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Introduction
Mainly in the last 80 years, the efforts spent on taxonomic studies (Santos 1944, 1945, 1950, Machado 1964, 1985, Machado et al. 1991, Lencioni 2005, 2006, 2017) and surveys (Dalzochio et al. 2011, Calvão et al. 2014, Vilela et al. 2016, Koroiva et al. 2017a, Koroiva et al. 2017b, Rodrigues & Roque 2017) helped to cover gaps on knowledge of Brazilian odonates. Of the over 6000 hitherto described Odonata species, about 1700 are known to occur in the Neotropical region, and over 800 are recorded for Brazil (von Ellenrieder 2009, Neiss 2012). Despite these efforts, there are many species yet to be described.
In the past five years, most of the surveys generated taxonomic and distributional novelties. In a recent expedition (Oct-Nov 2015) to Chapada dos Guimarães over 100 Odonata species, one of them a new species described and two larvae of Argia (Odonata:Coenagrionidae) were recorded (Vilela, et al. 2018a,b, Cezário et a. 2018). Furthermore, Calvão et al. (2014) recorded five species new to Mato Grosso State, Brazil; Dalzochio et al. (2018) reported 182 species and seven new records to the state of Rio Grande do Sul and Rodrigues & Roque (2017) recorded 33 species new for the state of Mato Grosso do Sul.
In the state of Minas Gerais (MG), however, few studies aimed to list Odonata species. Machado (1998) compiled records of 218 species for the entire state. Ferreira-Peruquetti & de Marco Jr (2002) found 17 species in the municipality of Viçosa. Souza et al. (2013) recorded 57 Odonata species in the Baú Forest, municipality of Barroso, and Bedê et al. (2015) collected 128 species in Serra de São José. More recently, Machado & Bedê (2015) described nine new species from a small area in Serra da Canastra, and Vilela et al. (2016) recorded three new species for the state, including one new to science (Guillermo-Ferreira et al. 2016).
Distributional studies are also important under a conservationist point of view since to determine if a given area needs conservational actions, the local fauna have to be taken into account. In this sense, the International Union for Conservation of Nature (IUCN 2019) relies on several types of species data, such as population size and distributional range, in order to assess the conservation status of species around the globe. IUCN is a global program for evaluating the conservation status of plant and animal species and providing some information on species habitat and biology can help to increase those conservationist actions (IUCN 2019). However, if there is a gap on the distributional data of a given species, it makes the IUCN work difficult or even impossible, disabling actions that may be taken towards the conservation of the species and its habitat. Furthermore, considering the last two great ecological disasters caused by mining in Minas Gerais (see Freitas et al. 2019), studies on the local biodiversity are crucial to know what we lose with such environmental crimes (e.g. MMA/IBAMA 2015).
Not only the species are understudied, but also some phytophisiognomies are almost unexplored. In the west of Minas Gerais, the predominant biome is the Cerrado, which presents a great array of freshwater ecosystems. Within this biome, the Veredas play an important role on the hidrology, being consisted by permanently water saturated soil and a dense vegetation, including the palm tree Mauritia flexuosa L (Fonseca 2005). Additionally, Veredas are an important source of shelter and food from insects to fishes and birds, playing an important role on the ecological balance in the Cerrado. Odonates are little explored within such areas, despite recent studies show that this ecosystem holds a great number of species of the order (Vilela et al. 2016; Barbosa et al. 2019; Borges et al. 2019).
The main objective of this study is to increase the knowledge of the geographical distribution of Odonata species.. For this, here we compile data on species from the western part of Minas Gerais State and make our database available to the academic community. In summary, our study aimed to contribute to the species conservation actions, such as the IUCN Red List of Threatened Species™ and the Latin-American Odonatological Society evaluations, which many times rely on species distributional data for assessments.
Material and methods
1. Field sampling sites and methodology
Altogether, we surveyed 49 localities in ten municipalities (Figure 1):
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Campina Verde (Altitude 494m): Chácara (CVC, -19.5476, -49.4936; small pond, formed by the damming of a Vereda, Figure 2d) and Granja (CVG, -19.5357, -49.5062; small pond connected to a Vereda), both sampled in February 2013.
Figure 2 Some of the sampled sites in this study: a. Cachoeira Rio do Peixe (CRP), São Roque de Minas; b. Cerradão (CER), São Roque de Minas; c. Ranchão (PAR), Paracatu; d. Chácara (CVC), Campina Verde; e. Ribeirão São Lourenço (ITBSL), Ituiutaba; f. Rio Tijuco (ITBT), Ituiutaba; g. Estância Paraíso (EPA), Ituiutaba; h. Landscape between Matutina 3 (MAT3) and Matutina 4 (MAT4), Matutina.
Guarda-Mor (Altitude 616m): Ponto 1 (GMP1, -17.7828, -47.1301; small stream) and Ponto 2 (GMP2, -17.7726, -47.1332; small stream), both sampled in December 2014.
Ituiutaba (Altitude 605m): Lagoa Temporária (ITBL, -18.9776, -49.4103; temporary pond) in April 2014; Ribeirão São Lourenço (ITBSL, -18.9853, -49.4179 and -19.0611,-49.3213; medium order stream, Figure 2e) sampled in December 2013 and April 2014; Rio Tijuco (ITBT, -18.9418,-49.4768; -18.9209,-49.5664 and -18.9525,-49.3377; medium sized river, Figure 2f) sampled in December 2011 and March 2012; Estância Paraíso (EPA, -18.9522, -49.3399; small stream connected to a river, Figure 2g) sampled in April 2014.
Matutina (Altitude 1060m, Figure 2h): Matutina 1 (MAT1, -19.1377, -46.0669; small stream connected with a Vereda), Matutina 2 (MAT2, -19.1251, -46.0568; small pond), Matutina 3 (MAT3, -19.1563, -46.0369; small stream) and Matutina 4 (MAT4, -19.1221, -46.0673; small stream) all sampled in March 2015.
Paracatu (Altitude 688m): Ranchão (PAR, -16.8608, -46.9811; stream connected with a Vereda, Figure 2c) sampled in July 2014.
Prata (Altitude 631m): Rodovia (PRA, -19.3210, -48.9454; stream connected with a Vereda) sampled in October 2012.
São Roque de Minas, Serra da Canastra (Altitude 819-1300m): Cachoeira Rio do Peixe (CRP, -20.2569, -46.4096; small stream by a waterfall, Figure 2a), Aqueduto (AQD, -20.2648, -46.3968; stream near to a Vereda), Riacho Costela de Vaca (RCV, -20.2658, -46.3957; stream near to a Vereda), Cerradão (CER, -20.1914, -46.3858; stream near to a Vereda, Figure 2b), Barro Sujo (BS, -20.2145, -46.3808; small stream), Cerca elétrica (CE, -20.2385, -46.3847; stream near to a Vereda), Espraiado (Subida) (ESS, -20.2411, -46.3857; stream near to a Vereda), Rio São Francisco (RSF, -20.3330, -46.4361; large river), Água Branca (AGB, -20.2922, -46.3658; small stream), Cachoeira da Chinela (CHI, -20.3002, -46.3955; stream by a waterfall), Casca D’anta (CCD, -20.2974, -46.5227; stream by a waterfall), Rolinho (RL, -20.1706, -46.5615; small stream near a Vereda), Nascente (NAS, -20.2433, -46.4463; small stream connected to a Vereda), Ponte 1 (PO1, -20.2402, -46.5875; small stream connected to a Vereda), Ponte 2 (PO2, -20.2325, -46.6083; small stream connected to a Vereda), Quintal da Canastra (QUI, -20.3282,-46.5038; large pond), Cachoeira da Capivara (CPV, -20.6287,-46.2863; large stream by a waterfall), Lago Tio Zezico (ZZC, -20.3121,-46.5319; small pond), Córrego Tio Zezico (ZZC, -20.3116,-46.5324; stream), Lago P31 (P31, -20.1656,-46.6877; two small ponds within a Vereda), Riacho Sr. Vicente (VCT, -20.3033,-46.5492; ; small stream connected to a Vereda), Córrego Guariba (GUA, -20.3017,-46.5483; small stream near a Vereda), Córrego Passageiro (PSG, -20.2668,-46.5554; small stream with rocky bottom), Córrego Rasga Canga (CAN, -20.1761,-46.5603; stream by a waterfall), Portaria Sul (SUL, -20.3063,-46.5236; small stream), Portaria 2 (PR2, -20.1523,-46.6605; small stream by a waterfall), Centro de Visitantes (VIS, -20.2541,-46.4168; small stream) sampled in May and October 2017, March and November 2018 and April 2019.
Uberlândia (Altitude 863m): Marileuza (UDI,-18.8712, -48.2481; small stream near a Vereda, which nowadays is turning to a habitation area) sampled in February 2015.
Unaí (Altitude 640m): Lagoa-Riacho (UNAI, -16.4822, -47.4149; medium order river and a pond) sampled in July 2014.
Varginha (Altitude 915m): Sítio Juriti (SRI, -21.5854, -45.4070; small pond), Carro Velho (CAR, -21.5901, -45.3721; small stream), Lagoa Soberana de Minas Jorge (JOR, -21.6289, -45.4342; small stream and pond), Faz. Tachos (TCH, -21.6154, -45.4803; small stream) sampled in June 2018.
The collections were made ad libitum in different hours of the day, for at least one hour per site, always between 8:00 to 14:00h. An entomological net was used to collect the specimens and they were placed in individual paper envelopes. The collection authorization process was issued by IBAMA, through SISBio system under the numbers 53026-3 and 54386-6. The identifications were made using the keys of Lencioni (Lencioni 2005, 2006, 2017) and Garrison et al. (2006, 2010). The specimens collected are deposited in the Laboratory of Ecological Studies on Ethology and Evolution (LESTES, UFSCar), São Carlos, São Paulo and in the personal collection of Frederico Lencioni in Jacareí, São Paulo.
During fieldwork, we observed and noted the following behavioral traits: (i) territoriality status, (ii) mate-guarding behavior and (iii) oviposition behavior. Such behavioral traits are important because they reveal important aspects of reproductive tactics, habitat occupancy and selection, competition and even larval development (Corbet 1999).
When not possible to notice behavioral traits on the moment of collections, we searched for such information on the literature on two online platforms (Google scholar: https://goo.gl/MtXqZN; Scopus: https://goo.gl/2F36gs) with the terms “species name”, “behavior”, “behaviour”. The terms were used all together, in different orders and separately along with the species names. For example: “Tigriagrion aurantinigrum” “behavior” “behaviour” or solely “Tigriagrion aurantinigrum”.For every sampled species, we checked the IUCN conservation status (iucnreslist.org) (IUCN 2019). When a species is assessed by specialists, a code is given to represent its conservational status. For example, if a species is lacking distributional data and nothing much is known of its conservational situation, it is given to it a data deficient status, or DD. Other classification status are: LC (least concern), when a species is well distributed, occurring in many localities, including protected areas; NT (near threatened), when its assessment indicates any degree of conservational threat, usually applied to species occurring near expanding urban areas, which can be threatened in a near future; VU (vulnerable), species facing extinction risk in nature, with a limited distributional range (<5000 km²) in areas threatened by human activities; CR (critically endangered), species inhabiting areas less than 100 km², occurring in usually one locality, often threatened by human activities.
The map was adapted from IBGE - Brazilian Geography and Statistics Institute (2019) using Adobe Photoshop 2017.1.1. Habitus scan of species was made using an EPSON V600 Perfection and posterior plate assembling using Adobe Photoshop 2017.1.1. Photographs of the sampling sites were all made by the authors, except for (Fig. 3g), courtesy of Edson Soares dos Santos.
Figure 3 Habitus scan of some species sampled in our expeditions: a. Argia lilacina Selys; b. Acanthagrion aepiolum Tennessen; c. Ischnura capreolus Hagen; d. Minagrion waltheri Selys; e. Minagrion franciscoi Machado & Bedê; f. Oxyagrion chapadense Costa; g. Heteragrion triangulare Hagen in Selys; h. Hetaerina hebe Selys; i. Neoneura waltheri Selys; j. Erythrodiplax ana Guillermo-Ferreira, Vilela, Del-Claro & Bispo; k. Zenithoptera lanei Santos; l. Perithemis mooma Kirby.
Results
1. Sampled species
We collected individuals from 11 families, 41 genera and 90 species (Table 1; Figure 3a−l). The family Libellulidae was the most representative of Anisoptera, with 13 genera and 27 species, whereas Coenagrionidae was the most specious family of Zygoptera, with 12 genera and 35 species (Table 1). PAR was the most speciose site with 16 records, whereas NAS recorded 15 species and ITBT and UNAI recorded 14 species each. On the other hand, VIS and RL were the least specious site with one species each (Table 2).
Table 1 Representation of each collected family by genera and species in 49 sampling points distributed in the State of Minas Gerais, Brazil.
| Anisoptera | Number of genera |
Number of species |
|
|---|---|---|---|
| Aeshnidae | 5 | 5 | |
| Corduliidae | 1 | 1 | |
| Gomphidae | 3 | 4 | |
| Libellulidae | 13 | 27 | |
| Zygoptera | |||
| Calopterygidae | 2 | 7 | |
| Coenagrionidae | 12 | 35 | |
| Dicteriadidae | 1 | 1 | |
| Heteragrionidae | 1 | 4 | |
| Lestidae | 1 | 3 | |
| Megapodagrionidae | 1 | 1 | |
| Protoneuridae | 1 | 2 | |
| Total | 11 Families | 41 | 90 |
Table 2 Sampled localities with the respective number of species collected on each site.
| Locality | Code | N. Species |
|---|---|---|
| Paracatu | PAR | 16 |
| Nascente | NAS | 15 |
| Rio Tijuco | ITBT | 14 |
| Unaí | UNAI | 14 |
| Ribeirão São Lourenço | ITBSL | 12 |
| Marileuza | UDI | 12 |
| Matutina Ponto 2 | MAT2 | 10 |
| Quintal da Canastra | QUI | 10 |
| Cachoeira da Capivara | CPV | 9 |
| Cachoeira Rio do Peixe | CRP | 9 |
| Aqueduto | AQD | 8 |
| Cerradão | CER | 8 |
| Rio São Francisco | RSF | 8 |
| Lago Tio Zezico | LZZ | 8 |
| Córrego Tio Zezico | ZZC | 7 |
| Carro Velho | CAR | 7 |
| Fazenda Sr. Gilberto | GIL | 7 |
| Granja | CVG | 7 |
| Guarda Mor Ponto 1 | GMP1 | 7 |
| Matutina Ponto 4 | MAT4 | 7 |
| Prata | PRA | 7 |
| Lago P31 | P31 | 6 |
| Guarda Mor Ponto 2 | GMP2 | 6 |
| Matutina Ponto 1 | MAT1 | 6 |
| Riacho Costela de Vaca | RCV | 6 |
| Espraiado (Subida) | ESS | 6 |
| Cachoeira da Chinela | CHI | 6 |
| Lagoa Temporária | ITBL | 5 |
| Matutina Ponto 3 | MAT3 | 5 |
| Ponte 1 | PO1 | 5 |
| Ponte 2 | PO2 | 5 |
| Riacho Sr. Vicente | VCT | 5 |
| Fazenda dos Tachos | TCH | 4 |
| Chácara | CVC | 4 |
| Barro Sujo | BS | 4 |
| Casca D'Anta | CCD | 4 |
| Cerca | CE | 4 |
| Córrego Guariba | GUA | 4 |
| Córrego Passageiro | PSG | 4 |
| Córrego Rasga Canga | CAN | 4 |
| Portaria Sul | SUL | 4 |
| Portaria 2 | PR2 | 3 |
| Sítio Juriti | SRI | 3 |
| Lagoa S. M. Jorge | JOR | 3 |
| Água Branca | AGB | 3 |
| Alojamento (P17) | P17 | 2 |
| Estância Paraíso | EPA | 2 |
| Centro de Visitantes | VIS | 1 |
| Rolinho | RL | 1 |
Furthermore, Machado (1998) reported 218 Odonata species to Minas Gerais state. Therefore, adding new species descriptions (n = 29) and new records (n = 22) since 1998, the number raises to 269 species to Minas Gerais (Table 3).
Table 3 List of new Odonata species and new records to the state of Minas Gerais, published after 1998.
| Author | Year | Type of data | Number of records |
|---|---|---|---|
| Machado | 2000 | New species | 1 |
| Costa et al. | 2000 | New species | 1 |
| Machado | 2002 | New species | 1 |
| Tennessen | 2004 | New species | 1 |
| Machado | 2005a | New species | 1 |
| Machado | 2005b | New species | 1 |
| Garrison | 2006 | New species | 1 |
| Machado | 2006 | New species | 1 |
| Machado & Bedê | 2006 | New species | 1 |
| Machado | 2007a | New species | 1 |
| Machado | 2007b | New species | 1 |
| Pessacq & Costa | 2007 | New species | 2 |
| Costa et al. | 2009 | New species | 1 |
| Santos et al. | 2010 | New species | 1 |
| Machado | 2010 | New species | 1 |
| Almeida et al. | 2013 | New record | 1 |
| Machado | 2014 | New species | 1 |
| Machado | 2015 | New species | 1 |
| Machado & Bedê | 2015 | New species | 7 |
| Bedê et al. | 2015 | New records | 6 |
| Guillermo-Ferreira et al. | 2016 | New species | 1 |
| Vilela et al. | 2016 | New record | 1 |
| Pinto & Almeida | 2016 | New species | 1 |
| Ávila-Júnior et al. | 2017 | New species | 1 |
| De Souza et al. | 2017 | New records | 7 |
| Borges et al. | 2018 | New records | 1 |
| Barbosa et al. | 2019 | New records | 6 |
| Vilela et al. | 2019 | New species | 1 |
| This study | New records | 2 | |
| Total | 51 |
2. First records and new species
We found two new records of odonate species for Minas Gerais, one belonging to Coenagrionidae and the other to Calopterygidae: Neoneura waltheri Selys and Hetaerina hebe Selys, respectively. Also, we found a new species from the Heteragrionidae family: Heteragrion denisye Vilela, Koroiva and Guillermo-Ferreira.
Hetaerina hebe Selys, three males (Figure 3h): This species is recorded in the states of Rio de Janeiro (Santos 1970) and São Paulo (Lencioni 2017). Our specimens of H. hebe were found in streams with channel shading by riparian vegetation and presence of rapids in sampling sites of Varginha and Serra da Canastra. We have noticed females perform endophytic oviposition.
Neoneura waltheri Selys, one male (Figure 3i): Neoneura waltheri occurs in Brazil and Argentina. There are records of this species in the states of Rio de Janeiro and Paraná. Specimens collected at the margins of the Tijuco River in Ituiutaba were perched in a clump of Elionurus sp. Jurzitza (1981), made some observations on specimens in National Park of Iguazu, Argentina. We have noticed females perform endophytic oviposition in tandem with the male, at running waters.
Heteragrion denisye Vilela, Koroiva and Guillermo-Ferreira, one male: this species was recently described (Vilela et al. 2019) inhabiting altitude fields within the Serra da Canastra National Park. We only found one specimen despite sampling the type location in two occasions. This species is peculiar because was found in small bushes in an open area, whereas other Heteragrion species are known to inhabit shaded areas with gallery woods.
3. Endemic and peculiar species
In addition to these records, we collected some of the endemic species of Serra da Canastra described by Machado & Bedê (2015), such as Franciscagrion franciscoi Machado & Bedê, Franciscagrion longispinum Machado & Bedê, Franciscobasis franciscoi Machado & Bedê, Franciscobasis sonia Machado & Bedê, Minagrion franciscoi Machado & Bedê (Fig. 2e) and Oxyagrion franciscoi Machado & Bedê. Despite the extensive sampling effort throughout western region of Minas Gerais, these species remain restricted to the Serra da Canastra National Park.
Given our large sampling size, some hard to sample species were collect in Serra da Canastra such as Limnetron debile Karsch and Neocordulia volxemi Selys, both at sites with running water and little shaded areas. Additionally, the larvae of Castoraeschna corbeti Carvalho, Pinto & Ferreira, a possible new Libellulidae species and the hitherto undescribed female of Mnesarete rhopalon Garrison were also sampled.
4. Behavioral and ecological traits
As most of the collections were made in the course of an hour, freshwater habitat selection was the only ecological trait that could be inferred for all species. We emphasize that some of the observed traits were already reported in the literature and the data observed by us are highlighted in bold (Table 4).
Table 4 Odonata species collected in this study with information about freshwater habitat selection, territoriality status, mate-guarding (MGB) and oviposition behavior. LE: lentic habitat; LO: lotic habitat; FA: female alone; IT: in tandem; NC: no contact; T: territorial; NT: non-territorial; EX: exofitic oviposition; END: endofitic oviposition; PE: percher; FL: flier; IUCN Status: LC (least concern), DD (data deficient); ND: no data recorded in the study area or literature. * First record to Minas Gerais State; ** Under evaluation by IUCN specialists. Traits in bold were observed in this study.
| Species | Habitat | Territoriality | MGB | Oviposition | IUCN status | Sampled localities | References other than this study |
|---|---|---|---|---|---|---|---|
| Acanthagrion aepiolum | LO | ND | ND | ND | ND | CER, GMP1, MAT2, MAT4, PAR, GIL | Tennessen, 2004; von Ellenrieder & Garrison, 2007; Muzón et al. 2008 |
| Acanthagrion gracile | LO, LE | ND | IT | END | ND | CER, CVG, ESS, GMP1, MAT3, PAR, SRI, CAR, TCH, QUI, LZZ | Paiva-Silva et al. 2010; Lencioni, 2011; Calvão et al. 2014; Nobre & Carvalho, 2014 |
| Acanthagrion lancea | LO, LE | NT | FA | END | ND | CVG, ITBSL, MAT4, PAR | von Ellenrieder & Garrison, 2007; von Ellenrieder, 2009b; Paiva-Silva et al. 2010; Vilela et al. 2016 |
| Acanthagrion temporale | LO, LE | ND | ND | ND | ND | RSF, P31 | Leonard, 1977; ouza, 2003 |
| Acanthagrion truncatum | LE | NT | IT | END | LC | MAT2, MAT4, QUI, LZZ | von Ellenrieder, 2009; Guillermo-Ferreira & Del-Claro, 2012a, 2013; Vilela et al. 2016 |
| Allopodagrion contortum | LO | ND | ND | ND | LC | GUA, VCT | von Ellenrieder, 2009 |
| Archaeogomphus infans | LE | ND | ND | ND | ND | ZZC | Barbosa et al. 2019 |
| Anax amazili | LE | ND | ND | END | ND | GMP2 | Convey, 1989; von Ellenrieder & Garrison, 2007 |
| Argia claussenii | LO | ND | ND | ND | ND | CPV, GUA, VCT | Lencioni, 2017 |
| Argia croceipennis | LO | ND | ND | ND | LC | NAS | Costa et al. 2008; von Ellenrieder, 2009; Dalzochio et al. 2011 |
| Argia lilacina | LO | ND | ND | ND | ND | AQD, RCV, CER, CE, CRP, CHI, CAR, PO2, PR2 | Machado, 1988; Machado & Bedê, 2015 |
| Argia modesta | LO | ND | ND | ND | ND | CPV, VIS, SUL, ZZC, CAN, VCT, PR2 | Ferreira-Peruquetti, & Trivinho-Strixino, 2003 |
| Argia mollis | LO | ND | ND | ND | ND | CAR, GIL, SUL, CAN | Cezário et al. 2018 |
| Argia reclusa | LO | T | IT, FA | END | ND | ITBT, MAT1, MAT2, PAR, EPA, GIL, TCH, CPV | Guillermo-Ferreira & Del-Claro, 2012b; Vilela et al. 2016 |
| Argia smithiana | LO | ND | ND | ND | ND | CRP, ITBSL, ITBT, MAT2, CPV | Vilela et al. 2016 |
| Argia tamoyo | LO, LE | ND | ND | ND | ND** | AQD, CRP, RCV, CER, ESS, RSF, AGB, CHI, CAN | Dalzochio et al. 2011 |
| Brechmorhoga praedatrix | LO | ND | ND | ND | LC | CPV | Fleck, 2004; von Ellenrieder, 2009 |
| Cacoides latro | LE | ND | ND | ND | ND | GMP2 | Borges et al. 2019 |
| Castoraeschna januaria | LO | ND | ND | ND | ND | CRP, P17 | Carvalho et al. 2009 |
| Coryphaeshna adnexa | LE | ND | ND | ND | LC | UNAI | von Ellenrieder & Garrison, 2007; Paulson, 2009; Vilela et al. 2016 |
| Cyanallagma ferenigrum | LE | T | ND | END | DD | MAT4 | von Ellenrieder, 2009a; Vilela et al. 2016 |
| Diastatops obscura | LE | T | NC | EXO | ND | CVC | Vilela et al. 2016 |
| Epipleoneura machadoi | LO | ND | IT | END | ND | UNAI | Rácenis, 1960; Lencioni, 2017 |
| Epipleoneura williamsoni | LO | NT | IT | END | LC | ITBSL, UNAI | von Ellenrieder, 2009; Vilela et al. 2016 |
| Erythemis credula | LE | ND | ND | ND | ND | ITBT | Vilela et al. 2016 |
| Erythemis vesiculosa | LE | ND | FA | END | LC | GMP2, UDI, UNAI | von Ellenrieder & Garrison, 2007; Paulson, 2009; Vilela et al. 2016 |
| Erythrodiplax ana | LE | T | NC | EXO | ND** | NAS, CCD | Guillermo-Ferreira et al. 2016; Vilela et al. 2016 |
| Erythrodiplax castanea | LE | ND | ND | ND | ND | AQD, CAR, JOR, CPV, ZZC, QUI | Brooks, 1989; von Ellenrieder, 2011; Monteiro-Junior et al. 2013; Borges et al. 2019 |
| Erythrodiplax fusca | LO, LE | T | NC | EXO | ND | GMP1, ITBL, ITBT, MAT3, PAR, UDI, AQD, CER, RSF | Convey, 1989; Louton et al. 1996; Paiva-Silva et al. 2010; Nobre & Carvalho, 2014; Vilela et al. 2016 |
| Erythrodiplax juliana | LE | T | NC | EXO | LC | ITBL, MAT2, PAR, CRP, AQD, RCV, ESS | von Ellenrieder, 2009; Vilela et al. 2016 |
| Erythrodiplax latimaculata | LE | T | NC | EXO | ND | GMP1, ITBT, MAT2, MAT3, UNAI | Calvão et al. 2013; Vilela et al. 2016 |
| Erythrodiplax lygaea | LE | ND | FA | EXO | LC | CAN | von Ellenrieder, 2009 |
| Erythrodiplax pallida | LO, LE | ND | ND | EXO | LC | NAS | Costa et al. 2001; von Ellenrieder, 2009; Del Palacio & Muzon, 2016 |
| Erythrodiplax umbrata | LE | T | FA | EXO | ND | CVG, PAR, PRA, UDI, UNAI | Garcia-Diaz, 1937; Convey, 1989; Paulson, 1998; Von Ellenrieder & Garrison, 2007; Vilela et al. 2016 |
| Franciscagrion franciscoi | LO, LE | NT | IT | END | ND** | NAS | Machado & Bedê, 2015 |
| Franciscagrion longispinum | LO, LE | NT | ND | ND | ND** | NAS | Machado & Bedê, 2015 |
| Franciscobasis franciscoi | LO, LE | NT | ND | ND | ND** | NAS | Machado & Bedê, 2015 |
| Franciscobasis sonia | LO, LE | NT | ND | ND | ND** | NAS | Machado & Bedê, 2015 |
| Heliocharis amazona | LO | T | ND | ND | ND | ITBSL, PAR | Dunkle, 1991; Louton et al. 1996 |
| Hetaerina hebe* | LO | T | ND | END | LC | TCH, SUL, VCT | von Ellenrieder, 2009 |
| Hetaerina longipes | LO | T | ND | ND | ND | ITBT, PAR | Pestana et al. 2018 |
| Hetaerina rosea | LO | T | IT, NC | EXO | ND | ITBSL, ITBT, MAT3, PAR, EPA, CAR, GIL, CPV, ZZC | Von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010; Guillermo-Ferreira & Del-Claro, 2011; Vilela et al. 2016 |
| Hetaerina simplex | LO | ND | ND | ND | ND** | CRP, AQD, RCV, CER, BS, ESS, RSF, AGB, CHI, PSG, CCD, QUI, NAS, PO2, PR2 | Garrison, 1990; Lencioni, 2017 |
| Heteragrion aurantiacum | LO | T | ND | ND | LC | MAT2, UNAI | Von Ellenrieder, 2009 |
| Heteragrion denisye | LO | ND | ND | ND | ND | PO2 | Vilela et al. 2019 |
| Heteragrion gracile | LO | ND | ND | ND | ND | CAR, SUL | Machado, 2006; Lencioni, 2013 |
| Heteragrion triangulare | LO | ND | ND | ND | ND | CER, GUA | This study |
| Homeoura chelifera | LO | NT | IT | ND | ND | MAT4, JOR | von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010; Vilela et al. 2016 |
| Idiataphe longipes | LE | ND | ND | ND | ND | ZZC, QUI | Oldrini & Mascarenhas, 2005; Borges et al. 2019 |
| Ischnura capreolus | LE | NT | FA | END | ND | MAT2, MAT4, PRA, UDI, UNAI, RSF, JOR, LZZ | Paiva-Silva et al. 2010; Nobre & Carvalho, 2014; von Ellenrieder & Garrison, 2007; Vilela et al. 2016; Vilela et al. 2017 |
| Ischnura fluviatilis | LE | NT | FA, NC | ND | LC | GMP1, PRA, UNAI | von Ellenrieder & Garrison, 2007; von Ellenrieder, 2009 |
| Lestes auritus | LE | ND | IT | ND | LC | UDI | von Ellenrieder, 2009; Renner et al. 2016a |
| Lestes bipupillatus | LE | NT | ND | ND | ND | P31 | Cardoso-Leite et al. 2014 |
| Lestes forficula | LE | NT | IT, FA | END | ND | CVG, ITBL, MAT2, PRA, UNAI, QUI, P31 | Garcia-Diaz, 1937; Von Ellenrieder & Garrison, 2007; Nobre & Carvalho, 2014 |
| Limnetron debile | LO | ND | ND | ND | ND | P17 | Santos, 1970; Assis et al. 2000 |
| Macrothemis imitans imitans | LO | ND | ND | ND | ND | SRI, GIL, CPV | Salgado et al. 2013 |
| Micrathyria hesperis | LE | ND | NC | ND | ND | CVG, GMP1, PAR | Von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010; Vilela et al. 2016 |
| Micrathyria ocellata dentiens | LO | ND | ND | ND | ND | SRI | Costa et al. 2002 |
| Micrathyria spuria | LO | ND | ND | ND | LC | QUI | Costa et al. 2002; von Ellenrieder, 2009 |
| Micrathyria stawiarskii | LO, LE | ND | ND | ND | LC | CPV, NAS, CCD, PO1 | Costa et al. 2002; von Ellenrieder, 2009 |
| Minagrion franciscoi | LO, LE | NT | ND | ND | ND** | NAS, PO1 | Machado & Bedê, 201 |
| Minagrion waltheri | LE | NT | ND | ND | LC | ITBSL, MAT1, PO1, PO2 | von Ellenrieder, 2009; Vilela et al. 2016 |
| Mnesarete guttifera | LO | T | ND | ND | LC | AQD, RCV, CER, BS, CE, ESS, RSF, AGB, CHI, CRP, ZZC, VCT | Garrison, 2006; von Ellenrieder, 2009 |
| Mnesarete pudica pudica | LO | T | FA | END | ND | CVG, ITBSL, ITBT | Guillermo-Ferreira & Bispo, 2012; Vilela et al. 2016 |
| Mnesarete rhopalon | LO | T | ND | ND | ND** | CHI, CRP, CCD | Garrison, 2006 |
| Neocordulia volxemi | LO | ND | ND | ND | DD | GUA | von Ellenrieder, 2009 |
| Neoneura sylvatica | LO, LE | ND | IT | END | ND | ITBSL, UNAI | Paiva-Silva et al. 2010; Nobre & Carvalho, 2014; Vilela et al. 2016 |
| Neoneura waltheri* | LO | ND | IT | END | ND | ITBT | Jurzitza, 1981; Garrison, 1999 |
| Oligoclada abbreviata | LE | ND | ND | ND | LC | CVG, GMP1, ITBL, PAR, QUI | von Ellenrieder, 2009 |
| Orthemis aequilibris | LE | ND | NC | EXO | ND | GMP2, ITBT | von Ellenrieder & Garrison, 2007 |
| Orthemis discolor | LE | T | NC | EXO | ND | GMP2, ITBL, ITBT, MAT1, MAT2, MAT3, PRA, UDI, ESS, LZZ | von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010 |
| Oxyagrion basale | LE | ND | ND | ND | ND | CRP, AQD, RCV, CER, BS, CE, ESS, RSF, CHI, GIL, PSG, LZZ | Hanauer et al. 2014 |
| Oxyagrion chapadense | LO | ND | ND | ND | ND | PAR, NAS | Calvão et al. 2014; Tennessen, 2015 |
| Oxyagrion franciscoi | LO | ND | ND | ND | ND** | PSG | Machado & Bedê, 2015 |
| Oxyagrion hempeli | LO | ND | ND | ND | ND | RL, PO1 | Costa et al. 2010; Renner et al. 2016b |
| Oxyagrion impunctatum | LO | ND | ND | ND | ND | NAS, PSG | Costa, 1981 |
| Oxyagrion microstigma | LE | T, NT | IT | END | ND | ITBT, UDI, UNAI, NAS, PO1, PO2, QUI, P31, LZZ | Vilela et al. 2016 |
| Oxyagrion santosi | LE | ND | ND | ND | ND | UDI, NAS, PO2 | Costa et al. 2000; Borges et al. 2019 |
| Oxyagrion terminale | LE | ND | ND | ND | ND | CER, CE, CAR, TCH | Capitulo, 1996; Von Ellenrieder & Garrison, 2006; Hanauer et al. 2014 |
| Pantala flavescens | LE, LO | T, NT | NC, FA, IT | EXO | LC | CVC, GMP2, ITBSL, PRA | Convey, 1989; Dumont, 1991; Mitra, 2000; von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010 |
| Perithemis lais | LE | T | ND | ND | LC | CVC, PAR | von Ellenrieder, 2009 |
| Perithemis mooma | LE | T | ND | ND | ND | CVC, ITBSL, MAT1, PAR, PRA, UDI, LZZ | Louton et al. 1996; Von Ellenrieder & Garrison, 2007; Paiva-Silva et al. 2010; Pinto et al. 2013 |
| Progomphus costalis | LO | ND | ND | ND | LC | NAS | Belle, 1973; von Ellenrieder, 2009 |
| Progomphus intricatus | LO | ND | ND | ND | LC | ZZC | Belle, 1973; von Ellenrieder 2009; Dalzochio et al. 2011 |
| Telebasis carmesina | LE | NT | IT | END | LC | ITBT, PAR, UDI, UNAI, BS, RSF, QUI, P31, LZZ | von Ellenrieder, 2009 |
| Telebasis coccinea | LE, LO | ND | ND | ND | ND | UDI | Garrison, 2009; Calvão et al. 2018; Borges et al. 2019 |
| Tholymis citrina | LE, LO | ND | ND | ND | LC | UNAI | Fleck et al. 2004; Paulson 2017 |
| Tigriagrion aurantinigrum | LO | T | ND | ND | ND | ITBSL, ITBT, MAT4, GIL | Paiva-Silva et al. 2010; Vilela et al. 2016 |
| Tramea calverti | LE | ND | ND | ND | LC | P31 | Paulson, 2017; Borges et al. 2019 |
| Zenithoptera lanei | LE | NT | IT, NC, FA | EXO | ND | ITBT, MAT1 | Vilela et al. 2016 |
Data on behavioral traits are derived from personal observations or from literature: territoriality status, mate-guarding behavior and oviposition behavior. Of all traits possible, however, 168 (63%) were not found on the literature or observed in our field study (i.e. ND). Moreover, 40 (45%) of all species had no behavioral information available. Territoriality was the most representative trait, determined for 38 (42%) of the species. Mate-guarding behavior (MGB) and oviposition were determined for 31 (35%) species (Table 4).
5. IUCN status
The IUCN conservational status was available for only 28 (31.5%) of the sampled species. Most of the evaluated species (26 out of 28) were assessed as being Least Concern by IUCN classification, whereas for Cyanallagma ferenigrum de Marmels and Neocordulia volxemi Selys are classified as Data Deficient (DD) (Table 4).
Discussion
After this study, with 269 recorded species, Minas Gerais is the Brazilian State with the second highest number of Odonata species, considering that Rio de Janeiro has recorded 280 species (Costa & Santos 2000), followed by São Paulo, 251 species (Costa, Machado, et al. 2000), Mato Grosso do Sul, 209 species (Rodrigues et al. 2018, Koroiva et al. 2017b), Rio Grande do Sul, 182 species (Dalzochio et al. 2018), and Goiás and the Distrito Federal, 152 species (Nóbrega & De Marco 2011). Obviously, such numbers reflect the areas where most Brazilian odonatologists explored, and the northern States of Brazil may concentrate an even higher number of species.
Concerning the sampled areas, 18 of the 49 sites were at or near Veredas, 24 areas were consisted of streams, rivers and waterfalls, and only seven of them were ponds. This consists, so far, in the most extensive (in number of species recorded and sampled areas) study of odonates in the State of Minas Gerais. Among the sampled sites, the Veredas were mostly well preserved, however few (five out of 18) of them were placed in protected areas. In fact, the Vereda in Uberlândia (UDI) is already threatened by a urbanization project with housing activities. Such anthropic pressure may permanently impact on the faunal composition of this and other unprotected areas, as Vereda areas do not regenerate after a severe disturbance (Guimarães et al. 2002; Oliveira et al. 2009).
Bearing in mind the information on species behavior, we provide a list of behavioral traits of our focal observations and the ones found in the literature. Despite being available for 55% of the sampled species, most of the information on species biology is underexplored, what requires data from more than one source per species to build a more complete behavioral perspective. Considering that only 40 of 90 species have biological information, it is important to know how much to we must advance in this sense. Although neglected, behavioral diversity is an important species feature, for it reveals a cryptic biodiversity (Cordero-Rivera 2017). For instance, for species that are morphologically similar but behave in different ways, the behavioral repertoire can be a distinguishable trait between them (Guillermo-Ferreira & Bispo 2012, Cordero-Rivera 2017).
Here, we also show that few species were assessed regarding their conservational status. Considering the current measures from the Latin-American Odonatological Society and IUCN to assess all Latin-American odonate species, our results make a great contribution to this effort. The IUCN evaluations require data of distribution ranges, altitude of occurrence, population status, and biological data (IUCN 2019). Therefore, the more information one can gather about a species, more precisely it can be evaluated. In our study, only 31.5% of the sampled species were already assessed by IUCN. Most of these species fit in the Least Concern category, which means that, although some of the species may be widespread and out of risk (IUCN 2019), others that were not assessed may be under threat. For Cyanallagma ferenigrum de Marmels and Neocordulia volxemi Selys, the status appears as Data Deficient, which means that, at the moment of the assessment, the data available for evaluation were insufficient to set a conservational status to these species (IUCN 2019). Our records for this two species and the others that were not assessed will help in ongoing assessments. The results presented here respect the guidelines of the Latin-American Odonatological Society and are being used to assess Neotropical species conservation status. Finally, we suggest that research focus on studies of natural history and distributional ranges that cover understudied species (Data Deficient and unevaluated species) in the Cerrado biome (mainly on Veredas) to ensure a broader perspective on the Neotropical odonate fauna.
