The Mosquitoes (Diptera: Culicidae) of Sonora: Distribution, Ecology, and the First Records of Aedes deserticola Zavortink and Toxorhynchites septentrionalis (Dyar and Knab) in México

Abstract

The diversity and distribution of mosquitoes from Sonora, Mexico, was documented through entomologic surveys conducted in the four physiographic regions and sub-regions of Sonora: the Sonoran Plain, the Sierra Madre Occidental, the Northern Mountains and Plains, and the Pacific Coastal Plain. Immature stages were collected from aquatic habitats, while adult mosquitoes were collected using Shannon traps, resting in vegetation, and by human landing collections. Overall, 11,316 specimens, which comprised 493 larvae, 224 larval exuviae, 400 pupal exuviae, 33 pupae, 4552 females, 5607 males, and seven male genitalia, were identified. Two subfamilies: Anophelinae and Culicinae, seven tribes, 10 genera, 23 subgenera, and 56 species are reported below. Of these, one tribe, one genus, five subgenera, and 15 species were recorded for the first time in Sonora. Two species, Aedes deserticola Zavortink and Toxorhynchites septentrionalis (Dyar and Knab), are the first recordings of their kind nationwide. Toxorhynchites septentrionalis was also barcoded with the cytochrome oxidase subunit 1 (COI) gene. The presence of Psorophora columbiae (Dyar and Knab) is confirmed in Mexico. Taxonomic notes, new geographic distribution limits of mosquitoes in Sonora, and information regarding their importance as disease vectors are provided. By adding Ae. deserticola and Tx. septentrionalis and confirming Ps. columbiae in Mexico, there are currently 251 mosquito species in the Country.

1. Introduction

Mosquitoes (Diptera: Culicidae) are biting insects that are of public and animal health interest as some species serve as disease vectors of pathogens for humans and livestock [1], such as dengue, Chikungunya, Japanese encephalitis, yellow fever, West Nile, Rift Valley fever, and Zika viruses, as well as of nematodes and protozoans, such as Dirofilaria spp. and Plasmodium spp. [1,2]. Hence, mosquitoes are often the focus of surveillance and control programs worldwide [3]. Mosquitoes also serve as ecological indicators of landscape degradation [4,5,6]. Eleven out of thirty-two states in Mexico have been systematically studied in terms of mosquito taxonomy, ecology, and distribution. There are 26 species in Tlaxcala [7], 29 in Mexico City [8,9,10], 50 in Querétaro [11], 51 in Mexico State [12], 57 in Hidalgo [13,14], 58 in Morelos [10], 68 in Nuevo León [15,16,17,18], 76 in Quintana Roo [18,19,20,21,22,23,24], 80 in Tamaulipas [18,25], 105 in Tabasco [18,26,27,28,29], and 139 in Veracruz [18,30,31,32].

Prior to this study, 248 species of Culicidae mosquitoes belonging to twenty-one genera have been reported in Mexico [10,18], of which nine genera and 41 species have been reported in the state of Sonora. However, most records were based on collections made in urban and sub-urban regions. Thus, eastern mountain regions of the Sierra Madre Occidental of Sonora remained an area to be surveyed. Sonora is the twelfth state of Mexico to have the list of mosquito species updated. Most specimens collected and examined were mostly deposited in the Culicidae Collection of the Parasitology Department of the Autonomous Agrarian University Antonio Narro, Laguna unit, Torreón, Coahuila, Mexico, while some other specimens were deposited in local entomological collections of Sonora.

2. Material and Methods

2.1. Study Area

Sonora is located in north-western Mexico (32°29′ and 26°14′ NL; 108°26′ and 205°2′ WL). This state, with an area of 179,355 km², is the second largest in Mexico after Chihuahua. To the north, Sonora borders Arizona, USA, to the south with Sinaloa and the Gulf of California, to the west with Baja California and the Gulf of California, and to the east with Chihuahua and Sinaloa. Sonora is divided into four physiographical regions and sub-regions indicated in parenthesis: Sonoran Plain (Altar Desert, Pinacate Mountains, Sonoran Mountains and Plains), Sierra Madre Occidental (Sierras and Valleys of the North, Sierras and Glens of the North, Foot of the Sierra, Great Chihuahuan Plateau and Canyons), Northern Mountains and Plains (Northern Dunes and Plains), and Pacific Coastal Plain (Coastal Plain and Deltas of Sonora and Sinaloa) [33] (Figure 1). The regions and sub-regions of Sonora, including the municipalities surveyed, are depicted in Table 1. Here, entomological surveys were conducted in four physiographical regions and sub-regions of Sonora during the dry and rainy seasons (Table 1), and a complete checklist of 56 species of Sonora is reported in Result section table.

2.2. Mosquito Collection and Identification

Immature stages (except eggs) and adult mosquitoes were collected in specific sites in the four physiographic regions of Sonora (Table 1) during the dry and rainy seasons from 2018 through 2022. Immature stages were collected from any available water body found in the collection site, and some environmental parameters were recorded, such as pH, salinity, and temperature, which were measured using a portable Hanna^® (mod. HI98129) tester. Immature stages were also collected using plastic ovitraps deployed in the branches of some trees to mimic tree holes containing rainwater. Live larvae and pupae were placed in plastic bags containing water from the original aquatic habitat and transported to the Laboratorio de Biología Molecular of the Parasitology Department of the Autonomous Agrarian University Antonio Narro Laguna unit (LBM-UAAAN-UL). A portion of individual fourth-instar larvae from each collection site was mounted on microscope slides containing Euparal for larval identification. Other individual larvae were placed into emergence tubes to obtain the adult stage and associated larval and pupal exuviae. Male genitalia were dissected solely when taxonomic identification required it. Host-seeking mosquitoes were collected using human landing collections and Shannon traps containing humans deployed between 10:00 and 15:00 h. and between 20:00 and 24:00 h. Resting mosquitoes were aspirated directly from caves and vegetation using mechanical aspirators. Adult mosquitoes were killed with triethylamine vapors, mounted on insect pins, and identified using taxonomic keys through a stereomicroscope Discovery V8 (Zeiss) (Oberkochen, Germany). Immature stages and exuviae were identified using a microscope Primostar (Zeiss). The Harbach and Knight [34] criteria for mosquito anatomy and Wilkerson et al. [35] for Aedini classification were followed in this study. Biological information for each species that are recorded for the first time in the state are noted; the northernmost and southernmost distributions of the species as well as the known medical importance of each species are noted (Table 2).

2.3. Genomic DNA Extraction and COI Amplification

A modified hotshot technique was used for genomic DNA extraction [36,37]. Briefly, two legs from a Tx. septentrionalis (Dyar and Knab) individual (serving as a voucher specimen) were placed into a micro-vial containing 50 µL of alkaline lysis buffer (25 mM NaOH, 0.2 mM Na₂EDTA) followed by sonication (Grant Ultrasonic 3L Bath) in a water bath for 20 min. Incubated afterwards in a thermocycler for 30 min at 94 °C and cooled for 5 min at 4 °C. The sample was centrifuged for 3 min at 3000 rpm, and 50 µL of neutralizing buffer (40 mMTri-HCL) were added. PCR amplification for the COI barcode gene [38,39] was performed using Folmer primers (LCO1490 and HCO2198) and the Qiagen PCR system using a reaction mix final volume of 50 μL containing 2 μL DNA template, 25 μL H₂O, 5 μL NH₄, 5 μL dNTPs (2 mM/μL), 2.5 μL MgCl₂ (25 mM/μL), 0.1 μL Bioline taq polymerase (Bioline Reagents Ltd., London, UK), 5 μL each primer at 10 pmol/μL, and 0.38 μL bovine serum albumin (20 mg/mL) [36,37]. PCR conditions included an initial denaturation step at 94 °C for 1 min, 5 pre-amplification cycles of 94 °C for 1 min, 45 °C for 1.5 min, and 72 °C for 1.5 min, followed by 35 amplification cycles of 94 °C for 1 min, 57 °C for 1.5 min, and 72 °C for 1 min. A final elongation step of 72 °C for 5 min was also performed. The PCR products were visualized through a 1.5% agarose gel, and the sample showed bands of the COI size bidirectionally sequenced using the ABI PRISM^® BigDye^® Terminator v3.1 Cycle Sequencing Kit (Applied Biosystems, Waltham, MA, USA). Toxorhynchites septentrionalis COI sequence and specimen data were uploaded in the Barcode of Life Database (BOLD) system with project named: Vectors of México Initiative.

2.4. Genetic Data Analysis

For molecular species identification using the COI DNA barcoding region, the protocols of Hebert et al. [38,39] and Hernández-Triana et al. [40,41] were followed. Amplicons were obtained using a Qiagen PCR system following the reaction mix of Hernández-Triana et al. [3]. The dataset was analyzed in MEGA v.6 [42]. Two sequences were retrived whose presumptive identity was Tx. Septentrionalis; additional Toxorhynchites spp. sequences were obtained from BOLD and/or GeneBank databases: six sequences of specimens identified as Tx. theobaldi (Dyar and Knab) (GeneBank: KY82650, KY86252–55) from specimens collected in Mexico; five sequences of Tx. moctezuma (Dyar and Knab) (BOLD:ADS6264) from specimens collected in Mexico; and five sequences of Tx. haemorrhoidalis (Fabricius) (BOLD:AXA1098) from specimens collected in Costa Rica were added to the dataset. One sequence of Cx. pipiens L. (Gene Bank:GU908074) was included as an outergroup for the phylogenetic analysis. A Bayesian inference was performed in BEASTv.2.6.7 [43] using the Hasegawa–Kishono–Yano (HKY) substitution model, assuming different nucleotide frequencies and transition/transversion rates, and estimated parameter priors with the Yule model. Three independent Markov chains of 10,000,000 replicates were run with sampling every 10,000. The sample parameters were combined with LogCombiner, and basic parameters were checked in Tracer v.1.7.1 [44]. The sampled trees were combined with LogCombiner and then summarized with TreeAnnotator. The posterior probability was calculated for each node in a maximum clade credibility tree with a burn-in of 10%. For barcode sequences larger than 500 bp, a barcode index number (BIN) was assigned, and each BIN was mapped according to species. The taxonomic discordance in the dataset [3] was analyzed using BOLD as detailed in Ratnasingham and Hebert [45].

2.5. Ethical Consideration

Mosquitoes were collected using human landing collections by co-author AIOM, brigades of the Ministry of Health (public health and vector control programs), and personnel from the Sonora Entomological and Bioassay Research Unit of Sonora (UIEB). The UIEB serves as a reference laboratory for the identification and examination of vectors collected by the State Ministry of Health. This study was approved by the Ethics Committee of the Antonio Narro Autonomous Agrarian University (UAAAN), the National Center of Preventive Programs and Disease Control (CENAPRECE), and the Sonora University (UNISON).

3. Results

3.1. Mosquitoes and New Records

A total of 11,316 specimens from 139 collection samples were studied. Of these, specimens were 493 fourth-instar larvae, 224 larval exuviae, 400 pupal exuviae, 33 pupae, 4552 adult females, and 5607 adult males. Seven genitalia of males were dissected. The mosquito fauna of Sonora consists of two subfamilies, seven tribes, 10 genera, 23 subgenera, and 56 species. Of these, one tribe (Uranotaeniini), one genus (Uranotaenia), five subgenera (Lewnielsenius, Janthinosoma, Anoedioporpa, Phenacomyia, and Uranotaenia), and 15 species: Anopheles franciscanus McCraken, Ae. deserticola Zavortink, Ae. muelleri Dyar, Ae. trivittatus (Coquillett), Haemagogus equinus Theobald, Psorophora longipalpus Randolph and O’Neill, Culex restrictor Dyar and Knab, Cx. erythrothorax Dyar, Cx. apicalis Adams, Cx. arizonensis Bohart, Cx. lactator Dyar and Knab, Toxorhynchites septentrionalis (Dyar and Knab), Uranotaenia geometrica Theobald, Ur. lowii Theobald, and Ur. sapphirina (Osten Sacken) are recorded for the first time in Sonora. Aedes deserticola and Tx. septentrionalis were first reported in Mexico (Table 3). Thus, the species accumulation curve of a total of 43 mosquito species collected is shown in Figure 2. In addition, medical importance of mosquitoes with known status as disease vectors in Sonora is provided in Table 4.

3.2. Biological and Ecological Notes for the New State Records

The collection sites, date of collection, larval habitat characteristics, aquatic parameters, and associated species are shown in Table 2.

3.2.1. Genus Anopheles

In Sonora, two subgenera and six species of Anopheles occurred; one species was recorded for the first time. Namely, immature stages of An. franciscanus were collected in northern Sonora, bordering with Arizona, USA; the aquatic habitats were stream margins and ground level ponds, always with abundant aquatic vegetation surrounding the habitats during the dry and rainy seasons. The municipalities where An. franciscanus was found were: Cumpas, Agua Prieta, and Imuris.

3.2.2. Genus Aedes

Three out of the eleven species of Aedes found in Sonora were recorded for the first time: Ae. deserticola, Ae. muelleri, and Ae. trivittatus; one subgenus Lewnielsenius was also recorded for the first time. One single Ae. deserticola and Ae. muelleri adult females were obtained from immature stages collected from a discarded tire with colored water and abundant leaves at the bottom in a mountain site of the Cananea municipality, near Arizona, USA; Ae. muelleri was collected using human landing collections in a forested site with partial shade of the Cananea municipality; and Ae. trivittatus adult females were usually collected when Shannon traps containing humans were deployed at day in several collection sites during the rainy season. All collection sites where these Aedes species were collected belong to the Sierra Madre Occidental region.

3.2.3. Genus Haemagogus

One of the two Haemagogus species in Sonora was recorded for the first time in the area, Hg. equinus adult females were collected during daylight hours in a sub-tropical site of southern Sonora, near Sinaloa. This collection site belongs to the Sierra Madre Occidental region. Another species of this genus reported in Sonora is Hg. Mesodentatus, which is potentially sympatric with Hg. equinus in southern Sonora.

3.2.4. Genus Psorophora

The subgenus Janthinosoma of Psorophora was recorded for the first time in the area. Psorophora longipalpus adult females were collected using human landing collection while intending to bite at day light hours in a site with partial shade in coastal Navojoa municipality. Specimens were deposited in the state public health mosquito collection, in Hermosillo municipality.

3.2.5. Genus Culex

Culex is the most diverse group in Sonora. Five out of twenty species were recorded for the first time, which also included the subgenera Anoedioporpa and Phenacomyia. Culex restrictor adults were collected resting in caves at ground level with total shade in a mountain collection site in the Sierra Madre Occidental; while Cx. erythrothorax females were caught using Shannon traps containing humans at night in another mountain collection site near Arizona, USA border; immature stages of Cx. apicalis were collected from discarded tires in a forested site in the Sierra Madre Occidental region; at this same collection site, immature stages of Cx. arizonensis where also found. In Sonora, species belonging to the subgenus Neoculex such as Cx. apicalis, Cx. arizonensis and Cx. territans are sympatric, thus, all those species were found at the same collection sites near the Yécora municipality in the Sierra Madre Occidental region. Finally, immature stages of Cx. lactator were usually found from temporal ground pools, at ground level, in different collection sites, while adult stages were collected resting in vegetation. This is a common species in Sonora.

3.2.6. Genus Toxorhynchites

Toxorhynchites septentrionalis is was recorded for the first time in Sonora and nationwide. Immature stages were collected from one discarded tire with colored water in one forest site of Álamos municipality in southern Sonora, a humid and sub-tropical area of the Sierra Madre Occidental region. Specimens were deposited at the UIEB mosquito collection in Navojoa municipalty. The other species previously reported in Sonora is Tx. moctezuma; as distribution limits of Tx. moctezuma remain unknown, thus, both species are sympatric and considered valid records.

3.2.7. Genus Uranotaenia

The tribe Uranotaeniini, genus Uranotaenia, subgenus Uranotaenia, and the species Ur. geometrica, Ur. lowii, and Ur. sapphirina were recorded for the first time in Sonora. Uranotaenia geometrica adults were collecting resting inside a discarded tire in Yécora municipality. These mosquitoes were observed before being caught using a mouth aspirator. Females were actively walking on the inner side of the tire; some refused to fly even if disturbed with the tip of the aspirator. A single female seemed to protect their freshly laid eggs, laid on the water surface using her mid legs. The eggs were collected, but did not hatch under the laboratory conditions. Ur. lowii is perhaps the most common species of the genus Uranotaenia in Mexico. Adults were found resting in vegetation in one collection site near Sinaloa, while immature stages were collected from small aquatic habitats in two collection sites near the coast area; finally, Ur. sapphirina immature stages were collected from natural and artificial breeding sites in a forested collection site in the Sierra Madre Occidental region of Yécora municipality. As Ur. sapphirina feeds on leenches [60], we pernocted overnight at the collection site where this species was found, but we did not observe leenches or other helminthes around, only earthworms. All Uranotaenia species were collected during the rainy season.

Table 4

Medical and veterinary importance as disease vectors of pathogens of Sonora. Pla: Plasmodium (Malaria). DI: Dirofilaria immitis. DENV: dengue virus. ZIKV: Zika virus. CHIKV: Chikungunya virus. YF: yellow fever virus. SLE: St. Louis encephalitis virus. WNV: West Nile virus. VEEV: Venezuelan equine encephalitis virus. EEEV: Eastern equine encephalitis virus. WEEV: Western equine encephalitis virus. LCV: La Crosse virus [61]. * Host-seeking mosquitoes caught using human landing collections at day light hours. ** Host-seeking mosquitoes caught using human landing collections at night hours.

Taxa	Pla	DI	DENV	ZIKV	CHKV	YF	SLE	WNV	VEEV	EEEV	WEEV	LCV
Anopheles freeborni	✓								✓
An. pseudopunctipennis	✓								✓
An. punctipennis	✓	✓
An. albimanus	✓				✓
Aedes vexans		✓		✓			✓	✓	✓	✓	✓
Ae. ramirezi *
Ae. epactius *
Ae. muelleri *
Ae. taeniorhynchus								✓	✓	✓
Ae. trivittatus		✓						✓		✓	✓	✓
Ae. podographicus *
Ae. aegypti			✓	✓	✓	✓		✓	✓	✓		✓
Haemagogus equinus *
Hg. mesodentatus					✓	✓		✓	✓
Psorophora columbiae								✓	✓
Ps. ciliata							✓	✓	✓		✓
Ps. howardii *
Culex erythrothorax **
Cx. interrogator *
Cx. nigripalpus							✓	✓	✓	✓
Cx. quinquefasciatus					✓		✓	✓	✓	✓
Cx. restuans							✓	✓		✓
Cx. salinarius							✓	✓		✓
Cx. stigmatosoma **
Cx. tarsalis								✓	✓		✓
Cx. erraticus								✓		✓
Culiseta inornata								✓			✓
Coquillettidia perturbans								✓		✓

3.3. Molecular Analysis, COI-Barcoding Identification, and Cryptic Diversity

Two DNA barcode sequences from Tx. septentrionalis were examined. One was from a single adult female (accession number No. 08210922-MN), and the other one was downloaded from the BOLD system associated with one specimen originally identified as Tx. rutilus (Coquillett) collected by UIEB-Sonora personnel (BOLD:AAK1973); both specimens were collected at the same collection site near the Álamos municipality. As a result of the phylogenetic analysis, both sequences were grouped together within the sequences NEONU980-12 and NEONU981-12, forming a clade for the Tx. septentrionalis species with a strong support value (0.998). In addition, the rest of the sequences analyzed clustered in well-defined clades for the Toxorhynchites species: Tx. theobaldi (0.9198), Tx. moctezuma (1), and Tx. haemorrhoidalis (1), also with strong support values. In contrast, Tx. rutilus sequences did not show a consistent pattern; however, ACMC010-04 diverged from Tx. theobaldi/Tx. moctezuma clade (0.5192), while GMCGG105-14 grouped with the Tx. theobaldi sequences (Figure 3).

3.4. Medical Importance of the Mosquitoes of Sonora

Some Culicidae species reported in Sonora are of medical and veterinary importance as disease vectors of pathogens (Table 4). Although the animal or human importance of some Culicidae species is still unknown, host-seeking species caught using human landing collection methods were noted.

4. Discussion and Conclusions

4.1. Mosquito Species-Groups Reaching Their Distributional Limits in Sonora

Based on collection records and the known distribution of mosquito species in Sonora, four groups of species can be recognized. Some species in each group have similar geographic distribution reaching either the northern or southern limits in Sonora, also sharing same or similar aquatic habitat characteristics and environmental parameters.

4.1.1. Group 1

Species occurring in the Nearctic region and extending into northern Mexico were reaching their southern limit in Sonora. The immature stages of these species develop in artificial or natural aquatic sites such as trees or rockholes containing rainwater. Species of Aedes subgenera: Kompia and Jarnellius. Aedes species within this group are desert-like mosquitoes distributed from the southwest of Arizona to northern Baja California and Sonora. In this group, Ae. purpureipes, Ae. deserticola, and Ae. monticola are included. Aedes purpureipes was originally described by Aitken [62] from specimens collected in Distrito del Sur, Baja California, and was designated as type material of the monobasic subgenus Kompia. More specimens were collected in Santa Cruz municipality of Arizona [58]; the larvae were collected and described from specimens collected from one tree hole in San Blas municipality of Sinaloa by Vargas [63]; some collection records in Sonora were prvodied by Henimann and Belkin [54]. Aedes purpureipes is known only in the southwestern United States, in California and Arizona [56,64], and in Baja California, Nayarit, Oaxaca, Sinaloa, and Sonora of Mexico [52]. Although records indicate the southernmost limit of Ae. purpureipes as southern Mexico (Oaxaca), this is doubtful, but this record remains valid until additional specimens for this species can be collected and examined from this region. Aedes deserticola was described by Zavortink [57] from specimens collected in Los Angeles municipality, California, United States. Here, immature stages were collected from a discarded tire in Cananea municipality, northern Sonora, near the Mexico–Arizona border. Thus, it is the first record of Ae. deserticola in Mexico and an outer record from the United States. This species occurs in southwestern and western Arizona [53,65,66] where it can be sympatric with local populations of Ae. monticola. Aedes deserticola adult females are distinguished from other species of Jarnellius subgenus by the following characters [58]: Postcoxal area without scales; subspiracular area without setae; base of hindtarsal segment IV without one broad ring of white scales, which is very narrow if is present; metameron without scales; and postpronotum without dark scales (Figure 4). Aedes monticola was described by Belkin and McDonald [67] from specimens collected in Santa Cruz county, Arizona, United States. This species is found in the southwestern United States, in Arizona and New Mexico [67,68], and northwestern Mexico in Sonora [52]. Toxorhynchites septentrionalis (Figure 5) is included in this group because it reaches the southernmost distributional limit in Sonora. This species was originally described as Megarhinus septentrionalis by Dyar and Knab [69] from specimens collected in the District of Columbia, United States. Afterwards, this taxon was reduced to sub-specific status of Tx. rutilus by Jenkins and Carpenter [70]. Further, Tx. septentrionalis returned to valid species status [71]. Toxorhynchites septentrionalis has been found in the North Atlantic and southeastern United States [59]. In Mexico, specimens of this species were collected from one discarded tire in Álamos municipality. Adults are also deposited in the UIEB of Sonora collection in the Navojoa municipality, where there is a common species (Med. entomol. of Sonora, pers. com.). Toxorhynchites septentrionalis is distributed from the North Atlantic of the United States, throughout the Mid-Atlantic and southeastern Atlantic, reaching its southernmost distribution limit in the Sierra Madre Occidental of Sonora. Our hypothesis is that this species is also present in New Mexico and Arizona, USA, and Chihuahua, Mexico, but this needs to be further investigated.

4.1.2. Group 2

Species occurring in the Nearctic region and extending into northern Mexico were reaching their southern limit in Sonora. Immature stages are in swamps and ponds at ground level; they include the species Anopheles freeborni, which was originally described by Aitken [72] as a sub-species of An. maculipennis Meigen from specimens collected in Davis, California, EUA, and further designated as valid species by King and Bradley [73]. Anopheles freeborni is found in southern Canada, the western arid and semi-arid regions of the United States, and northwestern Mexico in Baja California and Sonora. Although, in this study, we were not able to find this species, we reviewed the Collection of Insects and Mites of Medical Importance deposited at the Institute of Diagnosis and Epidemiological Reference (CAIM-INDRE) in Mexico City and observed large numbers of An. freeborni mounted on insect pins previously collected in Sonora.

4.1.3. Group 3

Species occurring in the neotropical region extend into northern Mexico were reaching their northern limit in Sonora. Immature stages are in artificial or natural aquatic sites such as trees or rockholes containing rainwater. Include Aedes ramirezi, Ae. podographicus, Haemagogus mesodentatus, and Culex restrictor. Although Ae. ramirezi is placed in this group, this Mexican endemic species is distributed mainly in sites related to the Nearctic region. Aedes ramirezi was originally described by Vargas and Downs [74] from specimens collected in Cuernavaca, Morelos; this species has also been found in Jalisco, Morelos, Sinaloa, Sonora, and Veracruz [52]. Adult females can bite humans during the day and dusk; however, the medical importance is still unknown. Aedes podographicus was originally described by Dyar and Knab [75] from specimens collected in Sonsonate, El Salvador. This species has also been found in Belize, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, and Panama [76]; in Mexico, Ae. podographicus is common within the Terrens group in the coastal and tropical regions of the south-east, being a frequent human-biting species during the day, but the medical importance of this species is still unknown. Haemagogus mesodentatus was originally described by Komp and Kumm [77] from specimens collected in San Jose, Costa Rica; this species occurs in Belize, Costa Rica, El Salvador, Guatemala, Mexico, and Panama [76]. In Mexico, this species has also been recorded in Campeche, Chiapas, Guerrero, Morelos, Nayarit, Oaxaca, Puebla, Sinaloa, Sonora, Tabasco, Veracruz, and Yucatán [52,78], being Sonora’s northernmost distribution limit, which apparently is restricted to the tropical, sub-tropical areas of the Pacific Coastal Plain and the Sierra Madre Occidental regions. Culex restrictor was originally described by Dyar and Knab [79] from specimens collected in Almoloya, Oaxaca, Mexico; this species has also been found in Belize, Brazil, Colombia, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama, and Venezuela [76]; in Mexico, this species has been collected in tree holes in several states of the neotropical region but have also been found in temperate and conserved areas of Mexico [12,15]; in Sonora, Cx. restrictor is a rare species, and its distribution is restricted to the forested region of the Sierra Madre Occidental. Our hypothesis is that all species included in this group could extend their distribution range further north than they were found, through the Sierra Madre Occidental region and they could reach and inhabit the forested area of northern Sonora, in the Northern Mountains and Plains region (Figure 1) where there is no physiographic disruption between both physiographic regions, but rather the ecotone between them allows woody trees to grow in whose tree holes these species could develop. If this were the case, species of this group could inhabit the isolated mountain ranges of southern Arizona.

4.1.4. Group 4

Species occurring in neotropical region extending into northern Mexico, where they reaching their northern distribution limit in Sonora. Immature stages are in swamps and ponds at ground level; include the species Culex bidens and Uranotaenia geometrica. Culex bidens was originally described by Dyar [80] from specimens collected in Rosario, Bolivia; this species occurs in Argentina, Bolivia, Brazil, Colombia, Costa Rica, Mexico, Nicaragua, Paraguay, Uruguay, and Venezuela [81]. Although this species was not found in this study, the occurence records of Cx. bidens are relatively common in Mexico, and the species is well known [82]. Uranotaenia geometrica was originally described by Theobald [83] from sepecimens collected in São Paulo, Brazil; this species occurs in the neotropical region from Brazil to Mexico, where it reaches its northernmost distribution limit. The previous group, Cx. bidens and Ur. geometrica could be present in northern Sonora, as well as southern Arizona.

4.2. Confirmation of Psorophora columbiae in Mexico

In the latest inventory of mosquito species in Mexico [18], Ps. confinnis (Arribálzaga) was removed from the species list because this taxon had historically been confused and misidentified with another species in the Confinnis group. Psorophora columbiae was not included in the Mexican species list until additional material of species within the Confinnis group could be further studied. In Sonora, immature stages of Ps. columbiae were collected from a temporal pond with partial shade and abundant aquatic vegetation in a sub-arid area of the Fronteras municipality; from a temporal pound with total shade in the Moctezuma municipality; and from a temporal pond in a site near the coast with partial shade and abundant aquatic vegetation in the Navojoa municipality. Adult Ps. columbiae mounted on insect pins and deposited in the UIEB mosquito collection were also examined. The presence of Ps. columbiae is confirmed in Mexico and this species occurs in Baja California, Baja California Sur, Campeche, Chiapas, Coahuila, Colima, Durango, Guerrero, Jalisco, Michoacán, Morelos, Nuevo León, Oaxaca, Puebla, Quintana Roo, San Luis Potosí, Sinaloa, Sonora, Tabasco, Tamaulipas, Veracruz, and Yucatán.

4.3. Species Not Collected in This Study

Nine species of this group were not found in this study, but records were obtained from the literature reports, including those from Vargas [48]: Anopheles freeborni and An. judithae. In Mexico, those Anopheles species have been reported previously in northwestern Mexico; An. freeborni was not found in this study, although data from labels of specimens mounted on insect pins from CAIM collection (see Group 2 of previous section) shows this species is common during the rainy season in the border municipalities of Agua Prieta and Nogales. Anopheles judithae is one of the rarest mosquito species in Mexico; this tree-hole Anopheles has been found from southern Arizona, New Mexico, and Texas [84,85] up to the Sierra Madre Occidental in Zacatecas [54]. Although surveys were conducted in the Imuris municipality, where An. judithae was previously reported [85], deploying ovitraps in tree trunks to mimic the aquatic habitat of the immature stages, this species was absent in our survey.

Martínez-Palacios [50]: Culex bidens. This tropical species has been previously reported from Argentina to Mexico. In Mexico, this species has been reported in different states, where apparently it seems a common mosquito in tropical regions of Mexico. However, it was not found in our study.

Zavortink [53]: Aedes burgeri. In Mexico, this species has been reported in Jalisco, Sinaloa, and Sonora [54]; immature stages of this species develop in tree holes containing rainwater, but as this habitat was not observed during our study, Ae. burgeri and other tree-hole species such as An. judithae, Ae. Monticola, and Hg. mesodentatus were not found.

Díaz-Nájera and Vargas [52]: Aedes monticola, Culiseta impatiens, Cs. incidens, and Cs. inornata. In Mexico, Ae. monticola is known only in Sonora; this may have been absent in our study due to the above reasons. Distribution in Mexico of Cs. impatiens was recently reviewed, and its presence was confirmed for Sonora and Nuevo León [18]. Cs. incidens has been reported in Baja California, Baja California Sur, Coahuila, Mexico State, Sonora, and Tlaxcala, while Cs. inornata has been reported in all previous states as well as in Aguascalientes, Durango, Hidalgo, Mexico City, Nuevo León, and Tamaulipas [10]. All these species of Culiseta are common during the winter and can be found in temporal ground ponds with cold water even when the surface of the water is frozen; our surveys were conducted only during the dry and rainy seasons, with no surveys conducted during the winter season.

Heinemann and Belkin [54]: Haemagogus mesodentatus. In Mexico, this tropical species has been reported in Chiapas, Colima, Guerrero, Mexico State, Morelos, Nayarit, Oaxaca, Puebla, Sinaloa, Tabasco, Veracruz, and Yucatán [12,52,78,86]. In Sonora, within the genus Haemagogus, only Hg. equnius was found.

4.4. Species from Adjacent Regions That May Occur in Sonora

Numerous species of mosquitoes that have not yet been reported from Sonora but occur in the adjacent states of Arizona and/or Chihuahua can logically be expected to occur within Sonora. Included among these 10 species in Arizona are Aedes papago Zavortink*, Ae. varipalpus (Coquillett)*, Aedes cataphylla Dyar*, Ae. fitchii (Felt and Young)*, Ae. implicates Vockeroth*, Ae. nigromaculis (Ludlow), Ae. ventrovittis Dyar*, Psorophora discolor (Coquillett), Orthopodomyia signifera (Coquillett), and Uranotaenia anhydor Dyar (*Species not reported in Mexico); three species in Chihuahua: Ae. canadensis (Theobald), Ae. mitchellae (Dyar), and Ae. thelcter Dyar; and three species reported in Arizona and Chihuahua: Ae. dorsalis (Meigen), Ae. melanimon Dyar, and Ae. sollicitans (Walker) [70,87,88].

4.5. Mosquito Diversity in Sonora and Mexico

With the addition of the two new mosquito records for Mexico (Ae. deserticola and Tx. septentrionalis) and the confirmation of the presence of Ps. columbiae in the country, there are currently 251 species known in Mexico. With 56 species, Sonora ranks eighth place with the greatest diversity of species out of the 12 Mexican states that have been systematically inventoried for mosquito species.

4.6. DNA Barcoding

The DNA barcodes sequences of the four species of Toxorhynchites: Tx. haemorrhoidalis, Tx. moctezuma, Tx. septenrtionalis, and Tx. theobaldi of specimens grouped in well-defined and clades. The specimens B12 and MEXAO072-14 were identified as Tx. septentrionalis based on the diversification pattern observed in the phylogenetic analysis. Toxorhynchites septentrionalis and Tx. moctezuma showed diversification patterns with a strong support value, suggesting a sympatric speciation. Phylogenetic evidence and morphological traits found in the adult female general coloration in external vestitrue and scales coloration support the specimens correspond to different species. Mexican specimens identified as Tx. Theobaldi, collected in the southern part of the country, including the Yucatán Peninsula, belong to an undescribed species, which presumably corresponds to a new species, and might be indicative of cryptic diversity within the genus Toxorhynchites in Mexico.

Author Contributions

A.I.O.-M. and F.W.-C.—study conceptualization; A.I.O.-M. and F.W.-C.—methodology; A.I.O.-M.—sampling; A.I.O.-M., J.M.Q.-B. and M.A.R.-P.—data curation; A.I.O.-M.—mosquito identification; L.M.H.-T., J.M.Q.-B. and M.A.R.-P.—molecular analysis; A.I.O.-M.—writing and original draft preparation; L.M.H.-T. and M.A.R.-P.—review and editing; A.I.O.-M., L.M.H.-T., M.A.R.-P. and F.C.-M.—project administration and funding acquisition. All authors have read and agreed to the published version of the manuscript.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

All mosquitoes (mounted on insect pins and/or microscope slides) collected in Sonora State were deposited in the Culicidae Collection (CC-UL), Parasitology Department, Universidad Autónoma Agraria Antonio Narro unidad laguna (Autonomous Agrarian University Antonio Narro laguna unit), Torreón, Coahuila, Mexico. Some specimens of seven species (An. franciscanus, Ae. ramirezi, Hg. equinus, Ps. longipalpus, Ps. howardii, Cs. particeps, and Tx. moctezuma) were deposited in the Medical Entomology collection of Sonora state in Hermosillo municipality and the UIEB of Sonora in Navojoa municipality.

Acknowledgments

We thank Alejandro Samaniego-Pesqueira, Antonio A. Sánchez-García, Carla P. Alvarez-Echeverría, Diego A. Barrales-Alcalá, Elia S. Castro-Lomeli, Fabiola J. Gandarilla-Aizpuro, Germán Quiros-Chavez, Gladis Moreno-Bello, Guillermo Morillón-Borjón, Isis J. Morales-Avitia, Juan B. Contreras-Reza, Juan P. Castillo-Munguía, Orlando Tomas-Cruz, Quetzaly K. Siller-Rodríguez, Rodrigo Cedillo-Osornio, Salvador Morales-Avitia, and Ximena Santana-Garduño for their valuable collaboration during our collection trips; and to Herón Huerta for letting us review the CAIM-INDRE collection.

Conflicts of Interest

The authors declare no conflicts of interest.

Footnotes

Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Figures and Tables

Figure 1. Physiography of Sonora state. Red box shows the study area. Map originally created for this article.

Figure 2. Species accumulation curve for 45 mosquito species collected at 139 site collections in Sonora from 2021 through 2022.

Figure 3. Bayesian phylogenetic tree of COI DNA barcoding for Toxorhynchites spp. Red box shows Tx. septentrionalis of this study.

Figure 4. Aedes deserticola. female. PSc: postcoxal area. SA: subspiracular area. Mem: metameron. Ppn: postpronotum.

Figure 5. Toxorhynchites septentrionalis, male.

Table 1

Description of physiography of Sonora and municipalities surveyed for mosquitoes [33].

Region (Sub-Region)	Municipalities Sampled	Description of Sub-Regions
Sonoran Plain (Altar Desert)	Caborca, Puerto Peñasco	This arid sub-region extends from Arizona to Sonora and is distinguished by having large plains and fields of sand dunes that reach up to 200 m.a.s.l. The Sonoran Plain has the largest desert in Mexico.
(Pinacate Mountains)	Puerto Peñasco	This small sub-region is a group of desertic volcanic mountains and craters, surrounded by a large field of sand dunes.
(Sonoran Mountains and Plains)	Imuris, Altar, Caborca, Cajeme, Guaymas, Hermosillo, La Colorada, Magdalena, Pitiquito, Puerto Peñasco, Santa Ana	This arid sub-region extends from Arizona to Sonora, having plains, hills, and isolated mountains between 200 and 400 m.a.s.l. In the summer, the daytime temperature is 30–50 °C.
Sierra Madre Occidental (Sierras and Valleys of the North)	Imuris, Agua Prieta, Bacoachi, Banámichi, Bavispe, Cajeme, Cananea, Cumpas, Divisaderos, Onavas, Fronteras, Granados, Guaymas, Huásabas, La Colorada, Magdalena, Moctezuma, Nacozari de García, Rosario, San Javier, Santa Ana, Tepache	The Sierra Madre Occidental is the largest mountain system in Mexico. This sub-region presents alluvial plains, steep mountain ranges, descents with hills, and fields of sand dunes.
(Sierras and Glens of the North)	Agua Prieta, Arivechi, Bacadéhuachi, Bacerac, Bavispe, Divisaderos, Onavas, Granados, Huachinera, Huásabas, Nacozari de García, Rosario, Sahuaripa, Tepache, Yécora	This sub-region is made of high mountain ranges with ravines. The mountains with elevations of 1000–2700 m.a.s.l.
(Foot of the Sierra)	Álamos, Cajeme, Huatabampo, Navojoa, Rosario	The climate in this sub-region is temperate, and the predominant vegetation is pine and oak forests. There are some small patches of sub-tropical forest.
(Great Chihuahuan Plateau and Canyons)	Álamos, Rosario, Yécora	This sub-region is composed by mountains and canyons, some mountains with elevations above 2000 m.a.s.l.; the climate is temperate humid in winter and warm humid in summer. Extensive areas of forest and grasslands.
Northern Mountains and Plains (Northern Dunes and Plains)	Imuris, Agua Prieta, Bavispe, Cananea, Fronteras, Nacozari de García	This small sub-region extends from Arizona to Sonora. The relief of this sub-region is not very rugged; it presents alluvial plains, steep mountain ranges with hills, and fields of dunes.
Pacific Coastal Plain (Coastal Plain and Deltas of Sonora and Sinaloa)	Álamos, Cajeme, Ciudad Obregón, Etchojoa, Guaymas, Huatabampo, Navojoa, Villa Juárez	This coastal sub-region is mostly covered by alluvium, which descends from the Sierra Madre Occidental to the sea. Its climate is warm and sub-humid.

Table 2

Collection data of first records of Culicidae species in Sonora. Aquatic habitat characteristics and environmental parameters (pH, temperature, and concentrations of dissolved salts), collection site and municipality of Sonora; coordinates and elevation as well as date of collection and associated species are shown.

Species	Habitat/Environmental Parameters	Environment	Collection Site and Municipality	Coordinates/Elevation	Collection Date	Associated Species
Anopheles franciscanus	Stream margin with abundant aquatic vegetation, without shade	Arid and mountain region	Carretera a Nacozari, Cumpas	30°16′28″ N–109°41′28″ W, 950 m.	15 April 2022	An. pseudopunctipennis
	Stream margin with abundant aquatic vegetation, partial shade, (pH 6.41, 329 PPM, 29 °C)	Arid and mountain region	El Diablo, Agua Prieta	31°18′52″ N–108°59′2″ W, 1490 m.	14 September 2022	Cx. thriambus
	Stream margin with abundant aquatic vegetation, partial shade, (pH 8.15, 408 PPM, 27.1 °C)	Mountain region	Rancho Aribabi, Imuris	30°51′48″ N–110°39′39″ W, 1000 m.	21 September 2022	Cx. coronator, Cx. stigmatosoma, Cx. thriambus
	Temporal ground-pool with clear water and total shade (pH 6.35, 301 PPM, 29 °C)	Mountain region	Rancho Aribabi, Imuris	30°51′48″ N–110°39′39″ W, 1000 m.	21 September 2022	Cx. stigmatosoma, Cx. thriambus
Aedes deserticola	Discarded tire with colored water and abundant leaves at bottom (pH 5.3, 74 PPM, 24.2 °C)	Mountain region	Cananea	31°0′57″ N–110°20′58″ W, 1730 m.	21 September 2022	Ae. epactius, Ae. aegypti, Cx. quinquefasciatus, Cx. stigmatosoma
Aedes muelleri	Human biting/approaching at day with partial shade	Mountain region	Cananea	31°0′57″ N–110°20′58″ W, 1730 m.	21 September 2022	Ae. epactius
Aedes trivittatus	Shannon trap baited with human at day	Arid and mountain region	El Diablo, Agua Prieta	31°18′52″ N–108°59′2″ W, 1490 m.	14 September 2022	No associated spp.
	Shannon trap baited with human at day	Mountain region	Yécora	28°21′40″ N–108°57′20″ W, 1650 m.	16 September 2022	Simulium spp.
	Human biting/approaching at day with total shade	Mountain region	Campanero, Yécora	28°22′15″ N–108°55′55″ W, 1960 m.	16 September 2022	Ae. epactius
	Human biting/approaching at dusk	Mountain region	Yécora	28°21′57″ N–108°55′31″ W, 1550 m.	16 September 2022	No associated spp.
	Shannon trap baited with human at night	Mountain region	Rancho Aribabi, Imuris	30°51′30″ N–110°39′55″ W, 985 m.	21 September 2022	Cx. erythrothorax, Cx. stigmatosoma
	Human biting/approaching at day	Mountain region	Villa Alpina, Cananea	31°0′19″ N–110°19′41″ W, 1690 m.	26 September 2022	No associated spp.
Haemagogus equinus	Human biting/approaching at day with partial shade	Sub-tropical region	Camino al Chinal, Álamos	26°42′24.1″ N–108°43′14.1″ W	28 September 2022	Ae. podographicus
Psorophora longipalpus	Human biting/approaching at day with partial shade	Arid coastal region	Loma del Refugio, Navojoa	27°7′34.14″ N–109°27′36.42″ W	12 September 2018	No associated spp.
Culex restrictor	Adults resting in caves, total shade	Mountain region	Campanero, Yécora	28°22′15″ N–108°55′55″ W, 1960 m.	16 September 2022	No associated spp.
Culex erythrothorax	Shannon trap baited with human at night	Mountain region	Rancho Aribabi, Imuris	30°51′30″ N–110°39′55″ W, 985 m.	21 September 2022	Ae. trivittatus, Cx. stigmatosoma
Culex apicalis	Discarded tire with clean water and abundant leaves at bottom	Mountain region	Yécora	28°21′40″ N–108°57′29″ W, 1650 m.	16 September 2022	Cx. arizonensis, Cx. territans, Cs. particeps, Ur. sapphirina
	Discarded tire with colored water and abundant leaves at bottom	Mountain region	Campanero, Yécora	28°22′15″ N–108°55′55″ W, 1960 m.	16 September 2022	Cx. restuans, Cx. thriambus, Cx. arizonensis, Cs. particeps
Culex arizonensis	Discarded tire with clean water and abundant leaves at bottom	Mountain region	Yécora	28°21′40″ N–108°57′29″ W, 1650 m.	16 September 2022	Cx. apicalis, Cx. territans, Cs. particeps, Ur. sapphirina
	Discarded tire with colored water and abundant leaves at bottom	Mountain region	Yécora	28°22′15″ N–108°55′55″ W, 1960 m.	16 September 2022	Ae. epactius, Cx. territans
	Discarded tire with colored water and abundant leaves at bottom	Mountain region	Campanero, Yécora	28°22′15″ N–108°55′55″ W, 1960 m.	16 September 2022	Cx. restuans, Cx. thriambus, Cx. apicalis, Cs. particeps
Culex lactator	Adults resting in vegetation, total shade	Mountain region	Rancho Aribabi, Imuris	30°51′48″ N–110°39′39″ W, 1000 m.	21 September 2022	Cx. quinquefasciatus
	Temporal ground-pool with colored water and abundant vegetation with partial shade (pH 8.6, 393 PPM, 30 °C)	Arid region	Magdalena de Khino	30°37′46″ N–110°57′52″ W, 760 m.	26 September 2022	Cx. interrogator, Cx. quinquefasciatus, Cx. stigmatosoma
	Temporal ground-pool with clear water and abundant vegetation with total shade (pH 8.6, 1,386 PPM, 28.6 °C)	Coastal region	Huatabampo	26°48′39″ N–109°37′50″ W, 10 m.	28 October 2022	Cx. interrogator, Cx. quinquefasciatus, Cx. erraticus
	Temporal ground-pool with colored water and abundant vegetation with total shade (pH 5.12, 678 PPM, 20 °C)	Mountain region	El Rosario	27°50′32″ N–109°22′28″ W, 430 m.	2 November 2022	Cx. interrogator, Cx. tarsalis, Cx. stigmatosoma
Toxorhynchites septentrionalis	Discarded tire with colored water and abundant leaves at bottom	Sub-tropical region	Minas nuevas, Álamos	27°1′53″ N–109°0′52″ W, 560 m.	22 September 2022	No associated spp.
Uranotaenia geometrica	Adults resting in a discarded tire with total shade	Mountain region	La Aduana, Yécora	28°21′40″ N–108°57′29″ W, 1650 m.	22 September 2022	No associated spp.
Uranotaenia lowii	Adults resting in vegetation	Semi-arid and hilly region	Ciudad Obregón	27°31′9″ N–109°55′36″ W, 40 m.	12 April 2022	Ae. vexans, Cx. interrogator, Cx. quinquefasciatus, Cx. salinarius, Or. kummi
	Water channel with abundant aquatic vegetation and total shade (pH 6.5, 1255 PPM, 26 °C)	Coastal region	Bocobampo, Villa Juárez	27°1′41″ N–109°44′56″ W, 10 m.	28 October 2022	An. pseudopunctipennis, An. albimanus, Cx. coronator, Cx. interrogator, Cx. thriambus
	Animal tracks with clear water, abundant aquatic vegetation and partial shade (pH 7.9, 450 PPM, 17.8 °C)	Coastal region	Pótam, Guaymas	27°41′46″ N–110°22′33″ W, 10 m.	29 October 2022	An. pseudopunctipennis, An. albimanus, Cx. coronator, Cx. nigripalpus, Cx. thriambus, Cx. tarsalis, Cx. erraticus
Uranotaenia sapphirina	Temporal ground-pool with colored water and abundant vegetation with partial shade	Mountain region	Yécora	28°21′49″ N–108°57′29″ W, 1650 m.	16 September 2022	Cx. coronator
	Discarded tire with clean water and abundant leaves at bottom	Mountain region	Yécora	28°21′40″ N–108°57′29″ W, 1650 m.	16 September 2022	Cx. apicalis, Cx. arizonensis, Cx. territans, Cs. particeps

Table 3

Checklist of mosquito species of Sonora. Ho: [46]. Ma¹: [47]. Va¹: [48]. Va²: [49]. Ma²: [50]. Va³: [51]. DV: [52]. Za: [53]. HB: [54]. Ma³: [55]. F.R.: first record. FSR: first state record. FNR: first nationwide record. First records in Sonora are indicated in bold letters. ^a Reported as Anopheles barberi Coquillett. ^b Reported as Psorophora confinnis (Lynch Arribálzaga). ^c Reported as Ps. virescens Dyar and Knab. ^d Reported as Culex virgultus Theobald. ^e Reported as Cx. peus Speiser. Species were identified using identification keys (Carpenter and LaCasse [56], Zavortink [57], Zavortink [53], Arnell and Nielsen [58], and Darsie and Ward [59].

Taxa	F.R.	Taxa	F.R.
Anopheles (Anopheles)		26. Cx. restrictor Dyar and Knab	FSR
1. An. franciscanus McCracken	FSR	Culex (Culex)
2. An. freeborni Aitken	Va1	27. Cx. bidens Dyar and Knab	Ma2
3. An. judithae Zavortink a	Va1	28. Cx. chidesteri Dyar	Va3
4. An. pseudopunctipennis Theobald	Ma1	29. Cx. coronator Dyar and Knab	Ma2
5. An. punctipennis (Say)	Va2	30. Cx. declarator Dyar and Knab d	DV
Anopheles (Nyssorhynchus)		31. Cx. erythrothorax Dyar	FSR
6. An. albimanus Wiedemann	Ho	32. Cx. interrogator Dyar and Knab	DV
Aedes (Aedimorphus)		33. Cx. nigripalpus Theobald	DV
7. Ae. vexans (Meigen)	HB	34. Cx. quinquefasciatus Say	Ma1
Aedes (Aztecaedes)		35. Cx. restuans Theobald	DV
8. Ae. ramirezi Vargas and Downs	DV	36. Cx. salinarius Coquillett	DV
Aedes (Georgecriagius)		37. Cx. stigmatosoma Dyar e	DV
9. Ae. epactius Dyar and Knab	DV	38. Cx. tarsalis Coquillett	Ma1
Aedes (Jarnellius)		39. Cx. thriambus Dyar	DV
10. Ae. deserticola Zavortink	FNR	Culex (Melanoconion)
11. Ae. monticola Belkin and McDonald	DV	40. Cx. erraticus (Dyar and Knab)	Ma2
Aedes (Kompia)		Culex (Neoculex)
12. Ae. purpureipes Aitken	DV	41. Cx. apicalis Adams	FSR
Aedes (Lewnielsenius)		42. Cx. arizonensis Bohart	FSR
13. Ae. muelleri Dyar	FSR	43. Cx. reevesi Wirth	DV
Aedes (Ochlerotatus)		44. Cx. territans Walker	DV
14. Ae. taeniorhynchus (Wiedemann)	DV	Culex (Phenacomyia)
15. Ae. trivittatus (Coquillett)	FSR	45. Cx. lactator Dyar and Knab	FSR
Aedes (Protomacleaya)		Culiseta (Culiseta)
16. Ae. burgeri Zavortink	Za	46. Cs. impatiens (Walker)	DV
17. Ae. podographicus Dyar and Knab	HB	47. Cs. incidens (Thomson)	DV
Aedes (Stegomyia)		48. Cs. inornata (Williston)	DV
18. Ae. aegypti (Linneaus)	Va3	49. Cs. particeps (Adams)	DV
Haemagogus (Haemagogus)		Coquillettidia (Coquillettidia)
19. Hg. equinus Theobald	FSR	50. Cq. perturbans (Walker)	Va3
20. Hg. mesodentatus Komp and Kumm	HB	Orthopodomyia
Psorophora (Grabhamia)		51. Or. kummi Edwards	DV
21. Ps. columbiae (Dyar and Knab) b	Ma1	Toxorhynchites (Lynchiella)
22. Ps. signipennis (Coquillett)	DV	52. Tx. moctezuma (Dyar and Knab)	Ma3
Psorophora (Janthinosoma)		53. Tx. septentrionalis (Dyar and Knab)	FNR
23. Ps. longipalpus Randolph and O’Neill	FSR	Uranotaenia (Uranotaenia)
Psorophora (Psorophora)		54. Ur. geometrica Theobald	FSR
24. Ps. ciliata (Fabricius)	DV	55. Ur. lowii Theobald	FSR
25. Ps. howardii Coquillett c	Va3	56. Ur. sapphirina (Osten Sacken)	FSR
Culex (Anoedioporpa)

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