Checklist of dragonflies and damselflies (Insecta: Odonata) of the Amazonas state, Brazil

Checklist das libélulas (Insecta: Odonata) do estado do Amazonas, Brasil

Abstract:

Here we provide a checklist of the odonates from Amazonas state, Brazil. We registered 324 species and 101 genera, making Amazonas the Brazilian state with the most Odonata species recorded. The families with the highest number of species were Coenagrionidae with 32 genera and 101 species, followed by Libellulidae with 28 genera and 100 species and Gomphidae with 12 genera and 45 species. Some regions of Amazonas state remain poorly explored, such as the southern area, and large municipalities, such as São Gabriel da Cachoeira. This work underlines the importance of the biological diversity from Amazonas state and the Amazonian Biome for Odonata species richness in Brazil and shows that many areas in the world’s largest tropical forest have not yet been sampled.

Keywords:
Aquatic insects; List of species; Anisoptera; Zygoptera; Amazon; Neotropical region

Resumo:

Aqui nós apresentamos a lista de espécies de libélulas presentes no estado do Amazonas, Brasil. Nós registramos 324 espécies e 101 gêneros, tornando o estado com o maior número de espécies de libélulas registradas no país. As famílias com maior número de espécies foram Coenagrionidae, com 32 gêneros e 101 espécies, seguido por Libellulidae, com 28 gêneros e 100 espécies e Gomphidae com 12 gêneros e 45 espécies. Regiões do estado do Amazonas permanecem pouco exploradas, como a parte sul e os municípios com grande extensão territorial, como São Gabriel da Cachoeira. Este trabalho reconhece a importância da diversidade biológica do estado do Amazonas e do bioma Amazônico para a riqueza de espécies de Odonata no Brasil e mostra que ainda existem muitas áreas não amostradas na maior floresta tropical do mundo.

Palavras-chave:
Insetos aquáticos; lista de espécies; Anisoptera; Zygoptera; Amazônia; região Neotropical

Introduction

Considered one of the most diverse regions for Odonata, the Neotropical region has more than 1,700 species, and in Brazil 749 species are registered (Olaya 2019). Dragonflies and damselflies (Insecta: Odonata) constitute an important group of aquatic insects characterized by their extensive predation, with larvae being important links for fish and other aquatic vertebrates in food webs. Besides their ecological importance, odonates have great artistic appeal, and have been used in environmental quality assessments using several environmental quality metrics (Abbott 2015).

Even with such importance, the taxonomic knowledge about Odonata in Brazil is still heterogeneous. As evidenced by Dalzochio et al. (2018), Brazilian studies concerning Odonata fauna are mainly concentrated in the south-center of Brazil, with species lists registered for the states of São Paulo (Costa et al. 2000), Rio de Janeiro (Costa & Santos 2000), Mato Grosso do Sul (Koroiva et al. 2017; Rodrigues et al. 2018), Rio Grande do Sul (Dalzochio et al. 2018; Pires et al. 2019), Espírito Santo (Costa & Oldrini 2005), Goiás and the Federal District (Nóbrega & De Marco 2011). Despite the increasing number of new species descriptions, the lack of species lists in other regions and states make the analysis of current sampling distributions and knowledge about Odonata species distribution inviable, which directly affects large-scale conservation assessments (Silveira et al. 2010).

The Amazonas state located in the northern region of Brazil, presents the largest territorial extension and stands out (IBGE 2019) as it encompasses Amazonian and Cerrado biomes (Parédio 2012). In recent years, political issues and increased deforestation rates in both biomes have raised concerns for the maintenance of their forested areas and associated fauna (Rochedo et al. 2018, Fearnside 2019). Regarding Odonata, several studies have been dedicated to increasing knowledge about its fauna in Amazonas state in the last 120 years, however, almost all of them are restricted to taxonomic descriptions and ecological analysis and do not provide information about the total number of species. In the most recent estimate, Neiss & Hamada (2014) indicated about 262 Odonata species in Amazonas, representing 35% of the known species from Brazil. Considering this information gap in Amazonas, the main objective of this work was to present the diversity of Odonata species in this state, as well as their known locations in the municipalities.

Materials and Methods

1. Study area

The state of Amazonas has an area of 1,559,168 km² (IBGE 2019) and stands out due to the predominant presence of the Amazonian Biome. The state shares borders with Pará, Mato Grosso, Rondônia, Acre and Roraima states and with the countries of Venezuela, Peru and Colombia (Figure 1). The geographical relief of Amazonas state is constituted by depressions (Western Amazon Depression, Northern Amazonian Depression and Southern Amazon Depression), plains, and plateaus (North-Amazonian Residual Plateau and Plateau of the Eastern Amazon). Its hydrology is formed mainly by the eastern basin of the Amazon River and is composed of most of the rivers from this hydrographic basin, whose tributaries are the rivers Negro, Japurá, Solimões, Juruá, Purus and Madeira. The northern and eastern regions of the state are located in the northern Amazonian plateau, which contain the highest altitudes (above 2,950 m) in Brazil: “Pico da Neblina” and “31 de Março” mountains (Parédio 2012). The vegetation types vary from Ombrophylous Forest to “Campinarana” and Savana in the Amazon and Cerrado biomes. According to Köppen’s climatic classification, the predominant climate is Af type, tropical without dry season, except in the southern part of the state, which is considered Am, tropical monsoon. The average annual precipitation varies between 1,900 mm in the extreme southwest and above 3,000 mm in other areas and average temperatures are above 23° C (Alvares et al. 2013).

Figure 1
Left, Map of South America highlighting the geopolitical division of Brazil and Amazonas State (grey) (Datum: WGS 84); Right, Municipalities with Odonata recorded in Amazonas State. For code numbers see Table 1.

2. Elaboration of the list

To prepare the list, we gathered information from the Invertebrates Collection at the Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil. Additional data were collected from 102 publications, published between 1897 and 2019, the sites SpeciesLink (http://splink.cria.org.br/) and “Catálogo Taxonômico da Fauna do Brasil” (http://fauna.jbrj.gov.br ; “Brazilian Fauna Taxonomic Catalog” in English; Pinto 2019) (see Table 1). We also gathered information about the municipalities where the species were collected (Table 1). For systematic classification, we followed Dijkstra et al. (2014, 2013) and Fleck (2018).

Table 1
Numbers and codes for municipalities and references used in the list of Odonata species from Amazonas state, Brazil. Where the municipality is unknown, only the state is provided.

Municipalities	Code	Number	References	Number
State of Amazonas (municipality not informed)	AMZ	-	Heckman 2006	1
Manaus	MAO	1	Pinto 2019	2
Presidente Figueiredo	PRF	2	Neiss 2012	3
Atalaia do Norte	ATA	3	Belle 1973	4
Barcelos	BAR	4	Belle 1983	5
Benjamin Constant	BEC	5	Belle 1984	6
Beruri	BER	6	Belle 1989	7
Borba	BOR	7	Belle 1994	8
Careiro/Careiro da Várzea	CAR	8	Belle 1998	9
Coari	COA	9	Bick & Bick 1985	10
Eirunepé	EIR	10	Bick & Bick 1986	11
Fonte Boa	FOB	11	Bick & Bick 1988	12
Guajará	GJR	12	Borror 1931	13
Humaitá	HMT	13	Borror 1942	14
Iranduba	IRN	14
Itacoatiara	ITA	15	Calvert 1908	15
Juruá	JUR	16	Costa & Oldrini 2003	16
Manacapuru	MPU	17	Costa & Santos 1997	17
Manicoré	MNE	18	Costa 1990	18
Maués	MAU	19	Costa et al. 2002	19
Nhamundá	NHA	20	Lencioni 2005	20
Novo Airão	NAI	21	De Marco 1998	21
Novo Aripuanã	NRP	22	De Marmels & Rácenis 1982	22
Parintins	PAR	23	De Marmels 1987	23
Rio Preto da Eva	RPE	24	De Marmels 1989	24
Santa Isabel do Rio Negro	SIR	25	De Marmels 2001	25
Santo Antônio do Içá	SAI	26	De Marmels & Neiss 2011	26
São Gabriel da Cachoeira	SGC	27	De Marmels & Neiss 2013	27
São Paulo de Olivença	SPO	28	Dunkle 1991	28
Tabatinga	TAB	29	Fleck & Neiss 2012	29
Tapauá	TAP	30	Fulan et al. 2015	30
Tefé	TEF	31	Garrison & Ellenrieder 2006	31
Tonantins	TON	32	Garrison & Ellenrieder 2009	32
			Garrison & Ellenrieder 2015	33
			Garrison 1990	34
			Garrison 1999	35
			Garrison 2004	36
			Garrison 2006	37
			Garrison 2009	38
			Garrison 2014	39
			Geijskes 1963	40
			Geijskes 1970	41
			Geijskes 1984	42
			Guillermo-Ferreira et al. 2014	43
			Hamada & Oliveira 2003	44
			Kirby 1897	45
			Lencioni 2006	46
			Lencioni 2013	47
			Leonard 1977	48
			Machado & Lacerda 2017	49
			Machado 2002	50
			Machado 2007	51
			Machado 2009a	52
			Machado 2009b	53
			Machado 2012	54
			Machet 1990	55
			May 1991	56
			Monteiro-Júnior et al. 2013	57
			Montgomery 1940	58
			Neiss & Hamada 2010	59
			Neiss & Hamada 2016	60
			Neiss et al. 2008	61
			Neiss et al. 2013	62
			Pessacq 2014	63
			Pessacq et al. 2012	64
			Pinto & Carvalho 2010	65
			Pinto & Lamas 2010	66
			Pinto & Lamas 2011	67
			Pinto 2013	68
			Pujol-Luz & Fonseca 1997	69
			Ris 1911	70
			Ris 1930	71
			Rodrigues et al. 2016	72
			Rodríguez et al. 2015	73
			Santos & Machado 1960	74
			Santos 1954	75
			Santos 1961	76
			Santos 1967	77
			Santos 1970a	78
			Santos 1970b	79
			Schmidt 1952	80
			Spindola et al. 2001	81
			St. Quentin 1973	82
			Tennessen 2015	83
			Torreias et al. 2008	84
			von Ellenrieder & Garrison 2003	85
			von Ellenrieder & Garrison 2008	86
			Vilela et al. 2018	87
			von Ellenrieder 2008	88
			von Ellenrieder 2009	89
			von Ellenrieder 2013	90
			Walker 2009	91
			Williamson & Williamson 1924b	92
			Williamson & Williamson 1924a	93
			Williamson 1923	94
			Species Link website	95
			Neiss & Hamada 2012	96
			Juen & De Marco 2011	97
			Juen 2011	98
			Vilaça 2017	99
			Heckman 2008	100
			De Marco-Júnior et al. 2015	101
			Ris 1910	102
			Invertebrates Collection (INPA)	103
			Neiss & De Marmels 2017	104
			Vilela et al. 2019	105

Results

The total number of Odonata species recorded in Amazonas is 324 (Table 2). Some species recorded in this study are shown in Figure 2. The total number of genera recorded for the state is 101, distributed in 16 families; this number include one genus from the Calopterygoidea “Incertae sedis group 3” and one genus from the Libelluloidea “incertae sedis”. Coenagrionidae is the family with the highest number of records, with 32 genera and 101 species, followed by Libellulidae with 28 genera and 100 species, Gomphidae with 12 genera and 45 species, and Aeshnidae with eight genera and 27 species. The other families are: Calopterygidae (two genera, 12 species), Corduliidae (two genera, four species), Dicteriadidae (two genera, two species), Heteragrionidae (two genera, seven species), Lestidae (one genus, one species), Megapodagrionidae (one genus, one species), Perilestidae (two genera, seven species), Philogeniidae (one genus, two species), Platystictidae (one genus, one species), Polythoridae (three genera, 10 species), Rimanellidae (one genus, one species) and Oxygastridae sensuFleck (2018) (one genus, one species). Calopterygoidea “incertae sedis group 3” was represented by one genus and one species and Libelluloidea incertae sedis was also represented by one genus and one species.

Table 2
Odonata species recorded for Amazonas State, Brazil. For references and municipalities codes, please check Table 1.

Species	Municipalities	References
ZYGOPTERA
Dicteriadidae
Dicterias atrosanguinea Selys, 1853	MPU; MAO; PRF; NAI; RPE	3; 95; 1; 57; 87; 97; 103
Heliocharis amazona Selys, 1853	PRF; TEF	3; 28; 103
Calopterygidae
Hetaerina amazonica Sjöstedt, 1918	ITA; BAR; MPU; MAO; PRF; NRP; RPE; NAI	3; 95; 1; 57; 97; 103
Hetaerina auripennis (Burmeister,1839)	AMZ	100
Hetaerina caja (Drury, 1773)	BAR	3; 103
Hetaerina laesa Hagen in Selys, 1853	BAR	3; 103
Hetaerina medinai Rácenis, 1968	BAR	3; 100; 103
Hetaerina moribunda Hagen in Selys,1853	MAO; NAI; PRF; RPE; MPU	3; 100; 57; 34; 97
Hetaerina sanguinea Selys, 1853	MAO	3; 95; 100; 57; 103
Hetaerina westfalli Rácenis,1968	BAR; MPU; MAO; RPE; NAI	3; 100; 34; 103
Mnesarete aenea (Selys,1853)	BER	37; 100
Mnesarete astrape De Marmels,1989	MPU; MAO; PRF; RPE; BAR; NAI	3; 95; 100; 37; 103
Mnesarete cupraea (Selys,1853)	BAR; PRF; RPE; SPO; MAO	3; 95; 100; 37; 103
Mnesarete loutoni Garrison,2006	SPO	1; 37; 46
Polythoridae
Chalcopteryx rutilans (Rambur 1842)	NRP	3
Chalcopteryx scintillans McLachlan, 1870	BAR; ITA; MPU; MAO; PRF; RPE; NAI; BEC; SGC; SPO	3; 95; 1; 98; 43; 74; 57; 103
Chalcopteryx seabrai Santos & Machado,1961	PRF	3; 103
Euthore fasciata (Hagen in Selys, 1853)	AMZ	100
Polythore aurora (Selys, 1879)	TEF	100; 11; 20
Polythore batesi (Selys, 1879)	SPO	100; 11; 20
Polythore beata (McLachlan, 1869)	FOB	100; 11; 20
Polythore picta (Rambur, 1842)	TEF	100; 10; 20
Polythore procera (Selys, 1869)	AMZ	100; 20
Polythore vittata (Selys, 1869)	CAR; PRF; BOR; FOB; MNE; SPO; TEF	3; 100; 11; 20
Philogeniidae
Philogenia margarita Selys,1862	TEF	12; 100
Philogenia silvarum Ris, 1918	AMZ	100
Rimanellidae
Rimanella arcana (Needham, 1933)	PRF	3; 100; 20; 44; 103
Calopterygoidea incertae sedis group 3
Dimeragrion percubitale Calvert, 1913	BAR	3; 103
Heteragrionidae
Heteragrion angustipenne Selys, 1886	TAP	100; 105
Heteragrion bariai De Marmels, 1989	PRF; MAO	3; 95; 103
Heteragrion ictericum Williamson,1919	MAO; RPE; MPU	95; 100; 47; 57
Heteragrion icterops Selys, 1862	MAO; NAI; RPE	3; 100; 23; 103
Heteragrion silvarum Sjöstedt,1918	MAO	95; 100; 23; 47; 97
Oxystigma cyanofrons Williamson, 1919	NAI; PRF; RPE; MAO	3; 95; 39; 103
Oxystigma petiolatum (Selys, 1862)	BAR; ITA; MPU; NAI; PRF; RPE; MAO	95; 3; 39; 57; 97
Megapodagrionidae
Megapodagrion megalopus (Selys, 1862)	BAR; TAP; SGC; COA	3; 100; 99; 25; 103
Lestidae
Lestes falcifer Sjöstedt, 1918	BAR	3; 100; 103
Perilestidae
Perilestes attenuatus Selys, 1886	BAR; MAO; NAI; PRF	3; 95; 100; 59; 92; 57; 103
Perilestes bispinus Kimmins, 1958	AMZ	100; 20
Perilestes fragilis Hagen in Selys, 1862	AMZ	100
Perilestes gracillimus Kennedy, 1941	MAO	95; 100
Perissolestes cornutus (Selys, 1886)	TEF; SPO	100; 20; 80
Perissolestes flinti De Marmels, 1988	BAR	3; 103
Perissolestes paprzyckii Kennedy, 1941	AMZ	100; 20
Platystictidae
Palaemnema brasiliensis Machado, 2009	BAR; PRF	3; 60; 103
Coenagrionidae
Acanthagrion adustum Williamson, 1916	MAO; SIR	95; 100; 48; 57; 46
Acanthagrion amazonicum Sjöstedt, 1918	MAO	95; 100; 48; 46
Acanthagrion apicale Selys, 1876	BAR; HMT; MAO	3; 30; 54; 103
Acanthagrion chararum Calvert, 1909	BAR	3; 103
Acanthagrion chicomendesi Machado, 2012	SIR; BEC; BAR; TEF; SPO	54
Acanthagrion cuyabae Calvert, 1909	MAO	3; 103
Acanthagrion flaviae Machado, 2012	ATA	54
Acanthagrion kaori Machado, 2012	MAO	54
Acanthagrion minutum Leonard, 1977	SGC	3; 103
Acanthagrion phallicorne Leonard, 1977	MAO	3; 103
Acanthagrion rubrifrons Leonard, 1977	MAO	100; 48; 46
Acanthallagma caeruleum Williamson & Williamson, 1924	TAP	100; 99; 93; 46
Aeolagrion dorsale (Burmeister, 1839)	MAO	2; 100; 57; 46
Aeolagrion inca (Selys, 1876)	AMZ	2; 100; 46
Amazoneura ephippigera (Selys, 1886)	TEF	100; 64
Argia bicellulata (Calvert, 1909)	MAO	3; 57; 103
Argia botacudo Calvert, 1909	MAO	95
Argia collata Selys, 1865	AMZ	100
Argia cuneifera Garrison & von Ellenrieder, 2015	BAR	2; 33
Argia euphorbia Fraser, 1946	BAR; NAI; MPU; TAP	2; 100; 33; 46
Argia fumigata Hagen in Selys, 1865	MAO; MPU; RPE; BAR; PRF	95; 2; 100; 33; 46
Argia gemella Garrison & von Ellenrieder, 2015	MAO; BAR; PRF; RPE	95; 2; 33
Argia hasemani Calvert, 1909	MAO	100; 46; 97
Argia impura Rambur, 1842	AMZ	100
Argia indicatrix Calvert, 1902	MAO; RPE; PRF	3; 95; 2; 100; 33; 57; 46; 97; 103
Argia infumata Selys, 1865	MPU; TEF; SPO; MNE; TON; NAI	3; 2; 100; 33; 46; 103
Argia insipida Hagen in Selys, 1865	MPU; MAO; PRF; BAR	3; 2; 100; 33; 46; 103
Argia loutoni Garrison & von Ellenrieder, 2015	TAP; BAR; MAU	2; 33
Argia meioura Garrison & von Ellenrieder, 2015	MAO; MPU; PRF	2; 33
Argia mollis Hagen in Selys, 1865	AMZ	100; 46
Argia oculata Hagen in Selys, 1865	BAR; PRF; NAI; MAO	2; 100; 33; 46; 97
Argia palmata Garrison & von Ellenrieder, 2015	MAO	2; 33
Argia pulla Hagen in Selys, 1865	MAO	2; 33
Argia reclusa Selys, 1865	AMZ	100; 46
Argia subapicalis Calvert, 1909	MAO	97
Argia thespis Hagen in Selys, 1865	MAO	97; 46
Argia tinctipennis Selys, 1865	AMZ	100; 46
Argia translata Hagen in Selys, 1865	AMZ	2;
Bromeliagrion rehni Garrison, 2005	MAO	3; 95; 100; 84; 46; 103
Calvertagrion charis Tennessen, 2015	AMZ	2; 83;
Calvertagrion minutissimum (Selys, 1876)	AMZ	2; 100; 46
Denticulobasis ariken Machado, 2009	MAO	3; 103
Dolonagrion fulvellum (Selys, 1876)	AMZ	46
Drepanoneura muzoni von Ellenrieder & Garrison, 2008	BAR	3; 103
Epipleoneura capilliformis (Selys, 1886)	MPU; MAO; NAI; PRF; RPE	3; 95; 63; 57; 103
Epipleoneura fuscaenea Williamson, 1915	NRP	3; 103
Epipleoneura haroldoi Santos, 1964	MPU; MAO; NAI; PRF	3; 100; 63; 103
Epipleoneura humeralis (Selys, 1886)	COA	3; 99; 63; 103
Epipleoneura kaxuriana Machado, 1985	MPU; MAO; NRP; PRF; RPE; NAI	3; 100; 63; 103
Epipleoneura lamina Williamson, 1915	BOR	63
Epipleoneura manauensis Santos, 1964	BAR; MPU; MAO; RPE; NAI	3; 95; 100; 98; 63; 57; 96; 103
Epipleoneura metallica Rácenis, 1955	MAO	97
Epipleoneura spatulata Rácenis, 1960	BAR	3; 103
Epipleoneura tariana Machado, 1985	BAR	3; 100; 98; 63; 103
Epipleoneura uncinata De Marmels, 1989	BAR	3; 103
Epipotoneura nehalennia Williamson, 1915	MAO	3; 95; 64; 86; 103
Forcepsioneura itatiaiae (Santos, 1970)	AMZ	99; 98
Homeoura nepos (Selys, 1876)	MAO	100; 88; ­46
Homeoura obrieni von Ellenrieder, 2008	MAO	88
Hylaeonympha magoi Rácenis, 1968	AMZ	2
Inpabasis machadoi Santos, 1961	MPU; MAO; NAI	3; 2; 100; 76; 46; 103
Inpabasis rosea (Selys, 1877)	MPU; MAO	3; 95; 2; 100; 76; 46; 103
Ischnura capreolus (Hagen, 1861)	MPU; MAO; SGC	3; 95; 57; 97; 103
Ischnura fluviatilis Selys, 1876	MAO; PRF	3; 91; 103
Leucobasis candicans Rácenis, 1959	AMZ	2
Mecistogaster linearis (Fabricius, 1777)	BAR; MAO; NAI; PRF	3; 72; 103
Mecistogaster lucretia (Drury, 1773)	MAO; PRF	3; 100; 97; 103
Mesoleptobasis acuminata Santos, 1961	IRN; CAR	95; 100; 46
Mesoleptobasis cantralli Santos, 1961	BAR; SGC	3; 2; 100; 98; 32; 103
Mesoleptobasis elongata Garrison & von Ellenrieder, 2009	CAR	2; 32
Mesoleptobasis incus Sjöstedt, 1918	MPU; NAI; BOR; TEF	3; 2; 100; 98; 99; 32; 46; 103
Metaleptobasis amazonica Sjöstedt, 1918	MAO	1; 32; 46
Metaleptobasis brysonima Williamson, 1915	PRF	2; 90
Metaleptobasis paludicola von Ellenrieder, 2013	MAO	95; 2; 90
Metaleptobasis tridentigera von Ellenrieder, 2013	NAI; MPU	2; 90
Microstigma anomalum Rambur, 1842	MAO	95; 100; 97
Microstigma maculatum Hagen in Selys, 1860	MAO; PRF	3; 100; 61; 103
Microstigma rotundatum Selys, 1860	BAR; MPU; MAO; PRF; SGC; TEF; SPO; NAI	3; 95; 100; 80; 103
Neoneura bilinearis Selys, 1860	BAR	3; 103
Neoneura desana Machado, 1989	COA	3; 100; 98; 103
Neoneura luzmarina De Marmels, 1989	BAR; NRP; PRF; RPE; SGC; MAO	3; 100; 35; 103
Neoneura mariana Williamson, 1917	MAO	95
Neoneura rufithorax Selys, 1886	COA; ATA	3; 100; 35; 50; 103
Neoneura sylvatica Hagen in Selys, 1886	AMZ	51
Oxyagrion terminale Selys, 1876	AMZ	2; 100; 4
Phasmoneura exigua (Selys, 1886)	ITA; MPU; MAO; NAI; PRF; RPE	3; 95; 100; 98; 57; 103
Phoenicagrion flammeum (Selys, 1876)	MAO; NAI	3; 95; 2; 100; 98; 103
Platystigma buckleyi (McLachlan, 1881)	AMZ	100; 49
Protoneura scintilla Gloyd, 1939	BAR	3; 103
Protoneura tenuis Selys, 1860	MAO; PRF	3; 103
Psaironeura bifurcata (Sjöstedt, 1918)	MPU; MAO; PRF; NAI	3; 100; 36; 97; 103
Psaironeura tenuissima (Selys, 1886)	MPU; MAO; PRF; RPE	3; 95; 100; 99; 103
Telebasis abuna Bick & Bick, 1995	IRN	95
Telebasis dunklei Bick & Bick, 1995	IRN; CAR	95
Telebasis griffinii (Martin, 1896)	IRN	95; 2
Telebasis inalata (Calvert, 1961)	AMZ	2
Telebasis obsoleta (Selys, 1876)	IRN; CAR	95; 2; 38; 46
Telebasis sanguinalis Calvert, 1909	AMZ	100; 46
Telebasis simulata Tennessen, 2002	PRF; MAO	3; 95; 2; 100; 38; 46; 103
Tuberculobasis inversa (Selys, 1876)	TEF	2; 99; 52
Tukanobasis corbeti Machado, 2009	SGC	2; 53
ANISOPTERA
Aeshnidae
Anax amazili (Burmeister, 1839)	AMZ	2
Anax concolor Brauer, 1865	BAR; RPE	3; 1; 103
Castoraeschna tepuica De Marmels, 1989	BAR	3; 103
Coryphaeschna adnexa (Hagen, 1861)	BEC	1; 78
Coryphaeschna amazonica De Marmels, 1989	PRF	3; 2; 103
Coryphaeschna viriditas Calvert, 1952	MAO	95
Gynacantha auricularis Martin, 1909	BAR; MAO; PRF	3; 26; 103
Gynacantha dryadula Neiss & De Marmels, 2017	BAR; MAO	104
Gynacantha gracilis (Burmeister, 1839)	BAR; MAO	3; 1; 103
Gynacantha klagesi Williamson, 1923	MPU	3; 103
Gynacantha litoralis Williamson, 1923	MAO; SGC	3; 1; 103
Gynacantha membranalis Karsch, 1891	MAO; PRF	3; 103
Gynacantha mexicana Selys, 1868	MAO	3; 103
Gynacantha nervosa Rambur, 1842	MAO	95
Gynacantha tenuis Martin, 1909	PRF	3; 103
Neuraeschna calverti Kimmins, 1951	PRF; MAO	3; 1; 7; 103
Neuraeschna capillata Machet, 1990	MAO	55
Neuraeschna claviforcipata Martin, 1909	BAR; MAO	1; 27; 7
Neuraeschna costalis (Burmeister, 1839)	AMZ	1
Neuraeschna harpya Martin, 1909	AMZ	1; 98
Rhionaeschna planaltica (Calvert, 1952)	BAR	3; 103
Staurophlebia gigantula Martin, 1909	AMZ	1
Staurophlebia wayana Geijskes, 1959	PRF	3; 103
Triacanthagyna dentata (Geijskes, 1943)	AMZ	85
Triacanthagyna ditzleri Williamson, 1923	MAO; SGC	3; 103
Triacanthagyna satyrus (Martin, 1909)	MAO	3; 103
Triacanthagyna septima (Selys in Sagra, 1857)	MAO	95; 85
Gomphidae
Agriogomphus sylvicola Selys, 1869	PRF; TEF	3; 99; 82; 103
Aphylla barbata Belle, 1994	AMZ	1
Aphylla brasiliensis Belle, 1970	AMZ	2; 1
Aphylla dentata Selys, 1859	AMZ	2
Aphylla edentata Selys, 1869	AMZ	2; 99
Aphylla molossus Selys, 1869	AMZ	2
Cacoides latro (Erichson in Schomburgk, 1848)	PRF	3; 103
Desmogomphus tigrivensis Williamson, 1920	PRF; RPE; MAO	3; 82; 103
Diaphlebia angustipennis Selys, 1854	MAO; PRF	3; 95; 2; 1; 57; 97; 103
Epigomphus hylaeus Ris, 1918	MAO; PRF; NAI	3; 1; 82; 103
Gomphoides infumata (Rambur, 1842)	MAO	82
Peruviogomphus bellei Machado, 2005	AMZ	1; 99
Phyllocycla armata Belle, 1977	MAO; RPE; PRF	3; 103
Phyllocycla diphylla (Selys, 1854)	AMZ	1
Phyllocycla modesta Belle, 1970	MAO	95
Phyllocycla neotropica Belle, 1970	PRF	3; 103
Phyllocycla ophis (Selys, 1869)	AMZ	1
Phyllocycla pegasus (Selys, 1869)	AMZ	1
Phyllocycla volsella (Calvert, 1905)	MAO	95
Phyllogomphoides andromeda (Selys, 1869)	AMZ	98
Phyllogomphoides angularis Belle, 1982	AMZ	1
Phyllogomphoides atlanticus (Belle, 1970)	PRF	3; 103
Phyllogomphoides major Belle, 1984	SGC	3; 103
Phyllogomphoides pedunculus Belle, 1984	AMZ	2; 1
Phyllogomphoides pseudangularis Belle, 1994	COA	2; 1; 99; 8
Phyllogomphoides selysi (Navás, 1924)	AMZ	95; 2; 1; 6
Phyllogomphoides undulatus (Needham, 1944)	MAO; PRF; SGC	3; 103
Progomphus amazonicus Belle, 1973	MPU	2; 1; 4
Progomphus angeloi Belle, 1994	AMZ	2; 1; 99
Progomphus approximatus Belle, 1966	MAO; PRF	3; 103
Progomphus cf. boliviensis Belle, 1973	RPE	3; 103
Progomphus cf. phyllochromus Ris, 1918	PRF	3; 103
Progomphus cf. tibialis Belle, 1973	SGC	3; 103
Progomphus delicatus Belle, 1973	NAI; PRF	3; 103
Progomphus fassli Belle, 1973	AMZ	1
Progomphus guyanensis Belle, 1966	MAO; PRF	3; 103
Progomphus intricatus Hagen in Selys, 1858	AMZ	2
Progomphus maculatus Belle, 1984	MAO	3; 103
Progomphus perpusillus Ris, 1918	AMZ	2; 1; 4
Progomphus pijpersi Belle, 1966	AMZ	2
Zonophora batesi Selys, 1869	AMZ	2; 1; 99
Zonophora calippus Selys, 1869	TEF	2; 1; 99; 5
Zonophora nobilis Belle, 1983	SGC	2; 1; 5
Zonophora supratriangularis Schmidt, 1941	MAO; TEF	95; 2; 1; 99; 5
Zonophora wucherpfennigi Schmidt, 1941	MAO; PRF; RPE	3; 2; 1; 99; 103
Oxygastridae
Neocordulia batesi (Selys, 1871)	SPO	56; 66
Corduliidae
Aeschnosoma auripennis Geijskes, 1970	MPU; MAO; PRF; NAI; BAR	66; 3; 2; 62; 21; 103
Aeschnosoma forcipula Hagen in Selys, 1871	BAR; MPU; MAO; NAI; PRF	3; 95; 2; 1; 41; 103
Aeschnosoma hamadae Fleck & Neiss, 2012	MAO; PRF; NAI	3; 2; 29; 103
Paracordulia sericea (Selys, 1871)	BAR	2; 29; 3; 103
Libelluloidea incertae sedis
Lauromacromia cf. dubitalis (Fraser, 1939)	MAO	65
Libellulidae
Anatya guttata (Hoffmansegg in Schomburgk, 1848)	COA	70
Argyrothemis argentea Ris, 1911	MAO; MPU; NAI	3; 16
Brachymesia herbida (Gundlach, 1889)	MAO; PRF	95; 98; 91; 21; 15
Brechmorhoga nubecula (Rambur, 1842)	BAR; MAO; PRF	3; 103
Brechmorhoga praedatrix Calvert, 1909	PRF	3; 103
Dasythemis esmeralda Ris, 1910	PRF; RPE; MAO	3; 95; 57; 103
Dasythemis essequiba Ris, 1919	AMZ	1
Diastatops estherae Montgomery, 1940	BAR; MAO; MAU; SGC; TON	3; 1; 45; 58; 101; 103
Diastatops maxima Montgomery, 1940	PRF; TEF	1; 99; 58; 101
Diastatops nigra Montgomery, 1940	BAR; MAO; MAU; PAR; PRF; SGC	3; 1; 98; 58; 101; 103
Diastatops obscura (Fabricius, 1775)	MAO	102
Diastatops pullata (Burmeister, 1839)	BAR; PRF; MAU; MAO; MPU; SPO	3; 1; 98; 99; 58; 103
Elasmothemis williamsoni (Ris, 1919)	MAO; PRF	3; 103
Elga leptostyla Ris, 1911	NAI; MAO; NRP; PRF	3; 103
Erythemis attala (Selys in Sagra, 1857)	NAI; PRF; MAO; BEC; PAR; SPO	3; 95; 2; 99; 73; 15; 68; 97; 103
Erythemis carmelita Williamson, 1923	BEC; TEF	1; 94; 68
Erythemis haematogastra (Burmeister, 1839)	MAO; BAR; PRF; PAR; MNE; BEC	3; 95; 21; 91; 73; 15; 68; 103
Erythemis mithroides (Brauer, 1900)	PAR	73
Erythemis peruviana (Rambur, 1842)	IRN; NRP; MAO; PAR; ITA	3; 95; 98; 45; 73; 15; 68; 103
Erythemis plebeja (Burmeister, 1839)	MAO	73
Erythemis vesiculosa (Fabricius, 1775)	IRN; MAO; NAI; PRF; MNE	3; 21; 91; 73; 15; 68; 103
Erythrodiplax amazonica Sjöstedt, 1918	BAR; PRF; MAO; MAO	3; 95; 2; 1; 14; 103
Erythrodiplax anatoidea Borror, 1942	MAO; PRF	3; 103
Erythrodiplax attenuata (Kirby, 1889)	MAO; MNE	1; 98; 99; 14; 70
Erythrodiplax basalis (Kirby, 1897)	COA; MPU; MAO; RPE; NRP	3; 95; 1; 57; 14; 103
Erythrodiplax castanea (Burmeister, 1839)	MAO; MPU; PRF; NAI; RPE	3; 95; 1; 57; 103
Erythrodiplax famula (Erichson in Schomburgk, 1848)	BAR; MAO; PRF; RPE	3; 57; 14; 103
Erythrodiplax fusca (Rambur, 1842)	MPU; MAO; PRF; NRP; RPE; ITA; NAI	3; 95; 1; 99; 57; 103
Erythrodiplax lativittata Borror, 1942	MPU; MAO; PRF; RPE; SGC; NAI; NRP	3; 98; 99; 21; 103
Erythrodiplax laurentia Borror, 1942	PRF	3; 103
Erythrodiplax longitudinalis (Ris, 1919)	AMZ	1
Erythrodiplax solimaea Ris, 1911	TEF	1; 99; 14
Erythrodiplax umbrata (Linnaeus, 1758)	MPU; MAO; PRF; SPO; COA	3; 95; 1; 99; 21; 14; 103
Erythrodiplax unimaculata (de Geer, 1773)	MAO; ITA; MNE; TEF; SPO	95; 1; 99; 14
Erythrodiplax venusta (Kirby, 1897)	RPE; MAO	3; 95; 1; 14; 103
Fylgia amazonica Kirby, 1889	MAO; NAI; PRF	3; 95; 97; 103
Gynothemis pumila (Karsch, 1890)	BAR; MPU; MAO; NRP; PRF; RPE; NAI	3; 57; 31; 95; 103
Idiataphe amazonica (Kirby, 1889)	AMZ	1
Idiataphe cubensis (Scudder, 1866)	PRF	3; 103
Libellula herculea Karsch, 1889	BAR; PRF; MAO	3; 95; 103
Macrothemis brevidens Belle, 1983	MPU; PRF	3; 103
Macrothemis idalia Ris, 1919	BAR; PRF; MPU	3; 103
Macrothemis newtoni Costa, 1990	MAO	1; 18
Macrothemis nobilis Rácenis, 1957	AMZ	1
Macrothemisrupicola Rácenis, 1957	BAR	3; 103
Macrothemis taurepan De Marmels, 2008	BAR	3; 103
Miathyria marcella (Selys in Sagra, 1857)	MPU; IRN; MAO	3; 98; 21; 15; 102; 103
Miathyria simplex (Rambur, 1842)	PRF	3; 21; 103
Micrathyria artemis Ris, 1911	MAO	1; 19
Micrathyria atra (Martin, 1897)	PRF; MAO; ITA	3; 95; 1; 19; 68; 103
Micrathyria cambridgei Kirby, 1897	AMZ	1
Micrathyria catenata Calvert, 1909	NHA; MAO	1; 19; 24
Micrathyria hippolyte Ris, 1911	MAO	3; 103
Micrathyria laevigata Calvert, 1909	MNE	99; 75
Micrathyria pseudeximia Westfall, 1992	MAO	1; 19
Micrathyria romani Sjöstedt, 1918	MAO	1; 40
Micrathyria spinifera Calvert, 1909	MAO	3; 1; 103
Micrathyria venezuelae De Marmels, 1989	BAR	3; 103
Misagria calverti Geijskes, 1951	NAI	3; 103
Misagria divergens De Marmels, 1981	BAR	3; 103
Misagria parana Kirby, 1889	MAO	3; 95; 1; 103
Oligoclada abbreviata (Rambur, 1842)	BAR; MAO	3; 95; 1; 98; 13; 103
Oligoclada amphinome Ris, 1919	NRP; MAO	3; 95; 1; 13; 103
Oligoclada monosticha Borror, 1931	NRP; BAR	3; 103
Oligoclada pachystigma Karsch, 1890	BAR; COA	3; 42; 103
Oligoclada risi Geijskes, 1984	PRF; MAO	3; 103
Oligoclada sylvia (Kirby, 1889)	BAR; PRF	3; 1; 98; 99; 13; 67; 103
Oligoclada walkeri Geijskes, 1931	PRF; BAR	3; 98; 99; 13; 103
Oligoclada xanthopleura Borror, 1931	MAO	1; 13
Orthemis aequilibris Calvert, 1909	MPU; MAO	3; 89; 103
Orthemis attenuata (Erichson in Schomburgk, 1848)	RPE	3; 103
Orthemis biolleyi Calvert, 1906	MAO; NRP; PRF; RPE	3; 57; 102; 103
Orthemis cultriformis Calvert, 1899	SPO	1; 77
Orthemis discolor (Burmeister, 1839)	MPU; MAO; RPE; NAI	3; 103
Orthemis ferruginea (Fabricius, 1775)	MAO	15; 57
Pantala flavescens (Fabricius, 1798)	MPU; MAO; PRF	3; 2; 21; 57; 103
Perithemis bella Kirby, 1889	MAO; SPO	3; 1; 102; 103
Perithemis cornelia Ris, 1910	TEF	1; 99; 71
Perithemis electra Ris, 1930	BEC	79
Perithemis lais (Perty, 1834)	BAR; MAO; SGC	3; 2; 1; 98; 99; 91; 71; 103
Perithemis mooma Kirby, 1889	MAO	3; 103
Perithemis thais Kirby, 1889	BAR; PRF; BEC	3; 1; 81; 103
Rhodopygia cardinalis (Erichson in Schomburgk, 1848)	MAO	1; 68;
Rhodopygia geijskesi Belle, 1964	MAO; SAI	9; 68
Rhodopygia hollandi Calvert, 1907	AMZ	1
Tauriphila australis (Hagen, 1867)	MAO	3; 98; 103
Tholymis citrina Hagen, 1867	BAR; ITA; MAO; NRP; PRF; MNE	3; 95; 103
Tramea binotata (Rambur, 1842)	BAR	3; 103
Tramea calverti Muttkowski, 1910	RPE; ITA; PRF; MAO; TEF	3; 95; 2; 1; 99; 21; 22; 103
Tramea minuta De Marmels & Rácenis, 1982	BAR; SPO	3; 2; 1; 22; 103
Tramea rustica De Marmels & Rácenis, 1982	MAO	2; 22
Uracis fastigiata (Burmeister, 1839)	PRF; SPO; MAO; MNE; JUR; GJR; EIR; BEC; TAB	3; 1; 98; 99; 70; 17; 103
Uracis imbuta (Burmeister, 1839)	MAO; NRP; CAR; ATA; MNE; SPO; BEC; TAB; SGC	3; 95; 1; 98; 99; 57; 70; 17; 103
Uracis infumata (Rambur, 1842)	PRF; MAO; SPO; EIR	3; 1; 17; 103
Uracis ovipositrix Calvert, 1909	NAI; MAO; SPO; MNE; SGC	3; 95; 1; 98; 99; 70; 17; 103
Uracis siemensi Kirby, 1897	PRF; BAR; MAO; NRP; PAR; MNE; EIR; SPO	3; 95; 1; 99; 17; 103
Zenithoptera anceps Pujol-Luz, 1993	MAO; ITA	2; 69
Zenithoptera fasciata (Linnaeus, 1758)	MPU; PRF; MAO; BOR; ITA; PAR	3; 2; 57; 95; 69; 102; 103
Zenithoptera lanei Santos, 1941	NAI; PRF; ITA; MAO; TAB	3; 2; 98; 69; 103
Zenithoptera viola Ris, 1910	ITA	95; 2

Figure 2
Some Odonata species recorded in Amazonas State, Brazil. A, Chalcopteryx rutilans (Rambur 1842); B, Chalcopteryx scintillans McLachlan, 1870; C, Chalcopteryx seabrai Santos & Machado, 1961; D, Polythore vittata (Selys, 1869); E, Rimanella arcana (Needham, 1933); F, Mecistogaster linearis (Fabricius, 1777); G, Dimeragrion percubitale Calvert, 1913; H, Aeschnosoma hamadae Fleck & Neiss, 2012; I, Elga leptostyla Ris, 1911; J, Erythrodiplax anatoidea Borror, 1942; K, Zenithoptera lanei Santos, 1941.

Considering the territorial divisions of the state, Manaus (code number 1) municipality has the most species registered, 163 species, followed by Presidente Figueiredo (code number 2), 100 species, and Barcelos (code number 4), 71 species. In addition, only 33 of the 62 municipalities in Amazonas have published Odonata records.

Discussion

The 324 species listed from Amazonas state represent about 43% of the known species in Brazil (749 species). This number increases the species records for the state by 23% (Neiss & Hamada 2014). With the results presented in this study, Amazonas state is the Brazilian state with the highest number of Odonata species recorded, followed by Rio de Janeiro, 280 species (Costa & Santos 2000), Minas Gerais, 269 species (Vilela et al. 2020), São Paulo, 251 species (Costa et al. 2000), Mato Grosso do Sul, 209 species (Koroiva et al. 2017; Rodrigues et al. 2018), Rio Grande do Sul, 183 species (Pires et al. 2019), Espírito Santo, 180 species (Costa & Oldrini 2005), and Goiás and the Federal District, 152 species (Nóbrega & De Marco 2011).

We highlight that five species were not included in the results due to imprecise information. Despite being cited for Amazonas state in Heckman (2008), in the original description of Planiplax machadoiSantos, 1949, Santos (1949) states that the species is present in “Amazonas”, but in the text the author explains that specimens came from Pará state. Similar problem occurs with Erythrodiplax julianaRis, 1911 and Erythrodiplax latimaculata Ris, 1911. Borror (1942) states that these species are present in Amazonas state; however, the specimens were collected in Monte Roraima and Uraricoera, respectively, which currently belong to Roraima state. Finally, Diastatops intensaMontgomery, 1940 and Diastatops dimidiata Montgomery, 1940 were registered for the floodplains of the Solimões-Amazonas rivers by Nessimian et al. (2008). Although most of their sampled sites were in the Amazonas state, there are some sites in Pará state as well, and authors do not present a detailed list of where each species was found.

Despite the significant increase in the number of species known from Amazonas, some regions remain poorly explored, such as municipalities in the southwest region of the state. Besides Guajará (code number 10) and Eirunepé (code number 12), there are no published species records from municipalities located on the border with Acre state. In addition, large municipalities also appear to be under-sampled. For example, São Gabriel da Cachoeira, the third largest municipality in Brazil (109,181 km²; IBGE 2019) with an area larger than Portugal, only has 20 recorded species. In this way, we suggest that new field expeditions should focus on such large municipalities and the 29 municipalities that presented no specimen records in our survey.

In general, knowledge about the Odonata order in Amazonas state has improved, especially in the last two decades. However, limitations on sampling still underestimate the number of odonates species, both for new species and new records. Therefore, this work recognizes the importance of the biological diversity in Amazonas state and the Amazonian Biome for species richness of Brazil and shows that we still do not entirely know the odonatofauna in the world’s largest tropical forest.

Acknowledgements

We are thankful to FAPEAM (Fundação de Amparo à Pesquisa do Estado do Amazonas) for a post-doctoral fellowship (FIXAM-I/FAPEAM; Proc.062.01503/2018) to RK. NH is a CNPq research fellow (Proc. 307849/ 2014-7). INPA/ MCTI and PRONEX-CNPq-FAPEAM provided partial financial support for this research.

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Publication Dates

• Publication in this collection
  31 Jan 2020
• Date of issue
  2020

History

• Received
  28 Aug 2019
• Reviewed
  05 Dec 2019
• Accepted
  09 Dec 2019