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Papéis Avulsos de Zoologia

Print version ISSN 0031-1049On-line version ISSN 1807-0205

Pap. Avulsos Zool. vol.59  São Paulo  2019  Epub Oct 03, 2019

https://doi.org/10.11606/1807-0205/2019.59.38 

ARTICLES

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

Lívia Pires do Prado1  6 
http://orcid.org/0000-0003-1819-8767

Rodrigo Machado Feitosa2  7
http://orcid.org/0000-0001-9042-0129

Stefania Pinzón Triana3  8
http://orcid.org/0000-0002-7160-0826

Jhonatan Andrés Muñoz Gutiérrez3  9
http://orcid.org/0000-0002-2090-2226

Guillaume Xavier Rousseau3  10
http://orcid.org/0000-0002-2482-4376

Raimunda Alves Silva4  11
http://orcid.org/0000-0002-0380-8190

Glécio Machado Siqueira4  12
http://orcid.org/0000-0002-3513-2658

Ciro Líbio Caldas dos Santos5  13
http://orcid.org/0000-0002-0987-3620

Francisco Veras Silva5  14
http://orcid.org/0000-0003-0057-6467

Thiago Sanches Ranzani da Silva2  15
http://orcid.org/0000-0002-4239-1500

Alexandre Casadei-Ferreira2  16
http://orcid.org/0000-0002-2977-8348

Rogério Rosa da Silva1  17
http://orcid.org/0000-0002-0599-2155

Joudellys Andrade-Silva1  18
http://orcid.org/0000-0002-5393-6502

1Museu 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.

2Universidade Federal do Paraná (UFPR), Departamento de Zoologia (DZOO), Laboratório de Sistemática e Biologia de Formigas (LSBF).Curitiba, PR, Brasil.

3Universidade Estadual do Maranhão (UEMA), Programa de Pós-Graduação em Agroecologia. São Luís, MA, Brasil.

4Universidade Federal do Maranhão (UFMA), Departamento de Geociências (DEGEO). São Luís, MA, Brasil.

5Universidade Federal do Maranhão (UFMA). Imperatriz, MA, Brasil.


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 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).

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

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 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 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.

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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)

Received: May 22, 2019; Accepted: July 15, 2019

6 E-mail: livia.pires7@gmail.com (corresponding author)

7

E-mail: rsmfeitosa@gmail.com

8

E-mail: stefaniapt@gmail.com

9

E-mail: energiaselvatica@gmail.com

10

E-mail: guilirous@yahoo.ca

11

E-mail: ray-234@hotmail.com

12

E-mail: gleciosiqueira@hotmail.com

13

E-mail: cirolb@gmail.com

14

E-mail: fveras_silva@hotmail.com

15

E-mail: tsranzanidasilva@gmail.com

16

E-mail: alexandrefrreira@gmail.com

17

E-mail: rogeriorosas@gmail.com

18

E-mail: joudellys@gmail.com

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