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An updated checklist of Anopheles (Diptera, Culicidae) of Colombia with new records and distribution data
expand article infoNelson Naranjo-Díaz, Margarita M. Correa
‡ Universidad de Antioquia, Medellín, Colombia
Open Access

Abstract

Several species of Anopheles mosquitoes (Arthropoda, Insecta, Diptera, Culicidae) are important in public health due to their role in malaria transmission. Of the more than 500 Anopheles species worldwide, 47 have been reported in Colombia, but only nine are known to transmit malaria. Taxonomic classification of these mosquitoes is complicated by the existence of species complexes and groups of closely related species that are difficult to distinguish based on morphological characteristics. However, molecular techniques have contributed to resolving taxonomic uncertainties, definition of molecular variants and facilitated the correction of erroneous taxonomic assignments. This study aimed to update the list of Anopheles species reported for Colombia. A species checklist was compiled by reviewing catalogs, publications, databases, and unpublished data. Only formally characterized species were included, along with their geographic range and ecological distribution. The final list includes 44 formally characterized Anopheles species belonging to five subgenera. The Nyssorhynchus subgenus constituted the largest group with 17 species and the widest distribution, occurring in 18 ecoregions. The Anopheles subgenus was the second largest group with 16 species and occurrences in 16 ecoregions. Sixty-six new presence records were added to the checklist. The updated Anopheles checklist, encompassing presence records and ecological distributions, enhances our understanding of Anopheles mosquito biodiversity. Furthermore, it contributes to improved public health by providing a foundation for targeted vector control interventions.

Key words:

Anopheles, biodiversity, Colombia, ecological distribution, inventory, malaria vectors

Introduction

The Anopheles genus is highly diverse, with 511 formally recognized species belonging to eight subgenera, including various species complexes. Some of these complexes still contain unnamed members (Harbach 2023). Anopheles mosquitoes thrive in a wide range of ecosystems, contributing to their broad geographical distribution. Additionally, some species have adapted to anthropically modified habitats (Hiwat and Bretas 2011; Harbach 2023). The primary importance of this genus lies in its role as a vector of human and animal pathogens, particularly, Plasmodium parasites, the causative agents of malaria (WHO 2022). Some Anopheles species also transmit Wuchereria bancrofti, the nematode that causes filariasis in Africa and Asia (Manguin et al. 2010), and O’nyong-nyong virus, which produces polyarthritis and fever in Africa (Brault et al. 2004).

Most of Colombia has ecological conditions that favor the widespread distribution of Anopheles mosquitoes (Olano et al. 2001; IGAC 2002; Hernández-Valencia et al. 2020). In the country, 47 species have been identified (Montoya-Lerma et al. 2011; Gómez et al. 2015), and nine of these are considered malaria vectors. Three species from the Nyssorhynchus subgenus play a significant role in malaria transmission: Anopheles darlingi, An. nuneztovari, and An. albimanus (Olano et al. 2001; Gutiérrez et al. 2008). Anopheles darlingi is the primary malaria vector in Latin America (Hiwat and Bretas 2011) and is predominantly found in the northwest, east and Amazon regions of Colombia. Anopheles nuneztovari is more common in the northwest, northeast and east, while An. albimanus is primarily present in coastal areas (Olano et al. 2001; Naranjo-Díaz et al. 2016a, b).

Knowledge of Anopheles species in Colombia is based primarily on catalogs published in the mid-20th century (Gast 1943; Barreto-Reyes 1955; Stone et al. 1959; Knight and Stone 1977) and older reports from the former government malaria control program “Servicio Nacional de Erradicación de la Malaria-SEM” (SEM 1957). Subsequent contributions by Carrejo and González (1992) and González and Carrejo (2007, 2009), focusing on the taxonomy and control of medically important insects, with an emphasis on population and taxonomic studies, have significantly improved our understanding of Anopheles in Colombia. While the number of reports on Anopheles species occurrence and distribution has increased, information remains scattered and, in some cases, inaccessible to the public.

Molecular tools have significantly improved the resolution of taxonomic ambiguities, particularly within species complexes and among closely related species; they have also facilitated the correction of erroneous taxonomic assignments (Brochero et al. 2007; Ruiz-Lopez et al. 2012; Escovar et al. 2014; Gómez et al. 2015). These findings underscore the need for an updated list of species occurrence. A comprehensive checklist of Anopheles species in Colombia, incorporating current ecological and geographical data, is essential for advancing medical entomological research and biodiversity assessment.

Materials and methods

To update the Anopheles species checklist, we reviewed various sources, including mosquito catalogs (Gast 1943; Barreto-Reyes 1955; Knight and Stone 1977; Heinemann and Belkin 1978; Faran 1980; Carrejo and Gonzalez 1992), taxonomic keys (Faran and Linthicum 1981; Gonzalez and Carrejo 2009), and government reports (SEM 1957). Additionally, we obtained data on Anopheles species occurrence, including geographical coordinates, from databases (Gaffigan et al. 2014; SIB Database 2020; GBIF 2024), scientific articles, and unpublished new records of specimens collected by members of our research group (named “new occurrence data”). The identities of the new records were previously verified using molecular methods, e.g., COI barcoding or ITS2 analysis (Zapata et al. 2007; Cienfuegos et al. 2011; Gómez et al. 2015) (Suppl. material 1).

To be included in the checklist, an Anopheles species was required to have a formal description and validation (Harbach 2023; ITIS 2024). Species variants originally described using only molecular methods were excluded. The checklist includes species listed in alphabetic order. Each entry contains the genus, subgenus, authorship, and year of description; (new occurrence data): This designation is used to indicate that the records are the result of previous work by our research group. Geographical distribution is presented at the level of Colombian administrative departments. An asterisk (*) indicates species occurrences with associated geographical coordinates.

Notes provide information on the sources of the data, changes in the taxonomic classification of species, molecular species designations, and new occurrence data, including the municipalities where specimens were collected. The checklist is accompanied by a map illustrating the distribution of Anopheles species with associated geographical coordinates (Figs 1, 2, Suppl. material 1).

Figure 1. 

Maps depicting the number of Anopheles species reported in Colombia A by administrative area (Department) B by ecological region.

Figure 2. 

Map indicating the location of species records, according to geographic coordinates. Blue: previously reported records; red: new occurrence data.

In addition to the list, a summary table of the Anopheles records by administrative departments is included (Table 1), and also, a table of species distribution per ecoregion (WWF 2015) (Table 2), which includes the following: Amazon-Orinoco-Southern Caribbean mangroves, Apure-Villavicencio dry forests, Caquetá moist forests, Catatumbo moist forests, Cauca Valley dry forests, Cauca Valley montane forests, Chocó-Darién moist forests, Cordillera Oriental montane forests, Eastern Cordillera Real montane forests, Guajira-Barranquilla xeric scrub, Llanos, Magdalena Valley montane forests, Magdalena-Urabá moist forests, Napo moist forests, Negro-Branco moist forests, Northwestern Andean montane forests, Sinú Valley dry forests, Solimões–Japurá moist forests, South American Pacific mangroves, Southwest Amazon moist forests, and Western Ecuador moist forests.

Table 1.

Summary of Anopheles records by Colombian administrative departments.

Department Subgenera Species New geographic coordinate registers* Total geographic coordinate registers£
Amazonas 3 8 - 11
Antioquia 3 22 40 155
Arauca 2 16 - 6
Atlántico 2 10 - 1
Bolívar 3 17 - -
Boyacá 3 17 - 4
Caldas 3 17 - 1
Caquetá 4 18 - 6
Casanare 2 16 - 2
Cauca 4 18 1 42
Cesar 2 15 - -
Chocó 4 23 - 31
Córdoba 2 18 21 80
Cundinamarca 5 23 - 2
Guainía 2 7 - -
Guaviare 3 13 3 5
Huila 3 13 - -
La Guajira 2 15 - 4
Magdalena 3 17 - 6
Meta 5 30 - 60
Nariño 4 15 - 35
Norte de Santander 4 24 - 14
Putumayo 3 18 - 36
Quindío 2 7 - -
Risaralda 2 7 - 3
San Andres y Providencia 1 1 - -
Santander 2 19 - 33
Sucre 2 13 - -
Tolima 3 18 - 1
Valle del Cauca 4 19 1 33
Vaupés 5 15 - -
Vichada 2 10 - 9
Table 2.

List of Anopheles recorded in Colombia by ecoregion.

Ecoregions* Species
Amazon-Orinoco-Southern Caribbean mangroves An. (Ano.) apicimacula, An. (Ano.) punctimacula, An. (Ano.) pseudopunctipennis±
Apure-Villavicencio dry forests An. (Ano.) costai, An. (Ano.) apicimacula, An. (Ano.) pseudopunctipennis±, An. (Ker) bambusicolus, An. (Ker) homunculus, An. (Ker) pholidotus±, An. (Nys.) albitarsis, An. (Nys.) argyritarsis, An. (Nys.) benarrochi±, An. (Nys.) braziliensis, An. (Nys.) darlingi¥, An. (Nys.) marajoara, An. (Nys.) nuneztovari¥, An. (Nys.) rangeli, An. (Nys.) triannulatus, An. (Ste.) nimbus
Caquetá moist forests An. (Nys.) darlingi¥, An. (Nys.) braziliensis
Catatumbo moist forests An. (Ano.) malefactor, An. (Ano.) neomaculipalpus, An. (Nys.) albitarsis, An. (Nys.) marajoara, An. (Nys.) nuneztovari¥
Cauca Valley dry forests An. (Ano.) calderoni±, An. (Ano.) punctimacula±, An. (Ano.) pseudopunctipennis±, An. (Ker) neivai±, An. (Nys.) albimanus¥
Cauca Valley montane forests An. (Ano.) apicimacula, An. (Ano.) calderoni±, An. (Ano.) pseudopunctipennis±, An. (Nys.) albimanus¥, An. (Nys.) albitarsis, An. (Nys.) nuneztovari¥, An. (Nys.) triannulatus
Chocó-Darién moist forests An. (Ano.) apicimacula, An. (Ano.) calderoni±, An. (Ano.) costai/forattinii, An. (Ano.) malefactor, An. (Ano.) punctimacula±, An. (Ker) neivai±, An. (Nys.) albimanus¥, An. (Nys.) darlingi¥, An. (Nys.) eiseni, An. (Nys.) nuneztovari¥, An. (Nys.) triannulatus
Cordillera Oriental montane forests An. (Ker) homunculus, An. (Nys.) darlingi¥, An. (Nys.) rangeli
Eastern Cordillera Real montane forests An. (Nys.) argyritarsis
Guajira-Barranquilla xeric scrub An. (Ano.) neomaculipalpus, An. (Ano.) punctimacula±
Llanos An. (Ano.) costai, An. (Ano.) peryassui, An. (Ano.) pseudopunctipennis±, An. (Ano.) shannoni, An. (Ker) bambusicolus, An. (Nys.) albitarsis, An. (Nys.) argyritarsis, An. (Nys.) braziliensis, An. (Nys.) darlingi¥, An. (Nys.) marajoara, An. (Nys.) nuneztovari¥, An. (Nys.) oswaldoi, An. (Nys.) rangeli, An. (Nys.) triannulatus
Magdalena Valley montane forests An. (Ano.) apicimacula, An. (Ano.) neomaculipalpus, An. (Ano.) pseudopunctipennis±, An. (Nys.) argyritarsis, An. (Nys.) darlingi¥, An. (Nys.) nuneztovari¥, An. (Nys.) parvus, An. (Nys.) rangeli, An. (Nys.) triannulatus
Magdalena-Urabá moist forests An. (Ano.) apicimacula, An. (Ano.) malefactor, An. (Ano.) mattogrossensis, An. (Ano.) neomaculipalpus, An. (Ano.) peryassui, An. (Ano.) pseudopunctipennis±, An. (Ano.) punctimacula±, An. (Ker) neivai±, An. (Lop) squamifemur,An. (Nys.) albimanus¥, An. (Nys.) albitarsis, An. (Nys.) aquasalis, An. (Nys.) argyritarsis, An. (Nys.) benarrochi±, An. (Nys.) braziliensis, An. (Nys.) darlingi¥, An. (Nys.) evansae, An. (Nys.) marajoara, An. (Nys.) nuneztovari¥, An. (Nys.) oswaldoi, An. (Nys.) rangeli, An. (Nys.) strodei, An. (Nys.) triannulatus, An. (Ste.) nimbus
Napo moist forests An. (Ano.) costai, An. (Ano.) mattogrossensis, An. (Ano.) neomaculipalpus, An. (Ano.) punctimacula±, An. (Nys.) albitarsis, An. (Nys.) bellator, An. (Nys.) benarrochi±, An. (Nys.) braziliensis, An. (Nys.) darlingi¥, An. (Nys.) marajoara, An. (Nys.) oswaldoi, An. (Nys.) rangeli, An. (Nys.) strode, An. (Nys.) triannulatus
Negro-Branco moist forests An. (Ano.) mattogrossensis, An. (Ano.) peryassui, An. (Nys.) braziliensis, An. (Nys.) darlingi¥, An. (Nys.) oswaldoi
Northwestern Andean montane forests An. (Ano.) calderoni±, An. (Ano.) pseudopunctipennis±, An. (Nys.) albimanus¥, An. (Nys.) argyritarsis, An. (Nys.) darlingi¥, An. (Nys.) nuneztovari¥, An. (Nys.) triannulatus
Sinú Valley dry forests An. (Ano.) punctimacula±
Solimões–Japurá moist forests An. (Ano.) costai, An. (Ano.) peryassui, An. (Nys.) darlingi¥, An. (Nys.) dunhami
South American Pacific mangroves An. (Ano.) apicimacula, An. (Ano.) calderoni±, An. (Ker) neivai±, An. (Nys.) albimanus¥
Southwest Amazon moist forests An. (Nys.) darlingi¥
Western Ecuador moist forests An. (Ano.) calderoni±, An. (Nys.) albimanus¥

Checklist of Anopheles mosquitos in Colombia

Order Diptera Linnaeus, 1758

Infraorder Culicomorpha Hennig, 1948

Superfamily Culicoidea Meigen, 1818

Family Culicidae Meigen, 1818

Subfamily Anophelinae Grassi, 1900

Genus Anopheles Meigen, 1818

Subgenus Anopheles Meigen, 1818

Anopheles (Anopheles) apicimacula Dyar & Knab, 1906

Distribution. Antioquia*, Arauca, Bolívar, Boyacá, Caldas, Casanare, Cauca, Cesar, Chocó*, Córdoba*, Cundinamarca, Guaviare, Huila, La Guajira, Magdalena, Meta*, Nariño*, Norte de Santander, Putumayo, Risaralda, Santander*, Sucre, Tolima, Valle del Cauca*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2005), González and Carrejo (2009), Parra-Henao et al. (2012), Montoya et al. (2017), SIB (2020), Zuleta-Ruiz et al. (2022).

Anopheles (Anopheles) calderoni Wilkerson, 1991

Distribution. Antioquia, Caldas, Chocó*, Huila, La Guajira, Magdalena, Nariño*, Norte de Santander, Quindio, Risaralda*, Tolima, Valle del Cauca*.

Notes. Local malaria vector. Reported by González and Carrejo (2009), González et al. (2010), Lucumi-Aragón et al. (2011), Orjuela et al. (2015), Montoya et al. (2017), Galeano-Castañeda et al. (2019), SIB Database (2020).

Anopheles (Anopheles) costai Da Fonseca & Ramos, 1939

Distribution. Amazonas*, Antioquia, Arauca, Bolívar, Boyacá, Caldas, Caquetá, Cesar, Chocó, Córdoba, Cundinamarca, Guainía, Guaviare, Huila, Meta*, Nariño, Putumayo*, Santander, Sucre, Valle del Cauca, Vaupés.

Notes. Reported by González and Carrejo (2009), Gutiérrez et al. (2009), Ahumada et al. (2013), Orjuela et al. (2013), SIB (2020). Anopheles costai was resurrected from synonymy with Anopheles mediopunctatus (Sallum et al. 1999), and it was previously erroneously reported in Colombia as An. mediopunctatus, as pointed out by Sallum et al. (1999) and Quiñones et al. (2001).

Anopheles (Anopheles) eiseni Coquillett, 1902

Distribution. Antioquia, Boyacá, Caldas, Casanare, Cauca, Chocó*, Cundinamarca, Huila, La Guajira, Magdalena, Meta, Nariño, Norte de Santander, Quindío, Risaralda, Santander, Tolima, Valle del Cauca*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), SIB (2020).

Anopheles (Anopheles) fluminensis Root, 1927

Distribution. Norte de Santander.

Notes. Reported by González and Carrejo (2009).

Anopheles (Anopheles) forattinii Wilkerson & Sallum, 1999

Distribution. Meta, Vaupés.

Notes. Reported by Wilkerson and Sallum (1999). It was indicated that An. forattinii was previously reported in Colombia as An. mediopunctatus (Sallum et al. 1999; Quiñones et al. 2001).

Anopheles (Anopheles) malefactor Dyar & Knab, 1907

Distribution. Antioquia*, Chocó*, Córdoba*, Meta, Norte de Santander*.

Notes. Reported by Wilkerson (1990), Álvarez et al. (2018), SIB Database (2020). New occurrence data from Monteria municipality, Córdoba Department.

Anopheles (Anopheles) mattogrossensis Lutz & Neiva, 1911

Distribution. Amazonas*, Arauca, Caquetá*, Cauca, Guainía, Guaviare, Meta, Norte de Santander, Putumayo, Vaupés, Vichada*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2006), Orjuela et al. (2013), Brochero and Conn (2015), Álvarez et al. (2018), Prado et al. (2019).

Anopheles (Anopheles) neomaculipalpus Curry, 1931

Distribution. Amazonas, Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Casanare, Caquetá, Cauca, Cesar, Chocó, Córdoba*, Cundinamarca, Guaviare, Huila, La Guajira*, Magdalena, Meta, Nariño, Norte de Santander*, Putumayo*, Santander*, Sucre, Tolima, Valle del Cauca, Vaupés.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2006), Parra-Henao and Alarcón (2008), González and Carrejo (2009), Parra-Henao et al. (2012), Orjuela et al. (2013), Álvarez et al. (2018). New occurrence data from Caceres Municipality, Antioquia Department, and Monteria and Valencia municipalities, Córdoba Department.

Anopheles (Anopheles) peryassui Dyar & Knab, 1908

Distribution. Amazonas*, Antioquia*, Caldas, Caquetá, Casanare, Cundinamarca, Guainía, Guaviare, Meta*, Putumayo, Santander*, Sucre, Vaupés, Vichada*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009), Brochero and Conn (2015), Álvarez et al. (2018), SIB (2020).

Anopheles (Anopheles) pseudopunctipennis Theobald, 1901

Distribution. Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó, Córdoba*, Cundinamarca, Guaviare, Huila, La Guajira, Magdalena*, Meta*, Nariño, Norte de Santander, Putumayo, Quindío, Risaralda, Santander*, Sucre, Tolima, Valle del Cauca*.

Notes. Local malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2006), Parra-Henao and Alarcón (2008), González and Carrejo (2009), Parra-Henao et al. (2012), Naranjo-Díaz et al. (2013), Montoya et al. (2017), SIB (2020). New occurrence data from Apartadó, Arboletes, Cáceres, Necoclí, San Juan de Urabá and Tarazá Municipalities in Antioquia Department, and Canalete Municipality in Córdoba Department.

Anopheles (Anopheles) punctimacula Dyar & Knab, 1906

Distribution. Antioquia*, Arauca, Atlántico, Bolívar, Boyacá*, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó*, Córdoba*, Cundinamarca, Guaviare, Huila, La Guajira*, Magdalena*, Meta, Nariño*, Norte de Santander, Putumayo*, Quindío, Risaralda, Santander, Sucre, Tolima, Valle del Cauca*, Vaupés, Vichada.

Notes. Local malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009), Gutiérrez et al. (2009), Orjuela et al. (2013), Naranjo-Díaz et al. (2013, 2014), Montoya et al. (2017), Álvarez et al. (2018), Galeano-Castañeda et al. (2019), SIB (2020). New occurrence data from Apartadó, Caucasia and Necoclí Municipalities in Antioquia Department, and Canalete Municipality in Córdoba Department.

Anopheles (Anopheles) shannoni Davis, 1931

Distribution. Vaupés, Vichada*.

Notes. Reported by González and Carrejo (2009), Brochero and Conn (2015).

Anopheles (Anopheles) vestitipennis Dyar & Knab, 1906

Distribution. Cesar, Valle del Cauca.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977).

Subgenus Kerteszia Theobald, 1905

Anopheles (Kerteszia) bambusicolus Komp, 1937

Distribution. Caquetá, Meta*.

Notes. Reported by Gast (1943), Barreto-Reyes (1955), Knight and Stone (1977), SIB Database (2020).

Anopheles (Kerteszia) bellator Dyar & Knab, 1906

Distribution. Putumayo*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), SIB Database (2020).

Anopheles (Kerteszia) boliviensis (Theobald, 1905)

Distribution. Caldas, Cauca, Chocó, Cundinamarca, Huila, Meta, Nariño, Tolima.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009).

Anopheles (Kerteszia) homunculus Komp, 1937

Distribution. Boyacá, Cauca, Chocó, Cundinamarca, Meta*, Norte de Santander, Tolima.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009), SIB Database (2020).

Anopheles (Kerteszia) neivai Howard, Dyar & Knab, 1913

Distribution. Antioquia, Bolívar, Boyacá, Cauca, Chocó*, Cundinamarca, Nariño*, Norte de Santander, Tolima, Valle del Cauca*, Vaupés.

Notes. Local malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Solarte et al. (1996), Gutierréz et al. (2008, 2009), González and Carrejo (2009), Naranjo-Díaz et al. (2014, 2023), SIB (2020).

Anopheles (Kerteszia) pholidotus Zavortink, 1973

Distribution. Caquetá, Cundinamarca, Magdalena, Meta*, Norte de Santander, Putumayo*, Tolima*, Valle del Cauca.

Notes. Local malaria vector. Reported by González and Carrejo (2009), Escovar et al. (2014), SIB (2020). Previously reported as Anopheles lepidotus (Escovar et al. 2014), a species that is not present in the country.

Subgenus Lophopodomyia Antunes, 1937

Anopheles (Lophopodomyia) gilesi (Peryassú, 1908)

Distribution. Meta.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977).

Anopheles (Lophopodomyia) oiketorakras Osorno-Mesa, 1947

Distribution. Cundinamarca, Nariño.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977).

Anopheles (Lophopodomyia) squamifemur Antunes, 1937

Distribution. Antioquia*, Cauca, Chocó, Norte de Santander, Valle del Cauca. Vaupés.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977). New occurrence data from Cáceres Municipality in Antioquia Department.

Subgenus Nyssorhynchus Blanchard, 1902

Anopheles (Nyssorhynchus) albimanus Wiedemann, 1820

Distribution. Antioquia*, Atlántico, Bolívar, Cauca*, Cesar, Chocó*, Córdoba*, La Guajira, Magdalena, Nariño*, Risaralda*, Sucre, San Andrés y Providencia, Valle del Cauca*.

Notes. Primary malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Faran (1980), Gutierrez et al. (Gutiérrez et al. 2008, 2009), Calle et al. (2008), González and Carrejo (2009), Naranjo-Díaz et al. (2013, 2014, 2023), Montoya et al. (2017), Galeano-Castañeda et al. (2019), Altamiranda-Saavedra et al. (2023), SIB (2020). New occurrence data from Apartadó, Mutatá, Necoclí, San Juan de Urabá and Tarazá Municipalities in Antioquia Department, Guapi Municipality in Cauca Department, Canalete, Monteria and Puerto Escondido Municipalities in Córdoba Department, and El Zarzal Municipality in Valle del Cauca.

Anopheles (Nyssorhynchus) albitarsis Lynch Arribálzaga, 1878

Distribution. Antioquia*, Meta*, Norte de Santander*, Putumayo*, Vichada*.

Notes. Reported by Calle et al. (2008), Gutiérrez et al. (2009), González and Carrejo (2009), Jiménez et al. (2012), Ahumada et al. (2013), Orjuela et al. (2013), Montoya et al. (2017), Galeano-Castañeda et al. (2019), SIB (2020). In Colombia, Anopheles albitarsis was previously reported as An. albitarsis s.l. or An. marajoara, both are part of the Albitaris Complex, which potentially comprises at least ten species, only five have been formally described (Bourke et al. 2021). Only the molecular variants An. albitarsis F and An. albitarsis I are reported in the country, An. marajoara could not be confirmed (Ruiz-Lopez et al. 2012). New occurrence data from Arboletes and Caucasia Municipalities in Antioquia Department, Moñitos and San Antero Municipalities in Córdoba Department.

Anopheles (Nyssorhynchus) aquasalis Curry, 1932

Distribution. Atlántico, Bolívar, Chocó, Córdoba*, La Guajira*, Magdalena

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Calle et al. (2008). New occurrence data from San Antero Municipality in Córdoba Department.

Anopheles (Nyssorhynchus) argyritarsis Robineau-Desvoidy, 1827

Distribution. Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó, Cordobá*, Cundinamarca, Huila, La Guajira, Magdalena, Meta*, Nariño*, Norte de Santander, Putumayo, Quindío, Risaralda, Santander*, Tolima, Valle del Cauca, Vaupés, Vichada*.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009), Jiménez et al. (2012), Parra-Henao et al. (2012) Naranjo-Díaz et al. (2013), SIB (2020). New occurrence data from Cáceres Municipality in Antioquia Department.

Anopheles (Nyssorhynchus) benarrochi Gabaldón, Cova García & Lopez, 1941

Distribution. Meta*, Putumayo*, Santander*.

Notes. Local malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Quiñones et al. (2001), Calle et al. (2008), González and Carrejo (2009), Parra-Henao et al. (2012), Orjuela et al. (2013). In Colombia, a molecular variant denominated An. benarrochi B was reported, distributed in the south of the country (Ruiz et al. 2005).

Anopheles (Nyssorhynchus) braziliensis Chagas, 1907

Distribution. Amazonas, Antioquia*, Arauca, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cesar, Chocó, Córdoba, Cundinamarca, Guainía, Guaviare*, Huila, La Guajira, Magdalena, Meta*, Norte de Santander, Putumayo*, Santander, Tolima, Valle del Cauca, Vaupés, Vichada*.

Notes. Reported by Barreto-Reyes (1955), Brochero et al. (2005), Calle et al. (2008), González and Carrejo (2009), Naranjo-Díaz et al. (2013, 2023), Jiménez et al. (2012, 2014), Ahumada et al. (2013), SIB (2020). New occurrence data from Arboletes, Cáceres, Caucasia and Tarazá Municipalities in Antioquia Department, San Jose del Guaviare Municipality in Guaviare Department.

Anopheles (Nyssorhynchus) darlingi Root, 1926

Distribution. Amazonas*, Antioquia*, Arauca, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cesar, Chocó*, Córdoba*, Cundinamarca, Guainía, Guaviare*, La Guajira, Magdalena, Meta*, Norte de Santander, Putumayo*, Santander, Sucre, Vaupés, Vichada*.

Notes. Primary malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2005), Calle et al. (2008), González and Carrejo (2009), Jiménez et al. (2012, 2014), Ahumada et al. (2013), Orjuela et al. (2013), Naranjo-Díaz et al. (2013, 2014, 2016a, 2019), Montoya et al. (2017), Pacheco et al. (2017), Galeano-Castañeda et al. (2019), SIB (2020). New occurrence data from Apartadó, Cáceres, Caucasia and Necoclí Municipalities in Antioquia Department, San Carlos and Valencia Municipalities in Córdoba Department, San Jose del Guaviare Municipality in Guaviare Department.

Anopheles (Nyssorhynchus) dunhami Causey, 1945

Distribution. Amazonas*.

Notes. Reported by Ruiz et al. (2010).

Anopheles (Nyssorhynchus) evansae Brèthes, 1926

Distribution. Córdoba*.

Notes. Reported by Knight and Stone (1977), Parra-Henao and Alarcon (2008).

Anopheles (Nyssorhynchus) marajoara Galvão & Damasceno, 1942

Distribution. Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Caquetá*, Casanare, Cauca, Cesar, Chocó, Córdoba, Cundinamarca, Guaviare, Huila, La Guajira, Magdalena*, Meta*, Norte de Santander*, Putumayo, Santander, Sucre, Tolima, Vaupés, Vichada*.

Notes. Reported by Knight and Stone (1977), González and Carrejo (2009), Brochero et al. (2010), Jiménez et al. (2012).

Anopheles (Nyssorhynchus) nuneztovari Gabaldón, 1940

Distribution. Antioquia*, Arauca*, Bolívar, Boyacá, Caldas, Caquetá, Casanare*, Cauca, Cesar, Chocó*, Córdoba*, Cundinamarca, Huila, Magdalena, Meta, Norte de Santander*, Putumayo, Santander*, Sucre, Tolima, Valle del Cauca*.

Notes. Primary malaria vector. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Calle et al. (2002, 2008), Brochero et al. (2006), Fajardo et al. (2008), Parra-Henao and Alarcón (2008), González and Carrejo (2009), Fonseca-González et al. (2009), Ruiz et al. (2010), Parra-Henao et al. (2012), Naranjo-Díaz et al. (2013, 2014, 2016b, 2023), Montoya et al. (2017), Galeano-Castañeda et al. (2019), Altamiranda-Saavedra et al. (2023), SIB (2020). New occurrence data from Apartadó, Caucasia, Necoclí and Tarazá Municipalities in Antioquia Department, Monteria and Moñitos Municipalities in Córdoba Department.

Anopheles (Nyssorhynchus) oswaldoi Peryassú, 1922

Distribution. Amazonas, Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó, Córdoba, Cundinamarca, Guainía, Guaviare*, Magdalena, Meta, Nariño, Norte de Santander, Putumayo*, Santander*, Sucre, Tolima, Valle del Cauca, Vaupés, Vichada*.

Notes. Reported by Barreto-Reyes (1955), González and Carrejo (2009), Ruiz et al. (2010), Parra-Henao et al. (2012), Jiménez et al. (2012, 2014), Orjuela et al. (2013). New occurrence data from Caucasia Municipality in Antioquia Department, San Jose del Guaviare Municipality in Guaviare Department.

Anopheles (Nyssorhynchus) parvus Chagas, 1907

Distribution. Arauca, Casanare, Meta, Santander*, Vichada.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Parra-Henao et al. (2012), SIB (2020).

Anopheles (Nyssorhynchus) rangeli Gabaldón, Cova García & Lopez, 1940

Distribution. Antioquia*, Arauca*, Bolívar, Boyacá*, Caldas*, Caquetá*, Casanare, Cauca, Cesar, Chocó, Córdoba*, Cundinamarca*, Guainía, Guaviare*, La Guajira, Magdalena, Meta*, Nariño, Norte de Santander, Putumayo*, Santander*, Tolima, Valle del Cauca.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2005, 2006), Calle et al. (2008), González and Carrejo (2009), Jiménez et al. (2012), Parra-Henao et al. (2012), Orjuela et al. (2013), SIB (2020). New occurrence data from Arboletes and Caucasia Municipalities in Antioquia Department, Moñitos Municipality in Córdoba Department.

Anopheles (Nyssorhynchus) strodei Root, 1926

Distribution. Antioquia, Arauca*, Bolívar, Boyacá, Caquetá, Casanare, Chocó, Córdoba, Cundinamarca, La Guajira, Meta*, Norte de Santander, Putumayo*, Santander*, Valle del Cauca.

Notes. Reported by Barreto-Reyes (1955), Calle et al. (2008), González and Carrejo (2009), Parra-Henao et al. (2012), SIB Database (2020).

Anopheles (Nyssorhynchus) triannulatus Neiva & Pinto, 1922

Distribution. Amazonas, Antioquia*, Arauca, Atlántico, Bolívar, Boyacá, Caldas, Caquetá, Casanare, Cauca, Cesar, Chocó*, Córdoba*, Cundinamarca, Guaviare, Huila, La Guajira, Magdalena*, Meta*, Nariño, Norte de Santander, Putumayo*, Quindío, Santander*, Sucre, Tolima, Valle del Cauca.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), Brochero et al. (2006), Calle et al. (2008), González and Carrejo (2009), Parra-Henao et al. (2012), Ahumada et al. (2013), Naranjo-Díaz et al. (2013, 2023), Orjuela et al. (2013), Montoya et al. (2017), Atencia-Pineda et al. (2018), SIB (2020). New occurrence data from Apartadó, Arboletes, Caucasia, Necoclí, and San Juan de Urabá Municipalities in Antioquia Department, Canalete, Monteria, Moñitos, San Antero, and Valencia Municipalities in Córdoba Department.

Anopheles (Nyssorhynchus) trinkae Faran, 1979

Distribution. Meta*.

Notes. Reported by González and Carrejo (2009), SIB Database (2020).

Subgenus Stethomyia Theobald, 1902

Anopheles (Stethomyia) kompi Edwards, 1930

Distribution. Caquetá.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977).

Anopheles (Stethomyia) nimbus (Theobald, 1902)

Distribution. Cundinamarca, Guaviare, Meta, Vaupés, Valle del Cauca.

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977), González and Carrejo (2009).

Anopheles (Stethomyia) thomasi Shannon, 1933

Notes. Reported by Barreto-Reyes (1955), Knight and Stone (1977).

Discussion

The genus Anopheles contains eight subgenera, five of which are present in the Neotropical region (Anopheles, Kerteszia, Lophopodomyia, Nyssorhynchus, and Stethomyia; Harbach 2023). All five subgenera are present in Colombia. The current checklist only includes formally described species and excludes molecularly identified variants. Therefore, a total of 44 species are listed as present in Colombia. Species of the subgenera Nyssorhynchus and Anopheles are the most widely distributed in the country, occurring in 32 and 31 Colombian departments, respectively. Nyssorhynchus is the subgenus with the highest species richness, comprising 17 species. Stethomyia has a limited distribution, being found in only three departments. Stethomyia and Lophopodomyia subgenera are represented by three species each. The highest species richness was detected in Meta Department, with a total of 30 species, followed by Norte de Santander with 24. San Andres and Providencia Department exhibited the lowest species richness, with only the main Colombian malaria vector, An. albimanus, being recorded in this insular region (Table 1) (Fig. 1).

It is well known that factors such as temperature, rainfall, and humidity affect the geographical distribution of Anopheles species (Abiodun et al. 2016). Colombia exhibits a wide variety of ecological conditions, and 34 ecoregions are described (WWF 2015). The current checklist reports the presence of Anopheles species in 21 of those ecoregions (Table 2). The Magdalena-Urabá moist forests ecoregion exhibits the highest richness with 23 species, dominated by the species of the subgenera Nyssorhynchus and Anopheles, 14 and 7 species, respectively. The characteristics of this ecoregion include consistently high temperatures exceeding 28 °C and average annual rainfall within the range of 2,000–4,000 millimeters (WWF 2015). This ecoregion forms part of the important Tumbes-Chocó-Magdalena biodiversity hotspot, which spans southern Panamá to northern Perú and encompasses coastal and lowland areas of the Magdalena River basin and the Urabá region in northwest Colombia. The diverse ecological requirements of Anopheles species, which thrive on a variety of landscapes ranging from mosaic tropical rainforests to wetlands, mangrove swamps, and coastal plains, contribute to the high species richness detected in these Colombian regions.

Several members of the genus Anopheles are important in public health due to their role as malaria vectors. In Colombia, the annual number of malaria cases has exceeded 70,000 in recent years (INS 2021, 2022, 2023). Of the nine Anopheles species implicated in malaria transmission (Olano et al. 2001; Gutiérrez et al. 2008; Orjuela et al. 2013; Naranjo-Díaz et al. 2013), at least one malaria vector species was registered in each of the 21 ecoregions with occurrence data, except for the Eastern Cordillera Real montane forests (Table 1). As the Magdalena-Urabá moist forests ecoregion exhibited the highest species richness, it is not surprising that it also contained most of the malaria vectors, including the three primary vectors (An. albimanus, An. darlingi, and An. nuneztovari) and four local vectors (An. pseudopunctipennis, An. punctimacula, An. neivai, and An. benarrochi). The Chocó-Darién moist forests ecoregion was second in the number of vectors present, including the three primary vectors and three local vectors (An. calderoni, An. punctimacula, and An. neivai). These two ecoregions encompass the most important Colombian malaria regions, the Urabá, Bajo Cauca, Alto Sinú, and Pacific regions, which report the highest annual number of malaria cases (INS 2023).

Several factors contribute to the discrepancies between previous catalogs and the current checklist. For example, An. mediopunctatus may be restricted to Brazil, and in Colombia, it was likely misidentified as An. costai or An. forattinii (Sallum et al. 1999; Quiñones et al. 2001). Additionally, Anopheles cruzii, although reported from Costa Rica to Argentina, has not been confirmed in Colombia (Wilkerson and Peyton 1991). Furthermore, Anopheles vargasi has been registered in neighboring Venezuela (Del Ventura et al. 2013), but there is no evidence of its presence in Colombia (Osorno-Mesa 1947). Mistaken taxonomic assignments, often due to the existence of cryptic species, have led to erroneous reports. For instance, An. pholidotus was previously reported as An. lepidotus (Escovar et al. 2014).

Conclusions

In recent decades, numerous studies and surveys conducted in the country have provided valuable data on the presence and distribution of Anopheles species. The current checklist attempts to compile the available information. The current list includes 44 formally characterized Anopheles species from five subgenera, with the subgenera Nyssorhynchus and Anopheles being the largest and most widely distributed species groups. Information for 66 new occurrence data is also provided. The incorporation of presence records and ecological distributions is essential for accurately estimating the Anopheles species diversity and assessing the malaria vectors. This is fundamental for the design and implementation of effective control interventions.

An interesting finding is that most of the new species records are concentrated in specific regions, likely reflecting research interest in the most malaria-endemic regions of Colombia. It is noteworthy that after 47 species had been recorded in the country, the current checklist only includes 44 species. For an Anopheles species to be included in the list, it was required to have a formal description and validation; species variants originally described using only molecular methods were excluded. Among the factors contributing to the discrepancies between previous catalogs and the current checklist are the existence of problematic species, or in some cases damaged specimens, which can lead to misidentifications; also, mistaken taxonomic assignments can occur due to the existence of cryptic species. In addition, for some species, there was no evidence of their presence in Colombia. Finally, while the use of molecular techniques has helped to clarify the taxonomic status of several problematic species, this has led to an increase in the number of molecular variants reported. However, it has also facilitated the correction of erroneous taxonomic assignments, which in turn, may lead to a decrease in the number of species formally described.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work received support from Escuela de Microbiología, Universidad de Antioquia, project code No. 2023-66350.

Author contributions

Nelson Naranjo-Díaz: specimen identification, data analysis, writing, review & editing. Margarita M. Correa: data analysis, project coordinator, critical revisions and editing of manuscript drafts.

Author ORCIDs

Margarita M. Correa https://orcid.org/0000-0003-2419-7269

Data availability

The dataset containing the information on Anopheles species occurrence in Colombia is available in: https://doi.org/10.5281/zenodo.13527884. (Data will be available upon publication).

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Supplementary material

Supplementary material 1 

Data on Anopheles species occurrence in Colombia

Nelson Naranjo-Díaz, Margarita M. Correa

Data type: xlsx

Explanation note: Geographic coordinates are shown in decimal degrees and were taken from database portals, scientific articles, and the research group unpublished new occurrence data.

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.
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