Analysis of Brazilian Ceratopogonidae (Diptera: Culicomorpha) species diversity and knowledge assessment

ABSTRACT

An updated list of the biting midges recorded from the 26 Brazilian states and the Federal District is provided based on the data available in the “Taxonomic Catalog of the Brazilian Fauna” (Portuguese CTFB). The Brazilian Ceratopogonidae fauna is represented by 529 known species, corresponding to 40% of the Neotropical fauna. A table showing the number of species of Ceratopogonidae genera in the Neotropical region, Brazil, and Brazilian Amazon is included. In accordance with the family knowledge worldwide, the subfamily Ceratopogoninae is the best represented in the number of species due to the study of Culicoides, which has sanitary importance. The analysis of the diversity in each region and in the Brazilian states indicates the need for further studies in the five Brazilian geopolitical regions, especially in the Midwest and Northeast, which are home to the natural biomes of Cerrado, Pantanal, and Caatinga.

Key words:
Biting midges; Brazil; distribution; taxonomy

INTRODUCTION

Ceratopogonidae Newman, 1834 includes small dipteran species that compose a diverse, abundant, and widely distributed group. Borkent and Dominiak (2020) recognize 6,206 valid species divided into three subfamilies and 112 genera. They are distributed throughout the world, except in Antarctica, occurring from coastal areas to the highest mountain peaks, with the uppermost record at 4,651 m from Lake Huacracocha in Peru (Tapia et al. 2018). The immature stages are abundant and present in almost all aquatic and subaquatic habitats, including phytotelmata and rock pools, seeps, streams, rivers, ponds, swamps and lakes, moist substrates such as mud, decaying plant material and dung. Some species are terrestrial, occurring under the bark of trunks, in damp mosses and fungi. Adult females have the broadest dietary repertoire of any group of biting insects, with many requiring extra proteins to develop their eggs (Borkent and Dominiak 2020). Vertebrate hosts include mammals, reptiles, birds, amphibians, and even fish (Wirth and Hubert 1989, Spinelli et al 2002). Most of the genera include predators of other small insects, usually other dipterans, which are captured and injected with a proteolytic enzyme; after liquefaction, the prey contents are sucked out. Another lineage of females feeds on hemolymph from other arthropods much larger than themselves, such as centipedes, spiders, phasmids, dragonflies, coleopterans, and other large insects (Downes 1978, Borkent and Spinelli 2007, Borkent et al. 2009, Borkent 2017). Some Ceratopogonidae are exclusively floral visitors, constituting important pollinators of tropical plants such as cocoa (Theobroma cacao L.), rubber tree (Hevea brasiliensis Müll. Arg.) and avocado (Persea americana Mill.), having great importance for agriculture (Wirth and Cavalieri 1977). In this case, males and females look for nectar as a source of energy.

Many species of Culicoides Latreille, 1809, a hyperdiverse genus with 1,347 species, act as vectors for a wide range of viruses, protozoa, and nematodes that affect humans and domestic and wild animals (Borkent 2004). Some species of Culicoides, Leptoconops Skuse, 1889, Austroconops Wirth & Lee, 1958 and Forcipomyia (Lasiohelea) Kieffer, 1921 occur in such large numbers that their hematophagic habit affects tourism (Hendry 2011). On the other hand, their presence protects several vulnerable habitats by restricting the distribution of humans in some locations, such as in Scotland, some beaches in the Caribbean, areas of Australia, and some mangroves that would otherwise be vulnerable to disturbance caused by humans (Borkent and Dominiak 2020). Furthermore, veterinary diseases such as Bluetongue, and human diseases such as Mansonellosis and Oropouche Fever, are transmitted by different species of Culicoides (Mellor et al. 2000).

Despite Brazil’s large size, diversity of biomes, and the agricultural and sanitary importance of some species, the Brazilian fauna of biting midges is still little studied; in some parts of the country, the diversity of these dipterans is completely unknown. Data from records of Ceratopogonidae species have been provided in different publications, making research into the knowledge of the specific distribution of the Brazilian fauna very laborious. In this context, catalogs and lists of species are important tools that contribute to the study of biodiversity in localities, states, countries, and zoogeographic regions. Along the years, several catalogs, as well as lists of Ceratopogonidae tribes and genera, have been created to update the species lists for family study facilitation. Ceratopogonidae catalogs and lists from different parts of the world are accessible at http://campus.belmont.edu/cienews/cie.html.

From the Neotropical region, the most recent Ceratopogonidae species catalogs have been elaborated by Borkent and Spinelli (2000, 2007). The Nicaraguan, Mexican, and Argentinian species were published sequentially by Maes and Wirth (1990), Ibáñez-Bernal et al. (1996), and Spinelli et al. (2023). The Ceratopogonidae fauna was also mentioned by Wirth (1952), Davies and Giglioli (1979), and Borkent (1991) for Juan Fernandez, Grand Cayman, and Galápagos islands, respectively. Culicoides species lists from Argentina, Colombia and Costa Rica are available in the publications by Spinelli et al. (2005, 2009a) and Spinelli and Borkent (2004). The Culicoides species from Panama, West Indies, Mexico, and Ecuador were published by Wirth and Blanton (1959), Wirth (1974), Mendez-Andrade and Ibáñez-Bernal (2023), and Mosquera et al. (2022). From Brazil, Wirth and Blanton (1973) and Castellón and Felippe-Bauer (2015) have published Culicoides species and their distribution in the Brazilian Amazon area. Since 2015, the Brazilian fauna has been updated on the website “Taxonomic Catalog of the Brazilian Fauna” (Catálago Taxonômico da Fauna Brasileira - CTFB, http://fauna.jbrj.gov.br/) and in the Fiocruz Collection website (Santarém and Felippe-Bauer 2023).

Considering the need for a reference publication of data on the distribution of midges in Brazil, this work aims to list the species of Ceratopogonidae recorded from the 26 Brazilian states and the Federal District based on the data available in the CTFB, as well as to carry out an analysis of the knowledge of the diversity of this important family in Brazil.

MATERIAL AND METHODS

In this work, we follow the species arrangement according to the “Catalog of the Biting Midges of the World” (Borkent and Dominiak 2020). The distribution of species was based on the specific literature, in addition to reliable localities of specimens deposited in the entomological collections of the following institutions: Faculdade de Saúde Pública, Universidade de São Paulo, São Paulo, Brazil (FSP); Coleção de Ceratopogonidae da Fundação Oswaldo Cruz, Rio de Janeiro, Brazil (CCER); National Museum of Natural History, Washington, D.C., USA (USNM); The Natural History Museum, London, United Kingdom (BMNH). In the case of doubtful species records, we chose to maintain them in the distribution and in the result analysis until further studies, such as revisions of genera, subgenera or species groups, are carried out to confirm or not their validity.

In the species list, the records are given in alphabetical order of Brazilian states after the indication of the type locality, considering the geopolitical regions. The states that comprise each region are listed: North - Acre (AC), Amapá (AP), Amazonas (AM), Pará (PA), Rondônia (RO), Roraima (RR), Tocantins (TO); Northeast - Alagoas (AL), Bahia (BA), Ceará (CE), Maranhão (MA), Paraíba (PB), Pernambuco (PE), Piauí (PI), Rio Grande do Norte (RN); Midwest - Distrito Federal (DF), Goiás (GO), Mato Grosso (MT), Mato Grosso do Sul (MS); Southeast - Espírito Santo (ES), Minas Gerais (MG), Rio de Janeiro (RJ), São Paulo (SP); and South - Paraná (PR), Rio Grande do Sul (RS), Santa Catarina (SC) (Fig. 1). In 1977, the state of Mato Grosso was split into the states of Mato Grosso and Mato Grosso do Sul; in 1989, the state of Goiás was divided into the states of Goiás and Tocantins. Localities mentioned in manuscripts before these dates had to be updated due to the newly created states. When the collection site of the species is not provided in the original description, we considered the cited geographic region of the species.

Figure 1
Political map of Brazil showing the number of Ceratopogonidae records from the 26 Brazilian States and the Federal District.

We have also counted the species recorded from the Neotropical region and Brazilian Amazon region, aiming to compare and update this data. To consider a species as recorded from the Neotropical region, we employed the biogeographical regionalization of this region proposed by Morrone (2014). For inclusion of a species in the Brazilian Amazon region, we considered the political map of the “Amazônia Legal Brasileira.” Brazilian municipalities and states included here can be accessed at http://www.cidades.ibge.gov.br/xtras/home.php.

The distribution data of Brazilian species of biting midges were uploaded to the CTFB website and to the Fiocruz Collections website (Santarém and Felippe-Bauer 2023). They were also listed herein in the ^{Supplementary file S1}.

RESULTS AND DISCUSSION

Herein, we recognized 1,329 extant valid Ceratopogonidae species from the Neotropical region, 529 from Brazil, and 303 from the Brazilian Amazon region (Table 1, ^{Supplementary file S1}).

Table 1
Number of species of Ceratopogonidae genera recorded from the Neotropical region, Brazil, and the Brazilian Amazon region.

Subfamily	Tribe	Genus	Subgenus	Neotropical region	Brazil	Brazilian Amazon region
Ceratopogoninae	Ceratopogonini	Allohelea		1	-	-
		Alluaudomyia		18	7	5
		Austrohelea		3	-	-
		Baeodasymyia		5	-	-
		Baeohelea		1	-	-
		Bahiahelea		1	1	-
		Borkenthelea		4	-	-
		Brachypogon	Brachypogon	23	8	6
		Brachypogon	Isohelea	7	1	-
		Bhachypogon	Out of subgenus	2	-	-
		Cacaohelea		1	-	-
		Ceratoculicoides		1	-	-
		Diaphanobezzia		4	-	-
		Downeshelea		46	19	9
		Echinohelea	Echinohelea	10	5	4
		Echinohelea	Echinoideshelea	1	1	1
		Fittkauhelea		1	1	1
		Isthmohelea		1	-	-
		Leptohelea		1	-	-
		Macrurohelea		17	-	-
		Monohelea		27	18	11
		Nannohelea		1	-	-
		Notiohelea		2	-	-
		Parabezzia		30	8	2
		Rhynchohelea		1	-	-
		Schizonyxhelea		9	4	2
		Spinellihelea		1	-	-
		Stilobezzia	Acanthohelea	41	4	4
		Stilobezzia	Eukraiohelea	5	5	3
		Stilobezzia	Stilobezzia	47	25	13
		Yungahelea		1	-	-
	Culicoidini	Culicoides	Amossovia	1	-	-
		Culicoides	Anilomyia	20	2	1
		Culicoides	Avaritia	8	3	2
		Culicoides	Beltranmyia	3	-	-
		Culicoides	Cotocripus	6	2	-
		Culicoides	Culicoides	6	-	-
		Culicoides	Diphaomyia	14	3	2
		Culicoides	Drymodesmyia	13	3	1
		Culicoides	Glaphiromyia	2	-	-
		Culicoides	Haematomyidium	38	25	19
		Culicoides	Hoffmania	46	37	35
		Culicoides	Macfiella	2	1	1
		Culicoides	Mataemyia	19	13	10
		Culicoides	Oecacta	5	2	2
		Culicoides	Psychophaena	2	1	-
		Culicoides	Out of subgenus	116	59	50
		Paradasyhelea		3	-	-
	Heteromyiini	Clinohelea	Ceratobezzia	1	1	1
		Clinohelea	Clinohelea	11	9	6
		Heteromyia		11	6	4
		Pellucidomyia		4	1	-
		Physohelea		2	-	-
	Johannsenomyiini	Groganhelea		1	1	1
		Johannsenomyia		2	1	-
		Lanehelea		2	-	-
		Mallochohelea		6	4	2
		Neobezzia		8	6	5
		Nilobezzia		3	3	2
	Palpomyiini	Amerohelea		13	4	3
		Bezzia		50	19	11
		Clastrieromyia		4	1	1
		Pachyhelea		2	1	1
		Palpomyia		55	32	6
		Phaenobezzia		2	2	1
	Sphaeromyiini	Austrosphaeromias		4	-	-
		Sphaerohelea		1	-	-
	Stenoxenini	Paryphoconus		42	30	18
		Stenoxenus		16	7	4
Forcipomyiinae	Dasyheleini	Dasyhelea		104	18	7
	Forcipomyiini	Atrichopogon	Atrichopogon	98	28	5
		Atrichopogon	Lophomyidium	5	2	2
		Atrichopogon	Psilokempia	17	6	3
		Forcipomyia	Blantonia	1	-	-
		Forcipomyia	Caloforcipomyia	11	9	4
		Forcipomyia	Euprojoannisia	16	5	3
		Forcipomyia	Forcipomyia	37	11	4
		Forcipomyia	Lasiohelea	11	5	1
		Forcipomyia	Lepidohelea	23	10	1
		Forcipomyia	Metaforcipomyia	23	1	-
		Forcipomyia	Microhelea	50	22	11
		Forcipomyia	Pedilohelea	5	3	2
		Forcipomyia	Phytohelea	11	5	1
		Forcipomyia	Pterobosca	3	2	-
		Forcipomyia	Rhynchoforcipomyia	7	2	1
		Forcipomyia	Saliohelea	2	1	1
		Forcipomyia	Schizoforcipomyia	2	1	1
		Forcipomyia	Synthyridomyia	4	1	-
		Forcipomyia	Thyridomyia	8	2	-
		Forcipomyia	Trichohelea	13	2	1
		Forcipomyia	Warkea	7	4	4
Leptoconopinae		Leptoconops	Brachyconops	1	-	-
		Leptoconops	Holoconops	3	1	-
		Leptoconops	Leptoconops	7	1	1
		Leptoconops	Megaconops	1	1	-
		Leptoconops	Proleptoconops	1	-	-
Unplaced	Unplaced	Ceratopogon		1	-	-
Total				1,329	529	303

It is possible to observe that almost 40% of the Neotropical Ceratopogonidae fauna has already been recorded from Brazil, highlighting the country’s importance in the distribution of these insects. It is also noteworthy that approximately 57% of the Brazilian fauna was recorded from the Amazon region, indicating that this biome has favorable environments for the development of biting midges and reinforcing its importance in the study of the country’s fauna. The characteristics of the local climate and the number of studies carried out in the Amazon, mainly due to the transmission of the Oropouche virus, are crucial factors in understanding the representativeness of the region in the knowledge of the Brazilian Ceratopogonidae fauna.

Ceratopogoninae Newman, 1834 is the best represented subfamily with 386 species recorded from Brazil (73%), followed by Forcipomyiinae Lenz, 1934 with 140 species (26.5%). The Leptoconopinae Noè, 1907is represented by only three species: Leptoconops (Leptoconops) brasiliensis (Lutz, 1913), Leptoconops (Holoconops) knowltoni Clastrier & Wirth, 1978, and the recently recorded Leptoconops (Meganocops) floridensis Wirth, 1951 (Santarém et al. 2023). These species have been recorded from the coastal areas of northern and southern Brazil. This data corroborates the knowledge of the family worldwide. According to Borkent and Dominiak (2020), Ceratopogoninae is the best-known subfamily in the world, with the number of species driven by the study of the genus Culicoides, which has sanitary importance. The authors recognize that there are genera that are clearly much more diverse than others and that the greatest species diversity is restricted to only four genera: Culicoides (n = 1,347), Forcipomyia Meigen, 1818 (n = 1,142), Dasyhelea Kieffer, 1911 (n = 617), and Atrichopogon Kieffer, 1906 (n = 513). Together, these genera represent 58% of the known extant diversity of the family in the world. This pattern is quite similar to the diversity found in Brazil, except for the genus Dasyhelea, which has only 18 species recorded from the country, behind several predator Ceratopogoninae midges from the tribes Ceratopogonini Newman, 1834, Palpomyiini Enderlein, 1936 and Stenoxenini Coquillett, 1899. This is certainly due to the lack of extensive studies involving this important pollinator genus. On the other hand, there are several studies on the diversity of predatory midges that are widely distributed in the Neotropical region and Brazil, boosting the number of known species for such genera as Bezzia Kieffer, 1899 (Dippolito et al. 1995), Downeshelea Wirth & Grogan, 1988 (Santarém et al. 2020), Palpomyia Meigen, 1818 (Spinelli et al. 2009b, Almeida et al. 2017), Paryphoconus Enderlein, 1912 (Spinelli and Wirth 1984) and Stilobezzia Kieffer, 1911 (Cazorla and Spinelli 2014, Cazorla 2016, Cazorla et al. 2017, Da Silva et al. 2023). These data corroborate the number of species known in Brazil from the six predatory tribes included in Ceratopogoninae. The tribes Culicoidini and Ceratopogonini, which include Culicoides and the main predatory genera, respectively, are those with the largest number of known and described species (151 and 107 species, respectively).

The Brazilian states with the highest number of records of Ceratopogonidae are Pará (176 species), Amazonas (165 species), and Rio de Janeiro (141 species) (Fig. 1). When records are considered by geographic region, those with the highest number are the North (296 species) and Southeast (199 species) regions. The representativeness of these regions can be explained by the existence of research groups in these areas that promoted the study of local faunas and, consequently, increased the number of records. In this sense, we highlight the studies carried out in these areas by Wirth and Blanton (1973), who described 15 new species of Culicoides from the Brazilian Amazon region, and the work of Castellón included in the book by Castellón and Veras (2015), in addition to the several species described and recorded by Adolpho Lutz, Osmar Tavares, John Lane, Oswaldo Forattini and Maria Luiza Felippe-Bauer for the North and Southeast regions.

The significant number of records from the state of Bahia (61 species) is the result of Saulo Soria’s research on cocoa pollinators from the “Recôncavo Baiano.” All specimens that he studied are deposited in the Ceratopogonidae Collection of Fiocruz (Soria et al. 2002). The high number of records from Santa Catarina (110 species) was driven by John William Scott Macfie, who published several important works describing species collected by Fritz Plaumann (Macfie 1939). Furthermore, there are several records of Forcipomyia by Jean Clastrier (Clastrier and Wirth 1995).

A closer look at the number of listed species indicates the need for future studies in the five Brazilian geopolitical regions, especially in the Midwest and Northeast, which are home to the natural biomes of Cerrado, Pantanal, and Caatinga. Despite the high number of records from the North region, some states are still poorly represented and certainly have a greater number of species, since the Amazon Forest biome, typical of this region, has characteristics that are favorable to the development of biting midges during all seasons of the year. Despite being politically included in the North region, the state of Tocantins has a Cerrado biome and has no records of Ceratopogonidae to date. In the Northeast, no species of Ceratopogonidae have been reported for the state of Sergipe, and just Culicoides insignis Lutz, 1913 was recorded from Alagoas (Rios et al. 2021). All other states have few records, despite this being a region that covers nine states in an area of 1,558,196 km² and includes the Atlantic Forest and Caatinga biomes. The Midwest region also has very few records for Ceratopogonidae and clearly requires broader studies. This region hosts two important Brazilian biomes, the Cerrado and the Pantanal, which have unique characteristics and currently are under great human pressure. The Southeast region is characterized by the presence of the Atlantic Forest and a strong influence of the coastal climate, which makes this region very conducive to the development of “maruins”, as is the Amazon. Therefore, the various states that politically make up the Southeast of Brazil are undersampled, even Rio de Janeiro. Finally, the fauna of the South region is basically represented by species from Santa Catarina, with few records from Paraná and Rio Grande do Sul, which indicates the need for broad studies involving these states that present important transition regions and the Atlantic Forest and Pampas biomes.

Therefore, further studies on the Ceratopogonidae fauna from Brazil are urgently needed, especially in states that are characterized by biomes such as the Caatinga, Cerrado, Pantanal, and Pampas. These biomes have very peculiar features, with alternating periods of drought and rain that certainly influence the composition of the fauna and the population dynamics of the “maruins” from these locations, which are mostly unknown to date.

We have been updating the species reported from Brazil and their records from Brazilian states in the CTFB and Fiocruz collections website. We hope that the cited data and references available online will aid the systematic, biodiversity, biogeographical, and epidemiological studies on the Ceratopogonidae fauna.

ACKNOWLEDGEMENTS

We are grateful to Tiago do Nascimento da Silva for figure edition. We further thank Fundação de Apoio à Fiocruz (Fiotec) for the research grant (VPPCB-004-FIO-21) to the first author.

LITERATURE CITED

• Almeida JF, Farias ES, Alencar RB, Pessoa FAC (2017) Description of two new species of Palpomyia Meigen (Diptera: Ceratopogonidae) from the Brazilian Amazon. EntomoBrasilis 10(2): 118-122. https://doi.org/10.12741/ebrasilis.v10i2.607
  » https://doi.org/10.12741/ebrasilis.v10i2.607
• Borkent A (1991) The Ceratopogonidae (Diptera) of the Galápagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Insect Systematics and Evolution 22(1): 97-122.
• Borkent A (2004) The Biting Midges, the Ceratopogonidae (Diptera). In: Marquardt WC (Ed.) Biology of Disease Vectors. Elsevier, Academic Press, 2^nd ed., 113-126.
• Borkent A (2017) Ceratopogonidae (Biting Midges). In: Kirk-Spriggs AH, Sinclair BJ (Eds) Manual of Afrotropical Diptera. South African National Biodiversity Institute, Pretoria, vol. 2, 733-812.
• Borkent A, Dominiak P (2020) Catalog of the Biting Midges of the World (Diptera: Ceratopogonidae). Zootaxa 4787(1): 1-377. https://doi.org/10.11646/zootaxa.4787.1.1
  » https://doi.org/10.11646/zootaxa.4787.1.1
• Borkent A, Spinelli GR (2000) Catalog of New World Biting Midges South of the United States (Diptera: Ceratopogonidae). Contributions on Entomology, International 4: 1-107.
• Borkent A, Spinelli GR (2007) Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis J, Arias JR, Rueda-Delgado G, Wnatzen KM (Eds) Aquatic Biodiversity in Latin America (ABLA). Pensoft, Sofia-Moscow, vol. 4, 1-198.
• Borkent A, Spinelli GR, Grogan WL (2009) Ceratopogonidae. In: Brown BV, Borkent A, Cumming JM, Wood DM, Woodley NE, Zumbado MA (Eds) Manual of Central American Diptera. NRC Research Press, Ottawa, vol. 1, 407-435.
• Castellón EG, Felippe-Bauer ML (2015) Classificação e distribuição das espécies de Culicoides (Diptera: Ceratopogonidae) na Amazônia Brasileira ou Amazônia Legal. In: Castellón EG, Veras RS (Eds) Maruins (Culicoides: Ceratopogonidae) na Amazônia Brasileira. Editora INPA, Manaus, 27-71.
• Castellón EG, Veras RS (2015) Maruins (Culicoides: Ceratopogonidae) na Amazônia Brasileira. Editora INPA, Manaus, 144 pp.
• Cazorla CG (2016) New records of “jejenes” on the subgenus Stilobezzia of Stilobezzia (Diptera: Ceratopogonidae) from the Neotropical Region. Revista de la Sociedad Entomológica Argentina 75(3-4): 186-190.
• Cazorla CG, Cardoso EA, Felippe-Bauer ML (2017) Contributions to the knowledge of predaceous midges of the subgenus Eukraiohelea Ingram & Macfie of Stilobezzia Kieffer (Diptera: Ceratopogonidae), from Brazil. Zootaxa 4324(3): 557-570. https://doi.org/10.11646/zootaxa.4324.3.9
  » https://doi.org/10.11646/zootaxa.4324.3.9
• Cazorla CG, Spinelli GR (2014) A revision of the Patagonian predaceous midges of the subgenus Acanthohelea of Stilobezzia excluding the S. (A.) edwardsi group (Diptera: Ceratopogonidae). Journal of Natural History 49(3-4): 155-209. https://doi.org/10.1080/00222933.2014.939728
  » https://doi.org/10.1080/00222933.2014.939728
• Clastrier J, Wirth WW (1995) Révision des Forcipomyia du sous-genre Microhelea de la région Néotropicale, parasites de phasmes (Diptera: Ceratopogonidae). Annales de la Société Entomologique de France 31(2): 97-150. https://doi.org/10.1080/21686351.1995.12277880
  » https://doi.org/10.1080/21686351.1995.12277880
• Da Silva IM, Ferreira-Kepler RL, Cazorla CG (2023) Stilobezzia Kieffer (Diptera: Ceratopogonidae) from the Brazilian Amazon: three new species, redescription of S. (Stilobezzia) maculata Lane and new records. Zootaxa 5249(4): 485-499. https://doi.org/10.11646/zootaxa.5249.4.6
  » https://doi.org/10.11646/zootaxa.5249.4.6
• Davies JE, Giglioli MEC (1979) The Ceratopogonidae (Diptera) of Grand Cayman, West Indies: species and ecological notes. Mosquito News 39(3): 586-594.
• Dippolito A, Spinelli GR, Wirth WW (1995) A report on a collection of Ceratopogonidae (Diptera) from Rondonia (Brazil) and Iquitos (Peru) 1. Tribes Palpomyiini and Stenoxenini. Insecta Mundi 9(1-2): 53-60.
• Downes JA (1978) Feeding and mating in the insectivorous Ceratopogoninae (Diptera). Memoirs of the Entomological Society of Canada 104: 1-62. https://doi.org/10.4039/entm110104fv
  » https://doi.org/10.4039/entm110104fv
• Hendry G (2011) Midges in Scotland. Bell and Bain Ltd., Glasgow, 5^th ed., 85 pp.
• Ibáñez-Bernal S, Wirth WW, Huerta H (1996) Ceratopogonidae. In: Bousquets JL, Garcia Aldrete NA, Gonzalez Soriano E (Eds) Biodiversidad, taxonomia y biogeografia de artropodos de México: hacia una sintesis de su conocimiento. Universidad Nacional Autonoma de México, Mexico City, 67-577.
• Macfie JWS (1939) A report on a collection of Brazilian Ceratopogonidae. Revista de Entomologia 10: 137-219.
• Maes JM, Wirth WW (1990) Catalogo de los Diptera de Nicarágua. 6. Ceratopogonidae (Nematocera). Revista Nicaraguense de Entolomogia 14: 1-17.
• Mellor PS, Boorman J, Baylis M (2000) Culicoides biting midges: their role as arbovirus vectors. Annual Review of Entomology 45: 307-340. https://doi.org/10.1146/annurev.ento.45.1.307
  » https://doi.org/10.1146/annurev.ento.45.1.307
• Mendez-Andrade A, Ibáñez-Bernal S (2023) An updated catalogue of biting midges of the genus Culicoides Latreille, 1809 (Diptera, Ceratopogonidae) of Mexico and their known distribution by state. Zookeys 1167: 1-47. https://doi.org/10.3897/zookeys.1167.102858
  » https://doi.org/10.3897/zookeys.1167.102858
• Morrone JJ (2014) Biogeographical regionalisation of the Neotropical region. Zootaxa 3782: 1-110. https://doi.org/10.11646/zootaxa.3782.1.1
  » https://doi.org/10.11646/zootaxa.3782.1.1
• Mosquera JD, Zapata S, Spinelli GR, Gualapuro M, León R, Augot D (2022) An updated list of the Culicoides (Diptera, Ceratopogonidae) fauna from Ecuador. Parasite 29(63): 1-13. https://doi.org/10.1051/parasite/2022061
  » https://doi.org/10.1051/parasite/2022061
• Rios RRS, Santarém MCA, Ribeiro Júnior KAL, de Melo BA, da Silva SGM, da Silva NC, et al. (2021) Culicoides insignis Lutz, 1913 (Diptera: Ceratopogonidae) biting midges in Northeast of Brazil. Insects 12(4): 1-6. https://doi.org/10.3390/insects12040366
  » https://doi.org/10.3390/insects12040366
• Santarém MCA, Borkent A, Felippe-Bauer ML (2020) Taxonomic Revision of Neotropical Downeshelea Wirth and Grogan predaceous midges (Diptera: Ceratopogonidae). Insects 11(9): 1-94. https://doi.org/10.3390/insects11010009
  » https://doi.org/10.3390/insects11010009
• Santarém MCA, Cremer MJ, Vieira JV, Lemos GG, Pecor DB, Felippe-Bauer ML (2023) Redescription of Leptoconops (Leptoconops) brasiliensis (Lutz) and Leptoconops (Megaconops) floridensis Wirth (Diptera: Ceratopogonidae) with its first Brazilian record. Zootaxa 5380(2): 173-183. https://doi.org/10.11646/zootaxa.5380.2.5
  » https://doi.org/10.11646/zootaxa.5380.2.5
• Santarém MCA, Felippe-Bauer ML (2023) Brazilian species of Biting Midges. FIOCRUZ, Rio de Janeiro, 69 pp. https://portal.fiocruz.br/documento/especies-maruins-do-brasil
  » https://portal.fiocruz.br/documento/especies-maruins-do-brasil
• Soria SJ, Felippe-Bauer ML, Oliveira SJ (2002) Lista das espécies de Ceratopogonidae (Diptera, Nematocera) do agro-ecossistema cacaueiro, depositadas na Coleção Entomologica do Instituto Oswaldo Cruz, Rio de Janeiro, Brasil. Entomologia e Vetores 9(3): 317-327.
• Spinelli GR, Borkent A (2004) New species of Central American Culicoides Latreille (Diptera: Ceratopogonidae) with a synopsis of species from Costa Rica. Proceedings of the Entomological Society of Washington 106(2): 361-395.
• Spinelli GR, Grogan WL, Ronderos MM (2009b) A revision of the Patagonian predaceous midges of the genus Palpomyia Meigen (Diptera: Ceratopogonidae). Insect Systematics and Evolution 40: 43-70. https://doi.org/10.1163/187631209x416705
  » https://doi.org/10.1163/187631209x416705
• Spinelli GR, Ronderos MM, Ayala MM, Diaz F (2023) Catalog of the biting midges of Argentina (Diptera: Ceratopogonidae). Zootaxa 5261(1): 1-83. https://doi.org/10.11646/ZOOTAXA.5261.1.1
  » https://doi.org/10.11646/ZOOTAXA.5261.1.1
• Spinelli GR, Ronderos MM, Diaz F, Marino PI (2005) The bloodsucking biting midges of Argentina (Diptera: Ceratopogonidae). Memórias do Instituto Oswaldo Cruz 100: 137-150. https://doi.org/10.1590/S0074-02762005000200006
  » https://doi.org/10.1590/S0074-02762005000200006
• Spinelli GR, Santamaría E, Cabrera OL, Ronderos MM, Suárez MF (2009a) Five new species of Culicoides Latreille described from Colombia, yielding a new species list and country records (Diptera: Ceratopogonidae). Memórias do Instituto Oswaldo Cruz 104(1): 81-92. https://doi.org/10.1590/S0074-02762009000100013
  » https://doi.org/10.1590/S0074-02762009000100013
• Spinelli GR, Schaefer EF, Kehr AI (2002) First record of biting midges (Diptera: Ceratopogonidae) attacking frogs in the Neotropical region. Proceedings of the Entomological Society of Washington 104(2): 527-528.
• Spinelli GR, Wirth WW (1984) A review of the Neotropical predaceous midge genus Paryphoconus (Diptera: Ceratopogonidae). Proceedings of the Biological Society of Washington 97(4): 882-908.
• Tapia L, Sánchez T, Baylón M, Jara E, Arteaga C, Maceda D, Salvatierra A (2018) Invertebrados bentónicos como bioindicadores de calidad de agua en Lagunas Altoandinas del Perú. Ecología Aplicada 17(2): 149-163. https://doi.org/10.21704/rea.v17i2.1235
  » https://doi.org/10.21704/rea.v17i2.1235
• Wirth WW (1952) Los insectos de las islãs Juan Fernandez. 7. Heleidae and Tendipedidae (Diptera). Revista Chilena de Entomología 2: 87-104.
• Wirth WW (1974) A catalogue of the Diptera of Americas South of the United States. 14. Family Ceratopogonidae. Museu de Zoologia, São Paulo, 89 pp.
• Wirth WW, Blanton FS (1959) Biting midges of the genus Culicoides from Panama (Diptera: Heleidae). Proceedings of the United States National Museum 109: 237-482.
• Wirth WW, Blanton FS (1973) A review of the maruins or biting midges of the genus Culicoides (Diptera: Ceratopogonidae) in the Amazon Basin. Amazoniana 4: 405-470.
• Wirth WW, Cavalieri F (1977) Ceratopogonidae. In: Hurlbert SH (Ed.) Biota acuática de Sudamérica Austral. San Diego University, California, 280-281.
• Wirth WW, Hubert AA (1989) The Culicoides of Southeast Asia (Diptera: Ceratopogonidae). Memoirs of the American Entomological Institute 44: 1-508.

ADDITIONAL NOTES

• ZooBank register

  https://zoobank.org/802AC2BC-04DE-4200-87B6-13CC3B459308

• How to cite this article

  Santarém MCA, Felippe-Bauer ML (2024) Analysis of Brazilian Ceratopogonidae (Diptera: Culicomorpha) species diversity and knowledge assessment. Zoologia 41: e23066. https://doi.org/10.1590/S1984-4689.v41.e23066

• Published by

  Sociedade Brasileira de Zoologia at Scientific Electronic Library Online (https://www.scielo.br/zool)

Supplementary material 1

Supplementary S1. An updated list of Brazilian species of Biting Midges (Diptera: Ceratopogonidae).

Authors: MCA Santarém, ML Felippe-Bauer

Data type: Catalogue.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/zoologia.41.e23066

Edited by

Editorial responsibility Gabriel L.F. Mejdalani

Publication Dates

• Publication in this collection
  07 June 2024
• Date of issue
  2024

History

• Received
  27 Sept 2023
• Accepted
  19 Mar 2024