório de Sistemática e Biologia de Formigas (LSBF). Curitiba, PR, Brasil. ³ Universidade Estadual do Maranhão (UEMA), Programa de Pós-Graduação em Agroecologia. São Luís, MA, Brasil. ⁴ Universidade Federal do Maranhão (UFMA), Departamento de Geociências (DEGEO). São Luís, MA, Brasil. ⁵ Universidade Federal do Maranhão (UFMA). Imperatriz, MA, Brasil. ⁶ ORCID: http://orcid.org/0000-0003-1819-8767. E-mail: livia.pires7@gmail.com (corresponding author) ⁷ ORCID: http://orcid.org/0000-0001-9042-0129. E-mail: rsmfeitosa@gmail.com ⁸ ORCID: http://orcid.org/0000-0002-7160-0826. E-mail: stefaniapt@gmail.com ⁹ ORCID: http://orcid.org/0000-0002-2090-2226. E-mail: energiaselvatica@gmail.com ¹⁰ ORCID: http://orcid.org/0000-0002-2482-4376. E-mail: guilirous@yahoo.ca ¹¹ ORCID: http://orcid.org/0000-0002-0380-8190. E-mail: ray-234@hotmail.com ¹² ORCID: http://orcid.org/0000-0002-3513-2658. E-mail: gleciosiqueira@hotmail.com ¹³ ORCID: http://orcid.org/0000-0002-0987-3620. E-mail: cirolb@gmail.com ¹⁴ ORCID: http://orcid.org/0000-0003-0057-6467. E-mail: fveras_silva@hotmail.com ¹⁵ ORCID: http://orcid.org/0000-0002-4239-1500. E-mail: tsranzanidasilva@gmail.com ¹⁶ ORCID: http://orcid.org/0000-0002-2977-8348. E-mail: alexandrefrreira@gmail.com ¹⁷ ORCID: http://orcid.org/0000-0002-0599-2155. E-mail: rogeriorosas@gmail.com ¹⁸ ORCID: http://orcid.org/0000-0002-5393-6502. E-mail: joudellys@gmail.com Abstract. The state of Maranhão, located in northeastern Brazil, comprises three biomes: Amazonian, Caatinga, and the Cerrado. To date, 99 ant species have been recorded in the literature from the state. In the present work, we provide for the first time a profile of the ant fauna in the state based on data from the historical literature and Brazilian institutional collections. The updated records on ant diversity for the state of Maranhão revealed a total of 279 species, belonging to 71 genera and 10 subfamilies. In total, 180 species are recorded for the first time in the state, of which four species recorded for the first time in Brazil. In summary, apart from documenting the ant fauna of the region, these results provide a basis for further studies and may contribute to future conservation efforts for the biomes present in this complex landscape. Key-Words. Distribution, Amazon, Caatinga, Cerrado, Checklist. INTRODUCTION Understanding the distribution of species is essential to determine regional and global patterns of biodiversity (Dalzochio et al., 2018). In this sense, taxonomic inventories contribute to characterize areas of endemism, reveal taxonomic novelties and improve scientific collections (Moura et al., 2014; Freitas et al., 2017). Further, the analysis of species distribution databases can Pap. Avulsos Zool., 2019; v.59: e20195938 http://doi.org/10.11606/1807-0205/2019.59.38 http://www.revistas.usp.br/paz http://www.scielo.br/paz Edited by: Helena Carolina Onody Received: 22/05/2019 Accepted: 15/07/2019 Published: 15/08/2019 help to identify gaps in sampling and species records, and can also be used in macroecological studies, species distribution modeling and to promote conservation strategies (Gasper et al., 2016). Maranhão is a northeastern state in Brazil and comprises a total area of 329,642.170 km² (IBGE, 2018). Its political boundaries are the Atlantic Ocean to the north, the state of Tocantins to the south, the state of Piauí to the east, and the state of Pará to the west (Chaves et al., 2016). The state ISSN On-Line: 1807-0205 ISSN Printed: 0031-1049 ISNI: 0000-0004-0384-1825 Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 2/14 is located in a heterogeneous landscape area under the influence of three biomes: Amazon, Cerrado, and the Caatinga. The vegetation cover – encompassing 14 different vegetation types – reflects the transition between super-humid and semi-arid climates (Santos et al., 2010; IBGE, 2018). Similar to other states in Brazil, Maranhão has suffered with high human impact, mainly from the early 1960s, through the construction of highways, agricultural and mining projects (Celentano et al., 2017). Impacts include large-scale forest conversion to pasture or by “babaçu” palm trees (Orbignya phalerata Mart.) (Santos et al., 2010), and the expansion of agroindustry has converted large natural areas into grain crops (Brasil, 2009; Santos et al., 2010). In addition, other human activities, such as occupation, recreation and tourism (Chaves et al., 2016) have also a negative impact and have caused severe loss of biodiversity, resulting in drastic changes of the landscape. The biodiversity of Maranhão is extremely diverse (Chaves et al., 2016; Desidério et al., 2017). Compared to other Brazilian states, however, the ant diversity is poorly known. The most recent information on ant species diversity in the state recorded 99 species, belonging to 37 genera and seven subfamilies (Janicki et al., 2016). This represents about ¼ of ant diversity in the state of Goiás and 35% of the ant species richness described for the state of Mato Grosso do Sul (Janicki et al., 2016), two other Brazilian states comparable in size to Maranhão. Since the end of the 20th century, collective efforts of several research groups, carring out inventories in different areas and employing complementary sampling methodologies, resulted in a significant increase in our knowledge about ant diversity in this state. Thus, the aim of this study is to present an updated list of the ant species in the state of Maranhão, considering recent field expeditions as well as material deposited in the main Brazilian ant collections. We also discuss some relevant aspects about the profile of the ant fauna, recovering the history of ant studies historically carried out in the state. Overall, our findings should be of great help in creating measures for species preservation and species recovery plans and represent the basis for future research. MATERIAL AND METHODS Data from collections and literature We listed material obtained from six Brazilian ant collections (Table 1), which have historically acted as main depositary institutions for samples collected in the state of Maranhão. We also compiled data from literature, including collection events focused on partial surveys of Maranhão ant fauna (Table 2). Identifications and taxonomic validation Ants were identified by the authors of the present study using taxonomic keys, comparing specimens with myrmecological collections, or by sending them to specialists (see “Acknowledgements”). The final list containing all specimens was verified by authors of this study (JAS, LPP and RMF). Species with dubious identification were carefully examined and, when necessary, have been removed from final data set. Table 1. List of ant collections consulted in this study. Repositories Coleção Entomológica, Museu Paraense Emílio Goeldi, Pará, Brazil Laboratório de Biologia, Universidade Federal do Maranhão, Campus Imperatriz, Maranhão, Brazil Laboratório de Entomologia e Vetores, Universidade Federal do Maranhão, Campus São Luís, Maranhão, Brazil Laboratório de Hymenoptera, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil Laboratório de Artrópodes e Microbiologia do Solo, Universidade Estadual do Maranhão, Campus São Luís, Maranhão, Brazil Laboratório de Solos e Meio Ambiente, Universidade Federal do Maranhão, Campus São Luís, Maranhão, Brazil Distribution and maps The biomes present in Maranhão are the Amazon Forest, characterized by tall trees and periodic to permanently flooded plains; this biome is present in the north and, essentially, in the west portion of the state. The Cerrado covers the south, central and northeast areas of the state, formed by open grasslands (Cerrado aberto) to patches of dense vegetation (Cerradão). Finally, Maranhão presents a small and fragmented portion of Table 2. References and their respective areas that have records of ants collected in the state of Maranhão. Context Behavioral Ecological Forensic Genetics Inventory Occurrence Sanitary Taxonomic Source Wheeler, 1922; Monnin et al., 2003; Silva, 2007; Pereira et al., 2014 Brandão et al., 2011; Ramos et al., 2015; Silva et al., 2017; Gutiérrez et al., 2017 Andrade-Silva et al., 2015; Pereira et al., 2017 Wauters et al., 2018 Dáttilo et al., 2012 Dáttilo et al., 2010 Carvalho et al., 2011; Silva et al., 2012; Lima et al., 2013 Forel, 1904; Mann, 1916; Gonçalves, 1942, 1947; Borgmeier, 1955; Kempf, 1959, 1960a, 1960b; Gonçalves, 1961; Kempf, 1964, 1968, 1971, 1972a, 1972b, 1975; Watkins, 1976; Ward, 1989; Brandão, 1991; De Andrade & Baroni Urbani, 1999; Ward, 1999; Cuezzo, 2000; Longino & Snelling, 2002; Ward & Downie, 2005; Shoemaker et al., 2006; Wild & Cuezzo, 2006; Feitosa et al., 2007; Ward, 2007; Wild, 2007; Feitosa et al., 2008; Lattke, 2011; Fernandes et al., 2014; Brandão et al., 2015; Johnson, 2015; Ulysséa et al., 2015, 2017; Jesovnik & Schultz, 2017; Dias & Lattke, 2019 Prado, L.P. et al.: Ants of the state of Maranhão, Brazil the Caatinga biome, in the extreme east of the state, characterized by the presence of bushy vegetation with deep roots, cacti and bromeliads (Spinelli-Araujo et al., 2016). We used shapefiles from the state of Maranhão made available by the Ministério do Meio Ambiente (MMA) (http://mapas.mma.gov.br/i3geo/datadownload.htm#). We used a classification in “meso-regions” pre-established by the government agency, in order to describe and discuss our results. We also used shapefiles provided by MMA for the three main biomes present in the state, to overlap sampling points and the main ecosystems in Maranhão. For the confirmation of sampled sites (Table 3) and maps preparation, the geographical coordinates, when not available on the specimens’ label, were obtained from the IBGE (2011) or georeferenced using Google Earth Pro. In those cases, because we did not have access to the exact point of the sample site, we adapted a classification by the IBGE. Whenever the IBGE classified a municipality covering two biomes, we used the “transition” term after the government classification. For instance, the municipality of Imperatriz, which is classified by IBGE as “Amazon/Cerrado” biomes, becomes for the purpose of this study, “Amazon-Cerrado transition”. For specific sites and localities for which names have been historically altered, we consulted Vanzolini & Papavero (1968) and Vanzolini (1992). The geographical records were mapped using QGIS v2.18.2 (QGIS Development Team, 2019). RESULTS Pap. Avulsos Zool., 2019; v.59: e20195938 3/14 W. Kempf. During the next three decades, collections by researchers, enthusiasts, and professional collectors had pursued the same goal – discovering new taxa and increasing the coverage of ants in scientific collections (Kempf, 1972a). Differently, from the 1980s until the beginning of the 21st century, the main purpose of the expeditions was to carry out environmental impact assessment programs (Brandão et al., 2011). On the other hand, from the late 20th century, with the hiring of researchers at universities in the state of Maranhão, several expeditions have been conducted focusing on ecological studies and reporting faunal inventories (Ramos et al., 2015; Gutiérrez et al., 2017; Silva et al., 2017). Museums, scientific collections, and historical published literature all contain important information on species distributions recorded as presence data (Newbold, 2010). The accuracy of the distribution data is important for several applications in biology and for species conservation planning (Graham et al., 2008). Despite the concern to accurately document of species distribution that began in the first half of the 19th century (Vanzolini, 2004), for the ants this occurred in the second half of the 20th century. In the case of the records analysed in this work, the specific localities and geographic coordinates became available in the late 20th century. Most of the ant records for the state of Maranhão remained unavailable to the specialized public for a long time, while many other records remained unidentified at a specific level. In this sense, our study has analyzed both the material deposited in Brazilian collections (Table 1) and the records in the published literature (Table 2), revealing that 64% of species were recorded in the state for the first time. Further, we made an additional effort Based on data from Brazilian collections (Table 1) and published literature (Table 2), we recorded a total of 279 ant species for the state of Maranhão, belonging to 71 genera and 10 subfamilies (Table 4), and sampled across 65 localities (Table 3). The subfamily Myrmicinae was the most diverse, with 126 species, followed by Ponerinae (36 species), Formicinae (35 species), Dolichoderinae (27 species), Ectatomminae (25 species), Pseudomyrmecinae (16 species), Dorylinae (10 species), Amblyoponinae (2 species), and Paraponerinae and Proceratiinae (1 species each). The majority of records (214 species) was concentrated along the Amazon region, followed by the Cerrado (129 species), the Amazon-Cerrado transition regions (80 species) and finally the Cerrado-Caatinga transition region where only one species was recorded (Fig. 1). A total of 180 ant species were recorded for the first time in the state, and four species were recorded for the first time in Brazil (Table 4). DISCUSSION The first expeditions focused on studying the ant fauna of the state of Maranhão were performed in the late 1940s, with collections in the Cerrado areas undertaken by the myrmecologists Cincinnato Gonçalves and Walter Figure 1. Map of the state of Maranhão showing its location in Brazil. Black points indicate the sampling sites within the state that were georeferenced and recorded in the literature and collections according to biomes. Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 4/14 Table 3. Information from the sampled sites for the state of Maranhão. The abbreviations are as follows: (Am) Amazon, (Ce) Cerrado, (ACT) Amazon-Cerrado transition, (CCT) Cerrado-Caatinga transition. (*) For the geographic coordinates attributed in this work. Locality 15 km E of Canindé, Aldeia Araçu, Igarapé Gurupi-Uma Açailândia Açailândia, Fazenda Pedro Maranhão Açailândia, Horto Fazenda Pompéia Alcântara Alcântara, Só Assim Aldeia do Ponto Alto Turiaçu Alto Turiaçu, Aldeia Gurupiuna Bacabal Balsas Balsas, Córrego Xupé Balsas, Gerais de Balsas Balsas, Gerais de Balsas, Rio Mandacaru Balsas, Mata do Capão do Catulé Barão de Grajaú, Bem Quer Barra do Corda Bom Jardim Buriticupu Buriticupu, Fazenda Cacique Cajazeiras Carolina Carolina, Pedra Caída Caxias Centro Novo do Maranhão Chapadinha, Anapurus Chapadinha, Fazenda Unha de Gato Codó Estreito Estreito, Fazenda Itaueiras Estreito, Fazenda Planalto Estreito, Ilha do Cabral, Rio Tocantins Grajaú, Rio Santana Gurupi Imperatriz Imperatriz, Bananal Imperatriz, Reserva do 50º Batalhão de Infantaria e Selva Imperatriz, Ribeirãozinho Imperatriz, Tocantinópolis Itinga do Maranhão João Lisboa Lago do Junco Lago dos Rodrigues Lago Verde Lagoa Grande do Maranhão Livramento Mirador, Parque Estadual do Mirador Pedreiras Pindaré-Mirim, Sítio Moisés Presidente Dutra Rio Maracaçumé Rosário Santa Inês Santa Luzia São Francisco do Brejão São José de Ribamar, Sítio Aguahy São Luís São Luís, Área de Preservação Ambiental do Itapiracó São Luís, Campus da Universidade Federal do Maranhão São Luís, Maracana, Sítio Mangalho São Luís, Mata do Quebra Pote São Luís, Reserva Alumar São Luís, Reserva da CAEMA São Mateus Zé Doca Coordinate 02°34’S, 46°02’W* 04°52’30”S, 47°17’40”W 04°56’48”S, 47°30’17”W 04°52’30”S, 47°17’40”W 02°20’56”S, 44°29’01”W 02°20’53.48”S, 44°28’50.71”W 06°07’01.21”S, 45°08’59.99”W* 01°39’46”S, 45°22’19”W* 02°51’44.66”S, 46°15’29.79”W* 04°13’30”S, 44°46’48”W* 08°34’19.6”S, 46°42’28.2”W 07°31’58”S, 46°02’09”W 08°34’06”S, 46°42’38”W 08°32’32”S, 46°36’18”W 09°22’53.8”S, 46°44’59.3”W 06°09’24.5”S, 42°58’02.0”W 05°30’21”S, 45°14’34”W* 03°55’32.90”S, 46°46’16.33”W 04°20’45”S, 46°24’03”W* 04°20’34”S, 46°24’06”W* 02°50’43”S, 42°12’20”W* 07°19’58”S, 47°28’08”W* 07°02’30.39”S, 47°26’35.95”W* 04°51’32”S, 43°21’21”W 03°40’55.70”S, 46°46’40.73”W 03°40’19”S, 43°06’57”W 03°41’42.3”S, 43°11’48.1”W 04°27’18”S, 43°53’09”W* 06°50’45”S, 47°23’00”W 06°31’54.4”S, 47°22’16.0”W 06°35’59.3”S, 47°24’50.4”W 06°31’54.1”S, 47°26’32.6”W 05°49’08”S, 46°08’20”W* 04°22’09.04”S, 46°56’16.05”W 05°30’38”S, 47°28’46”W* 05°39’26”S, 47°13’07”W* 05°30’37.60”S, 47°28’46.11”W 05°45’11.15”S, 47°21’36.36”W* 05°31’33”S, 47°28’33”W* 04°02’05.50”S, 46°52’51.00”W* 05°19’46.30”S, 47°19’13.00”W 04°36’32”S, 45°02’56”W* 04°36’32”S, 44°58’48”W* 03°57’25”S, 44°49’19”W* 04°57’38.77”S, 45°16’27.88”W* 02°25’33.19”S, 44°25’26.59”W 06°24’38.62”S, 44°28’42.24”W 04°34’26”S, 44°35’49”W* 03°36’44.00”S, 45°19’59.90”W 05°17’24”S, 44°29’24”W 01°27’09”S, 45°42’19”W 02°51’11.8”S, 44°09’02.6”W 03°40’01”S, 45°22’48”W 04°04’08”S, 45°41’24”W 05°17’19.0”S, 47°15’01.7”W 02°38’59.30”S, 44°08’49.63”W 02°31’48”S, 44°18’10”W* 02°31’28.81”S, 44°12’00.33”W 02°33’36”S, 44°18’33”W 02°36’13.39”S, 44°17’56.78”W 02°42’26.75”S, 44°15’23.87”W 02°42’39.31”S, 44°17’47.40”W 02°34’01.01”S, 44°15’04.03”W 04°02’24”S, 44°28’12”W 03°14’34”S, 45°49’26”W* Physiognomy Am Am Am Am Am Am Ce Am Am ACT Ce Ce Ce Ce Ce CCT Ce Am Am Am Ce Ce Ce Ce Am Ce Ce Ce Ce Ce Ce Ce ACT Am ACT Ce ACT Ce ACT Am Am ACT Ce Am Ce Am Ce Ce Am Ce Am Am Am ACT Am Am Am Am Am Am Am Am Am ACT Am Code 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 5/14 Table 4. List of taxa recorded in the state of Maranhão and the occurrence data of the species in the literature and localities and biome present in the state. The codes of localities follow Table 3. (*) new record for Maranhão, (**) new record for Brazil, (Am) Amazon, (Ce) Cerrado, (ACT) Amazon-Cerrado transition, (CCT) CerradoCaatinga transition. Taxon name Amblyoponinae Forel, 1893* Fulakora Mann, 1919* Fulakora degenerata (Borgmeier, 1957)* Prionopelta Mayr, 1866* Prionopelta antillana Forel, 1909* Dolichoderinae Forel, 1878 Azteca Forel, 1878* Azteca alfari Emery, 1893* Azteca chartifex Emery, 1896* Azteca schimperi Emery, 1893* Dolichoderus Lund, 1831 Dolichoderus abruptus (Smith, 1858)* Dolichoderus attelaboides (Fabricius, 1775)* Dolichoderus bidens (Linnaeus, 1758)* Dolichoderus bispinosus (Olivier, 1792)* Dolichoderus debilis Emery, 1890* Dolichoderus diversus Emery, 1894* Dolichoderus imitator Emery, 1894* Dolichoderus lamellosus (Mayr, 1870) Dolichoderus lutosus (Smith, 1858) Dolichoderus quadridenticulatus (Roger, 1862) Dorymyrmex Mayr, 1866 Dorymyrmex biconis Forel, 1912* Dorymyrmex brunneus Forel, 1908 Dorymyrmex goeldii Forel, 1904* Dorymyrmex jheringi Forel, 1912 Dorymyrmex pyramicus (Roger, 1863) Dorymyrmex spurius Santschi, 1929 Dorymyrmex thoracicus Gallardo, 1916 Forelius Emery, 1888 Forelius brasiliensis (Forel, 1908) Forelius maranhaoensis Cuezzo, 2000 Forelius pusillus Santschi, 1922* Gracilidris Wild & Cuezzo, 2006 Gracilidris pombero Wild & Cuezzo, 2006 Linepithema Mayr, 1866 Linepithema cerradense Wild, 2007 Linepithema neotropicum Wild, 2007 Tapinoma Foerster, 1850* Tapinoma melanocephalum (Fabricius, 1793)* Dorylinae Leach, 1815 Acanthostichus Mayr, 1887* Acanthostichus brevicornis Emery, 1894* Eciton Latreille, 1804 Eciton burchellii (Westwood, 1842)* Eciton mexicanum Roger, 1863* Eciton quadriglume (Haliday, 1836) Eciton rapax Smith, 1855* Labidus Jurine, 1807 Labidus coecus (Latreille, 1802)* Labidus mars (Forel, 1912)* Labidus praedator (Smith, 1858) Neocerapachys Borowiec, 2016* Neocerapachys splendens (Borgmeier, 1957)* Nomamyrmex Borgmeier, 1936* Nomamyrmex esenbeckii (Westwood, 1842)* Ectatomminae Emery, 1895 Ectatomma Smith, 1858 Locality — — 4, 41 — 46 — — 13, 37 57 57 — 8 1, 2, 56 19 5, 24, 35, 37, 39 19 5, 43 5, 13, 31, 40, 52, 57, 50 3, 10, 31, 45 14, 57, 59 5, 24, 45, 57, 64 — 52 5, 13, 34, 40, 52, 57, 59, 61 61 11, 13 11 11, 13 4, 10, 11, 13, 45, 57 — 11, 13 11, 13, 35, 37, 57, 62 13 — 11, 13 — 11 11, 13, 45 — 5, 34, 40, 52, 57, 60 — — 34, 52 — 1, 21, 31, 33 60 1, 35, 45 3, 31 — 5, 11, 13, 14, 37, 41, 52, 56, 57 34 45 — 5, 31, 40 — 23 — — Biome — — Am — Am — — Ce, ACT Am Am — Am Am Am Am, Ce, ACT Am Am, Ce Am, Ce Am, Ce, ACT Am, Ce Am, Ce, ACT — Am Am, Ce Am Ce Ce Ce Am, Ce, ACT — Ce Am, Ce, ACT Ce — Ce — Ce Ce — Am — — Am — Am, Ce, ACT Am Am, Ce, ACT Am, Ce — Am, Ce, ACT Am Ce — Am, Ce — Ce — — Source — — Collection — Collection — — Collection Collection Collection — Collection Collection Collection Collection Collection Collection Collection Kempf, 1972a, 1972b; Collection Andrade-Silva et al., 2015; Collection Kempf, 1972a; Collection — Collection Andrade-Silva et al., 2015; Pereira et al., 2017; Collection Collection Brandão et al., 2011; Collection Brandão et al., 2011; Collection Brandão et al., 2011; Collection Brandão et al., 2011; Collection — Brandão et al., 2011; Collection Cuezzo, 2000; Brandão et al., 2011; Ulysséa et al., 2017; Collection Collection — Wild & Cuezzo, 2006; Brandão et al., 2011; Collection — Brandão et al., 2011; Collection Wild, 2007; Brandão et al., 2011; Collection — Collection — — Collection — Colletion Collection Kempf, 1972a; Watkins, 1976; Collection Collection — Brandão et al., 2011; Collection Collection Borgmeier, 1955; Watkins, 1976; Kempf, 1972a; Collection — Collection — Collection — — Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 6/14 Taxon name Ectatomma brunneum Smith, 1858 Ectatomma edentatum Roger, 1863 Ectatomma lugens Emery, 1894* Ectatomma muticum Mayr, 1870 Ectatomma opaciventre (Roger, 1861) Ectatomma permagnum Forel, 1908* Ectatomma planidens Borgmeier, 1939 Ectatomma ruidum (Roger, 1860)* Ectatomma suzanae Almeida Filho, 1986* Ectatomma tuberculatum (Olivier, 1792) Gnamptogenys Roger, 1863 Gnamptogenys acuminata (Emery, 1896)* Gnamptogenys ammophila Lattke, 1990 Gnamptogenys annulata (Mayr, 1887)* Gnamptogenys caelata Kempf, 1967 Gnamptogenys haenschi (Emery, 1902)* Gnamptogenys horni (Santschi, 1929)* Gnamptogenys lanei Kempf, 1960* Gnamptogenys mina (Brown, 1956)* Gnamptogenys minuta (Emery, 1896) Gnamptogenys moelleri (Forel, 1912)* Gnamptogenys rastrata (Mayr, 1866)* Gnamptogenys striatula Mayr, 1884* Gnamptogenys sulcata (Smith, 1858)* Gnamptogenys triangularis (Mayr, 1887)* Typhlomyrmex Mayr, 1862* Typhlomyrmex rogenhoferi Mayr, 1862* Formicinae Latreille, 1809 Acropyga Roger, 1862* Acropyga goeldii Forel, 1893* Acropyga smithii Forel, 1893* Brachymyrmex Mayr, 1868 Brachymyrmex australis Forel, 1901 Brachymyrmex heeri Forel, 1874* Brachymyrmex patagonicus Mayr, 1868 Camponotus Mayr, 1861 Camponotus arboreus (Smith, 1858) Camponotus atriceps (Smith, 1858) Camponotus balzani Emery, 1894* Camponotus bidens Mayr, 1870* Camponotus blandus (Smith, 1858) Camponotus cameranoi Emery, 1894* Camponotus chartifex (Smith, 1860)* Camponotus crassus Mayr, 1862 Camponotus fastigatus Roger, 1863* Camponotus femoratus Fabricius, 1804)* Camponotus godmani Forel, 1899* Camponotus latangulus Roger, 1863* Camponotus leydigi Forel, 1886 Camponotus melanoticus Emery, 1894 Camponotus novogranadensis Mayr, 1870* Camponotus personatus Emery, 1894 Camponotus rectangularis Emery, 1890* Camponotus renggeri Emery, 1894 Camponotus rufipes (Fabricius, 1775) Camponotus senex (Smith, 1858) Camponotus sexguttatus (Fabricius, 1793)* Camponotus silvestrii Emery, 1906 Camponotus substitutus Emery, 1894* Locality 2, 4, 5, 10, 11, 12, 13, 17, 19, 21, 34, 35, 37, 43, 48, 49, 54, 56, 57, 59 11, 13, 44 3, 19, 34, 40 11, 13, 22, 24, 35, 45 11, 13 35, 37 11, 13 5 37, 56 2, 3, 5, 21, 37, 40, 41, 56, 57, 59 — 31, 40, 57, 58 11, 13 60 34 5, 6, 34, 40 5, 37, 40, 52 40 34, 57 30, 41, 52 30, 41, 52, 57, 63 30 30, 32, 34, 41 34, 37 40 — 1 — — 57 4 — 11, 13 5, 34, 37, 40, 46, 52, 56, 57 11, 13 — 10, 45 9, 29, 37, 38, 53, 57 37 5, 31, 34 5, 11, 17, 24, 28, 31, 34, 35, 40, 43, 56, 57, 59, 65 34 5 5, 10, 11, 30, 31, 35, 37, 40, 45, 52, 56, 57 37, 56 4 5, 34, 60 5, 34, 40, 52, 58 5, 10, 17, 40, 43, 44, 45, 47, 53, 56 37, 56, 57, 59 5, 35, 37, 57 11 62 11, 19, 27, 37, 42, 43, 45, 53, 56, 64 57, 59, 57, 59 58 35 4, 10, 30, 41, 57 Biome Am, Ce, ACT Am, Ce Am Ce, C Ce ACT Ce Am Am, ACT Am, Ce, ACT — Am, Ce Ce Am Am Am Am, ACT Am Am Am, Ce Am, Ce, ACT Ce Ce, ACT Am, ACT Am — Am — — Am Am — Ce Am, ACT Ce — Ce, ACT Am, Ce, ACT AST Am, Ce Am, Ce, ACT Source Kempf, 1972a; Brandão et al., 2011; Dáttilo et al., 2012; Andrade-Silva et al., 2015; Pereira et al., 2017; Collection Brandão et al., 2011; Collection Collection Kempf, 1972a; Brandão et al., 2011; Collection Brandão et al., 2011; Collection Collection Brandão et al., 2011; Collection Collection Collection Andrade-Silva et al., 2015; Collection — Collection Brandão et al., 2011; Collection Collection Dias & Lattke, 2019 Collection Collection Collection Collection Collection Dias & Lattke, 2019 Collection Collection Collection Collection Collection Collection — Collection — — Collection Collection — Brandão et al., 2011; Collection Collection Brandão et al., 2011; Collection — Mann, 1916; Kempf, 1972a; Collection Dáttilo et al., 2012; Collection Collection Collection Brandão et al., 2011; Andrade-Silva et al., 2015; Collection Am Am Am, Ce, ACT Collection — Kempf, 1972a; Brandão et al., 2011; Collection Am Am Am Am Am, Ce, ACT Am, ACT Am, ACT Ce Am Am, Ce, ACT Am Am Am ACT Am, Ce, ACT Collection Collection Collection Collection Kempf, 1972a; Collection Andrade-Silva et al., 2015; Collection Collection Brandão et al., 2011; Collection Collection Kempf, 1972a; Brandão et al., 2011; Collection Andrade-Silva et al., 2015; Pereira et al., 2017 Andrade-Silva et al., 2015 Collection Janicki et al., 2016 Collection Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Taxon name Locality Camponotus tenuiscapus Roger, 1863* 34 Camponotus trapeziceps Forel, 1908* 5, 34, 46 Camponotus trapezoideus Mayr, 1870* 5, 34, 40 Gigantiops Roger, 1863 — Gigantiops destructor (Fabricius, 1804) 12, 13, 14, 45, 57 Nylanderia Emery, 1906* — Nylanderia fulva (Mayr, 1862)* 5, 10, 13, 52, 57 Nylanderia guatemalensis (Forel, 1885)* 34, 40 Paratrechina Motschoulsky, 1863* — Paratrechina longicornis (Latreille, 1802)* 10, 13, 37, 57, 58 Myrmicinae Lepeletier de Saint-Fargeau, 1835 — Acromyrmex Mayr, 1865 — Acromyrmex hystrix (Latreille, 1802) 29 Acromyrmex landolti (Forel, 1885) 11, 45, 57 Acromyrmex laticeps (Emery, 1905)* 22 Acromyrmex nigrosetosus (Forel, 1908) 22, 45 Acromyrmex rugosus (Smith, 1858) 4, 11, 22, 45, 57, 59 Acromyrmex subterraneus (Forel, 1893)* 40 Apterostigma Mayr, 1865* — Apterostigma robustum Emery, 1896* 34, 40, 52 Atta Fabricius, 1804 — Atta cephalotes (Linnaeus, 1758) 45 Atta laevigata (Smith, 1858) 45 Atta opaciceps Borgmeier, 1939* 35, 37, 63 Atta sexdens (Linnaeus, 1758) 11, 31, 45, 57 Basiceros Schulz, 1906 — Basiceros militaris (Weber, 1950) 4, 34, 55 Basiceros scambognathus (Brown, 1949) 30 Blepharidatta Wheeler, 1915 — Blepharidatta conops Kempf, 1967 11, 13, 30, 32 Cardiocondyla Emery, 1869* — Cardiocondyla emeryi Forel, 1881* 10, 13 Cardiocondyla obscurior Wheeler, 1929* 52 Carebara Westwood, 1840* — Carebara arabara Fernández, 2010* 4 Carebara brevipilosa Fernández, 2004* 4 Carebara urichi (Wheeler, 1922)* 34, 40 Cephalotes Latreille, 1802 — Cephalotes atratus (Linnaeus, 1758) 3, 5, 9, 13, 17, 21, 27, 33, 51, 52, 57, 60 Cephalotes clypeatus (Fabricius, 1804) 5, 3, 30 Cephalotes cordatus (Smith, 1853) 45, 57 Cephalotes eduarduli (Forel, 1921)* 10 Cephalotes grandinosus (Smith, 1860)* 63 Cephalotes maculatus (Smith, 1876)* 5, 57 Cephalotes marginatus (Fabricius, 1804) 38 Cephalotes minutus (Fabricius, 1804) 5, 38, 40, 41, 52, 61 Cephalotes pavonii (Latreille, 1809) 11, 13, 45, 52 Cephalotes pilosus (Emery, 1896)* 35, 43, 64 Cephalotes pusillus (Klug, 1824) 1, 10, 11, 13, 17, 29, 31, 34, 35, 37, 43, 45, 57, 61, 64 Cephalotes serraticeps (Smith, 1858) 9 Cephalotes umbraculatus (Fabricius, 1804)* 57 Crematogaster Lund, 1831 — Crematogaster abstinens Forel, 1899* 13, 56 Crematogaster acuta (Fabricius, 1804)* 13 Crematogaster brasiliensis Mayr, 1878* 63 Crematogaster curvispinosa Mayr, 1862* 10 Crematogaster erecta Mayr, 1866 5, 13, 34, 40, 45, 52, 57 Crematogaster evallans Forel, 1907* 13, 57 Crematogaster limata Smith, 1858* 34, 35, 37, 40, 52, 57, 63 Crematogaster tenuicula Forel, 1904* 5, 34, 35, 37, 52, 57, 60 Crematogaster victima Smith, 1858 57, 59 Pap. Avulsos Zool., 2019; v.59: e20195938 7/14 Biome Am Am Am — Am, Ce — Am, Ce, ACT Am — Am, Ce, ACT — — Ce Am, Ce Ce Ce Am, Ce Am — Am — Ce Ce Am, ACT Am, Ce — Am Ce — Ce — Ce, ACT Am — Am Am Am — Am, Ce, ACT Am, Ce Am, Ce ACT Am Am Ce Am, Ce, ACT Am, Ce Ce, ACT Am, Ce, ACT Source Collection Collection Collection — Forel, 1904; Wheeler, 1922; Kempf, 1972a; Collection — Collection Collection — Collection — — Dáttilo et al., 2010 Gonçalves, 1961; Kempf, 1972a; Brandão et al., 2011; Collection Collection Gonçalves, 1961; Kempf, 1972a; Collection Gonçalves, 1961; Brandão et al., 2011; Andrade-Silva et al., 2015; Collection Collection — Collection — Kempf, 1972a; Collection Kempf, 1972a; Collection Collection Gonçalves, 1942, 1947; Kempf, 1972a; Brandão et al., 2011; Collection — Janicki et al., 2016; Collection Feitosa et al., 2007; Collection — Silva, 2007; Brandão et al., 2011; Pereira et al., 2014; Brandão et al., 2015; Collection — Collection Collection — Collection Collection Collection — De Andrade & Baroni Urbani, 1999; Brandão et al., 2011; Collection De Andrade & Baroni Urbani, 1999; Collection Kempf, 1972a; Kempf, 1960a; Brandão, 1991; Collection Collection Collection Collection De Andrade & Baroni Urbani, 1999; Collection De Andrade & Baroni Urbani, 1999; Collection Kempf, 1972a; Kempf, 1960a; Brandão, 1991; Collection Collection Kempf, 1972a; Kempf, 1960a; Brandão, 1991; De Andrade & Baroni Urbani, 1999; Collection Am Am — Am, Ce Ce Am ACT Am, Ce Am, Ce Am, ACT Am, ACT Am De Andrade & Baroni Urbani, 1999; Collection Collection — Collection Collection Collection Collection Kempf, 1968; Kempf, 1972a; Collection Collection Collection Collection Andrade-Silva et al. 2015; Collection Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 8/14 Taxon name Cyphomyrmex Mayr, 1862* Cyphomyrmex laevigatus Weber, 1938* Cyphomyrmex major Forel, 1901* Cyphomyrmex minutus Mayr, 1862* Cyphomyrmex peltatus Kempf, 1966* Cyphomyrmex transversus Emery, 1894* Daceton Perty, 1833* Daceton armigerum (Latreille, 1802)* Hylomyrma Forel, 1912* Hylomyrma balzani (Emery, 1894)* Hylomyrma immanis Kempf, 1973* Hylomyrma longiscapa Kempf, 1961* Hylomyrma praepotens Kempf, 1973* Hylomyrma reginae Kutter, 1977* Megalomyrmex Forel, 1885* Megalomyrmex drifti Kempf, 1961* Monomorium Mayr, 1855* Monomorium floricola (Jerdon, 1851)* Monomorium pharaonis (Linnaeus, 1758)* Mycetarotes Emery, 1913* Mycetarotes parallelus (Emery, 1906)* Mycocepurus Forel, 1893 Mycocepurus goeldii (Forel, 1893) Mycocepurus smithii (Forel, 1893)* Nesomyrmex Wheeler, 1910 Nesomyrmex asper (Mayr, 1887)* Nesomyrmex brasiliensis (Kempf, 1958) Nesomyrmex spininodis (Mayr, 1887)* Nesomyrmex wilda (Smith, 1943)* Ochetomyrmex Mayr, 1878 Ochetomyrmex neopolitus Fernández, 2003* Ochetomyrmex semipolitus Mayr, 1878 Octostruma Forel, 1912 Octostruma balzani (Emery, 1894) Octostruma iheringi (Emery, 1888)* Oxyepoecus Santschi, 1926* Oxyepoecus vezenyii (Forel, 1907)* Pheidole Westwood, 1839 Pheidole allarmata Wilson, 2003* Pheidole diligens (Smith, 1858)* Pheidole dolon Wilson, 2003** Pheidole fallax Mayr, 1870* Pheidole fimbriata Roger, 1863* Pheidole flavens Roger, 1863 Pheidole fracticeps Wilson, 2003* Pheidole gauthieri Forel, 1901* Pheidole impressa Mayr, 1870* Pheidole jeannei Wilson, 2003* Pheidole microps Wilson, 2003** Pheidole midas Wilson, 2003* Pheidole obscurithorax Naves, 1985 Pheidole radoszkowskii Mayr, 1884 Pheidole scolioceps Wilson, 2003* Pheidole sensitiva Borgmeier, 1959** Pheidole susannae Forel, 1886* Pheidole synarmata Wilson, 2003 Pheidole transversostriata Mayr, 1887* Pogonomyrmex Mayr, 1868 Pogonomyrmex naegelii Emery, 1878 Procryptocerus Emery, 1887 Procryptocerus goeldii Forel, 1899 Procryptocerus hylaeus Kempf, 1951 Locality — 4, 40 5, 40 30, 31 4, 5, 30, 34, 37, 40, 57, 60 5, 37, 40, 62 — 4, 19 — 34, 37, 40, 57 4 4 55 55 — 60 — 5, 40, 43, 52 37, 57 — 57 — 11, 37, 52, 56, 57 4, 63 — 52 11, 13 5, 52 60 — 4, 37 11, 12, 13, 45 — 30, 45, 60 34, 40 — 15 — 34, 52 37 34 5, 37 4 45 31 34 10, 57 37 41 41, 52 29 5, 34, 35, 37, 40, 52, 56, 57, 59 4, 37 30 37 57, 59 37 — 14 — 45, 10, 45 Biome — Am Am Ce Am, Ce, ACT Am, ACT — Am — Am, ACT Am Am Am Am — Am — Am, Ce Am, ACT — Am — Am, Ce, ACT Am — Am Ce Am Am — Am, ACT Ce — Am, Ce Am — Ce — Am ACT Am Am, ACT Am Ce Ce Am Am, ACT ACT Am Am ACT Am, ACT Am, ACT Ce ACT Am ACT — Ce — Sa Ce, ACT Source — Collection Collection Collection Collection Collection — Collection — Collection Collection Collection Collection Collection — Collection — Collection Collection — Collection — Brandão et al., 2011; Collection Collection — Collection Brandão et al., 2011; Collection Collection Collection — Collection Brandão et al., 2011; Collection — Janicki et al., 2016; Collection Collection — Collection — Collection Collection Collection Collection Collection Janicki et al., 2016; Collection Collection Collection Collection Collection Collection Collection Dáttilo et al., 2012 Andrade-Silva et al., 2015; Collection Collection Collection Collection Andrade-Silva et al., 2015; Pereira et al., 2017; Collection Collection — Johnson, 2015 — Kempf, 1972a Longino & Snelling, 2002; Collection Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Taxon name Locality Procryptocerus pictipes Emery, 1896* 5, 40 Rogeria Emery, 1894 — Rogeria alzatei Kugler, 1994* 5, 34, 52, 57 Rogeria besucheti Kugler, 1994* 40 Rogeria germaini Emery, 1894* 40 Rogeria lirata Kugler, 1994* 58 Rogeria scobinata Kugler, 1994 11 Sericomyrmex Mayr, 1865 — Sericomyrmex mayri Forel, 1912 18, 29, 35, 37, 45 Sericomyrmex parvulus Forel, 1912 18, 25, 45 Sericomyrmex saussurei Emery, 1894 18, 29, 45 Solenopsis Westwood, 1840 — Solenopsis geminata (Fabricius, 1804) 5, 34, 37, 52, 57, 58 Solenopsis globularia (Smith, 1858) 5, 45, 52, 57, 59, 61 Solenopsis pollux Forel, 1893* 37 Solenopsis saevissima (Smith, 1855) 37, 57, 59, 61, 65 Solenopsis substituta Santschi, 1925* 57, 61 Solenopsis virulens (Smith, 1858)* 5, 34, 40, 52 Stegomyrmex Emery, 1912 — Stegomyrmex olindae Feitosa, Brandão & Diniz, 2008 2, 29, 45 Strumigenys Smith, 1860* — Strumigenys alberti Forel, 1893* 4 Strumigenys crassicornis Mayr, 1887* 30, 32, 55 Strumigenys denticulata Mayr, 1887* 5, 13, 30, 31, 34, 40, 52, 55 Strumigenys eggersi Emery, 1890* 4, 30, 31, 32, 41, 60 Strumigenys elongata Roger, 1863* 4, 13, 30, 31, 41 Strumigenys gytha Bolton, 2000* 41 Strumigenys hyphata (Brown, 1953)* 4, 30 Strumigenys metopia (Brown, 1959)* 4 Strumigenys mirabilis Mann, 1926* 4 Strumigenys orchibia (Brown, 1953)** 60 Strumigenys schmalzi Emery, 1906* 30, 31 Strumigenys subedentata Mayr, 1887* 30, 31, 40 Strumigenys trudifera Kempf & Brown, 1969* 4, 5, 40 Strumigenys urrhobia (Bolton, 2000)* 56 Strumigenys villiersi (Perrault, 1986)* 30 Strumigenys zeteki (Brown, 1959)* 12 Tetramorium Mayr, 1855* — Tetramorium simillimum (Smith, 1851) 10 Trachymyrmex Forel, 1893 — Trachymyrmex bugnioni (Forel, 1912) 11, 30 Trachymyrmex relictus Borgmeier, 1934* 34, 35, 37, 40, 52, 57, 61 Tranopelta Mayr, 1866* Tranopelta gilva Mayr, 1866* 5, 57 Wasmannia Forel, 1893 — Wasmannia auropunctata (Roger, 1863) 5, 11, 12, 13, 15, 30, 31, 32, 34, 35, 37, 40, 52, 55, 57, 61 Paraponerinae Emery, 1901 — Paraponera Smith, 1858 — Paraponera clavata (Fabricius, 1775) 2, 3, 7, 19, 36, 38, 42, 45, 50 Ponerinae Lepeletier de Saint-Fargeau, 1835 — Anochetus Mayr, 1861* — Anochetus bispinosus (Smith, 1858)* 5, 34 Anochetus diegensis Forel, 1912* 5, 34, 40, 57, 60 Anochetus horridus Kempf, 1964* 4, 27, 37 Anochetus mayri Emery, 1884* 5, 34, 52, 57 Centromyrmex Mayr, 1866* — Centromyrmex brachycola (Roger, 1861)* 5, 34, 57, 61 Cryptopone Emery, 1893* — Cryptopone guianensis (Weber, 1939)* 34 Dinoponera Roger, 1861 — Pap. Avulsos Zool., 2019; v.59: e20195938 9/14 Biome Am — Am Am Am Am Ce — Am, Ce, ACT Am, Ce Am, Ce — Am, ACT Am, Ce ACT Am, ACT Am Am — Am, Ce — Am Am, Ce Am, Ce Am, Ce, ACT Am, Ce Am Am, Ce Am Am Am Ce Am, Ce Am Am Ce Ce — ACT — Ce Am — Am — Am, Ce, ACT Source Collection — Collection Collection Collection Collection Brandão et al., 2011; Collection — Jesovnik & Schultz, 2017; Collection Jesovnik & Schultz, 2017; Collection Jesovnik & Schultz, 2017; Collection — Wauters et al., 2018; Collection Kempf, 1972a; Andrade-Silva et al., 2015; Collection Collection Shoemaker et al., 2006; Andrade-Silva et al., 2015; Pereira et al., 2017; Collection Collection Collection — Feitosa et al., 2008; Ulysséa et al., 2015; Collection — Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection Collection — Kempf, 1972a, 1975; Brandão, 1991; Collection — Brandão et al., 2011; Collection Collection — Collection — Brandão et al., 2011; Collection — — Am, Ce, ACT — — Am Am Am, Ce, ACT Am — Am — Am — — Ward & Downie, 2005; Ward, 2007; Collection — — Collection Collection Collection Collection — Collection — Collection — Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Pap. Avulsos Zool., 2019; v.59: e20195938 10/14 Taxon name Dinoponera gigantea (Perty, 1833) Hypoponera Santschi, 1938* Hypoponera distinguenda (Emery, 1890)* Hypoponera opacior (Forel, 1893)* Hypoponera trigona (Mayr, 1887)* Leptogenys Roger, 1861 Leptogenys guianensis Wheeler, 1923* Leptogenys unistimulosa Roger, 1863 Mayaponera Schmidt & Shattuck, 2014* Mayaponera constricta (Mayr, 1884)* Neoponera Emery, 1901 Neoponera commutata (Roger, 1860) Neoponera marginata (Roger, 1861)* Neoponera striatinodis (Emery, 1890)* Neoponera unidentata (Mayr, 1862)* Neoponera verenae Forel, 1922* Neoponera villosa (Fabricius, 1804) Odontomachus Latreille, 1804 Odontomachus bauri Emery, 1892* Odontomachus brunneus (Patton, 1894)* Odontomachus chelifer (Latreille, 1802)* Odontomachus haematodus (Linnaeus, 1758) Odontomachus meinerti Forel, 1905* Odontomachus opaciventris Forel, 1899 Odontomachus scalptus Brown, 1978* Pachycondyla Smith, 1858* Pachycondyla crassinoda (Latreille, 1802)* Pachycondyla harpax (Fabricius, 1804)* Pachycondyla impressa (Roger, 1861)* Pachycondyla lenis Kempf, 1961* Platythyrea Roger, 1863 Platythyrea angusta Forel, 1901 Platythyrea pilosula (Smith, 1858)* Pseudoponera Emery, 1900* Pseudoponera gilberti (Kempf, 1960)* Pseudoponera stigma (Fabricius, 1804)* Rasopone Schmidt & Shattuck, 2014* Rasopone arhuaca (Forel, 1901)* Rasopone ferruginea (Smith, 1858)* Proceratiinae Emery, 1895* Discothyrea Roger, 1863* Discothyrea sexarticulata Borgmeier, 1954* Pseudomyrmecinae Smith, 1952 Pseudomyrmex Lund, 1831 Pseudomyrmex curacaensis (Forel, 1912) Pseudomyrmex elongatus (Mayr, 1870) Pseudomyrmex ethicus (Forel, 1911)* Pseudomyrmex filiformis (Fabricius, 1804)* Pseudomyrmex flavidulus (Smith, 1858) Pseudomyrmex gracilis (Fabricius, 1804) Pseudomyrmex kuenckeli (Emery, 1890) Pseudomyrmex oculatus (Smith, 1855) Pseudomyrmex penetrator (Smith, 1877)* Pseudomyrmex pupa (Forel, 1911)* Pseudomyrmex schuppi (Forel, 1901) Pseudomyrmex sericeus (Mayr, 1870)* Pseudomyrmex tenuis (Fabricius, 1804) Pseudomyrmex tenuissimus (Emery, 1906) Pseudomyrmex termitarius (Smith, 1855) Pseudomyrmex urbanus (Smith, 1877) Locality 1, 11, 13, 33, 45 — 34 61 5, 34, 52, 57 57 30, 63 — 5, 30, 35, 37, 40, 60 — 7, 21, 45, 56 57 5 34 34, 37 11, 13, 19, 60 — 11, 22, 35, 37, 57, 59, 60 22 57, 26 1, 45, 57 34, 52 29 5 — 5, 30, 32, 35, 37, 52, 57, 60 5, 30, 31, 32, 34, 35, 37, 40, 46, 52, 55, 56, 57 5 34 — 45 60 5, 34, 40, 52, 61 41 — 5, 34, 41, 52, 55, 57 5, 30, 31, 34, 40, 55 — — 4, 5 — — 5, 10, 40, 45, 52 10, 45 4 5, 34 11, 13 5, 16, 38, 40, 45, 46, 56, 57, 63 35, 43 5, 34, 40, 45, 56, 57, 63 3 56, 40 52, 57, 59 5, 40 5, 13, 20, 27, 34, 37, 40, 45, 47, 56, 58, 60, 61, 62, 63 5, 10, 45 5, 13, 27, 43, 45, 47 13, 45 Biome Am, Ce, ACT — Am Am Am — Am Am, Ce — Am, Ce, ACT — Am, Ce Am Am Am Am, ACT Am, Ce — Am, Ce, ACT Ce Am, Ce Am, Ce Am ACT Am — Am, Ce, ACT Am, Ce, ACT Source Kempf, 1971, 1972a; Monnin et al., 2003; Brandão et al., 2011; Collection — Collection Collection Collection — Collection Lattke, 2011; Collection — Collection — Kempf, 1959; Kempf, 1972a; Collection Collection Collection Collection Collection Brandão et al., 2011; Fernandes et al., 2014; Collection — Brandão et al., 2011; Andrade-Silva et al., 2015; Collection Collection Collection Janicki et al., 2016; Collection Collection Dáttilo et al. 2012 Collection — Collection Collection Am Am — Ce Am — Am Am — Am, ACT Am, Ce — — Am — — Am, Ce, ACT Ce, ACT Am Am Ce Am, Ce, CCT Am, ACT Am, Ce Am Am Am Am Am, Ce, ACT Collection Collection — Forel, 1904; Kempf, 1964, 1972a; Collection Collection — Collection Collection — Collection Collection — — Collection — — Ward, 1989; Brandão, 1991; Collection Kempf, 1972a; Ward, 1989; Collection Collection Collection Brandão et al., 2011; Collection Janicki et al., 2016; Collection Ward, 1999; Collection Kempf, 1972a; Collection Collection Collection Andrade-Silva et al., 2015; Collection Collection Forel, 1904; Kempf, 1960b, 1972a; Ward & Downie, 2005; Collection Am, Ce, ACT Am, Ce Ce Mann, 1916; Kempf, 1972a; Ward, 1989; Brandão, 1991; Collection Kempf, 1972a; Brandão et al., 2011; Collection Ward, 1989; Brandão, 1991; Collection Prado, L.P. et al.: Ants of the state of Maranhão, Brazil to identify the morphospecies in ant collections. For instance, 73 ant morphospecies, belonging to 31 ant genera and two subfamilies were here identified at the specific level for the first time (Table 4). In our data compilation, we found a number of species that were recorded for the first time in the state of Maranhão, but are widely distributed in Brazil (Janicki et al., 2016), as is the case of Dolichoderus imitator Emery, 1894 and Gnamptogenys striatula Mayr, 1884, among others (Table 4). On the other hand, some hyperdiverse and taxonomically challenging genera, such as Pheidole, had a considerable increase in the number of new records. Of the 19 species of Pheidole known to the state, 12 were recorded for the first time in the state of Maranhão, and three species were recorded for the first time in Brazil. Not surprisingly, the data obtained from the ant literature clearly indicates that taxonomy is the discipline that most contributed to the knowledge of the ant fauna in the state. This is especially true for taxonomic revisions, which deal with large numbers of specimens (e.g., De Andrade & Baroni Urbani, 1999; Lattke, 2011). The high number of taxonomic publications in our survey is justified by the fact that this discipline was the first area of myrmecology to be developed in Brazil, allowing the formation of large repositories. However, although taxonomy is the discipline with the greatest number of published studies in relation to other areas, in the last 20 years the potential of ant fauna data has been explored in different study areas (Table 2). Other factors that have contributed to increasing our knowledge of the ant fauna in the state of Maranhão are online tools, which provide high definition images of species (AntWeb, 2019), taxonomic literature (Bolton, 2019), geographic distribution of ant specimens (Janicki et al., 2016), and general information on ant taxa (AntWiki, 2019). These tools facilitate the identification of specimens and provide a fast and effective access to information. In addition, the improvement and development of collection methodologies (Figueiredo et al., 2013) has made the sampling more efficient. Despite the increased understanding of biodiversity in this region, sampling coverage of ant fauna in Maranhão is strongly irregular (Fig. 1). Our study showed that the Amazon is the better sampled biome and also houses the largest number of species recorded in the state (Table 4). Most collection points are concentrated in the northern region of the state (Fig. 1), which corresponds to the Coastal region of Maranhão, with the highest population density (Chaves et al., 2016), and where the main research centers are located. While the Cerrado, which corresponds to the biome with the highest coverage in the state (64%) (MMA, 2011; Stella, 2011), remains poorly sampled with extremely sparse collections (Fig. 1). In relation to this biome, it is in the southern part of the state where most of the collection points are concentrated, which in most cases came from samples derived from environmental impact assessment programs (e.g., Brandão et al., 2011). The Amazon-Cerrado transition regions are also undersampled in the state, with few records available from Pap. Avulsos Zool., 2019; v.59: e20195938 11/14 taxonomic papers (Kempf, 1972a; Brandão, 1991) and collections. If we want to understand the association between species and forest formations it is essential to characterize species diversity in ecotones, as already observed by other groups (Santos et al., 2010; MaracahipesSantos et al., 2018). The Caatinga biome remains largely unknown in Maranhão, represented in our study by a single record in the Cerrado-Caatinga transition region (Fig. 1). Although the biome presents a small and fragmented spatial coverage (1% of the state territorial area) (Stella, 2011), the scarcity of information about the ant fauna in the Caatinga has also been observed in other regions of Brazil (Santos et al., 1999; Ulysséa & Brandão, 2013; Leal et al., 2017). This result illustrates the need for greater collection effort to understand and preserve biodiversity in the Caatinga and, consequently, in the state of Maranhão. One of the main limitations of the data available to date on the ant fauna in Maranhão was a strong sampling bias, with most samples being collected near the main roads (Fig. 2). This pattern of biased sampling near highways, rivers, coasts, and cities has been reported in several taxonomic groups (Hijmans et al., 2000; Kadmon et al., 2003; Reddy & Dávalos, 2003; Newbold, 2010; Santos & Hoppe, 2018), which is explained by the ease access, researchers’ interest in certain areas or taxa, and limited financial resources. However, further studies are required to reduce this sampling bias by using different collection methodologies and accessing previously unexplored sites. Low levels of sampling in conservation areas of the state were also observed (Fig. 3). Conservation areas (i.e., national parks, ecological stations, extractive reserves, Figure 2. Map of the state of Maranhão emphasizing the main highways and sampling sites of ant species within the state. Pap. Avulsos Zool., 2019; v.59: e20195938 12/14 Prado, L.P. et al.: Ants of the state of Maranhão, Brazil Ulysséa (Museu de Zoologia da Universidade de São Paulo), Phil Ward (University of California) and, Ricardo Vicente (Universidade do Estado de Mato Grosso) for their help with the identification/confirmation of the ant species. ACF and RMF were financed by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (processes 140260/2016-1 and 1302462/2016-3, respectively). LPP, JAS and SPT were financed by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brasil (CAPES) – Finance Code 001. TSRS was financed by CAPES (process 40001016005P5). JAMG and GXR were financed by the Fundação de Amparo à Pesquisa e ao Desenvolvimento Científico e Tecnológico do Maranhão (FAPEMA) - Finance Code 03135/13. REFERENCES Figure 3. Map of the state of Maranhão emphasizing the Priority Conservation Areas and sampling sites of ant species within the state. national forests, biological reserves, among others) are of fundamental importance for biodiversity conservation (Peres, 2005) and preserving ecosystem (Hallmann et al., 2017). To the best of our knowledge, this is the first compilation focused on studying the ant fauna of Maranhão, one of the largest geopolitical regions of Brazil. Our study significantly increase the number of ant species recorded in the state and demonstrates the importance of carrying out planned inventories for a more detailed understanding of the regional ant fauna. Finally, our data provide the baseline information to further explore the ant fauna in Maranhão, to improve current knowledge and to accurately determine the occurrence of several species. CONCLUSION This paper represents an updated record of the ant species occurring in the state of Maranhão, with numbers increasing from 99 to 279 species. Further collection efforts in different biomes are essential for a better understanding of the biodiversity of the state, and for planning long-term conservation action. Ongoing studies on taxonomy, natural history, and ecology are certainly expected to contribute to this. ACKNOWLEDGMENTS We thank the researchers Emília Z. 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