Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records

Abstract

Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. The Caatingas occur predominantly in northeastern Brazil and comparatively it is the biome that received less attention than any other ecosystem in Brazil, representing the region where invertebrate groups are less known. We present here the first list of ant species of the Caatingas, compiling information from the literature, from a study of samples preserved in alcohol in the Laboratory of Entomology (Universidade Estadual de Feira de Santana), and from a field survey conducted in Milagres, Bahia, submitting standardized 1-m² samples of the leaf-litter to Winkler extractors. Summing all information, 11 subfamilies, 61 genera and 173 species (plus one subspecies) of ants are recognized in the biome. This species number does not consider morphospecies that could not be named due to the lack of reliable recent taxonomic information for some Neotropical ant genera. The list presented here for ant species of the Caatingas is therefore underestimated, but it is relevant because it allows the identification of areas to be sampled in order to improve our knowledge of the diversity of ants in this biome.

Caatinga; Formicidae; Insecta; list of species; SDTF

Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records

Mônica A. Ulysséa^{I, II}; Carlos R. F. BrandãoII, ^*

^IPrograma de Pós-Graduação em Sistemática, Taxonomia Animal e Biodiversidade. [e-mail]

^IILaboratório de Hymenoptera, Museu de Zoologia Universidade de São Paulo, Avenida Nazaré, nº 481, Ipiranga. 04263 - 000. São Paulo-SP, Brasil

ABSTRACT

Ant species (Hymenoptera, Formicidae) from the seasonally dry tropical forest of northeastern Brazil: a compilation from field surveys in Bahia and literature records. The Caatingas occur predominantly in northeastern Brazil and comparatively it is the biome that received less attention than any other ecosystem in Brazil, representing the region where invertebrate groups are less known. We present here the first list of ant species of the Caatingas, compiling information from the literature, from a study of samples preserved in alcohol in the Laboratory of Entomology (Universidade Estadual de Feira de Santana), and from a field survey conducted in Milagres, Bahia, submitting standardized 1-m² samples of the leaf-litter to Winkler extractors. Summing all information, 11 subfamilies, 61 genera and 173 species (plus one subspecies) of ants are recognized in the biome. This species number does not consider morphospecies that could not be named due to the lack of reliable recent taxonomic information for some Neotropical ant genera. The list presented here for ant species of the Caatingas is therefore underestimated, but it is relevant because it allows the identification of areas to be sampled in order to improve our knowledge of the diversity of ants in this biome.

Keywords: Caatinga; Formicidae; Insecta; list of species; SDTF.

Studies about local or regional species biodiversity and the publication of taxonomic lists represent the base of biological knowledge on areas or biomes. This data is scientifically relevant (Vanzolini et al. 1980) as it is the basic element for the understanding of the functional relationships among organisms living in each biome (Marques et al. 2002; Lewinsohn et al. 2005).

The study of certain invertebrate groups is more appropriate than others for determining the local biological diversity, as well to assess or monitor them (Kremen et al. 1993). Furthermore, some of them can be used as environmental indicators (Freitas et al. 2004; Ribas et al. 2012). Ants, with 12,649 described species (Agosti & Johnson 2005), stand out in this scenario due to comparatively wide distribution (Brühl et al. 1999), strong interactions with several groups of plants and animals (Wilson 1987; Floren et al. 2002), sensitivity to habitat changes (Kaspari & Majer 2000) and their relatively short life cycles. Ants present also many specialized taxa, are relatively easy to collect and to separate into morphospecies, and account for more than 10% of animal biomass in tropical forests, savannas and other terrestrial biomes (Majer 1983; Hölldobler & Wilson 1990; Brown Jr. 1997; Folgarait 1998; Alonso 2000; Agosti et al. 2000; Leal 2003; Delsinne et al. 2008), except for the Arctic and the Antarctic.

Despite the crucial importance of species lists and the promising results generated by the studies of ants, there are still regions in Brazil where these studies are insufficient or absent (Brandão 1995). Comparatively, the Caatingas received less attention than any other Brazilian biome and represent the region where invertebrate groups, such as ants, are less known and studied (Brandão & Yamamoto 2004).

The Caatingas can be characterized as a seasonally dry tropical forest (SDTF), and may be also referred as dry tropical forest (DTF) (Leal et al. 2005) - a biome which occurs in tropical and subtropical environments in large and smaller areas of South America, Central America, Africa, Asia and Oceania (Pennington et al. 2000; Cardoso & Queiroz 2010; Espírito-Santo et al. 2010; Werneck 2011).

The Caatingas are associated to relatively dry areas (Ab'Saber 2003), characterized by minimal limits of annual precipitation (until 1000 mm), irregular distribution of rains with six to nine months of drought (dry winters and rainy summers), high average annual temperature (between 26 - 28°C), and rates of relative humidity and cloudiness among the lowest in the country (Aquad 1986; Prado 2003). The Caatingas occur in Brazil (Santana & Souto 2006), covering some 750,000 km² of the country (Ab'Sáber 2003), from the Jequitinhonha river valley (Minas Gerais) to the north, covering large areas in northeastern Brazilian states: Bahia, Ceará, Rio Grande do Norte and Pernambuco, west of Alagoas and Sergipe, southeast of Piauí and east of Maranhão (Fig. 1) (Ab'Saber 2003; Prado 2003; Leal et al. 2005; Zappi 2008). Furthermore, the Caatingas occur as enclaves in the Amazon and Atlantic Forest (Cabo Frio/Rio de Janeiro) (Ab'Saber 2003; Ibraimo et al. 2004; Werneck 2011). The Caatingas include diverse physiognomies that can be recognized locally, as the caatinga-de-lajedo, dense dry and low thorny forests. This is why we prefer to call the biome as the Caatingas. However, these physiog-nomies are not always clearly limited and discernible, preventing the clear mapping of these types.

The present study was carried out with the objective to present the first list of ant species of the Caatingas, in an attempt to overcome the lack of basic knowledge about one of the most important insects groups living in a much stressed environment, due to a long history of human occupation.

MATERIAL AND METHODS

Study site. The surveys, authorized by the Sistema de Autorização e Informação em Biodiversidade/SISBIO (license nr. 24005 - 2), were carried out in the municipality of Milagres, Bahia, defined as an area of extremely high priority for conservation of the Caatingas biome by PROBIO (Velloso et al. 2002; MMA 2003; MMA 2007), in three private areas: A1) 12°54'32.52"S 39°51'16.74"W, A2) 12°54'17.64"S 39°52'04.98"W and A3) 12°54'24.66"S 39°50'51.78"W, all covered by arboreal caatinga vegetation (Figs. 2A-B).

Data collecting. Ants were collected in three field trips, comprising dry and wet seasons from July/2010 to January/2011. In each trip we went through all three study areas and collected a total of fifty 1-m² samples of leaf-litter in points distant at least 50 m one from each other, along irregular transects, due to fact that the Caatingas vegetation is rich in lianas, thorny and urticant plants). The samples were submitted to individual Winkler extractors for 48 h.

In addition, we actively searched for ants in the study areas, like in the soil, plants, fungus, rotten wood, fallen leaves and under rocks, to complement the list of species.

Study of samples preserved in alcohol. The samples preserved in alcohol in the Laboratory of Entomology/LENT, in the Universidade Estadual de Feira de Santana (UEFS), were revised in search of ants. These samples came from surveys on the beetle fauna in distinct municipalities under Caatinga vegetation in the Bahia state (Euclides da Cunha, Milagres, Mucururé and Tucano), using baited pitfall traps as sampling technique (Campos 2012; Medina 2012).

Data from literature. Data on ant species recorded in the Caatingas available in literature were compiled from distinct sources: taxonomic reviews (Kempf 1975; Brandão 1990; Mackay 1993; De Andrade & Baroni-Urbani 1999; Bolton 2000; Albuquerque & Brandão 2004; Dietz 2004; Klingenberg & Brandão 2009), description of new species (Forel 1911), ecological investigations (Brandão 1995; Santos et al. 1999; Leal 2002; Leal 2003; Quinet & Tavares 2005; Leal et al. 2007; Tabatinga-Filho & Leal 2007; Oliveira et al. 2009; Neves et al. 2010; Silva et al. 2010; Brito 2011; Carvalho et al. 2011; Garro et al. 2011; Marques & Soares 2011; Nunes et al. 2011; Silva 2011a; Silva 2011b; Macêdo 2012) and an ethological study (Quinet et al. 2005). The main sampling techniques used in the compiled ecological investigations were sardine or honey baits and pitfall traps. Further, two studies used active search in flowers/plants (Tabatinga-Filho & Leal 2007; Brito 2011) and one submitted leaf-litter samples to Berlese-Tullgren funnels and extracted insects from vegetation with entomological umbrella (Santos et al. 1999).

RESULTS AND DISCUSSION

We recorded 68 ant species, 32 genera and 10 subfamilies in the 150 m² of leaf-litter sampled in the arboreal caatinga in Milagres, Bahia. Active search for ants in the study areas resulted in 23 species, 12 genera and five subfamilies. In the samples preserved in alcohol in the Laboratory of Entomology (UEFS), we recognized 42 species, 22 genera and eight subfamilies. Summing the information from the 28 surveys available in literature, we identified 151 species of 63 genera in 11 ant subfamilies.

In recent years, submission of standardized samples of the leaf-litter to Winkler extractors have been adopted worldwide to study this microhabitat, little investigated although explored by many relatively small and cryptic ant species (Delabie et al. 2000). In areas of arboreal caatinga, this technique has been applied only recently (Silva 2011a; Macêdo 2012). Our Winkler extracted samples in Milagres allowed us to add 23 new records of species for the biome (highlighted in Table I with an asterisk), which means an increase of almost 14% in the number of previously recognized species for the Caatingas, including a recently described species of Oxyepoecus (Myrmicinae, Solenopsidini), O. regularis Ulysséa & Brandão, 2012.

It is important to note that updated global and regional catalogues of ant species were published in the last decade (Bolton 2003; Bolton et al. 2006; Palácio & Fernández 2003), new mechanisms were made available for online identification (Longino 2005; Sarnat 2008), and tribes as well as widely distributed genera and subfamilies were partially or totally revised: Amblyoponinae (Yoshimura & Fisher 2012); Crematogaster of Costa Rica (Longino 2005); Dacetini (Baroni-Urbani & De Andrade 2007); Neotropical Gnampto-genys (Lattke et al. 2007); Linepithema (Wild 2007); Megalomyrmex (Brandão 1990, 2003); Mycetophylax, Paramycetophylax and Kalathomyrmex (Klingenberg & Brandão 2009), Oxyepoecus (Albuquerque & Brandão 2004, 2009); Pheidole of the Americas (Wilson 2003); Prenolepis (LaPolla et al. 2010), Trachymyrmex species groups (Mayhé-Nunes & Brandão 2002, 2005, 2007) and Wasmannia (Longino & Fernández 2007). These revisions, associated with the direct comparison with specimens deposited in the collection of Hymenoptera in the MZSP, allowed the identification at species-level of most of the analyzed material.

Given the difficulty in assigning names to several Neotropical ant species, however, and the lack of recent taxonomic revisions for many of its richest genera - Atta, Azteca, Brachymyrmex, Cyphomyrmex, Hypoponera, Nylanderia and Solenopsis - most studies taken from the literature shows a high number of morphospecies. As it was not possible to match these morphospecies, housed in different institutions and collections, to the Milagres morphospecies and to the alcohol LENT/UEFS samples, we listed the genera identified only with morphospecies (i.e. Carebara, Hypoponera, Myrmelachista, Myrmicocrypta, Nesomyrmex, Nylanderia, Ochetomyrmex and Trachymyrmex; see Table I) and all cases where confident assignments of specific names have been possible. Currently 11 subfamilies, 61 genera and 173 species of Formicidae are recognized as occurring in the Caatin-gas biome (Tables I and ^II).

Further, we are fully aware that this first list of ant species from the Caatingas represent an underestimation. Pheidole, for instance, is a worldwide genus, dominant in different habitats and with the largest number of described species (about 1,000; see Wilson 2003), of which 462 are recorded for the Neotropical region (Fernández & Sendoya 2004). Because there is no appropriate identification key for the Neotropical species (Lach et al. 2010), most specimens of Pheidole collected are in general separated into morpho-species. In Milagres, we sampled 13 morphospecies of this genus, not accounted in the list.

Our study shows that the Bahia state presents the highest number of sampled areas in Caatinga vegetation for ants (16), followed by Pernambuco (8), Ceará (4), Alagoas (3), Minas Gerais (2), Paraíba (2), Piauí (2), Sergipe (1) and Rio Grande do Norte (1) (Fig. 1 and Table III). However, these 39 studied areas represent an insufficient sampling effort when one considers the dimension of the biome. Paraiba and Rio Grande do Norte states, both with significant areas of Caatingas (around 5.000.000 ha), and Caatingas enclaves in the Amazon and Atlantic Forests show the most critical situation. Nothing is known about the ant fauna in these regions that should be prioritized accordingly in order to improve our knowledge of the diversity of ants in the Caatingas.

The paucity of knowledge on ants occurring in the Caatingas reflects other aspects. For instance, the Caatinga is the less protected biome by the Sistema Nacional de Unidades de Conservação/SNUC (completely protected areas = 1.03% and areas of sustainable use = 6.3%) and indigenous territories (0.22%) (MMA, ICMBIO & TNC 2008). Historically, it has suffered from intensive anthropogenic disturbances, mainly due to deforestation and agriculture, presenting nowadays significant degradation and an intense process of desertification in different places (Garda 1996; MMA 2002; Leal et al. 2005). Tropical and subtropical dry forests are among the most threatened biomes in the world, but comparatively received less attention from conservationists and ecologists (Pennington et al. 2000; Quesada and Stoner 2004; Lopes 2006; Santos et al. 2011).

Our survey, although circumscribed to one region of arboreal caatinga, revealed a significant number of ant species, in comparison with similar efforts in other biomes. However, obtaining more samples in adjacent areas and in other regions covered by the same vegetation type shall most probably reveal further species. Beyond that, in order to improve the list of Caatinga's ant fauna, more surveys employing diverse collecting techniques are needed in all Brazilian states that present this vegetation and in all phytophysiognomies of this biome.

ACKNOWLEDGMENTS

We would like to thank Mr. Antônio, owner of the area in Milagres, for allowing us to survey ants in his properties; the Transport Sector of the UEFS; Anderson M. Medina and Elkiaer M. Campos for their help in the field trips, with support provided by CNPq (grant nr. 0013/2009 to Dr. Gilberto M. de Mendonça Santos and scholarship nr. 148056/2010 - 5) and CAPES. We also thank Rodrigo M. Feitosa, Rogério R. da Silva and Nicolas L. de Albuquerque (MZSP) for their help in the identification of some of the material.

Received 4 October 2012;

Accepted 3 April 2013

Associate Editor: Maria Cristina Gaglianone

• Ab'Saber, A.N. (ed.). 2003. Os domínios da natureza no Brasil: potencialidades paisagísticas São Paulo, Ateliê Editorial, 160 p.
• Agosti, D., Majer, J.D., Alonso, L.E. & Schultz, T.R. (eds.). 2000. Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, Biological Diversity Handbook Series, Smithsonian Institution Press, 280 p.
• Agosti, D. & Johnson, N.F. 2005. Antbase. World Wide Web electronic publication. Available at: antbase.org (accessed 03 September 2012).
• Albuquerque, N.L. de & Brandão, C.R.F. 2004. A revision of the Neotropical Solenopsidini ant genus Oxyepoecus Santschi, 1926 (Hymenoptera: Formicidae: Myrmicinae). 1. The Vezenyii species-group. Papéis Avulsos de Zoologia 44: 55 - 80.
• Albuquerque, N.L. de & Brandão, C.R.F. 2009. A revision of the Neotropical Solenopsidini ant genus Oxyepoecus Santschi, 1926 (Hymenoptera: Formicidae: Myrmicinae). 2. Final. Key for species and revision of the Rastratus species-group. Papéis Avulsos de Zoologia 49: 289 - 309.
• Alonso, L.E. 2000. Ants as indicators of diversity, p. 80 - 88. In: Agosti, D., Majer, J.D., Alonso, L.E. & Schultz, T.R. (eds.). Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, Biological Diversity Handbook Series, Smithsonian Institution Press, 280 p.
• Aquad, M. dos S. 1986. Clima da Caatinga, p. 37 - 48. In: Anais do 1ş Simpósio sobre a Caatinga e sua exploração racional 1 Bahia, Universidade Estadual de Feira de Santana.
• Baroni-Urbani, C. & De Andrade, M.L. 2007. The ant tribe Dacetini: limits and constituent genera, with descriptions of new species (Hymenoptera, Formicidae). Annali del Museo Civico di Storia Naturale "Giacomo Doria" 95: 1 - 191.
• Bolton, B. 2000. The Ant Tribe Dacetini. Memoirs of the American Entomological Institute 65: 1 - 1028.
• Bolton, B. 2003. Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute 71: 1 - 370.
• Bolton, B., Alpert, G., Ward, P.S. & Naskrecki, P. 2006. Bolton's Catalogue of Ants of the World 1758 - 2005 Cambridge, Harvard University Press. CD-ROM.
• Brandão, C.R.F. 1990. Systematic revision of the Neotropical ant genus Megalomyrmex, Forel (Hymenoptera: Formicidae: Myrmicinae), with the description of thirteen new species. Arquivos de Zoologia 31: 411 - 481.
• Brandão, C.R.F. 1995. Formigas dos Cerrados e Caatingas. Habilitation thesis, Universidade de São Paulo, 147 p.
• Brandão, C.R.F. 2003. Further revisionary studies on the ant genus Megalomyrmex Forel (Hymenoptera: Formicidae: Myrmicinae: Solenopsidini). Papéis Avulsos de Zoologia 43: 145 - 159.
• Brandão, C.R.F & Yamamoto, C.I. 2004. Invertebrados da Caatinga, p. 135 - 140. In: Cardoso, J.M., Tabarelli, M., Fonseca, M.T. & Lins, L.V. (orgs.) Biodiversidade da Caatinga: áreas e ações prioritárias para a conservação. Brasília, MMA/EFPE/Conservação Internacional do Brasil/EMBRAPA, 382 p.
• Brito, A.F. 2011. Riqueza e Estrutura de Comunidades de Formigas (Hymenoptera: Formicidae) Associadas a Flores em Fragmentos de Caatinga na Região de Milagres, Bahia. M. Sc. thesis, Universidade Estadual de Feira de Santana, 60 p.
• Brown Jr., K.S. 1997. Diversity, disturbance, and sustainable use of Neotropical forests: insects as indicators for conservation monitoring. Journal of Insect Conservation 1: 25 - 42.
• Brühl, C.A., Mohamed, V. & Linsenmair, K.E. 1999. Altitudinal distribution of leaf-litter ants along a transect in primary forests on Mount Kinabalu, Sabah, Malaysia. Journal of Tropical Ecology 15: 265 - 277.
• Campos, E.M. 2012. Scarabaeidae (Coleoptera) na Caatinga baiana: Distribuição e Planejamento de Pesquisas. M. Sc. thesis, Universidade Estadual de Feira de Santana, 50 p.
• Cardoso, D.B.O.S. & Queiroz, L.P. de. 2010. Caatinga no contexto de uma Metacomunidade: Evidências da Biogeografia, Padrões Filogenéticos e Abundância de Espécies em Leguminosas, p. 241 - 260. In: Carvalho, C.J.B. de & Almeida, E.A.B. (orgs.) Biogeografia da América do Sul: padrões e processos. São Paulo, Roca, 328 p.
• Carvalho, K.S., Nascimento, I.C., Koch, E.B.A., Souza, A.S., Santos, V.S., Ramos, D.L., Carneiro, M.A.F., Souza, A.L.B., Dourado, E.A.S., Santos Jr., J.G. & Zina, J. 2011. Diversidade de Formigas da Caatinga e Mata de Cipó (Domínio da Mata Atlântica) no Semi-árido Baiano. In: Anais do XX Simpósio de Mirmecologia e I Encuentro de Mirmecologistas de las Américas. Petrópolis.
• De Andrade, M.L. & Baroni-Urbani, C. 1999. Diversity and adaptation in the ant genus Cephalotes, past and present. Stuttgarter Beiträge zur Naturkunde, Serie B 271: 1 - 889.
• Delabie, J.H.C., Agosti, D. & Nascimento, I.C. 2000. Litter ant communities of the Brazilian Atlantic rain forest region, p. 1 - 17. In: Agosti, D., Majer, J.D., Alonso, L.T. & Schultz, T.R. (eds.). Sampling Ground-dwelling Ants: Case Studies from the World's Rain Forests. Perth, Curtin University School of Environmental Biology Bulletin 18, 75 p.
• Delsinne, T., Leponce, M., Theunis, L., Braet, Y. & Roisin, Y. 2008. Rainfall influences ant sampling in dry forests. Biotropica 40: 590 - 596.
• Dietz, B.H. 2004. Uma revisão de Basicerotini Brown, 1948 (Formicidae: Myrmicinae), suas relações filogenéticas internas e com outras tribos dacetíneas (Dacetini e Phalacromyrmecini). Ph. D. Dissertation, Universidade de São Paulo, 245 p.
• Espirito-Santo, M.M. do, Barbosa, R.S., Melo Junior, A.F. de, Medeiros dos Santos, A., Oliveira D.A., Anaya, F.C., Neves, F.S., Fernandes, G.W., Sanchez-Azofeifa, G.A., Valério, H.M., Liete, L.O., Zaza Borges, M.A., Magalhães Veloso, M. das D., Fagundes, M., Pimenta, M.A., Lopes de Faria, M., Reis Junior, R. & Ferreira Nunes, Y.R. 2010. Nota técnica: Esclarecimentos sobre a situação ecológica e sócio-econômica das Matas Secas do norte-mineiras. Available at: http://tropi-dry.eas. ualberta.ca/3_data pub_publications.html (accessed 10 August 2011).
• Fernández, F. & Sendoya, S. 2004. List of Neotropical ants (Hymenoptera: Formicidae). Biota Colombiana 5: 3 - 93.
• Floren, A., Biun, A. & Linsenmair, K.E. 2002. Arboreal ants as key predators in tropical lowland rainforest trees. Oecologia 131: 137 - 144.
• Folgarait, P.J. 1998. Ant biodiversity and its relationship to ecosystem functioning: A review. Biodiversity and Conservation 7: 1221 - 1244.
• Forel, A. 1911. Ameisen des Hern Prof. v. Ihering aus Brasilien (São Paulo usw.) nebst einigen anderen aus Südamerika und Afrika (Hyme-noptera). Deutsche Entomologische Zeitschrift 1911: 285 - 312.
• Freitas, A.V.L., Francini, R.B. & Brown Jr., K.S. 2004. Insetos como indicadores ambientais, p. 125 - 151. In: Cullen Jr., L., Rudran, R. & Valladares-Padua, C. (orgs.). Métodos de estudos em biologia da conservação e manejo da vida silvestre. Curitiba, Editora UFPR, 665 p.
• Garda, E.C. 1996. Atlas do meio ambiente do Brasil. Brasília, Editora Terra Viva, 160 p.
• Garro, R.N., Campos, R.I., Castro, F.S. & Neves, F.S. 2011. Dominância e co-ocorrência de espécies de formigas atraídas por iscas em Floresta Estacional Decidual. In: Anais do XX Simpósio de Mirmecologia e I Encuentro de Mirmecologistas de las Américas. Petrópolis.
• Hölldobler, B. & E. O. Wilson. 1990. The Ants. Cambridge, Harvard University Press, 732 p.
• Ibraimo, M.M., Schaefer, C.E.G.R., Ker, J.C., Lani, J.L., Rolim-Neto, F.C., Albuquerque, M.A. & Miranda, V.J. 2004. Gênese e micromorfologia de solos sob vegetação xeromórfica (Caatinga) na região dos Lagos (RJ). Revista Brasileira de Ciência do Solo 28: 695 - 712.
• Ihering, H. von & Ihering, R. von 1911. O Museu Paulista nos annos de 1906 a 1909. Revista do Museu Paulista VIII: 1 - 22.
• Kaspari, M. & Majer, J.D. 2000. Using Ants to Monitor Environmental Change, p. 89 - 98. In: Agosti, D., Majer, J.D., Alonso, L.E. & Schultz, T.R. (eds.). Ants: Standard Methods for Measuring and Monitoring Biodiversity. Washington, Biological Diversity Handbook Series, Smithsonian Institution Press, 280 p.
• Kempf, W.W. 1971. A preliminary review of the Ponerine ant genus Dinoponera Roger (Hymenoptera: Formicidae). Studia Entomologica 14: 368 - 394
• Kempf, W.W. 1975. A revision of the Neotropical ponerine ant genus Thaumatomyrmex Mayr (Hymenoptera: Formicidae). Studia Entomologica 15: 95 - 126.
• Klingenberg, C. & Brandão, C.R.F. 2009. Revision of the fungus-growing ant genera Mycetophylax Emery and Paramycetophylax Kusnezov rev. stat., and description of Kalathomyrmex n. gen. (Formicidae: Myrmicinae: Attini). Zootaxa 2052: 1 - 31.
• Kremen, C., Cowell, R.K., Erwin, T.L., Murphy, D.D., Noss, R.F. & Sanjayan, M.A. 1993. Terrestrial arthropod assemblages - Their use in conservation planning. Conservation Biology 7: 796 - 808.
• Lach, L., Parr, C.L. & Abbott, K.L. (eds.). 2010. Ant Ecology. New York, Oxford University Press, 429 p.
• LaPolla, J.S., Brady, S.G. & Shattuck, S.O. 2010. Phylogeny and taxonomy of the Prenolepis genus-group of ants (Hymenoptera: Formicidae). Systematic Entomology 35: 118 - 131.
• Lattke, J.E., Fernández, F. & Palacio, E.E. 2007. Identification of the species of Gnamptogenys Roger in the Americas, p. 254 - 270. In: Snelling, R.R., Fisher, B.L. & Ward, P.S. (eds.). Advances in ant systematics (Hymenoptera: Formicidae): homage to E. O. Wilson - 50 years of contributions. Memoirs of the American Entomological Institute, 690 p.
• Leal, I.R. 2002. Diversidade de Formigas do Estado de Pernambuco, p. 483 - 492. In: Tabarelli, M. & Silva, J.M.C. (eds.). Diagnóstico da diversidade de Pernambuco. Recife, Editora Massangana, 722 p.
• Leal, I.R. 2003. Diversidade de formigas em diferentes unidades de paisagem na Caatinga, p. 435 - 462. In: Leal, I.R., Tabarelli, M. & Silva, J.M.C. (eds.). Ecologia e Conservação da Caatinga. Recife, Editora UFPE, 804 p.
• Leal, I.R., Silva, J.M.C., Tabarelli, M. & Lacher Jr., T.E. 2005. Changing the course of biodiversity conservation in the Caatinga of Northeastern Brazil. Conservation Biology 19: 701 - 703.
• Leal, I.R., Wirth, R. & Tabarelli, M. 2007. Seed Dispersal by Ants in the Semi-arid Caatinga of North-east Brazil. Annals of Botany 99: 885 - 894.
• Lewinsohn, T.M., Freitas, A.V.L. & Prado, P.I. 2005. Conservation of terrestrial invertebrates and their habitats in Brazil. Conservation Biology 19: 640 - 645.
• Longino, J.T. 2005. Key to Crematogaster of Costa Rica. Available at: http://academic.evergreen.edu/projects/ants/genera/crematogaster/key.html (accessed 17 March 2011).
• Longino, J.T. & Fernández, F. 2007. Taxonomic review of the genus Wasmannia, p. 271 - 289. In: Snelling, R.R., Fisher, B.L. & Ward, P.S. (eds.). Advances in ant systematic (Hymenoptera: Formicidae) homage to E. O. Wilson - 50 years of contributions. Memoirs of the American Entomological Institute, 690 p.
• Lopes, P.P. 2006. Besouros no Semi-árido Brasileiro: Ilustres, mas quase desconhecidos, p. 95 - 98. In: Queiroz, L.P. de, Rapini, A. & Giulietti, A.M. (eds.). Rumo ao amplo conhecimento da biodiversidade do Semi-árido Brasileiro. Brasília, Ministério da Ciência e Tecnologia, 144 p.
• Macêdo, L.S.S.R. 2012. Riqueza e Padrões de Atividade em Comunidades de Formigas (Hymenoptera: Formicidae) numa Área de Caatinga. M. Sc. thesis, Universidade Estadual de Feira de Santana, 69 p.
• Mackay, W.P. 1993. A review of the New World ants of the genus Dolichoderus (Hymentoptera: Formicidae). Sociobiology 22: 1 - 148.
• Majer, J.D. 1983. Ants: bio-indicators of minesite rehabilitation, land use, and land conservation. Environmental Management 7: 375 - 383.
• Marques, A.A.B., Fontana, C.S., Velez, E., Bencke, G.A., Schneider, M. & Reis, R.E. 2002. Lista de Referência da Fauna Ameaçada de Extinção no Rio Grande do Sul - Decreto Nş 41.672, de 11 de Junho de 2002. Porto Alegre, FZB/MCT-PUCRS/PANGEA, 52 p.
• Marques, L.C. & Soares, I.M.F. 2011. Caracterização da Mirmecofauna (Hymenoptera: Formicidae) da Mata da Pororoca e Adjacências na Reserva Ecológica do Raso da Catarina, Bahia, Brasil. In: Anais do XX Simpósio de Mirmecologia e I Encuentro de Mirmecologistas de las Américas. Petrópolis.
• Mayhé-Nunes, A.J. & Brandão, C.R.F. 2002. Revisionary studies on the attine ant genus Trachymyrmex Forel. Part 1: Definition of the genus and the Opulentus group (Hymenoptera: Formicidae). Sociobiology 40: 667 - 698.
• Mayhé-Nunes, A.J. & Brandão, C.R.F. 2005. Revisionary studies on the attine ant genus Trachymyrmex Forel. Part 2: The Iheringi group (Hymenoptera: Formicidae). Sociobiology 45: 271 - 305.
• Mayhé-Nunes, A.J. & Brandão, C.R.F. 2007. Revisionary studies on the attine ant genus Trachymyrmex Forel. Part 3: The Jamaicensis group (Hymenoptera: Formicidae). Zootaxa 1444: 1 - 27.
• Medina, A. M. 2012. Variação Temporal e Preferência Alimentar em uma comunidade de Scarabaeinae em um fragmento de Caatinga (Milagres, BA). M. Sc. thesis, Universidade Estadual de Feira de Santana, 94 p.
• MMA, Ministério do Meio Ambiente. 2002. Biodiversidade Brasileira: Avaliação e Identificação de Áreas Prioritárias para Conservação, Utilização Sustentável e Repartição de Benefícios da Biodiversidade Brasileira. Série Biodiversidade. Brasília, 404 p.
• MMA, Ministério do Meio Ambiente. 2003. Mapa das Áreas Prioritárias para a Conservação, Utilização Sustentável e Repartição de Benefícios da Biodiversidade Brasileira. Secretaria de Biodiversidade e Florestas, PROBIO. Brasília. Available at: http://www.mma.gov.br/estruturas/chm/_arquivos/maparea.pdf (accessed 13 September 2011).
• MMA, Ministério do Meio Ambiente. 2007. Áreas Prioritárias para Conservação, Uso Sustentável e Repartição de Benefícios da Biodiversidade Brasileira: Atualização - Portaria MMA n°9, de 23 de janeiro de 2007. Brasília, Secretaria de Biodiversidade e Florestas, 300 p.
• MMA, Ministério do Meio Ambiente, ICMBIO, Instituto Chico Mendes de Conservação da Biodiversidade & TNC, The Nature Conservancy. 2008. Mapa de Unidades de Conservação e Terras Indígenas do bioma Caatinga. Brasília. Available at: http://www.mma.gov.br/estruturas/203/arquivos/mapa_das_ucs.pdf (accessed 23 April 2011).
• Neves, F.S., Braga, R.F., Espírito-Santo, M.M. do, Delabie, J.H.C., Fernandes, G.W. & Sánchez-Azofeifa, G.A. 2010. Diversity of Arboreal Ants in a Brazilian Tropical Dry Forest: Effects of Seasonality and Successional Stage. Sociobiology 56: 177 - 194.
• Nunes, F.A., Martins Segundo, G., Vasconcelos, Y.B., Azevedo, R. & Quinet, Y. 2011. Ground-foraging ants (Hymenoptera: Formicidae) and rainfall effect on pitfall trapping in a deciduous thorn woodland (Caatinga), Northeastern Brazil. Revista de Biología Tropical 59: 1637 - 1650.
• Oliveira, G.V., Freire, C.B., Silva, K.S., Martins, F.R.S. & Corrêa, M.M. 2009. Diversidade de formigas de áreas preservadas e em regeneração de Caatinga da Floresta Nacional Contendas do Sincorá, sudoeste da Bahia. In: Anais do IX Congresso de Ecologia do Brasil. São Lourenço.
• Palácio, E.E. & Fernández, F. 2003. Claves para las subfamilias y géneros, p. 233 - 260. In: Fernández, F. (ed.). Introducción a las hormigas de la región Neotropical. Bogotá, Instituto de Investigación de Recursos Biológicos Alexander Von Humboldt, xxvi+398 p.
• Pennington, R.T., Prado, D.A. & Pendry, C. 2000. Neotropical seasonally dry forests and quaternary vegetation changes. Journal of Biogeography 27: 261 - 273.
• Prado, D.E. 2003. As Caatingas da América do Sul, p. 3 - 73. In: Leal, I.R., Tabarelli, M. & Silva, J.M.C. (eds.). Ecologia e conservação da Caatinga. Recife, Editora UFPE, 804 p.
• Quesada, M. & Stoner, K.E. 2004. Threats to the conservation of the tropical dry forest in Costa Rica, p. 266 - 280. In: Frankie, G.W., Mata, A. & Vinson, S.B. (eds.). Biodiversity conservation in Costa Rica: Learning the lessons in a seasonal dry forest. Berkeley, University of California Press, 352 p.
• Quinet, Y.P. & Tavares, A.A. 2005. Formigas (Hymenoptera: Formicidae) da área Reserva Serra das Almas, Ceará, p. 327 - 348. In: Araújo, F.S., Rodal, M.J.N. & Barbosa, M.R.V. (eds.). Análise das variações da Biodiversidade do Bioma da Caatinga: Suporte a estratégias regionais de conservação. Brasília, Ministério do Meio Ambiente, 434 p.
• Quinet, Y., Tekule, N. & Biseau, J.C. de 2005. Behavioural interactions between Crematogaster brevispinosa rochai Forel (Hymenoptera: Formicidae) and two Nasutitermes species (Isoptera: Termitidae). Journal of Insect Behavior, 18: 17 p. doi: 10.1007/s10905 - 005 - 9343 - y.
• Ribas, C.R., Campos, R.B.F., Schmidt, F.A. & Solar, R.R.C. 2012. Ants as Indicators in Brazil: A Review with Suggestions to Improve the Use of Ants in Environmental Monitoring Programs. Psyche 2012: ID 636749, 23 p. doi:10.1155/2012/636749.
• Santana, J.A. da S. & Souto, J.S. 2006. Diversidade e Estrutura Fitossociológica da Caatinga na Estação Ecológica do Seridó-RN. Revista de Biologia e Ciências da Terra 6: 232 - 242.
• 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 Semidecídua) em Itatim, Bahia, Brasil. Sitientibus 20: 33 - 43.
• Santos, J.C., Leal, I.R., Almeida-Cortez, J.S., Fernandes, G.W. & Tabarelli, M. 2011. Caatinga: the scientific negligence experienced by a dry tropical forest. Tropical Conservation Science 4: 276 - 286.
• Sarnat, E.M. 2008. PIAkey: Identification guide to ants of the Pacific Islands, Edition 2.0, Lucid v. 3.4. USDA/APHIS/PPQ Center for Plant Health Science and Technology and University of California - Davis. Available at: http://keys.lucidcentral.org/keys/v3/piakey/piakeyv2.html (accessed 12 March 2011).
• Silva, E.M. 2011a. Estrutura da comunidade de formigas (Hymenoptera-Formicidae) associada a duas paisagens de Caatinga em Milagres-Bahia. M. Sc. thesis, Universidade Estadual de Feira de Santana, 47 p.
• Silva, T.G.M. 2011b. Diversidade de Formicidae em Florestas Estacionais Decíduas em diferentes escalas temporais e espaciais. Ph. D. dissertation, Universidade Federal de Viçosa, 82 p.
• Silva, G.L. de, Santos, E.M. dos & Gomes, J.P. 2010. Predação de ovos de Corythomantis greeningi Boulenger, 1896 (Anura, Hylidae) por Solenopsis invicta Buren, 1972 (Formicidae: Myrmicinae). Biotemas 23: 153 - 156.
• Tabatinga-Filho, G.M. & Leal, I.R. 2007. Influência da presença de formigas na ocorrência de visitantes florais em Calotropis procera (Ait) R. Br, p. 1 - 2. In: Anais do VIII Congresso de Ecologia do Brasil. Caxam-bu.
• Ulysséa, M.A. & Brandão, C.R.F. 2012. A new ant species of Oxyepoecus (Hymenoptera: Formicidae: Myrmicinae), with the description of Oxyepoecus browni gyne and new records for the genus. Papéis Avulsos de Zoologia 52: 167 - 173.
• Vanzolini, P.E., Ramos-Costa, A.M.M. & Vitt, L.J. 1980. Répteis das Caatingas. Rio de Janeiro, Academia Brasileira de Ciências, 161 p.
• Velloso, A.L., Sampaio, E.V.S. B. & Pareyn, F.G.C. (eds.) 2002. Ecorregiões propostas para o Bioma Caatinga. Recife, Associação Plantas do Nordeste; Instituto de Conservação Ambiental The Nature Conservancy do Brasil, 80 p.
• Werneck, F.P. 2011. The diversification of eastern South American open vegetation biomes: Historical biogeography and perspectives. Quaternary Science Reviews 30: 1 - 19.
• Wild, A.L. 2007. Taxonomic revision of the ant genus Linepithema (Hymenoptera: Formicidae). University of California Publications in Entomology 126: 1 - 151.
• Wilson, E.O. 1987. Causes of ecological success: The case of the ants. The Journal of Animal Ecology 56: 1 - 9.
• Wilson, E.O. 2003. Pheidole in the New World: A dominant, hyperdiverse ant genus. Cambridge, Harvard University Press, 794 p.
• Yoshimura, M. & Fisher, B. 2012. A Revision of male ants of the Malagasy Amblyoponinae with resurrections of the genera Stigmatomma and Xymmer PLoS ONE 7: e33325. doi:10.1371/journal.pone.0033325.
• Zappi, D. 2008. Fitosifionomia da Caatinga à Cadeia do Espinhaço. Megadiversidade 4: 33 - 37.

Publication Dates

• Publication in this collection
  28 May 2013
• Date of issue
  June 2013

History

• Received
  04 Oct 2012
• Accepted
  03 Apr 2013