ABSTRACT INTRODUCTION. Universidade Estadual de Goiás, Brazil, Faculdade Araguaia, Goiânia-GO, Brazil,

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1 Termite (Isoptera) Assemblages in Some Regions of the Goiás State, Brazil by Hélida Ferreira da Cunha 1, Diogo Andrade Costa 2, Divino Brandão 3 ABSTRACT Termites are the dominant fauna in tropical soils, having an important role in nutrient cycling processes and soil formation. They are a dominant component of the Cerrado fauna, reaching high densities in some areas. Quantitative and qualitative assessment of the termite assemblages were carried out in natural (Riparian Forests, Mesophytic Interfluvial Forest, Cerrado and Cerradão ) and impacted areas (pastures and plantations of sugar cane, banana, Eucalyptus trees, Bahia-coconut and Guariroba) in eleven municipalities of the Goiás state in Brazil. In each environment, 8 quadrats of 25 m 2 were studied. The quadrats were located 50 m apart from the habitat edge and 30 m between each other. Seventy-seven species belonging to the Families Rhinotermitidae (Subfamilies Coptotermitinae, Rhinotermitinae and Heterotermitinae) and Termitidae (Subfamilies Apicotermitinae, Nasutitermitinae and Termitinae) were collected. The species with the highest number of colonies were Anoplotermes sp. 4 (68), Anoplotermes sp. 2 (57), Nasutitermes kemneri (46) and Ruptitermes sp. (34). Anoplotermes sp. 4 and N. kemneri were found in 10 of the 11 sampled municipalities while Anoplotermes sp. 2 was found in all of them. All the sampled localities have high equitability ranging form 0.85 to 0.94 because all the species were registered once or twice. Most of the species found feed upon decomposing organic matter and/or are litter foragers. Of the 77 species sampled, we were not able to identify 26 (designed as morphospecies). Some of them might be new species. Keywords: Cerrado, diversity, Isoptera, pasture, plantation, termite. INTRODUCTION Termites (Order Isoptera) are social insects with ca described species (Cancello & Schlemmermeyer 1999; Constantino 1998, 1999). Around Universidade Estadual de Goiás, Brazil, cunhahf@yahoo.com.br 2 Faculdade Araguaia, Goiânia-GO, Brazil, diogoacosta@yahoo.com.br 3 Universidade Federal de Goiás, Goiânia-GO, Brazil, dbrandao@icb.ufg.br

2 2 Sociobiology Vol. 47, No. 2, 2006 species occur in the Neotropics, a diversity only overcome by Oriental and Ethiopian regions where most studies are concentrated (Cancello & Schlemmermeyer 1999; Constantino 1998, 1999). Well known by their economic importance as agricultural and wood pests, the termites are detritivorous and represent one of the dominant groups in the fauna of tropical ecosystems, having an important role in nutrient cycling processes and soil formation (Black & Okwakol 1997; Cancello & Schlemmermeyer 1999; Constantino 2002; Davies et al. 1999; Eggleton et al. 1996). According to Eggleton & Bignell (1995) and Overal (2001), the group represents almost 10% of the animal biomass at the tropics. In tropical ecosystems, from open vegetation areas such as the Cerrado sensu sticto to rainforests, the ecological importance of termites has been poor studied (Cancello & Schlemmermeyer 1999), and the inventories that exist concentrate in few localities (Constantino in press). In Latin America, Brazil is the country where the termite fauna has been best studied, having ca. 280 described species (Constantino 1998, 1999). Termites are a dominant and conspicuous component of the Cerrado fauna, reaching overwhelming densities in some areas. Considering only the open areas, termite richness in the Cerrado can reach ca. 150 species (Constantino in press). Some of the species, such as Cornitermes cumulans can be considered keystone species because of their great abundance and environmental impact (Redford 1984). This work presents a quantitative and qualitative assessment of the termite fauna in some municipalities in Goiás state (Brazil), designed to represent the different meso-regions. The study intended to improve the knowledge on the termite fauna of the state, by collecting in regions where sampling was never carried out before. Study Area The study area includes natural and disturbed habitats in 11 municipalities of the state of Goiás (Fig. 1). This is the only Brazilian state that is totally embedded in the Cerrado Biome. The Cerrado is the second largest of Brazil s major biomes, occupying 22% of the national territory (Oliveira Filho & Ratter 2002). The biome has altitudes varying from 300 m to 1800 m, with a predominance of latosols (Oliveira Filho & Ratter 2002). The climate is characterized by a rainy season from October to March followed by a dry period from April to September. Rain fall ranging from 800 to 2000 mm and

3 da Cunha, H.F. et al. Termite Assemblages in Goiás 3 Fig. 1. Sampling Municipalities in Goiás State (Brazil).

4 4 Sociobiology Vol. 47, No. 2, 2006 annual temperature from 18ºC to 28ºC along the year (Dias 1992). Several vegetal formations occur at the Cerrado biome, including grasslands, savanna and forests (Eiten 1972; Coutinho 1978) like the cerrado sensu lato (represented by a physiognomic gradient that includes the campo limpo, campo sujo, the cerrado sensu stricto, and the cerradão), riparian forest, and the mesophytic interfluvial forest. Sample collection in disturbed habitats was made in grasslands and in plantations of sugar cane, banana, Eucalyptus trees, Bahia-coconut and Guariroba trees, where available in each municipality, and sample collection in natural areas was made in the three vegetation types cited above, remnant in the legal protected area of private properties. Some of these areas presented light degrees of habitat alteration such as selective wood extraction, presence of cattle or trails. Areas with more heavy alterations were not used. MATERIALS & METHODS In each environment, 50 m away from the edge, eight 25 m 2 quadrats were defined, 30 m apart from each other. In each quadrats, a one-hour exhaustive search was carried out in all the places where termites can be found (50 cm deep in the ground, soil surface, dead wood, hypogean nests, epigean nests, arboreal nests and foraging galleries). Samples were identified to species or morphospecies level using taxonomic identification keys, taxonomic notes and revisions (Araujo 1954; Emerson & Banks 1957; Emerson & Banks 1965; Krishna & Araujo 1968; Mathews 1977; Fontes 1985; Cancello 1989; Constantino 1990; Constantino 1995; Cancello et al. 1996; Constantino & De Souza 1997; Constantino 1999; Cancello & Myles 2000; Constantino 2001), and by comparison with the Isoptera collections at the Federal University of Goiás (UFG) and the Zoology Museum of the São Paulo University (MZUSP). Specimens from all the identified species and morphospecies were deposited at the termite collection of the UFG. Richness estimates and diversity indexes were calculated for the sample as a whole and for each sampled municipality, using the software EstimatS (Colwell 2005) using 500 permutations. Parameters such as mean species richness, uniques (mean number of species that occur in one sample), duplicates (mean number of species that occur in two samples), Michaelis-

5 da Cunha, H.F. et al. Termite Assemblages in Goiás Menten (richness estimator that predicts the stabilization of the number of species using the observed richness), Bootstrap (richness estimator based on the species incidence) and Shannon-Wiener diversity index (H) were calculated. Bootstrap is a non-parametric method (can be used in samples with different abundance distributions) that uses the total number of species to estimate the total richness, not being restricted to rare species (Santos 2003). The transformation e H makes possible to analyze Shannon-Wiener diversity in species richness units. The equitability was calculated from Shannon diversity (H) and species richness (S) using the formula E=H/ln(S) (Krebs 1989). Cumulative species curves (collector s curve) were done using mean richness and number of species. To estimate the stabilization point of the curve, the Michaelis-Menten equation was used (Colwell 2005). This curve represents the total number of species registered during data collection when new individuals or samples are added to the entire data set previously obtained (Gotelli & Colwell 2001). Only the quadrats where termite species were found were included in the analysis because some of the methods can not be implemented when there are zero values. Thus, the total number of analyzed quadrants was 208 while the total number of quadrats studied was 244. Colony number was interpreted as the abundance when used to calculate the diversity index. Total number of colonies in each municipality varied according to the habitat availability near sampling locations. The species were classified according to the feeding habit registered for the genus, following Constantino (1999). Trophic guilds used were: humivorous, that feed on humus and soil organic matter; xylophagous, that feed mainly on wood; litter-feeders, for the ones that forage at the surface, cutting leafs from litter or grasses; and intermediate for the ones that do not have a specialized feeding habit. RESULTS Seventy-seven species were collected (Appendix 1) in 244 quadrats (the coconut plantation at the Matrinchã municipality had enough area for just four quadrats), for a total of 694 colonies belonging to the families Rhinotermitidae (Subfamilies Coptotermitinae, Rhinotermitinae and Heterotermitinae) and Termitidae (Subfamilies Apicotermitinae, Nasutitermitinae and Termitinae). 5

6 6 Sociobiology Vol. 47, No. 2, 2006 The species that had more colonies were: Anoplotermes sp.4 (68), Anoplotermes sp.2 (57), Nasutitermes kemneri (46) and Ruptitermes sp (34). Anoplotermes sp.2 was the only species that was present in all sampled municipalities, while Anoplotermes sp.4 and N. kemneri were present in ten out of eleven municipalities. The mean number of colonies for a species was 9.01 ± and for 40% of the species, there were just one or two colonies found (Fig. 2). In 225 quadrats (91%) one to five species were found (3.34 ± 1.8) and no termites were found in 38 of the sampled quadrants (15%) (Fig. 3). The collector s curve (Fig. 4) began to stabilize with one third of the samples. The stabilization of the richness was predicted to occur after 54 samples, when just 51 species were collected. Richness estimators of Michaelis-Menten and Bootstrap overlapped when 80 species were collected in a total of 142 samples. Richness and diversity estimates are found in Table 1. Comparing H with e H it was verified that the expected diversity was higher than the observed diversity. The equitability was high in all the municipalities since most of the species were registered only once or twice, rendering a more homogeneous distribution. The number of unique and duplicate species together represented more than half of the total species richness in the majority of the sampled municipalities (Table 1). This makes the termites a very particular group since most of the species have restricted distributions. For example, in Matrinchã and Alto Paraíso just three species presented more than two colonies. The majority of the species in all the studied municipalities feed on soil and decomposing organic matter and/or foraged on the litter (Fig. 5). DISCUSSION This work is the first study carried in the Goiás State, with an extensive sampling that included eleven municipalities in different regions. Previous studies were all done in only one locality, generally in areas near Goiânia, the capital of the State. Compared to these previous works, the present study emerges as the greatest inventory of termite species (77) in Goiás, excluding the Federal District. Brandão & Souza (1998) collected 33 species in the Cerrado sensu stricto, mesophytic forest and grasslands in the Goiânia municipality, while in this same municipality Ferreira Junior (1999) registered 60 species

7 da Cunha, H.F. et al. Termite Assemblages in Goiás in 15 fragments of mesophytic forest. Oliveira (2003) found 22 species in an altitudinal gradient from Cerrado to Campo Rupestre at the Morro Feio, in the municipality of Hidrolândia. Freitas et al. (2004) collected 29 species in Cerrado, Campo Rupestre and riparian forests at the Serra de Caldas Novas State Park (PESCAN), in the Caldas Novas Municipality. Costa (2005) found 25 inquiline termite species of C. cumulans in 2.25 hectares of Campo Limpo at the Emas National Park (PNE) in the Mineiros municipality. Lima Filho (2005) collected 32 species in an altitude gradient from Cerrado to Campo Rupestre at the Serra dos Pirineus State Park in the Pirenópolis municipality. Most of the species collected by the authors cited above were also registered during the present work and only 24 previously registered species were not present at the eleven municipalities sampled. In this work, from the 77 collected species, 26 morphospecies were not identified, some of them might even be new species. The Apicotermitinae are grouped in a genus because of identification limitations, thus, the real number of species found may be higher. Noirotitermes noiroti was recently described in the Cerrado of the Piauí State (Cancello & Myles 2000) so it does not appear in the New World Isoptera Catalog of Constantino (1998). This species was not collected in Goiás during previous studies. Ferreira Junior (1999) was the only researcher that collected termites of the family Kalotermitidae at Goiás, generally underestimated in rapid inventories, because of their habit of using dry dead wood for shelter. If we consider all the lists in these studies, Nasutitermitinae is the dominant group in richness and abundance, and almost all the epigean and arboreal nests in the Cerrado are made by species from this subfamily. An important characteristic of the Cerrado termite fauna is the great amount of humivorous litter-feeding species, confirming the ecological importance of these species in the processes of ecosystem maintenance, being responsible for the organic matter cycling. The great amount of litter produced at Cerrado areas and the existence of some selective extraction of wood where the species were collected, explain the abundance of humivorous and litter feeders found. A taxonomic revision will probably increase the humivorous proportion since almost all the members of the subfamily Apicotermitinae are characterized as humivorous species. 7

8 8 Sociobiology Vol. 47, No. 2, 2006 Fig. 2. Number of species of termites by frequency of the colonies. Fig. 3. Distribution of the number of quadrats by number of species of the termites. Constantino (1998, 1999) and Cancello & Schlemmermeyer (1999) highlighted that one of the main obstacles for neotropical termitology development has been the lack of taxonomic information. Few groups have been comprehensively revised and there are few well trained taxonomy specialists, a fact that hampers the identification work. Thus, a lot of works with

9 da Cunha, H.F. et al. Termite Assemblages in Goiás 9 Fig. 4. Collector's curve of termites in eleven municipalities of the State of Goiás, Brazil. Fig. 5. Distribution of species of termites according trophic guild. erroneous or incomplete classifications have been published, hindering the process of production and storage of organized information, and making the comparisons between the results of different works difficult.

10 10 Sociobiology Vol. 47, No. 2, 2006 Table 1. Richness and diversity estimators for termite assemblages found in eleven municipalities of the State of Goiás, Brazil. Municipalities quadrats colonies Richness Uniques Duplicates H e H E Aruanã Cocalzinho Cristalina Cristianópolis Hidrolândia Iporá Matrinchã Nova Crixás Pontalina Alto Paraíso Serranópolis All In addition, the comparison of data from published studies is difficult because there is a lack of standardized sampling protocols. The species richness obtained in an inventory is dependent on the local features but also on the sampling effort that can be defined by using the collector s curve. But in most tropical ecosystems this curve rarely stabilizes (Santos 2003). Constantino (1992) and Ferreira Junior (1999) are some of the authors whose studies about diversity of termites presented the collector s curve. Despite the fact that our curve presented a tendency to stabilization when considering the eleven municipalities together, more sampling would be needed to appropriately sample all kinds of habitats considered in this study. There are regions and municipalities in Goiás state where inventories of the termites assemblages were never done. It is believed that the original fauna of these regions will never be completely known since most of them have been significantly disturbed. Most of the area of the state has been and continues to be cleared because of the expansion of the agricultural frontiers. Today s rate of clearance for the Cerrado varies from to km 2 per year and is believed that 55% of the original area of Cerrado was already converted into grasslands, agricultural fields, planted forests and urban areas, among others (Klink & Machado 2005). ACKNOWLEDGMENTS We are grateful to the colleagues that helped during data collection (Lorena, Kleber, Leandro, Gabriel, Guilherme, Danilo, Natália, Luiz). For the logistic

11 da Cunha, H.F. et al. Termite Assemblages in Goiás support and permits granted to enter their properties, we thank Eduardo Luiz Mesquita, Reginaldo Freire, Mozart Martins de Araújo, Sandra, Paulo Roberto de Carvalho, Cacique Arumani of Karajás Indigenous Society, Carlos Alberto Guimarães, André Ferreira da Cunha, Ozório Vilela Neto, Paulo Henrique and Carol. Dr Eliana M. Cancello allowed us to access the MZUSP collection for material comparisons. We also wish to thank FUNAPE-UFG for the financial support. please spell out acronym REFERENCES Araujo, R.L Notes on the genus Paracornitermes Emerson, 1949, with the description of two new species (Isoptera, Termitidae, Nasutitermitinae). Revista Brasileira de Entomologia, 1: Black, H.I.J. & M.J.N. Okwakol Agricultural intensification, soil biodiversity and agroecosystem function in the tropics: the role of termites. Applied Soil Ecology, 6: Brandão, D. & R.F. Souza Effects of deforestation and implantation of pastures on the termite fauna in the Brazilian Cerrado region. Tropical Ecology, 39: Cancello, E.M. & T.G. Myles Noirotitermes noiroti (Isoptera, Termitidae, Nasutitermitinae): a New Genus and New Species from Northeastern Brazil. Sociobiology, 36: Cancello, E.M. & T. Schlerrermeyer Isoptera. In: Brandão, R.F. & Cancello, E.M. (Eds.), Biodiversidade do Estado de São Paulo, Brasil: Síntese do conhecimento ao final do século XX. FAPESP, São Paulo, p Cancello, E.M., D. Brandão, & S.T.P. Amarante Two new Angularitermes species (Isoptera, Termitidae, Nasutitermitinae) from Brazil with a discussion of the cephalic microsculpture of the soldier. Sociobiology, 27: Cancello, E.M Revisão de Cornitermes Wasmann (Isoptera, Termitidae, Nasutitermitinae). Thesis, USP, São Paulo, 151 pp. Colwell, R.K EstimateS: Statistical estimation of species richness and shared species from samples. Versão 7. User s Guide and application published at: org/estimates. Constantino, R The pest termites of South America: taxonomy, distribution and status. Journal Applied Entomology, 126: Constantino, R Key to the soldiers of South American Heterotermes with a new species from Brazil (Isoptera: Rhinotermitidae). Insect. Syst. Evol., 31: Constantino, R Chave ilustrada para identificação dos gêneros de cupins (Insecta: Isoptera) que ocorrem no Brasil. Papéis Avulsos de Zoologia, 40: Constantino, R Catalog of the living termites of the New World (Insecta: Isoptera). Arquivos de Zoologia, 35 (2):

12 12 Sociobiology Vol. 47, No. 2, 2006 Constantino, R. & O.F.F. de Souza Key to the soldiers of Atlantitermes Fontes, with a new species from Brazil (Isoptera, Termitidae, Nasutitermitinae). Tropical Zoology, 10: Constantino, R Revision of the Neotropical termite genus Syntermes Holmgren (Isoptera: Termitidae). The University of Kansas Science Bulletin, 55 (13): Constantino, R Abundance and diversity of termites (Insecta: Isoptera) in two sites of primary rain forest in Brazilian Amazonia. Biotropica, 24: Constantino, R Notes on Cyranotermes araujo, with description of a new species (Isoptera, Termitidae, Nasutitermitinae). Goeldiana Zoologia, 2: Constantino, R. (in press). Padrões de diversidade e endemismo de térmitas no Bioma Cerrado. In: Scariot, A.; Felfili, J.M. & Souza Silva, J.C. (Eds.), Ecologia e biodiversidade do Cerrado. Embrapa, Brasília. Costa, D.A Inquilinos associados a ninho de Cornitermes cumulans (Isoptera: Termitidae) em uma área de campo no Parque Nacional das Emas, GO. MSc. Dissertation, UFG, Goiânia, 45 p. Coutinho, L.M O conceito de Cerrado. Revista Brasileira de Botânica, 1: Davies, R.G., P. Eggleton, L. Dibog, L.H. Lawton, D.E. Bignell, C.H. Brauman, L. Nunes, J. Holt, & C. Rouland Successional response of a tropical forest termite assemblage to experimental habitat perturbation. Journal of Applied Ecology, 36: Dias, B.F.S Alternativas de desenvolvimento dos Cerrados: manejo e conservação dos recursos naturais renováveis. Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis (Ibama), Fundação Pró-Natureza (Funatura), Brasília. Eggleton, P. & D.E. Bignell Monitoring the response of tropical insects to changes in the environment: troubles with termites. In: Harrington, R. & Stork, N. (Eds.), Insects in a changing environment. Academic Press, London, p Eggleton, P., D.E. Bignell, W.A. Sands, N.A. Mawdsley, J.H. Lawton, T.G. Wood, & N.C. Bignel The diversity, abundance and biomass of termites under differing levels of disturbance in the Mbalmayo Forest Reserve, Southern Cameroon. Philosophical Transactions of the Royal Society of London, 351: Eiten, G The Cerrado vegetation of Brazil. Bot. Rev. v. 38, p Emerson, A.E. & F.A. Banks The Neotropical genus Labiotermes (Holmgren): its phylogeny, distribution and ecology (Isoptera, Termitidae, Nasutitermitinae). American Museum Novitates, 2208: Emerson, A.E. & F.A. Banks Five new species and one redescription of the Neotropical genus Armitermes Wasmann (Isoptera, Termitidae, Nasutitermitinae). American Museum Novitates, 1841: Ferreira Junior, E Efeitos da fragmentação da Floresta Mesofítica de Interflúvio sobre a fauna de cupins. MSc. Dissertation, UFG, Goiânia, p. 73. Fontes, L.R New genera and new species of Nasutitermitinae from Neotropical Region (Isoptera: Termitidae). Revista Brasileira de Zoologia, 3: 7-25.

13 da Cunha, H.F. et al. Termite Assemblages in Goiás 13 Freitas, G.A., T. Santos, L.S. Crispim, F.G.F. Lima, R.A. Carvalho, D.A. Costa, & D. Brandão Padrões de diversidade de cupins no Parque Estadual da Serra de Caldas Novas, Goiás. In: Resumos do XXV Congresso Brasileiro de Zoologia. Sociedade Brasileira de Zoologia, Brasília, 523 p. Gotelli, N.J. & R.K. Colwell Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters, 4: Lima Filho, G.F Padrões de variação na fauna de cupins (Insecta: Isoptera) ao longo de um gradiente ambiental no Pico dos Pirineus, Goiás. MSc. Dissertation, 41 p. Klink, C.A. & R.B. Machado Conservation of the Brazilian Cerrado. Conservation Biology, 19 (3): Krebs, C.J Ecological Methodology. Harper Collins, New York, 654 p. Krishna, K. & R.L. Araújo A revision of the Neotropical termite genus Neocapritermes (Isoptera, Termitidae, Termitinae). Bulletin American Museum of Natural History, 38: Mathews, A.G.A Studies on termites from the Mato Grosso State, Brazil. Academia Brasileira de Ciências, Rio de Janeiro, 267 p. Oliveira Filho, A.T. & J.A. Ratter Vegetation physiognomies and woody flora of the Cerrado biome. In: Oliveira, P.S. & Marques, R.J. The Cerrados of Brazil: Ecology and natural history of a neotropical savanna. Columbia University Press, New York, p Oliveira, A.R.G Variação na composição de espécies e no padrão de riqueza e abundancia de cupins ao longo de uma transição cerrado-campo rupestre, no Morro Feio, município de Hidrolândia, GO. MSc. Dissertation, UFG, Goiânia, 26 p. Overal, W.L O peso dos invertebrados na balança de conservação biológicas da Amazônia. In: Capobianco, J.P.R. (coord.), Biodiversidade na Amazônia Brasileira: Avaliação e ações prioritárias para a conservação, uso sustentável e repartição de benefícios. Instituto SocioAmbiental / Estação Liberdade, p Redford, K The termitaria of Cornitermes cumulans (Isoptera, Termitidae) and their role in determining a potential keystone species. Biotropica, 16 (2): Santos, A.J Estimativas de riqueza em espécies. In: Cullen Jr., L.; Rudran, R. & Valladares-Pádua, C. (Eds.), Métodos de estudos em Biologia da Conservação e Vida Silvestre. Editora da UFPR, Curitiba, p

14 14 Sociobiology Vol. 47, No. 2, 2006 Appendix 1. List of species of termite of the eleven municipalities in Goiás State (Brazil). Family Rhinotermitidae Subfamily Coptotermitinae Coptotermes sp Wasmann Subfamily Rhinotermitinae Dolichorhinotermes sp Emerson Subfamily Heterotermitinae Heterotermes longiceps Snyder Heterotermes tenuis Hagen Family Termitidae Subfamily Apicotermitinae Anoplotermes sp1 Fr. Müller Anoplotermes sp2 Fr. Müller Anoplotermes sp3 Fr. Müller Anoplotermes sp4 Fr. Müller Aparatermes sp1 Fontes Aparatermes sp2 Fontes Grigiotermes sp Mathews Ruptitermes spp Mathews Tetimatermes sp Fontes Subfamily Nasutitermitinae Agnathotermes glaber Snyder Angularitermes pinochio Cancello & Brandão Araujotermes sp Fontes Armitermes cerradoensis Mathews Armitermes euamignathus Silvestri Armitermes lanei Snyder Armitermes sp1 Wasmann Armitermes sp2 Wasmann Atlantitermes stercophilus Constantino & De Souza Coatitermes sp Fontes Constrictotermes cyphergaster Silvestri Cornitermes bequaerti Emerson Cornitermes cumulans Kollar Cornitermes pilosus Holmgren Cornitermes snyderi Emerson Cornitermes villosus Emerson Curvitermes minor Silvestri Curvitermes odontognathus Silvestri Cyranotermes timuassu Araújo Diversitermes diversimiles Silvestri Embiratermes festivellus Silvestri Embiratermes silvestrii Emerson Embiratermes sp Fontes Labiotermes longilabius Silvestri Nasutitermes aff minor Holmgren Nasutitermes corniger Motschulsky Nasutitermes ephratae Holmgren Nasutitermes kemneri Snyder & Emerson Nasutitermes sp Dudley Nasutitermes unduliceps Mathews Noirotitermes noiroti Cancello & Myles Paracornitermes emersoni Araújo Paracornitermes hirsutus Araújo Paracornitermes orthocephalus Silvestri Parvitermes bachanalis Mathews Procornitermes araujoi Emerson Procornitermes triacifer (Silvestri) Rhynchotermes diphyes Mathews Rhynchotermes nasutissimus (Silvestri) Subulitermes aff microsoma (Silvestri) Syntermes molestus (Burmeister) Syntermes nanus Constantino Syntermes wheeleri Emerson Velocitermes glabrinotus Mathews Velocitermes heteropterus (Silvestri) Velocitermes paucipilis Mathews Velocitermes velox (Holmgren) Subfamily Termitinae Amitermes sp Silvestri Cylindrotermes sp1 Holmgren Cylindrotermes sp2 Holmgren Dentispicotermes globicephalus (Silvestri) Dentispicotermes sp Emerson Dihoplotermes inusitatus Araújo Microcerotermes sp Silvestri Microcerotermes exiguus (Hagen) Microceroteremes strunckii (Sörensen) Neocapritermes araguaia Krishna & Araújo Neocapritermes opacus (Hagen) Neocapritermes talpa (Holmgren) Orthognatotermes sp Holmgren Spinitermes sp1 Wasmann Spinitermes sp2 Wasmann Spinitermes trispinosus (Hagen & Bates) Termes sp Linnaeus