Research Article |
Corresponding author: Ludson Neves de Ázara ( ludsonazara@yahoo.com.br ) Academic editor: Oana Teodora Moldovan
© 2016 Ludson Neves de Ázara, Leopoldo Bernardi, Rodrigo Lopes Ferreira.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
de Ázara LN, Bernardi LFO, Ferreira RL (2016) The first survey on harvestmen in Brazilian artificial cavities, with notes on distribution and natural history. Subterranean Biology 17: 31-53. https://doi.org/10.3897/subtbiol.17.6762
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Several species of harvestmen occur in natural subterranean cavities using these habitats as shelters and sites of oviposition. Many species have evolved in these environments, thus becoming cave-dwellers. In a few cases harvestmen have been reported in artificial cavities (mines), but without details about their distribution or natural history. Based on faunal inventories carried out in 111 artificial cavities in the state of Minas Gerais, Brazil, this work aimed to register species of harvestmen that are associated with these artificial cavities. Seventeen species were found in 12 municipalities, in addition to new occurrences for 14 of these species and some new behavioral notes. Two undescribed species were also recorded. This study highlights that artificial cavities may be used as shelters by harvestmen, mainly in human-modified landscapes.
Opiliones , artificial subterranean cavities, mines, new records, aggregation, Neotropical, Brazil
Subterranean cavities can be formed by the action of environmental agents (e.g. water action and eruptions) (
Natural subterranean cavities (karstic caves) are originated mainly by the dissolution of rock through water action, thus forming galleries (
Although natural and artificial cavities present different genesis, these habitats may have similar environmental characteristics, which are primarily determined by the permanent absence of light, limited food, constantly high relative humidity and thermic stability (
The order Opiliones Sundevall, 1833 is divided into four extant suborders, comprising 49 families and 6519 species worldwide (
It has been shown by several studies that caves are natural shelters for many epigean harvestmen species (
Considering the small number of faunal surveys performed in artificial subterranean cavities, the present study aimed to: (i) register harvestmen species associated with artificial cavities in Minas Gerais state, Brazil and (ii) present data regarding the species distribution, new occurrences and presence of aggregations in these environments.
A total of 111 subterranean artificial cavities were sampled in 13 municipalities in the state of Minas Gerais, Southeast Brazil. Each municipality is shown in Figure
Among the sampled municipalities, Alagoa, Ataléia, Caeté, Caraí, Mariana, Mateus Leme, Medina, Nova Lima, Novo Oriente de Minas, Ouro Preto, Padre Paraíso and São José da Safira are in the Atlantic Forest biome; Caeté and Nova Lima are situated in the transition between Cerrado and Atlantic Forest, while Vazante is located in the Cerrado (
The original names of the artificial cavities, most of them given by local residents, were maintained since there is no agency in Brazil responsible for the registration of these habitats. Besides, many names refer to the location of the cavities, what might be a helpful tool to find the locality. Since all the artificial cavities of this study are mines we will use further the term mines.
The specimens found in the mines were sampled with the aid of tweezers, after a detailed and extensive search. In order to assess the position of the species in the cavities, each observed individual was plotted in a schematic map of the mine according to the methodology proposed by
Individuals were identified whenever possible. In cases of nymphs, females and undescribed species, the individuals were just separated into morphospecies. However these morphospecies were theoretically considered as species. When there was more than one morphospecies in the same genus, they were separated using correlative numbers.
To determine the extension of the aphotic zone, a luximeter was positioned 1.2 m above the ground with its reception cell directed to the cavity entrance. The device was then conducted from the entrance to the interior of the subterranean system. The location where the luximeter presented luminosity equal to zero was considered the transition between the aphotic and the photic zone.
Specimens were deposited in the Collection of Subterranean Invertebrates of Lavras (ISLA) at the Federal University of Lavras, Lavras, Minas Gerais, and in the
The occurrences of harvestmen species are indicated in alphabetical order in the results with the abundance of each species in each locality indicated in parentheses.
From all the 111 sampled mines 52 contained harvestmen distributed in 12 municipalities (Figure
Distribution of harvestmen species in the municipalities and mines with information about their total abundance and presence (P)/absence (A) in aphotic and/or photic zones. *The abbreviated name of the species will be used to refer to each species in Table
Abreviated name of the species | Species | N | Family | Municipality | Mine | Aphotic zone | Photic zone |
---|---|---|---|---|---|---|---|
Ab. b. | Abaetetuba bahiensis (Mello-Leitão, 1931) | 66 | Sclerosomatidae | g | 4 | A | P |
Gr. p. | Gryne perlata Mello-Leitão, 1936 | 5298 | Cosmetidae | d, g, k | 11 | A | P |
Pa. 1 | Paratricommatus sp. 1 | 1 | Cryptogeobiidae | c | 1 | - | - |
Sp. 1 | Spinopilar sp. 1 | 3 | Cryptogeobiidae | f | 1 | - | - |
Go. c. | Goniosoma carum (Mello-Leitão, 1936) | 1 | Gonyleptidae | j | 1 | - | - |
Go. v. | Goniosoma vatrax Koch, 1848 | 14 | Gonyleptidae | e, h | 5 | P | P |
Mi. i. | Mitogoniella indistincta Mello-Leitão, 1936 | 555 | Gonyleptidae | c, e, j, l | 21 | P | P |
Mi. m. |
Mitogoniella
mucuri
|
12 | Gonyleptidae | a, d | 2 | P | P |
Li. i. | Liogonyleptoides inermis (Mello-Leitão, 1922) | 4 | Gonyleptidae | i | 1 | A | P |
Ps. m. | Pseudopucrolia mutica (Perty, 1833) | 1 | Gonyleptidae | d | 1 | A | P |
Lo. t. | Longiperna trembao Pinto-da-Rocha & Bragagnolo, 2010 | 26 | Gonyleptidae | c, e, j | 5 | P | P |
Mi. 1 | Mitobatula sp. 1 | 22 | Gonyleptidae | a | 2 | P | P |
Eu. a. | Eusarcus aduncus (Mello-Leitão, 1942) | 24 | Gonyleptidae | c, l | 7 | P | P |
Eu. c. | Eusarcus cavernicola Hara & Pinto-da-Rocha, 2010 | 23 | Gonyleptidae | m | 1 | P | A |
Eu. 1 | Eusarcus sp. 1 | 14 | Gonyleptidae | e, j, l | 6 | P | P |
Eu. 2 | Eusarcus sp. 2 | 1 | Gonyleptidae | f | 1 | - | - |
Pl. 1 | Planiphalangodus sp. 1 | 1 | Gonyleptidae | a | 1 | P | A |
The following species and genera were new occurrences for the state of Minas Gerais (letters in brackets correspond to the municipality): Abaetetuba bahiensis (g), Gryne perlata (d, g, k), Paratricommatus sp. 1 (c), Spinopilar sp. 1 (f), Goniosoma carum (j), Goniosoma vatrax (h), Liogonyleptoides inermis (i), Pseudopucrolia mutica (d), Longiperna trembao (c, e), Mitobatula sp. 1 (a), Eusarcus aduncus (c, l), Eusarcus cavernicola (m), Eusarcus sp. 1 (e, l), Eusarcus sp. 2 (f) and Planiphalangodus sp. 1 (a). Additionally, this study improved the information about the distribution of some species, such as Goniosoma vatrax with about 40 km, and Gryne perlata with 1000 km. It was also registered for the first time an aggregation behavior for Abaetetuba bahiensis and Gryne perlata, besides the record of two undescribed species.
The mines with the highest number of species were Mina dos Canelas VII in Mariana, and Mina Turística do Chico Rei in Ouro Preto, each with 3 species (Table
List of mines containing harvestmen, with the geographic coordinates (in decimal degrees, DATUM WGS 84), age (in years), use (not in use = D., daily tourism = D.T., mineral extraction = M.E., scientific experiment = S.E., sparse tourism = S.T., sparse visitation = S.V., urban zone = U.Z.), type of the surrounding vegetation/rock (Atlantic Forest = A.F., Eucalyptus forest = E., pasture = P., ferriferous formations = R.F.) on a 30 m (S. 30 m) and 250 m radius (S. 250 m), extension and identified harvestmen species (see Table
Municipality | Mine | Latitude | Longitude | Age | Usage | S. 30 m | S. 250 m | Extension | Species |
---|---|---|---|---|---|---|---|---|---|
Alagoa | Mina da Companhia | -22.17812 | -44.70546 | > 100 | S.T. | A.F. | P. - A.F. | 46 | Mi. 1, Pl. 1 |
Alagoa | Túnel do Garrafão | -22.18221 | -44.71956 | > 100 | S.T. | A.F. | P. - A.F. | 73 | Mi. m., Mi. 1 |
Caeté | Mina do Morro Vermelho I | -19.96525 | -43.71271 | < 30 | S.V. | E. | P. - E. | 76 | Mi. i. |
Caeté | Mina do Morro Vermelho II | -19.96520 | -43.71364 | < 30 | S.V. | E. | P. - E. | 12 | Pa. 1, Eu. a. |
Caeté | Mina do Morro Vermelho III | -19.96538 | -43.68757 | < 30 | S.V. | E. | P. - E. | 60 | Mi. i., Mi. 1 |
Caeté | Mina dos Matarelli | -19.94719 | -43.74017 | < 50 | D. | E. | P. - E - A.F. | 62 | Lo. t. |
Caraí | Túnel da Br116 | -17.22967 | -41.49569 | < 5 | D. | P. | P. | 8 | Ps. m. |
Caraí | Túnel do Noel I | -17.11623 | -41.51527 | < 5 | M.E. | P. | P. | 69 | Gr. p. |
Caraí | Túnel do Noel II | -17.11727 | -41.51627 | < 5 | M.E. | P. | P. | 38 | Gr. p. |
Caraí | Túnel da Fazenda Cilindro I | -17.11920 | -41.32715 | < 20 | D. | P. | P. | 10.2 | Mi. m. |
Mariana | Mina dos Canelas Casa | -20.35245 | -43.43990 | > 100 | S.V. | R.F. | P. | 49 | Go. v., Eu. 1 |
Mariana | Minas dos Canelas VI | -20.35315 | -43.44090 | > 100 | S.V. | R.F. | R.F. | 68 | Go. v. |
Mariana | Mina dos Canelas VII | -20.54384 | -43.81015 | > 100 | S.V. | R.F. | R.F. | 229 | Go. v., Lo. t., Eu. 1 |
Mariana | Mina dos Canelas XI | -20.35256 | -43.43936 | > 100 | S.V. | R.F. | R.F. | 68 | Go. v., Mi. i. |
Mariana | Mina dos Canelas XII | -20.35377 | -43.43932 | > 100 | S.V. | R.F. | R.F. | 98 | Go. v., Mi. i. |
Mariana | Mina do Meio do Mato | -20.34518 | -43.44805 | > 100 | S.V. | R.F. | R.F. | 28 | Mi. i. |
Mariana | Mina da Cachoeira | -20.34507 | -43.44678 | > 100 | S.V. | R.F. | R.F. | 26 | Mi. i., Eu. 1 |
Mateus Leme | Mina do Aqueduto I | -19.95938 | -44.42231 | > 100 | S.V. | R.F. | R.F. | 11 | Eu. 2 |
Mateus Leme | Mina do Aqueduto II | 19.959223 | -44.42196 | > 100 | S.V. | R.F. | R.F. | 8 | Sp. 1 |
Medina | Mina da Fazenda do Sol V | -16.34362 | -41.45138 | < 30 | D. | P. | P. | 27 | Ab. b. |
Medina | Mina da Fazenda Serra Azul I | -16.20885 | -41.47934 | < 30 | D. | P. | P. | 14 | Gr. p. |
Medina | Mina da Fazenda Serra Azul II | -16.21157 | -41.47938 | < 30 | D. | P. | P. | 74 | Gr. p. |
Medina | Mina do Bloco I | -16.21425 | -41.46832 | < 30 | D. | P. | P. | 126 | Ab. b., Gr. p. |
Medina | Mina do Bloco II | -16.21610 | -41.46872 | < 30 | D. | P. | P. | 56 | Ab. b., 2 |
Medina | Mina do Bloco III | -16.21460 | -41.46815 | < 30 | D. | P. | P. | 31 | Ab. b., 2 |
Nova Lima | Mina de Capão Xavier | -20.0469 | -43.9798 | <5 | S.C/S.V. | R.F. | R.F> | 15 | Go. v. |
Novo Oriente de Minas | Mina do João Bufinha | -17.4137 | -41.2097 | - | S.V. | P. | P. | - | Li. i. |
Ouro Preto | Mina da Volta do Córrego | -20.37922 | -43.51066 | > 100 | S.V. | A.F. | A.F. - P. | 22 | Go. c. |
Ouro Preto | Mina de Vila Rica | -20.38757 | -43.49346 | > 100 | D.T. | U.Z. | U.Z. | 96 | Lo. t. |
Ouro Preto | Mina do Chico Rei | -20.38608 | -43.49930 | > 100 | D.T. | U.Z. | U.Z. | 157 | Mi. i., Lo. t., Eu. 1 |
Ouro Preto | Mina Velha | -20.38838 | -43.49163 | > 100 | D.T. | U.Z. | U.Z. | 174 | Mi. i. |
Padre Paraiso | Túnel dos Meninos I | -17.06916 | -41.46849 | < 20 | D. | P. | P. | 42 | Gr. p. |
Padre Paraiso | Túnel dos Meninos II | -17.06856 | -41.46903 | < 20 | M.E. | P. | P. | 29.8 | Gr. p. |
Padre Paraíso | Túnel do Hotel Entre Vales I | -17.06380 | -41.48198 | < 20 | D. | P. | P. - A.F. | 12.5 | Gr. p. |
Padre Paraíso | Túnel do Hotel Entre Vales II | -17.06380 | -41.48198 | < 20 | D. | P. | P. - A.F. | 39 | Gr. p. |
São José da Safira | Mina do Chiá Donizete I | -18.29455 | -42.18781 | < 50 | D. | A.F. | A.F. | 72 | Mi. i., Eu. a. |
São José da Safira | Mina do Chiá Donizete II | -18.29064 | -42.18758 | < 50 | M.E. | A.F. | A.F. | 68 | Mi. i., Eu. a. |
São José da Safira | Mina do Chiá Donizete III | -18.29163 | -42.18846 | < 50 | D. | A.F. | A.F. | 15.5 | Mi. i., Eu. a. |
São José da Safira | Mina do Milto Godinho I | -18.29614 | -42.18276 | < 50 | M.E. | P. | P. | 33 | Eu. a. |
São José da Safira | Mina do Milto Godinho II | -18.29631 | -42.18286 | < 50 | D. | P. | P. | 12 | Mi. i. |
São José da Safira | Mina do Milto Godinho III | -18.29639 | -42.18329 | < 50 | D. | P. | P. | 52 | Mi. i., Eu. 1 |
São José da Safira | Mina do Milto Godinho IV | -18.29598 | -42.18292 | < 50 | M.E. | P. | P. | 72 | Mi. i., Eu. a. |
São José da Safira | Mina do Milto Godinho VII | -18.29697 | -42.18121 | < 50 | D. | P. | P. | 37 | Mi. i. |
São José da Safira | Mina do Milto Godinho VIII | -18.29447 | -42.18075 | < 50 | D. | P. | P. | 84 | Mi. i., Eu. a. |
São José da Safira | Túnel do Milto Godinho IX | -18.29395 | -42.18040 | < 50 | D. | A.F. | P. - A.F. | 12 | Mi. i. |
São José da Safira | Túnel do Milto Godinho X | -18.29268 | -42.18421 | < 50 | D. | A.F. | P. - A.F. | 67.6 | Mi. i. |
São José da Safira | Túnel do Milto Godinho XI | -18.29268 | -42.18421 | < 50 | D. | A.F. | P. - A.F. | 31 | Mi. i. |
São José da Safira | Túnel do Milto Godinho XII | -18.29292 | -42.18485 | < 50 | D. | A.F. | P. - A.F. | 48.5 | Mi. i., Eu. 1 |
São José da Safira | Túnel do Milto Godinho XIII | -18.29250 | -42.18418 | < 50 | D. | A.F. | P. - A.F. | 100.3 | Mi. i. |
Vazante | Mina da Vmetais | -17.95928 | -46.82371 | < 30 | S.V. | P. | P. | 600 | Eu. c. |
Bahia: Ilhéus (
Medina: 1 female (ISLA 16035) from Mina da Fazenda do Sol V (50), 08/IV/2009; 1 male (ISLA 16048) from Mina do Bloco I (8), 14/VII/2009; 1 male (
This species was observed forming aggregations of 8, 19 and 26 individuals at a maximum distance of 5 meters from the entrance representing the first record of an aggregation behavior for this species. In other cavities, sparsely distributed individuals were observed, but always located in the photic zone no more than 10 meters from the entrance. These records represent a new occurrence of this species for this municipality, thus expanding its distribution with 250 km from the southwest of Juçari, Bahia state.
Pernambuco: municipality not specified (
Caraí: 1 male (ISLA 16033) from Túnel do Noel I (1), 13/VII/2008; 1 female (
This species was observed forming aggregations of 28, 45, 49, 103, 200, 400 and 2200 individuals at the entrance of seven cavities at a maximum distance of 5 m from the entrance. This was the first record of an aggregation behavior for this species. In other cavities, sparsely distributed individuals were also recorded, but always located in the photic zone at a maximum distance of 10 m from the entrance. During one of the fieldtrips, three aggregations with about 1000 individuals were observed. These aggregations were found in the epigean environment, next to the mine Túnel do Hotel Entre Vales I in holes and cracks in a ravine of exposed soil. Such shelters had a longitudinal form of ~ 80 cm in depth, 20 cm width and 1.5 cm height. The three municipalities represent new occurrences for this species, expanding its distribution ~ 1000 km from the south of Pernambuco state.
Alto Paraná (Paraguai), Espírito Santo, Rio de Janeiro and São Paulo (
Caeté: 1 female (
Only one female was found. This is a new occurrence in the municipality, expanding the genus distribution with about 300 km from the southwest of Colatina, Espírito Santo state.
Alto Paraná (Paraguai), Argentina (province not specified), Espírito Santo, Minas Gerais, Rio de Janeiro, Santa Catarina (
Mateus Leme: 1 female (
Only one female was found and Mateus Leme corresponds to a new occurrence for the genus.
Minas Gerais: Viçosa (
Ouro Preto: 1 male (ISLA 1468) from Mina da Volta do Córrego (1), 13/VI/2009, new record for locality.
Ouro Preto corresponds to a new occurrence for this species, expanding its distributions with about 100 km from the northwest of Viçosa, Minas Gerais state.
Minas Gerais: Catas Altas (
Mariana: 1 male (ISLA 16019) from Mina dos Canelas Casa (4), 01/V/2009; 1 male (
Sparse individuals were registered in the photic and aphotic zones of the mines at a maximum distance of 45 meters from the entrance. Nova Lima corresponds to a new occurrence for this species, expanding its distribution with about 40 km from the east of Santa Bárbara, Minas Gerais. However, the Mina de Capão Xavier was destroyed by the expansion of mining activities, and its fauna was translocated to another artificial mine constructed in the same region of the original one (Ferreira, R.L. pers. observation).
Minas Gerais: Alto Caparaó, Baependi, Barão de Cocáis (
Caeté: 1 female (ISLA 16038) from Mina do Morro Vermelho I (6), 17/VI/2009; 1 male (
In seven cavities, this species was observed forming aggregations of 10, 15, 18, 22, 41 and 43 individuals up to 5 meters from the entrance. In other cavities, sparsely distributed individuals were also recorded both in the photic and aphotic zones at a maximum distance of 60 m from the entrance. In some cases, individuals were found guarding their eggs and nymphs. This species also occurs in natural cavities in Minas Gerais State (
Minas Gerais: Alagoa, Caraí, Novo Oriente de Minas, Padre Paraíso (
Alagoa: 1 male (ISLA 3983) from Túnel do Garrafão (10), 31/VIII/2008 (
Individuals of this species were recorded sparsely distributed inside the mines both in the photic and aphotic zones at a maximum distance of 36 m from the entrance. In some mines, specimens were found guarding their eggs and nymphs. This species also occurs in natural cavities in Minas Gerais state (
Minas Gerais: Tapira (H.
Novo Oriente de Minas: 1 male (
Individuals were sparsely distributed inside the mines, both in the photic and aphotic zones at a maximum distance of 10 m from the entrance. Novo Oriente de Minas corresponds to a new occurrence for this species, expanding its distribution with about 700 km from the northeast of Petrópolis, Rio de Janeiro state.
Bahia: Salvador (
Caraí: 1 male (
Only one individual was recorded in the photic zone of the cavity, at less than 10 m from the entrance. The record at Caraí expands the distribution of this species with about 750 km from the southwest of Salvador, Bahia.
Minas Gerais: Belo Horizonte, Ouro Preto (
Caeté: 1 male (ISLA 16059) from Mina do Morro Vermelho III (1), 17/VI/2009; 1 male (
Individuals of this species were sparsely distributed in the photic and aphotic zones of the mine at a maximum distance of 30 m from the entrance. The records at Caeté and Mariana expand its distribution with about 60 km from the northwest of Ouro Preto, Minas Gerais state.
(the genus is monospecific). Santa Catarina: Serra Azul (
Alagoa: 1 male (
Individuals of this morphospecies were sparsely distributed in the photic and aphotic zones at a maximum distance of 12 m from the entrance. The record at Alagoa expands the distribution of the genus with about 150 km from the west of Teresópolis, Rio de Janeiro state.
Bahia: Carinhanha, Itagibá, Una (
Caeté: 1 male (
Individuals of this species were found sparsely distributed in the photic and aphotic zones at a maximum distance of 60 m from the entrance. The records at Caeté and São José da Safira expand its distribution with about 100 km from the south of Carinhanha, Bahia. This species also occurs in Minas Gerais and São Paulo state in natural cavities (
Bahia: Santana; Goiás: Anápolis, São Domingos; Minas Gerais: Itacarambi, Montes Claros, Unaí (
Vazante: 1 male (
Individuals of this species were observed sparsely distributed inside the mine. They were located in the photic and aphotic zones at a maximum distance of 500 m from the entrance. The record at Vazante expands its distribution with about 200 km from the south of Unaí, Minas Gerais. This species also occurs in natural cavities in Bahia, Goiás and Minas Gerais state (
From the northeast to the southeast of Brazil, northeast of Argentina, east of Paraguai and Uruguai (
Mariana: 1 male (ISLA 16061) from Mina dos Canelas I (1), 11/VI/2009; 1 male (ISLA 16021) from Mina dos Canelas VII (2), 11/VI/2009; 1 male (ISLA 16024) from Mina da Cachoeira (4), 12/VI/2009 (
Individuals were observed sparsely distributed inside the cavities both in the photic and aphotic zones, at a maximum distance of 22 m from the entrance. Individuals were also observed next to the touristic path. The records at Mariana and São José da Safira correspond to new occurrences for the genus.
From the northeast to the southeast of Brazil, northeast of Argentina, east of Paraguai and Uruguai. (
Mateus Leme: 1 male (ISLA 16062) from Mina do Aqueduto I (1), 05/IV/2008, new record for locality.
Only one female was found and the record of Mateus Leme corresponds to a new occurrence for this genus.
(the genus is monospecific). Santa Catarina: municipality not specified (
Alagoa: 1 male (
Only one male was found and the record of Alagoa corresponds to a new occurrence for this genus.
This study presented the occurrence of two undescribed species (Eusarcus sp. 1 and Mitobatula sp. 1) and new records for states and localities in the case of nine species and four genera of harvestmen. The results obtained demonstrate the importance of observations in unusual environments, like mines for the potential of finding new species, provide more information about the occurrence of species and register new behavioral patterns, thus improving the knowledge about the group.
Although there are studies in Brazil and worldwide about the fauna associated to mines (
The only record of harvestmen in artificial subterranean environments outside the state of Minas Gerais was for the species Pararezendesius luridus H. Soares, 1972, which was found in mines of Iporanga and also in natural cavities of Apiaí, Iporanga and Ribeira municipalities, São Paulo state (
The age of the majority of the mines in our study varied from less than five years to more than 100 years (Table
As shown by
The results of this work show that harvestmen can use mines as shelters during the day and for reproduction. These mines are important because many of the studied regions are fragmented, with forest areas replaced by pastures and agricultural land. Future studies can demonstrate whether harvestmen feed in the pastures or use the organic material inside the mines.
The presence of Eusarcus cavernicola individuals, a species commonly found in caves (
Further studies should be concentrated on understanding the dynamics of species inside mines and the taxonomic identification. Artificial cavities can contribute to the conservation of some harvestmen species in many areas where the original vegetation was removed and replaced by pastures, monocultures or constructions.
We are grateful to Marcus Paulo de Oliveira, Amanda M. Teixeira and Matheus Brajão for the field assistance and Adriano B. Kury for identifying the specimens. We also thank Rafaela Bastos Pereira and Lívia D. Audino for the English translation and review. This study was supported by grants from the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (Fapemig Processo No: APQ 4189 5 03-07). LFBO acknowledges a CAPES/Brazil scholarship. RLF is also grateful to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq grant nr. 304682/2014-4), and for the partial support of Vale S. A., allowing laboratory maintenance. We are grateful to the two anonymous reviewers that improved considerably the quality of the manuscript.