An overview of the ant fauna (Hymenoptera: Formicidae) of the state of Maranhão, Brazil

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 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 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 20^th 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).

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

Table 2
References and their respective areas that have records of ants collected in the state of Maranhão.

Context	Source
Behavioral	Wheeler, 1922; Monnin et al., 2003; Silva, 2007; Pereira et al., 2014
Ecological	Brandão et al., 2011; Ramos et al., 2015; Silva et al., 2017; Gutiérrez et al., 2017
Forensic	Andrade-Silva et al., 2015; Pereira et al., 2017
Genetics	Wauters et al., 2018
Inventory	Dáttilo et al., 2012
Occurrence	Dáttilo et al., 2010
Sanitary	Carvalho et al., 2011; Silva et al., 2012; Lima et al., 2013
Taxonomic	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

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.

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 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).

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	Coordinate	Physiognomy	Code
15 km E of Canindé, Aldeia Araçu, Igarapé Gurupi-Uma	02°34’S, 46°02’W*	Am	1
Açailândia	04°52’30”S, 47°17’40”W	Am	2
Açailândia, Fazenda Pedro Maranhão	04°56’48”S, 47°30’17”W	Am	3
Açailândia, Horto Fazenda Pompéia	04°52’30”S, 47°17’40”W	Am	4
Alcântara	02°20’56”S, 44°29’01”W	Am	5
Alcântara, Só Assim	02°20’53.48”S, 44°28’50.71”W	Am	6
Aldeia do Ponto	06°07’01.21”S, 45°08’59.99”W*	Ce	7
Alto Turiaçu	01°39’46”S, 45°22’19”W*	Am	8
Alto Turiaçu, Aldeia Gurupiuna	02°51’44.66”S, 46°15’29.79”W*	Am	9
Bacabal	04°13’30”S, 44°46’48”W*	ACT	10
Balsas	08°34’19.6”S, 46°42’28.2”W	Ce	11
Balsas, Córrego Xupé	07°31’58”S, 46°02’09”W	Ce	12
Balsas, Gerais de Balsas	08°34’06”S, 46°42’38”W	Ce	13
Balsas, Gerais de Balsas, Rio Mandacaru	08°32’32”S, 46°36’18”W	Ce	14
Balsas, Mata do Capão do Catulé	09°22’53.8”S, 46°44’59.3”W	Ce	15
Barão de Grajaú, Bem Quer	06°09’24.5”S, 42°58’02.0”W	CCT	16
Barra do Corda	05°30’21”S, 45°14’34”W*	Ce	17
Bom Jardim	03°55’32.90”S, 46°46’16.33”W	Am	18
Buriticupu	04°20’45”S, 46°24’03”W*	Am	19
Buriticupu, Fazenda Cacique	04°20’34”S, 46°24’06”W*	Am	20
Cajazeiras	02°50’43”S, 42°12’20”W*	Ce	21
Carolina	07°19’58”S, 47°28’08”W*	Ce	22
Carolina, Pedra Caída	07°02’30.39”S, 47°26’35.95”W*	Ce	23
Caxias	04°51’32”S, 43°21’21”W	Ce	24
Centro Novo do Maranhão	03°40’55.70”S, 46°46’40.73”W	Am	25
Chapadinha, Anapurus	03°40’19”S, 43°06’57”W	Ce	26
Chapadinha, Fazenda Unha de Gato	03°41’42.3”S, 43°11’48.1”W	Ce	27
Codó	04°27’18”S, 43°53’09”W*	Ce	28
Estreito	06°50’45”S, 47°23’00”W	Ce	29
Estreito, Fazenda Itaueiras	06°31’54.4”S, 47°22’16.0”W	Ce	30
Estreito, Fazenda Planalto	06°35’59.3”S, 47°24’50.4”W	Ce	31
Estreito, Ilha do Cabral, Rio Tocantins	06°31’54.1”S, 47°26’32.6”W	Ce	32
Grajaú, Rio Santana	05°49’08”S, 46°08’20”W*	ACT	33
Gurupi	04°22’09.04”S, 46°56’16.05”W	Am	34
Imperatriz	05°30’38”S, 47°28’46”W*	ACT	35
Imperatriz, Bananal	05°39’26”S, 47°13’07”W*	Ce	36
Imperatriz, Reserva do 50º Batalhão de Infantaria e Selva	05°30’37.60”S, 47°28’46.11”W	ACT	37
Imperatriz, Ribeirãozinho	05°45’11.15”S, 47°21’36.36”W*	Ce	38
Imperatriz, Tocantinópolis	05°31’33”S, 47°28’33”W*	ACT	39
Itinga do Maranhão	04°02’05.50”S, 46°52’51.00”W*	Am	40
João Lisboa	05°19’46.30”S, 47°19’13.00”W	Am	41
Lago do Junco	04°36’32”S, 45°02’56”W*	ACT	42
Lago dos Rodrigues	04°36’32”S, 44°58’48”W*	Ce	43
Lago Verde	03°57’25”S, 44°49’19”W*	Am	44
Lagoa Grande do Maranhão	04°57’38.77”S, 45°16’27.88”W*	Ce	45
Livramento	02°25’33.19”S, 44°25’26.59”W	Am	46
Mirador, Parque Estadual do Mirador	06°24’38.62”S, 44°28’42.24”W	Ce	47
Pedreiras	04°34’26”S, 44°35’49”W*	Ce	48
Pindaré-Mirim, Sítio Moisés	03°36’44.00”S, 45°19’59.90”W	Am	49
Presidente Dutra	05°17’24”S, 44°29’24”W	Ce	50
Rio Maracaçumé	01°27’09”S, 45°42’19”W	Am	51
Rosário	02°51’11.8”S, 44°09’02.6”W	Am	52
Santa Inês	03°40’01”S, 45°22’48”W	Am	53
Santa Luzia	04°04’08”S, 45°41’24”W	ACT	54
São Francisco do Brejão	05°17’19.0”S, 47°15’01.7”W	Am	55
São José de Ribamar, Sítio Aguahy	02°38’59.30”S, 44°08’49.63”W	Am	56
São Luís	02°31’48”S, 44°18’10”W*	Am	57
São Luís, Área de Preservação Ambiental do Itapiracó	02°31’28.81”S, 44°12’00.33”W	Am	58
São Luís, Campus da Universidade Federal do Maranhão	02°33’36”S, 44°18’33”W	Am	59
São Luís, Maracana, Sítio Mangalho	02°36’13.39”S, 44°17’56.78”W	Am	60
São Luís, Mata do Quebra Pote	02°42’26.75”S, 44°15’23.87”W	Am	61
São Luís, Reserva Alumar	02°42’39.31”S, 44°17’47.40”W	Am	62
São Luís, Reserva da CAEMA	02°34’01.01”S, 44°15’04.03”W	Am	63
São Mateus	04°02’24”S, 44°28’12”W	ACT	64
Zé Doca	03°14’34”S, 45°49’26”W*	Am	65

RESULTS

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).

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) Cerrado-Caatinga transition.

Taxon name	Locality	Biome	Source
Amblyoponinae Forel, 1893*	—	—	—
Fulakora Mann, 1919*	—	—	—
Fulakora degenerata (Borgmeier, 1957)*	4, 41	Am	Collection
Prionopelta Mayr, 1866*	—	—	—
Prionopelta antillana Forel, 1909*	46	Am	Collection
Dolichoderinae Forel, 1878	—	—	—
Azteca Forel, 1878*	—	—	—
Azteca alfari Emery, 1893*	13, 37	Ce, ACT	Collection
Azteca chartifex Emery, 1896*	57	Am	Collection
Azteca schimperi Emery, 1893*	57	Am	Collection
Dolichoderus Lund, 1831	—	—	—
Dolichoderus abruptus (Smith, 1858)*	8	Am	Collection
Dolichoderus attelaboides (Fabricius, 1775)*	1, 2, 56	Am	Collection
Dolichoderus bidens (Linnaeus, 1758)*	19	Am	Collection
Dolichoderus bispinosus (Olivier, 1792)*	5, 24, 35, 37, 39	Am, Ce, ACT	Collection
Dolichoderus debilis Emery, 1890*	19	Am	Collection
Dolichoderus diversus Emery, 1894*	5, 43	Am, Ce	Collection
Dolichoderus imitator Emery, 1894*	5, 13, 31, 40, 52, 57, 50	Am, Ce	Collection
Dolichoderus lamellosus (Mayr, 1870)	3, 10, 31, 45	Am, Ce, ACT	Kempf, 1972a, 1972b; Collection
Dolichoderus lutosus (Smith, 1858)	14, 57, 59	Am, Ce	Andrade-Silva et al., 2015; Collection
Dolichoderus quadridenticulatus (Roger, 1862)	5, 24, 45, 57, 64	Am, Ce, ACT	Kempf, 1972a; Collection
Dorymyrmex Mayr, 1866	—	—	—
Dorymyrmex biconis Forel, 1912*	52	Am	Collection
Dorymyrmex brunneus Forel, 1908	5, 13, 34, 40, 52, 57, 59, 61	Am, Ce	Andrade-Silva et al., 2015; Pereira et al., 2017; Collection
Dorymyrmex goeldii Forel, 1904*	61	Am	Collection
Dorymyrmex jheringi Forel, 1912	11, 13	Ce	Brandão et al., 2011; Collection
Dorymyrmex pyramicus (Roger, 1863)	11	Ce	Brandão et al., 2011; Collection
Dorymyrmex spurius Santschi, 1929	11, 13	Ce	Brandão et al., 2011; Collection
Dorymyrmex thoracicus Gallardo, 1916	4, 10, 11, 13, 45, 57	Am, Ce, ACT	Brandão et al., 2011; Collection
Forelius Emery, 1888	—	—	—
Forelius brasiliensis (Forel, 1908)	11, 13	Ce	Brandão et al., 2011; Collection
Forelius maranhaoensis Cuezzo, 2000	11, 13, 35, 37, 57, 62	Am, Ce, ACT	Cuezzo, 2000; Brandão et al., 2011; Ulysséa et al., 2017; Collection
Forelius pusillus Santschi, 1922*	13	Ce	Collection
Gracilidris Wild & Cuezzo, 2006	—	—	—
Gracilidris pombero Wild & Cuezzo, 2006	11, 13	Ce	Wild & Cuezzo, 2006; Brandão et al., 2011; Collection
Linepithema Mayr, 1866	—	—	—
Linepithema cerradense Wild, 2007	11	Ce	Brandão et al., 2011; Collection
Linepithema neotropicum Wild, 2007	11, 13, 45	Ce	Wild, 2007; Brandão et al., 2011; Collection
Tapinoma Foerster, 1850*	—	—	—
Tapinoma melanocephalum (Fabricius, 1793)*	5, 34, 40, 52, 57, 60	Am	Collection
Dorylinae Leach, 1815	—	—	—
Acanthostichus Mayr, 1887*	—	—	—
Acanthostichus brevicornis Emery, 1894*	34, 52	Am	Collection
Eciton Latreille, 1804	—	—	—
Eciton burchellii (Westwood, 1842)*	1, 21, 31, 33	Am, Ce, ACT	Colletion
Eciton mexicanum Roger, 1863*	60	Am	Collection
Eciton quadriglume (Haliday, 1836)	1, 35, 45	Am, Ce, ACT	Kempf, 1972a; Watkins, 1976; Collection
Eciton rapax Smith, 1855*	3, 31	Am, Ce	Collection
Labidus Jurine, 1807	—	—	—
Labidus coecus (Latreille, 1802)*	5, 11, 13, 14, 37, 41, 52, 56, 57	Am, Ce, ACT	Brandão et al., 2011; Collection
Labidus mars (Forel, 1912)*	34	Am	Collection
Labidus praedator (Smith, 1858)	45	Ce	Borgmeier, 1955; Watkins, 1976; Kempf, 1972a; Collection
Neocerapachys Borowiec, 2016*	—	—	—
Neocerapachys splendens (Borgmeier, 1957)*	5, 31, 40	Am, Ce	Collection
Nomamyrmex Borgmeier, 1936*	—	—	—
Nomamyrmex esenbeckii (Westwood, 1842)*	23	Ce	Collection
Ectatomminae Emery, 1895	—	—	—
Ectatomma Smith, 1858	—	—	—
Ectatomma brunneum Smith, 1858	2, 4, 5, 10, 11, 12, 13, 17, 19, 21, 34, 35, 37, 43, 48, 49, 54, 56, 57, 59	Am, Ce, ACT	Kempf, 1972a; Brandão et al., 2011; Dáttilo et al., 2012; Andrade-Silva et al., 2015; Pereira et al., 2017; Collection
Ectatomma edentatum Roger, 1863	11, 13, 44	Am, Ce	Brandão et al., 2011; Collection
Ectatomma lugens Emery, 1894*	3, 19, 34, 40	Am	Collection
Ectatomma muticum Mayr, 1870	11, 13, 22, 24, 35, 45	Ce, C	Kempf, 1972a; Brandão et al., 2011; Collection
Ectatomma opaciventre (Roger, 1861)	11, 13	Ce	Brandão et al., 2011; Collection
Ectatomma permagnum Forel, 1908*	35, 37	ACT	Collection
Ectatomma planidens Borgmeier, 1939	11, 13	Ce	Brandão et al., 2011; Collection
Ectatomma ruidum (Roger, 1860)*	5	Am	Collection
Ectatomma suzanae Almeida Filho, 1986*	37, 56	Am, ACT	Collection
Ectatomma tuberculatum (Olivier, 1792)	2, 3, 5, 21, 37, 40, 41, 56, 57, 59	Am, Ce, ACT	Andrade-Silva et al., 2015; Collection
Gnamptogenys Roger, 1863	—	—	—
Gnamptogenys acuminata (Emery, 1896)*	31, 40, 57, 58	Am, Ce	Collection
Gnamptogenys ammophila Lattke, 1990	11, 13	Ce	Brandão et al., 2011; Collection
Gnamptogenys annulata (Mayr, 1887)*	60	Am	Collection
Gnamptogenys caelata Kempf, 1967	34	Am	Dias & Lattke, 2019 Collection
Gnamptogenys haenschi (Emery, 1902)*	5, 6, 34, 40	Am	Collection
Gnamptogenys horni (Santschi, 1929)*	5, 37, 40, 52	Am, ACT	Collection
Gnamptogenys lanei Kempf, 1960*	40	Am	Collection
Gnamptogenys mina (Brown, 1956)*	34, 57	Am	Collection
Gnamptogenys minuta (Emery, 1896)	30, 41, 52	Am, Ce	Dias & Lattke, 2019 Collection
Gnamptogenys moelleri (Forel, 1912)*	30, 41, 52, 57, 63	Am, Ce, ACT	Collection
Gnamptogenys rastrata (Mayr, 1866)*	30	Ce	Collection
Gnamptogenys striatula Mayr, 1884*	30, 32, 34, 41	Ce, ACT	Collection
Gnamptogenys sulcata (Smith, 1858)*	34, 37	Am, ACT	Collection
Gnamptogenys triangularis (Mayr, 1887)*	40	Am	Collection
Typhlomyrmex Mayr, 1862*	—	—	—
Typhlomyrmex rogenhoferi Mayr, 1862*	1	Am	Collection
Formicinae Latreille, 1809	—	—	—
Acropyga Roger, 1862*	—	—	—
Acropyga goeldii Forel, 1893*	57	Am	Collection
Acropyga smithii Forel, 1893*	4	Am	Collection
Brachymyrmex Mayr, 1868	—	—	—
Brachymyrmex australis Forel, 1901	11, 13	Ce	Brandão et al., 2011; Collection
Brachymyrmex heeri Forel, 1874*	5, 34, 37, 40, 46, 52, 56, 57	Am, ACT	Collection
Brachymyrmex patagonicus Mayr, 1868	11, 13	Ce	Brandão et al., 2011; Collection
Camponotus Mayr, 1861	—	—	—
Camponotus arboreus (Smith, 1858)	10, 45	Ce, ACT	Mann, 1916; Kempf, 1972a; Collection
Camponotus atriceps (Smith, 1858)	9, 29, 37, 38, 53, 57	Am, Ce, ACT	Dáttilo et al., 2012; Collection
Camponotus balzani Emery, 1894*	37	AST	Collection
Camponotus bidens Mayr, 1870*	5, 31, 34	Am, Ce	Collection
Camponotus blandus (Smith, 1858)	5, 11, 17, 24, 28, 31, 34, 35, 40, 43, 56, 57, 59, 65	Am, Ce, ACT	Brandão et al., 2011; Andrade-Silva et al., 2015; Collection
Camponotus cameranoi Emery, 1894*	34	Am	Collection
Camponotus chartifex (Smith, 1860)*	5	Am	—
Camponotus crassus Mayr, 1862	5, 10, 11, 30, 31, 35, 37, 40, 45, 52, 56, 57	Am, Ce, ACT	Kempf, 1972a; Brandão et al., 2011; Collection
Camponotus fastigatus Roger, 1863*	37, 56	Am	Collection
Camponotus femoratus Fabricius, 1804)*	4	Am	Collection
Camponotus godmani Forel, 1899*	5, 34, 60	Am	Collection
Camponotus latangulus Roger, 1863*	5, 34, 40, 52, 58	Am	Collection
Camponotus leydigi Forel, 1886	5, 10, 17, 40, 43, 44, 45, 47, 53, 56	Am, Ce, ACT	Kempf, 1972a; Collection
Camponotus melanoticus Emery, 1894	37, 56, 57, 59	Am, ACT	Andrade-Silva et al., 2015; Collection
Camponotus novogranadensis Mayr, 1870*	5, 35, 37, 57	Am, ACT	Collection
Camponotus personatus Emery, 1894	11	Ce	Brandão et al., 2011; Collection
Camponotus rectangularis Emery, 1890*	62	Am	Collection
Camponotus renggeri Emery, 1894	11, 19, 27, 37, 42, 43, 45, 53, 56, 64	Am, Ce, ACT	Kempf, 1972a; Brandão et al., 2011; Collection
Camponotus rufipes (Fabricius, 1775)	57, 59,	Am	Andrade-Silva et al., 2015; Pereira et al., 2017
Camponotus senex (Smith, 1858)	57, 59	Am	Andrade-Silva et al., 2015
Camponotus sexguttatus (Fabricius, 1793)*	58	Am	Collection
Camponotus silvestrii Emery, 1906	35	ACT	Janicki et al., 2016
Camponotus substitutus Emery, 1894*	4, 10, 30, 41, 57	Am, Ce, ACT	Collection
Camponotus tenuiscapus Roger, 1863*	34	Am	Collection
Camponotus trapeziceps Forel, 1908*	5, 34, 46	Am	Collection
Camponotus trapezoideus Mayr, 1870*	5, 34, 40	Am	Collection
Gigantiops Roger, 1863	—	—	—
Gigantiops destructor (Fabricius, 1804)	12, 13, 14, 45, 57	Am, Ce	Forel, 1904; Wheeler, 1922; Kempf, 1972a; Collection
Nylanderia Emery, 1906*	—	—	—
Nylanderia fulva (Mayr, 1862)*	5, 10, 13, 52, 57	Am, Ce, ACT	Collection
Nylanderia guatemalensis (Forel, 1885)*	34, 40	Am	Collection
Paratrechina Motschoulsky, 1863*	—	—	—
Paratrechina longicornis (Latreille, 1802)*	10, 13, 37, 57, 58	Am, Ce, ACT	Collection
Myrmicinae Lepeletier de Saint-Fargeau, 1835	—	—	—
Acromyrmex Mayr, 1865	—	—	—
Acromyrmex hystrix (Latreille, 1802)	29	Ce	Dáttilo et al., 2010
Acromyrmex landolti (Forel, 1885)	11, 45, 57	Am, Ce	Gonçalves, 1961; Kempf, 1972a; Brandão et al., 2011; Collection
Acromyrmex laticeps (Emery, 1905)*	22	Ce	Collection
Acromyrmex nigrosetosus (Forel, 1908)	22, 45	Ce	Gonçalves, 1961; Kempf, 1972a; Collection
Acromyrmex rugosus (Smith, 1858)	4, 11, 22, 45, 57, 59	Am, Ce	Gonçalves, 1961; Brandão et al., 2011; Andrade-Silva et al., 2015; Collection
Acromyrmex subterraneus (Forel, 1893)*	40	Am	Collection
Apterostigma Mayr, 1865*	—	—	—
Apterostigma robustum Emery, 1896*	34, 40, 52	Am	Collection
Atta Fabricius, 1804	—	—	—
Atta cephalotes (Linnaeus, 1758)	45	Ce	Kempf, 1972a; Collection
Atta laevigata (Smith, 1858)	45	Ce	Kempf, 1972a; Collection
Atta opaciceps Borgmeier, 1939*	35, 37, 63	Am, ACT	Collection
Atta sexdens (Linnaeus, 1758)	11, 31, 45, 57	Am, Ce	Gonçalves, 1942, 1947; Kempf, 1972a; Brandão et al., 2011; Collection
Basiceros Schulz, 1906	—	—	—
Basiceros militaris (Weber, 1950)	4, 34, 55	Am	Janicki et al., 2016; Collection
Basiceros scambognathus (Brown, 1949)	30	Ce	Feitosa et al., 2007; Collection
Blepharidatta Wheeler, 1915	—	—	—
Blepharidatta conops Kempf, 1967	11, 13, 30, 32	Ce	Silva, 2007; Brandão et al., 2011; Pereira et al., 2014; Brandão et al., 2015; Collection
Cardiocondyla Emery, 1869*	—	—	—
Cardiocondyla emeryi Forel, 1881*	10, 13	Ce, ACT	Collection
Cardiocondyla obscurior Wheeler, 1929*	52	Am	Collection
Carebara Westwood, 1840*	—	—	—
Carebara arabara Fernández, 2010*	4	Am	Collection
Carebara brevipilosa Fernández, 2004*	4	Am	Collection
Carebara urichi (Wheeler, 1922)*	34, 40	Am	Collection
Cephalotes Latreille, 1802	—	—	—
Cephalotes atratus (Linnaeus, 1758)	3, 5, 9, 13, 17, 21, 27, 33, 51, 52, 57, 60	Am, Ce, ACT	De Andrade & Baroni Urbani, 1999; Brandão et al., 2011; Collection
Cephalotes clypeatus (Fabricius, 1804)	5, 3, 30	Am, Ce	De Andrade & Baroni Urbani, 1999; Collection
Cephalotes cordatus (Smith, 1853)	45, 57	Am, Ce	Kempf, 1972a; Kempf, 1960a; Brandão, 1991; Collection
Cephalotes eduarduli (Forel, 1921)*	10	ACT	Collection
Cephalotes grandinosus (Smith, 1860)*	63	Am	Collection
Cephalotes maculatus (Smith, 1876)*	5, 57	Am	Collection
Cephalotes marginatus (Fabricius, 1804)	38	Ce	De Andrade & Baroni Urbani, 1999; Collection
Cephalotes minutus (Fabricius, 1804)	5, 38, 40, 41, 52, 61	Am, Ce, ACT	De Andrade & Baroni Urbani, 1999; Collection
Cephalotes pavonii (Latreille, 1809)	11, 13, 45, 52	Am, Ce	Kempf, 1972a; Kempf, 1960a; Brandão, 1991; Collection
Cephalotes pilosus (Emery, 1896)*	35, 43, 64	Ce, ACT	Collection
Cephalotes pusillus (Klug, 1824)	1, 10, 11, 13, 17, 29, 31, 34, 35, 37, 43, 45, 57, 61, 64	Am, Ce, ACT	Kempf, 1972a; Kempf, 1960a; Brandão, 1991; De Andrade & Baroni Urbani, 1999; Collection
Cephalotes serraticeps (Smith, 1858)	9	Am	De Andrade & Baroni Urbani, 1999; Collection
Cephalotes umbraculatus (Fabricius, 1804)*	57	Am	Collection
Crematogaster Lund, 1831	—	—	—
Crematogaster abstinens Forel, 1899*	13, 56	Am, Ce	Collection
Crematogaster acuta (Fabricius, 1804)*	13	Ce	Collection
Crematogaster brasiliensis Mayr, 1878*	63	Am	Collection
Crematogaster curvispinosa Mayr, 1862*	10	ACT	Collection
Crematogaster erecta Mayr, 1866	5, 13, 34, 40, 45, 52, 57	Am, Ce	Kempf, 1968; Kempf, 1972a; Collection
Crematogaster evallans Forel, 1907*	13, 57	Am, Ce	Collection
Crematogaster limata Smith, 1858*	34, 35, 37, 40, 52, 57, 63	Am, ACT	Collection
Crematogaster tenuicula Forel, 1904*	5, 34, 35, 37, 52, 57, 60	Am, ACT	Collection
Crematogaster victima Smith, 1858	57, 59	Am	Andrade-Silva et al. 2015; Collection
Cyphomyrmex Mayr, 1862*	—	—	—
Cyphomyrmex laevigatus Weber, 1938*	4, 40	Am	Collection
Cyphomyrmex major Forel, 1901*	5, 40	Am	Collection
Cyphomyrmex minutus Mayr, 1862*	30, 31	Ce	Collection
Cyphomyrmex peltatus Kempf, 1966*	4, 5, 30, 34, 37, 40, 57, 60	Am, Ce, ACT	Collection
Cyphomyrmex transversus Emery, 1894*	5, 37, 40, 62	Am, ACT	Collection
Daceton Perty, 1833*	—	—	—
Daceton armigerum (Latreille, 1802)*	4, 19	Am	Collection
Hylomyrma Forel, 1912*	—	—	—
Hylomyrma balzani (Emery, 1894)*	34, 37, 40, 57	Am, ACT	Collection
Hylomyrma immanis Kempf, 1973*	4	Am	Collection
Hylomyrma longiscapa Kempf, 1961*	4	Am	Collection
Hylomyrma praepotens Kempf, 1973*	55	Am	Collection
Hylomyrma reginae Kutter, 1977*	55	Am	Collection
Megalomyrmex Forel, 1885*	—	—	—
Megalomyrmex drifti Kempf, 1961*	60	Am	Collection
Monomorium Mayr, 1855*	—	—	—
Monomorium floricola (Jerdon, 1851)*	5, 40, 43, 52	Am, Ce	Collection
Monomorium pharaonis (Linnaeus, 1758)*	37, 57	Am, ACT	Collection
Mycetarotes Emery, 1913*	—	—	—
Mycetarotes parallelus (Emery, 1906)*	57	Am	Collection
Mycocepurus Forel, 1893	—	—	—
Mycocepurus goeldii (Forel, 1893)	11, 37, 52, 56, 57	Am, Ce, ACT	Brandão et al., 2011; Collection
Mycocepurus smithii (Forel, 1893)*	4, 63	Am	Collection
Nesomyrmex Wheeler, 1910	—	—	—
Nesomyrmex asper (Mayr, 1887)*	52	Am	Collection
Nesomyrmex brasiliensis (Kempf, 1958)	11, 13	Ce	Brandão et al., 2011; Collection
Nesomyrmex spininodis (Mayr, 1887)*	5, 52	Am	Collection
Nesomyrmex wilda (Smith, 1943)*	60	Am	Collection
Ochetomyrmex Mayr, 1878	—	—	—
Ochetomyrmex neopolitus Fernández, 2003*	4, 37	Am, ACT	Collection
Ochetomyrmex semipolitus Mayr, 1878	11, 12, 13, 45	Ce	Brandão et al., 2011; Collection
Octostruma Forel, 1912	—	—	—
Octostruma balzani (Emery, 1894)	30, 45, 60	Am, Ce	Janicki et al., 2016; Collection
Octostruma iheringi (Emery, 1888)*	34, 40	Am	Collection
Oxyepoecus Santschi, 1926*	—	—	—
Oxyepoecus vezenyii (Forel, 1907)*	15	Ce	Collection
Pheidole Westwood, 1839	—	—	—
Pheidole allarmata Wilson, 2003*	34, 52	Am	Collection
Pheidole diligens (Smith, 1858)*	37	ACT	Collection
Pheidole dolon Wilson, 2003**	34	Am	Collection
Pheidole fallax Mayr, 1870*	5, 37	Am, ACT	Collection
Pheidole fimbriata Roger, 1863*	4	Am	Collection
Pheidole flavens Roger, 1863	45	Ce	Janicki et al., 2016; Collection
Pheidole fracticeps Wilson, 2003*	31	Ce	Collection
Pheidole gauthieri Forel, 1901*	34	Am	Collection
Pheidole impressa Mayr, 1870*	10, 57	Am, ACT	Collection
Pheidole jeannei Wilson, 2003*	37	ACT	Collection
Pheidole microps Wilson, 2003**	41	Am	Collection
Pheidole midas Wilson, 2003*	41, 52	Am	Collection
Pheidole obscurithorax Naves, 1985	29	ACT	Dáttilo et al., 2012
Pheidole radoszkowskii Mayr, 1884	5, 34, 35, 37, 40, 52, 56, 57, 59	Am, ACT	Andrade-Silva et al., 2015; Collection
Pheidole scolioceps Wilson, 2003*	4, 37	Am, ACT	Collection
Pheidole sensitiva Borgmeier, 1959**	30	Ce	Collection
Pheidole susannae Forel, 1886*	37	ACT	Collection
Pheidole synarmata Wilson, 2003	57, 59	Am	Andrade-Silva et al., 2015; Pereira et al., 2017; Collection
Pheidole transversostriata Mayr, 1887*	37	ACT	Collection
Pogonomyrmex Mayr, 1868	—	—	—
Pogonomyrmex naegelii Emery, 1878	14	Ce	Johnson, 2015
Procryptocerus Emery, 1887	—	—	—
Procryptocerus goeldii Forel, 1899	45,	Sa	Kempf, 1972a
Procryptocerus hylaeus Kempf, 1951	10, 45	Ce, ACT	Longino & Snelling, 2002; Collection
Procryptocerus pictipes Emery, 1896*	5, 40	Am	Collection
Rogeria Emery, 1894	—	—	—
Rogeria alzatei Kugler, 1994*	5, 34, 52, 57	Am	Collection
Rogeria besucheti Kugler, 1994*	40	Am	Collection
Rogeria germaini Emery, 1894*	40	Am	Collection
Rogeria lirata Kugler, 1994*	58	Am	Collection
Rogeria scobinata Kugler, 1994	11	Ce	Brandão et al., 2011; Collection
Sericomyrmex Mayr, 1865	—	—	—
Sericomyrmex mayri Forel, 1912	18, 29, 35, 37, 45	Am, Ce, ACT	Jesovnik & Schultz, 2017; Collection
Sericomyrmex parvulus Forel, 1912	18, 25, 45	Am, Ce	Jesovnik & Schultz, 2017; Collection
Sericomyrmex saussurei Emery, 1894	18, 29, 45	Am, Ce	Jesovnik & Schultz, 2017; Collection
Solenopsis Westwood, 1840	—	—	—
Solenopsis geminata (Fabricius, 1804)	5, 34, 37, 52, 57, 58	Am, ACT	Wauters et al., 2018; Collection
Solenopsis globularia (Smith, 1858)	5, 45, 52, 57, 59, 61	Am, Ce	Kempf, 1972a; Andrade-Silva et al., 2015; Collection
Solenopsis pollux Forel, 1893*	37	ACT	Collection
Solenopsis saevissima (Smith, 1855)	37, 57, 59, 61, 65	Am, ACT	Shoemaker et al., 2006; Andrade-Silva et al., 2015; Pereira et al., 2017; Collection
Solenopsis substituta Santschi, 1925*	57, 61	Am	Collection
Solenopsis virulens (Smith, 1858)*	5, 34, 40, 52	Am	Collection
Stegomyrmex Emery, 1912	—	—	—
Stegomyrmex olindae Feitosa, Brandão & Diniz, 2008	2, 29, 45	Am, Ce	Feitosa et al., 2008; Ulysséa et al., 2015; Collection
Strumigenys Smith, 1860*	—	—	—
Strumigenys alberti Forel, 1893*	4	Am	Collection
Strumigenys crassicornis Mayr, 1887*	30, 32, 55	Am, Ce	Collection
Strumigenys denticulata Mayr, 1887*	5, 13, 30, 31, 34, 40, 52, 55	Am, Ce	Collection
Strumigenys eggersi Emery, 1890*	4, 30, 31, 32, 41, 60	Am, Ce, ACT	Collection
Strumigenys elongata Roger, 1863*	4, 13, 30, 31, 41	Am, Ce	Collection
Strumigenys gytha Bolton, 2000*	41	Am	Collection
Strumigenys hyphata (Brown, 1953)*	4, 30	Am, Ce	Collection
Strumigenys metopia (Brown, 1959)*	4	Am	Collection
Strumigenys mirabilis Mann, 1926*	4	Am	Collection
Strumigenys orchibia (Brown, 1953)**	60	Am	Collection
Strumigenys schmalzi Emery, 1906*	30, 31	Ce	Collection
Strumigenys subedentata Mayr, 1887*	30, 31, 40	Am, Ce	Collection
Strumigenys trudifera Kempf & Brown, 1969*	4, 5, 40	Am	Collection
Strumigenys urrhobia (Bolton, 2000)*	56	Am	Collection
Strumigenys villiersi (Perrault, 1986)*	30	Ce	Collection
Strumigenys zeteki (Brown, 1959)*	12	Ce	Collection
Tetramorium Mayr, 1855*	—	—	—
Tetramorium simillimum (Smith, 1851)	10	ACT	Kempf, 1972a, 1975; Brandão, 1991; Collection
Trachymyrmex Forel, 1893	—	—	—
Trachymyrmex bugnioni (Forel, 1912)	11, 30	Ce	Brandão et al., 2011; Collection
Trachymyrmex relictus Borgmeier, 1934*	34, 35, 37, 40, 52, 57, 61	Am	Collection
Tranopelta Mayr, 1866*		—	—
Tranopelta gilva Mayr, 1866*	5, 57	Am	Collection
Wasmannia Forel, 1893	—	—	—
Wasmannia auropunctata (Roger, 1863)	5, 11, 12, 13, 15, 30, 31, 32, 34, 35, 37, 40, 52, 55, 57, 61	Am, Ce, ACT	Brandão et al., 2011; Collection
Paraponerinae Emery, 1901	—	—	—
Paraponera Smith, 1858	—	—	—
Paraponera clavata (Fabricius, 1775)	2, 3, 7, 19, 36, 38, 42, 45, 50	Am, Ce, ACT	Ward & Downie, 2005; Ward, 2007; Collection
Ponerinae Lepeletier de Saint-Fargeau, 1835	—	—	—
Anochetus Mayr, 1861*	—	—	—
Anochetus bispinosus (Smith, 1858)*	5, 34	Am	Collection
Anochetus diegensis Forel, 1912*	5, 34, 40, 57, 60	Am	Collection
Anochetus horridus Kempf, 1964*	4, 27, 37	Am, Ce, ACT	Collection
Anochetus mayri Emery, 1884*	5, 34, 52, 57	Am	Collection
Centromyrmex Mayr, 1866*	—	—	—
Centromyrmex brachycola (Roger, 1861)*	5, 34, 57, 61	Am	Collection
Cryptopone Emery, 1893*	—	—	—
Cryptopone guianensis (Weber, 1939)*	34	Am	Collection
Dinoponera Roger, 1861	—		—
Dinoponera gigantea (Perty, 1833)	1, 11, 13, 33, 45	Am, Ce, ACT	Kempf, 1971, 1972a; Monnin et al., 2003; Brandão et al., 2011; Collection
Hypoponera Santschi, 1938*	—	—	—
Hypoponera distinguenda (Emery, 1890)*	34	Am	Collection
Hypoponera opacior (Forel, 1893)*	61	Am	Collection
Hypoponera trigona (Mayr, 1887)*	5, 34, 52, 57	Am	Collection
Leptogenys Roger, 1861		—	—
Leptogenys guianensis Wheeler, 1923*	57	Am	Collection
Leptogenys unistimulosa Roger, 1863	30, 63	Am, Ce	Lattke, 2011; Collection
Mayaponera Schmidt & Shattuck, 2014*	—	—	—
Mayaponera constricta (Mayr, 1884)*	5, 30, 35, 37, 40, 60	Am, Ce, ACT	Collection
Neoponera Emery, 1901	—	—	—
Neoponera commutata (Roger, 1860)	7, 21, 45, 56	Am, Ce	Kempf, 1959; Kempf, 1972a; Collection
Neoponera marginata (Roger, 1861)*	57	Am	Collection
Neoponera striatinodis (Emery, 1890)*	5	Am	Collection
Neoponera unidentata (Mayr, 1862)*	34	Am	Collection
Neoponera verenae Forel, 1922*	34, 37	Am, ACT	Collection
Neoponera villosa (Fabricius, 1804)	11, 13, 19, 60	Am, Ce	Brandão et al., 2011; Fernandes et al., 2014; Collection
Odontomachus Latreille, 1804	—	—	—
Odontomachus bauri Emery, 1892*	11, 22, 35, 37, 57, 59, 60	Am, Ce, ACT	Brandão et al., 2011; Andrade-Silva et al., 2015; Collection
Odontomachus brunneus (Patton, 1894)*	22	Ce	Collection
Odontomachus chelifer (Latreille, 1802)*	57, 26	Am, Ce	Collection
Odontomachus haematodus (Linnaeus, 1758)	1, 45, 57	Am, Ce	Janicki et al., 2016; Collection
Odontomachus meinerti Forel, 1905*	34, 52	Am	Collection
Odontomachus opaciventris Forel, 1899	29	ACT	Dáttilo et al. 2012
Odontomachus scalptus Brown, 1978*	5	Am	Collection
Pachycondyla Smith, 1858*	—	—	—
Pachycondyla crassinoda (Latreille, 1802)*	5, 30, 32, 35, 37, 52, 57, 60	Am, Ce, ACT	Collection
Pachycondyla harpax (Fabricius, 1804)*	5, 30, 31, 32, 34, 35, 37, 40, 46, 52, 55, 56, 57	Am, Ce, ACT	Collection
Pachycondyla impressa (Roger, 1861)*	5	Am	Collection
Pachycondyla lenis Kempf, 1961*	34	Am	Collection
Platythyrea Roger, 1863	—	—	—
Platythyrea angusta Forel, 1901	45	Ce	Forel, 1904; Kempf, 1964, 1972a; Collection
Platythyrea pilosula (Smith, 1858)*	60	Am	Collection
Pseudoponera Emery, 1900*		—	—
Pseudoponera gilberti (Kempf, 1960)*	5, 34, 40, 52, 61	Am	Collection
Pseudoponera stigma (Fabricius, 1804)*	41	Am	Collection
Rasopone Schmidt & Shattuck, 2014*	—	—	—
Rasopone arhuaca (Forel, 1901)*	5, 34, 41, 52, 55, 57	Am, ACT	Collection
Rasopone ferruginea (Smith, 1858)*	5, 30, 31, 34, 40, 55	Am, Ce	Collection
Proceratiinae Emery, 1895*	—	—	—
Discothyrea Roger, 1863*	—	—	—
Discothyrea sexarticulata Borgmeier, 1954*	4, 5	Am	Collection
Pseudomyrmecinae Smith, 1952	—	—	—
Pseudomyrmex Lund, 1831	—	—	—
Pseudomyrmex curacaensis (Forel, 1912)	5, 10, 40, 45, 52	Am, Ce, ACT	Ward, 1989; Brandão, 1991; Collection
Pseudomyrmex elongatus (Mayr, 1870)	10, 45	Ce, ACT	Kempf, 1972a; Ward, 1989; Collection
Pseudomyrmex ethicus (Forel, 1911)*	4	Am	Collection
Pseudomyrmex filiformis (Fabricius, 1804)*	5, 34	Am	Collection
Pseudomyrmex flavidulus (Smith, 1858)	11, 13	Ce	Brandão et al., 2011; Collection
Pseudomyrmex gracilis (Fabricius, 1804)	5, 16, 38, 40, 45, 46, 56, 57, 63	Am, Ce, CCT	Janicki et al., 2016; Collection
Pseudomyrmex kuenckeli (Emery, 1890)	35, 43	Am, ACT	Ward, 1999; Collection
Pseudomyrmex oculatus (Smith, 1855)	5, 34, 40, 45, 56, 57, 63	Am, Ce	Kempf, 1972a; Collection
Pseudomyrmex penetrator (Smith, 1877)*	3	Am	Collection
Pseudomyrmex pupa (Forel, 1911)*	56, 40	Am	Collection
Pseudomyrmex schuppi (Forel, 1901)	52, 57, 59	Am	Andrade-Silva et al., 2015; Collection
Pseudomyrmex sericeus (Mayr, 1870)*	5, 40	Am	Collection
Pseudomyrmex tenuis (Fabricius, 1804)	5, 13, 20, 27, 34, 37, 40, 45, 47, 56, 58, 60, 61, 62, 63	Am, Ce, ACT	Forel, 1904; Kempf, 1960b, 1972a; Ward & Downie, 2005; Collection
Pseudomyrmex tenuissimus (Emery, 1906)	5, 10, 45	Am, Ce, ACT	Mann, 1916; Kempf, 1972a; Ward, 1989; Brandão, 1991; Collection
Pseudomyrmex termitarius (Smith, 1855)	5, 13, 27, 43, 45, 47	Am, Ce	Kempf, 1972a; Brandão et al., 2011; Collection
Pseudomyrmex urbanus (Smith, 1877)	13, 45	Ce	Ward, 1989; Brandão, 1991; Collection

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).

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.

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 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 21^st 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 20^th 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 19^th century (Vanzolini, 2004), for the ants this occurred in the second half of the 20^th century. In the case of the records analysed in this work, the specific localities and geographic coordinates became available in the late 20^th 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 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 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; Maracahipes-Santos 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.

Figure 2
Map of the state of Maranhão emphasizing the main highways and sampling sites of ant species within the state.

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, national forests, biological reserves, among others) are of fundamental importance for biodiversity conservation (Peres, 2005) and preserving ecosystem (Hallmann et al., 2017).

Figure 3
Map of the state of Maranhão emphasizing the Priority Conservation Areas and sampling sites of ant species within the state.

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. Albuquerque (Arizona State University), Fabrício Baccaro (Universidade Federal do Amazonas), Jacques Delabie (Comissão Executiva do Plano da Lavoura Cacaueira), Mônica 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

• Andrade-Silva, J.; Pereira, E.K.C.; Silva, O.; Santos, C.L.C.; Delabie, J.H.C. & Rebelo, J.M.M. 2015. Ants (Hymenoptera: Formicidae) associated with pig carcasses in an urban area. Sociobiology, 62(4): 527-532.
• AntWeb. 2019. AntWeb. Available at: Available at: http://www.antweb.org Access in: 03/01/2019.
  » http://www.antweb.org
• AntWiki. 2019. AntiWiki. Available at: Available at: http://www.antwiki.org Access in: 19/01/2019.
  » http://www.antwiki.org
• Bolton, B. 2019. An online catalog of the ants of the world. Available at: Available at: http://www.antcat.org Access in: 10/02/2019.
  » http://www.antcat.org
• Borgmeier, T. 1955. Die Wanderameisen der neotropischen Region. Studia Entomologica, 3: 1-720.
• Brandão, C.R.F. 1991. Adendos ao catálogo abreviado das formigas da região neotropical (Hymenoptera: Formicidae). Revista Brasileira de Entomologia, 35: 319-412.
• Brandão, C.R.F.; Feitosa, R.M. & Diniz, J.L.M. 2015. Taxonomic revision of the Neotropical Myrmicinae ant genus Blepharidatta Wheeler. Zootaxa, 4012(1): 33-56.
• Brandão, C.R.F.; Silva, R.R. & Feitosa, R.M. 2011. Cerrado ground-dwelling ants (Hymenoptera: Formicidae) as indicators of edge effects. Zoologia, 28(3): 379-387.
• Brasil. 2009. Ministério do Meio Ambiente. Relatório técnico de monitoramento do desmatamento no bioma Cerrado, 2002 a 2008: dados revisados. Brasília: MMA, 67p. Available at: Available at: http://www.mma.gov.br/estruturas/sbf_chm_rbbio/_arquivos/relatorio_tecnico_monitoramento_desmate_bioma_cerrado_csr_ibama_2002_2008_rev_72.pdf Access in: 08/12/2018.
  » http://www.mma.gov.br/estruturas/sbf_chm_rbbio/_arquivos/relatorio_tecnico_monitoramento_desmate_bioma_cerrado_csr_ibama_2002_2008_rev_72.pdf
• Carvalho, A.P.R.; Silva, C.G. & Fonseca, A.R. 2011. Diversidade de formigas em um hospital público no município de Chapadinha, Maranhão, Brasil. Revista de Biologia e Ciências da Terra, 11: 67-73.
• Celentano, D.; Rousseau, G.X.; Engel, V.L.; Zelarayán, M.; Oliveira, E.C.; Araújo, A.C.M. & De Moura, E.G. 2017. Degradation of riparian forest affects soil properties and ecosystem services provision in eastern amazon of Brazil. Land Degradation & Development, 28: 482-493.
• Chaves, L.P.F.A.; Silva, R.A.; Amaral, Y.T.; Costa, M.K.L. & Siqueira, G.M. 2016. Biogeographical diversity of north mesoregion of the Maranhão state (Brazil). Journal of Geospatial Modelling, 1: 19.
• Cuezzo, F. 2000. Revisión del género Forelius (Hymenoptera: Formicidae: Dolichoderinae). Sociobiology, 35: 197-275.
• Dalzochio, M.S.; Renner, S.; Sganzerla, C.; Prass, G.; Ely, G.J.; Salvi, L.C.; Dametto, N. & Périco, E. 2018. Checklist of Odonata (Insecta) in the state of Rio Grande do Sul, Brazil with seven new records. Biota Neotropica, 18: 1-14.
• Dáttilo, W.; Vicente, R.E.; Nunes, R.V. & Carvalho, M.S.G. 2010. First Record of the Quenquém cisco-da-Amazônia Acromyrmex hystrix (Latreille) (Formicidae: Myrmicinae) for Maranhão State, Brazil. EntomoBrasilis, 3(3): 92-93.
• Dáttilo, W.; Vicente, R.E.; Nunes, R.V. & Feitosa, R.M. 2012. Influence of cave size and presence of bat guano on ant visitation. Sociobiology, 59(2): 549-559.
• De Andrade, M.L. & Baroni Urbani, C. 1999. Diversity and Adaptation in the ant genus Cephalotes, past and present. Stuttgarter Beitrage zur Naturkunde Serie B, 271: 893.
• Desidério, G.R.; Barcelos-Silva, P.; De Souza, W.R.M.; Pes, A.M. & Azevêdo, C.A.S. 2017. Caddisflies (Insecta: Trichoptera) from Maranhão State, Northeast Region, Brazil: A new species, checklist, and new geographical records. Zootaxa, 4221: 151-171.
• Dias, A.M. & Lattke, J.E. 2019. A new species and new records of minuta-group Gnamptogenys from Brazil (Hymenoptera: Formicidae). Revista Brasileira de Entomologia, 63(1): 30-34.
• Feitosa, R.M.; Brandão, C.R.F. & Dietz, B.H. 2007. Basiceros scambognathus (Brown, 1949) n. comb., with the first worker and male descriptions, and a revised generic diagnosis (Hymenoptera: Formicidae: Myrmicinae). Papéis Avulsos de Zoologia, 47(2): 31-42.
• Feitosa, R.M.; Brandão, C.R.F. & Diniz, J.L.M. 2008. Revisionary studies on the enigmatic Neotropical ant genus Stegomyrmex Emery, 1912 (Hymenoptera: Formicidae: Myrmicinae), with the description of two new species. Journal of Hymenoptera Research, 17: 64-82.
• Fernandes, I.O.; Oliveira, M.L. & Delabie, J.H.C. 2014. Description of two new species in the Neotropical Pachycondyla foetida complex (Hymenoptera: Formicidae: Ponerinae) and taxonomic notes on the genus. Myrmecological News, 19: 133-163.
• Figueiredo, C.J.; Silva, R.R.; Munhae, C.B. & Morini, M.S.C. 2013. Fauna de formigas (Hymenoptera: Formicidae) atraídas a armadilhas subterrâneas em áreas de Mata Atlântica. Biota Neotropica, 13: 1-7.
• Forel, A. 1904. Miscellanea myrmécologiques. Revue Suisse de Zoologie, 12: 1-52.
• Freitas, M.A.; Vieira, R.S.; Entiauspe-Neto, O.M.; Sousa, S.O.; Farias, T.; Sousa, A.G. & Moura, G.J.B. 2017. Herpetofauna of the Northwest Amazon forest in the state of Maranhão, Brazil, with remarks on the Gurupi Biological Reserve. ZooKeys, 643: 141-155.
• Gasper, A.L.; Eisenlohr, P.V. & Salino, A. 2016. Improving collection efforts to avoid loss of biodiversity: lessons from comprehensive sampling of lycophytes and ferns in the subtropical Atlantic Forest. Acta Botanica Brasilica, 30: 166-175.
• Gonçalves, C.R. 1942. Contribuição para o conhecimento do gênero Atta Fabr., das formigas saúvas. Boletim da Sociedade Brasileira de Agronomia, 5: 333-358.
• Gonçalves, C.R. 1947. Saúvas do sul e centro do Brasil. Boletim Fitossanitário, 2: 183-218.
• Gonçalves, C.R. 1961. O gênero Acromyrmex no Brasil (Hym. Formicidae). Studia Entomologica, 4: 113-180.
• Graham, C.H.; Elith, J.; Hijmans, R.J.; Guisan, A.; Townsend Peterson, A. & Loiselle, B.A. 2008. The influence of spatial errors in species occurrence data used in distribution models. Journal of Applied Ecology, 45: 239-247.
• Gutiérrez, J.A.M.; Rousseau, G.X.; Andrade-Silva, J. & Delabie, J.H.C. 2017. Taxones superiores de hormigas como sustitutos de la riqueza de especies, en una cronosecuencia de bosques secundarios, bosque primario y sistemas agroforestales en la Amazonía Oriental, Brasil. Revista de Biología Tropical, 65(1): 279-291.
• Hallmann, C.A.; Sorg, M.; Jongejans, E.; Siepel, H.; Hofland, N.; Schwan, H.; Stenmans, W.; Müller, A.; Sumser, H.; Hörren, T.; Goulson, D. & de Kroon, H. 2017. More than 75 percent decline over 27 years in total flying insect biomass in protected areas. Plos One, 12(10): e0185809.
• Hijmans, R.J.; Garrett, K.A.; Huama’n, Z.; Zhang, D.P.; Schreuder, M. & Bonierbale, M. 2000. Assessing the geographic representativeness of genebank collections: the case of Bolivian wild potatoes. Conservation Biology, 14: 1755-65.
• Instituto Brasileiro de Geografia e Estatística (IBGE). 2011. Índice de Nomes Geográficos, Escala 1:1.000.000. Base Cartográfica Contínua do Brasil ao Milionésimo. Disponível em: Disponível em: http://www.ibge.gov.br/geociencias/cartas-e-mapas/bases-cartograficas-continuas/15759-brasil.html?=&t=sobre Acesso em: 03/12/2018.
  » http://www.ibge.gov.br/geociencias/cartas-e-mapas/bases-cartograficas-continuas/15759-brasil.html?=&t=sobre
• Instituto Brasileiro de Geografia e Estatística (IBGE). 2018. Maranhão. Available at: Available at: http://cidades.ibge.gov.br/brasil/ma/panorama Access in: 02/02/2019.
  » http://cidades.ibge.gov.br/brasil/ma/panorama
• Janicki, J.; Narula, N.; Ziegler, M.; Guénard, B. & Economo, E.P. 2016. Visualizing and interacting with large-volume biodiversity data using client-server web-mapping applications: The design and implementation of antmaps.org. Ecological Informatics, 32: 185-193.
• Jesovnik, A. & Schultz, T.R. 2017. Revision of the fungus-farming ant genus Sericomyrmex Mayr (Hymenoptera, Formicidae, Myrmicinae). ZooKeys, 670: 1-109.
• Johnson, R.A. 2015. A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I. Zootaxa, 4029(1): 1-142.
• Kadmon, R.; Farber, O. & Danin, A. 2003. A systematic analysis of factors affecting the performance of climatic envelope models. Ecological Applications, 13: 853-67.
• Kempf, W.W. 1959. Insecta Amapaensia. Hymenoptera: Formicidae. Studia Entomologica, 2: 209-218.
• Kempf, W.W. 1960a. Insecta Amapaensia. Hymenoptera: Formicidae (segunda contribuição). Studia Entomologica, 3: 385-400.
• Kempf, W.W. 1960b. Estudo sôbre Pseudomyrmex I. (Hymenoptera: Formicidae). Revista Brasileira de Entomologia, 9: 5-32.
• Kempf, W.W. 1964. Uma nova Platythyrea do Brasil (Hym., Formicidae). Revista Brasileira de Entomologia, 11: 141-144.
• Kempf, W.W. 1968. Miscellaneous studies on Neotropical ants. IV. (Hymenoptera, Formicidae). Studia Entomologica, 11: 369-415.
• Kempf, W.W. 1971. A preliminary review of the ponerine ant genus Dinoponera Roger (Hymenoptera: Formicidae). Studia Entomologica, 14: 369-394.
• Kempf, W.W. 1972a. Catálogo abreviado das formigas da região Neotropical (Hym. Formicidae). Studia Entomologica, 15: 1-4.
• Kempf, W.W. 1972b. A new species of the Dolichoderine ant genus Monacis Roger, from the Amazon, with further remarks on the genus (Hymenoptera, Formicidae). Revista Brasileira de Biologia, 32: 251-254.
• Kempf, W.W. 1975. Miscellaneous studies on neotropical ants. VI. (Hymenoptera, Formicidae). Studia Entomologica, 18: 341-380.
• Lattke, J.E. 2011. Revision of the New World species of the genus Leptogenys Roger (Insecta: Hymenoptera: Formicidae: Ponerinae). Arthropod Systematics & Phylogeny, 69: 127-264.
• Leal, I.R.; Leal, L.; Oliveira, F.P.; Arcoverde, G.B. & Andersen, A.N. 2017. Effects of human disturbance and climate change on myrmecochory in Brazilian Caatinga. In: Oliveira, P.S. & Koptur, S. (Eds.). Ant-plant interactions. Impacts of human on terrestrial ecosystems. Cambridge, UK, Cambridge University Press. p. 112-132.
• Lima, W.R.S.; Marques, S.G.; Rodrigues, F.S. & Rebelo, J.M.M. 2013. Ants in a hospital environment and their potential as mechanical bacterial vectors. Revista da Sociedade Brasileira de Medicina Tropical, 46(5): 637-640.
• Longino, J.T. & Snelling, R.R. 2002. A taxonomic revision of the Procryptocerus (Hymenoptera: Formicidae) of Central America. Contributions in Science, 495: 1-30.
• Mann, W.M. 1916. The Stanford Expedition to Brazil, 1911, John C. Branner, director. The ants of Brazil. Bulletin of the Museum of Comparative Zoology, 60: 399-490.
• Maracahipes-Santos, L.; Santos, J.O.; Reis, S.M. & Lenza, E. 2018. Temporal changes in species composition, diversity, and woody vegetation structure of savannas in the Cerrado-Amazon transition zone. Acta Botanica Brasilica, 32(2): 254-263.
• Ministério do Meio Ambiente (MMA). 2011. Plano de Ação para prevenção e controle do desmatamento e das queimadas: Cerrado. Brasília, MMA. 200p.
• Monnin, R.; Ratnieks, F.L.W. & Brandão, C.R.F. 2003. Reproductive Conflict in Animal Societies: Hierarchy Length Increases with Colony Size in Queenless Ponerine Ants. Behavioral Ecology and Sociobiology, 54(1): 71-79.
• Moura, C.C.M.; Moura, G.J.B.; Lisboa, E.B.F. & Luz, V.L.F. 2014. Distribuição geográfica e considerações ecológicas sobre a fauna de Testudines da Região Nordeste do Brasil. Sitientibus serie Ciencias Biologicas, 14: 1-20.
• Newbold, T. 2010. Applications and limitations of museum data for conservation and ecology, with particular attention to species distribution models. Progress in Physical Geography, 34: 3-22.
• Pereira, E.K.C.; Andrade-Silva, J.; Silva, O.; Santos, C.L.C.; Moraes, L.S.; Bandeira, M.C.A.; Silva, C.R.R. & Rebêlo, J.M.M. 2017. Solenopsis saevissima (Smith) (Hymenoptera: Formicidae) activity delays vertebrate carcass decomposition. Sociobiology, 64: 369.
• Pereira, J.C.; Delabie, J.H.C.; Zanette, L.R.S. & Quinet, Y. 2014. Studies on an Enigmatic Blepharidatta Wheeler Population (Hymenoptera: Formicidae) from the Brazilian Caatinga. Sociobiology, 61(1): 52-59.
• Peres, C.A. 2005. Why we need mega-reserves in Amazonian forests. Conservation Biology, 19: 728-733.
• QGIS Development Team. 2019. QGIS geographic information system. Open source geospatial foundation project. Disponível em: http://qgis.org/en/site/forusers/download.html
  » http://qgis.org/en/site/forusers/download.html
• Ramos, A.S.J.C.; Lemos, R.N.S.; Vale, A.M.S; Batista, M.C.; Moreira, A.A.; Harada, H.Y. & Mesquita, M.L.R. 2015. Ant diversity in agro ecosystems and secondary forest. African Journal of Agricultural Research, 10: 4449-4454.
• Reddy, S. & Dávalos, L.M. 2003: Geographical sampling bias and its implications for conservation priorities in Africa. Journal of Biogeography, 30: 1719-27.
• Santos, B.F. & Hoppe, J.P.M. 2018. Filling gaps in species distributions through the study of biological collections: 415 new distribution records for Neotropical Cryptinae (Hymenoptera, Ichneumonidae). Revista Brasileira de Entomologia, 62(4): 288-291.
• Santos, G.M.M.; Delabie, J.H.C. & Resende, J.J. 1999. Caracterização da Mirmecofauna (Hymenoptera: Formicidae) associada à vegetação periférica de inselbergs (Caatinga-arbórea-estacional-semi-descídua) em Itatim-Bahia-Brasil. Sitientibus. Revista da Universidade Estadual de Feira de Santana, 20: 33-43.
• Santos, M.P.D.; Cerqueira, P.V. & Soares, L.M.S. 2010. Avifauna em seis localidades no centro-sul do Estado do Maranhão, Brasil. Ornithologia, 4(1): 49-65.
• Shoemaker, D.D.; Ahrens, M.E. & Ross, K.G. 2006. Molecular phylogeny of fire ants of the Solenopsis saevissima species-group based on mtDNA sequences. Molecular Phylogenetics and Evolution, 38: 200-215.
• Silva, E.F.; Corá, J.E.; Harada, A.Y. & Sampaio, I.B.M. 2017. Association of the Occurrence of Ant Species (Hymenoptera: Formicidae) with Soil Attributes, Vegetation, and Climate in the Brazilian Cerrado Northeastern Region. Sociobiology, 64(4): 442-450.
• Silva, G.M.; Carmo, M.S.; Moraes, L.S.; Moraes, F.C.; Barnabé, A.S. & Figueiredo, P.M.S. 2012. Formigas (Hymenoptera: Formicidae) como Vetores de Bactéria em Ambiente Hospitalar na Cidade de São Luis, Maranhão. Revista de Patologia Tropical, 41(3): 348-355.
• Silva, P.R. 2007. Biologia de algumas espécies de Blepharidatta. Biológico, 69(2): 161-164.
• Spinelli-Araujo, L.; Bayma-Silva, G.; Torresan, F.E.; Victoria, D.; Vicente, L.E.; Bolfe, E.L. & Manzatto, C. 2016. Conservação da biodiversidade do estado do Maranhão: cenário atual em dados geoespaciais. Jaguariúna, EMBRAPA Meio Ambiente. 28p.
• Stella, A. 2011. Plano de prevenção e controle do desmatamento e queimadas do Maranhão. São Luís, SEMA. 120p.
• Ulysséa, M.A. & Brandão, C.R.F. 2013. Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. Revista Brasileira de Entomologia, 57: 217-224.
• Ulysséa, M.A.; Prado, L.P. & Brandão, C.R.F. 2015. Type specimens of the traditional Myrmicinae (Hymenoptera: Formicidae) ant tribes deposited in the Museu de Zoologia da Universidade de São Paulo, Brazil: Adelomyrmecini, Basicerotini, Blepharidattini, Crematogastrini, Formicoxenini, Lenomyrmecini, Myrmicini, Phalacromyrmecini, Pheidolini, Stegomyrmecini, Stenammini and Tetramoriini. Papéis Avulsos de Zoologia, 55: 175-204. Disponível em: Disponível em: http://www.revistas.usp.br/paz/article/view/106374 Acesso em 25.04.2019.
  » http://www.revistas.usp.br/paz/article/view/106374
• Ulysséa, M.A.; Prado, L.P. & Brandão, C.R.F. 2017. Catalogue of the Dolichoderinae, Formicinae and Martialinae (Hymenoptera: Formicidae) types deposited at the Museu de Zoologia da Universidade de São Paulo, Brazil. Papéis Avulsos de Zoologia , 57(23): 295-311. Disponível em: Disponível em: http://www.revistas.usp.br/paz/article/view/125647 Acesso em: 25.04.2019
  » http://www.revistas.usp.br/paz/article/view/125647
• Vanzolini, P.E. 1992. A Supplement to the Ornithological Gazetteer of Brazil. São Paulo, Museu de Zoologia da Universidade de São Paulo. 252p.
• Vanzolini, P.E. 2004. Episódios da Zoologia Brasílica. São Paulo, Hucitec. 212p.
• Vanzolini, P.E. & Papavero, N. 1968. Índice dos Topônimos Contidos na Carta do Brasil 1:1.000.000 do IBGE. São Paulo, Fundação de Amparo à Pesquisa do Estado de São Paulo. 201p.
• Ward, P.S. 1989. Systematic Studies on Pseudomyrmecine Ants: Revision of the Pseudomyrmex oculatus and P. subtilissimus species groups with taxonomic comments on other species. Questiones Entomologicae, 25: 393-468.
• Ward P.S. 1999. Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris- and Tachigali-inhabiting ants. Zoological Journal of the Linnean Society, 126: 451-540.
• Ward, P.S. 2007. The ant genus Leptanilloides: discovery of the male and evaluation of phylogenetic relationships based on DNA sequence data. Memoirs of the American Entomological Institute, 80: 637-649.
• Ward, P.S. & Downie, D.A. 2005. The ant subfamily Pseudomyrmecinae: phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology, 30: 310-335.
• Watkins, J.F. 1976. The identification and distribution of New World army ants (Dorylinae: Formicidae). Waco, Texas, Baylor University Press. 102p.
• Wauters, N.; Dekoninck, W. & Fournier, D. 2018. Introduction history and genetic diversity of the invasive ant Solenopsis geminata in the Galapagos Islands. Biological Invasions, 20(11): 3207-3226.
• Wheeler, W.M. 1922. Observations on Gigantiops destructor Fabricius and other leaping ants. Biological Bulletin (Woods Hole), 42: 185-201.
• Wild, A.L. 2007. Taxonomic revision of the ant genus Linepithema (Hymenoptera: Formicidae). University of California Publications in Entomology, 126: 1-151.
• Wild, A.L. & Cuezzo, F. 2006. Rediscovery of a fossil dolichoderine ant lineage (Hymenoptera: Formicidae: Dolichoderinae) and a description of a new genus from South America. Zootaxa, 1142: 57-68.

• Edited by: Helena Carolina Onody
• Published with the financial support of the Committee of "Programa de Apoio às Publicações Científicas Periódicas da USP" (SIBi-USP)

Data availability

Data citations

AntWeb. 2019. AntWeb. Available at: Available at: http://www.antweb.org Access in: 03/01/2019.

Publication Dates

• Publication in this collection
  03 Oct 2019
• Date of issue
  2019

History

• Received
  22 May 2019
• Accepted
  15 July 2019