Aedes (Howardina) quadrivittatus (Coquillett).

Both the subgenus Howardina and the species Ae. quadrivittatus are recorded for the first time in Nuevo León. Females were collected approaching humans in La Camotera, Santiago County, in association with Ae. amateuri, Ae. trivittatus, Ae. brelandi, and Wyeomyia n. sp.; biting during the day in association with Ae. muelleri and Ae. lewnielseni; and with Ae. brelandi in oak forest in Santiago County. Immature stages were collected from epiphytic bromeliad axils in Sierra la Camotera, and in La Encantada, Zaragoza County; and from a large tree hole filled with rain water in La Camotera in association with Ae. lewnielseni and Cx. thriambus.

Aedes (Lewnielsenius) muelleri Dyar.

This Nearctic species has been recorded in the United States and Mexico. This is the first record in Nuevo León state for the subgenus and the species. Females were collected biting during the day in La Camotera, Santiago County, in a pine forest in association with Ae. quadrivittatus and Ae. lewnielseni; and immature stages were collected in the same location from a tree hole in association with Cx. restrictor.

Aedes (Ochlerotatus) bimaculatus (Coquillett).

This species was collected in only one location in Dulces Nombres, Pesquería County. One female was collected at night in a CDC light trap (baited with CO₂) located near a pond with abundant emergent and floating vegetation, such as water hyacinths, in a region of shrubs. Associated species were An. pseudopunctipennis, An. quadrimaculatus, An. albimanus, Ae. scapularis, Ae. taeniorhynchus, Ps. ciliata, Cx. coronator, Cx. quinquefasciatus, Cx. erraticus, Ma. titillans, and Ur. lowii.

Aedes (Ochlerotatus) trivittatus (Coquillett).

Although Ae. trivittatus is widely distributed in Mexico, this is the first record for Nuevo León state. Females were collected biting humans during the day in oak forest in La Camotera, Santiago County, in association with Ae. quadrivittatus, Ae. amateuri, Ae. brelandi, and Wyeomyia n. sp.; in an area of shrubs in Mina County in association with Ps. cyanescens; and in caves in Lampazos del Naranjo County. Females were collected at night in Lagunita de Taberna, Mier y Noriega County; and in Ciénega de González, Santiago County, in association with Ae. amateuri, Ae. amabilis, Ae. triseriatus/Ae. brelandi, Ps. ferox, Cs. particeps, and Ur. coatzacoalcos. Specimens were also collected at night in octenol-baited CDC light traps in Lagunita de Taberna; in a CDC light trap without bait in Ciénega de González, in association with Ae. amabilis and Cs. particpes; and in a CDC light trap without octenol located in a shrubby area in Las Labores del Ojo, Mina County, in association with An. pseudopunctipennis, Ps. columbiae, Ps. signipennis, and Ps. cyanescens.

Aedes (Protomacleaya) amabilis Schick.

The presence of Ae. amabilis in Nuevo León is the most unexpected record in this study. Since the species was originally described from females collected in Cueva del Nacimiento del Agua, Veracruz state, Mexico, by Schick [11], it has been recorded only in Tamaulipas state. Females were captured biting collectors at dusk in El Salto, Zaragoza County, and biting at night in Ciénega de González, Santiago County. Associated species were Ae. amateuri, Ae. trivittatus, Ae. triseriatus/brelandi, Ps. ferox, and Cs. particeps. Additional females were collected at night in CDC light traps in the latter location; associated species were Ae. trivittatus and Cs. particeps. All of the collection sites were in oak forest. No immature stages of Ae. amabilis were collected.

Haemagogus (Haemagogus) equinus Theobald.

The genus Haemagogus, subgenus Haemagogus, and Hg. equinus are recorded for the first time in Nuevo León. Females were collected in only one location in Belisario Domínguez, Linares County, during the day in a region of shrubs in association with Ae. aegypti, Ae. albopictus, and Ps. ferox.

Psorophora (Psorophora) cilipes (Fabricius).

The presence of Ps. cilipes in Nuevo León is another unexpected record in the state. This species was collected in only one location in Apodaca County, by aspirating in shaded vegetation in a suburban area near a large pond with abundant emergent vegetation. Dissection of male genitalia was required to identify this species. Associated species were Ae. vexans, Ps. columbiae, Ps. cyanescens, and Ps. ciliata.

Culex (Anoedioporpa) restrictor Dyar and Knab.

This species was collected only in a tree hole filled with rain water in an oak forest in Sierra la Camotera, Santiago County, in association with Ae. muelleri. Culex restrictor has been recorded as far north as Tamaulipas, and this is the first report in Nuevo León state.

Culex (Culex) chidesteri Dyar.

This is one of two species of the subgenus Culex that are new records for Nuevo León state. Immature stages were collected from a ground-level water channel in an oak forest in San Juan Bautista, Santiago County, in association of An. franciscanus, Cx. thriambus, Cx. restuans, and Cs. particeps; and from an artificial container in Belisario Domínguez, Linares County. Adults were collected at night in CDC light traps in Rancho las Flores, Dr. Coss County, in a shrubby area in association with An. pseudopunctipennis and Ps. cyanescens.

Culex (Culex) erythrothorax Dyar.

This is the first report of Cx. erythrothorax in Nuevo León state, where it was found in just one location in Bustamante County. Females were taken at night using CDC light traps in a shrubby area of Bustamante in association with An. pseudopunctipennis, Ae. vexans, Cx. coronator, and Cx. tarsalis; and in the same locality where females were collected biting humans at night in association with Ae. vexans, Ps. signipennis, and Cx. coronator.

Culex (Neoculex) arizonensis Bohart.

This is the first report of Cx. arizonensis and the first record for the subgenus Neoculex in the state. Immature stages were collected from discarded tires with colored water in La Encantada, Zaragoza County, in association with Cx. thriambus and Cs. particeps; from a rock hole with colored water in Lagunita de Taberna, Mier y Noriega County; and from springs in La Camotera, Santiago County, in association with An. pseudopunctipennis and Cs. particeps.

Orthopodomyia kummi (Coquillett).

In Mexico, the genus Orthopodomyia is represented by Or. alba Baker, Or. kummi, and Or. signifera (Coquillett). This is the first record of Or. kummi in Nuevo León state. Females were collected at night in CO₂-baited CDC light traps located in oak forest in Rincon de la Silla, Guadalupe County; adults were found resting in vegetation in La Camotera, Santiago Coiunty; and immatures were collected from a tree hole filled with rain water and abundant litter in oak forest.

Wyeomyia (Wyeomyia) n. sp.

The genus Wyeomyia includes 13 species in Mexico, mostly distributed in the southeastern region of the country where tropical forests are located. Two species occur in the Nearctic forest of northeastern Mexico: Wy. mitchellii (Theobald) is known from Biosphere Reserve El Cielo, in Tamaulipas state, and this undescribed species collected in Nuevo León in La Camotera, Santiago County. Two females of the new species were collected biting humans at dusk in oak forest in association with Ae. quadrivittatus, Ae. amateuri, Ae. trivittatus, and Ae. brelandi. These specimens are in poor condition; hence, this species will be formally described when more speciemens, including immature stages and adult males, are obtained.

Toxorhynchites (Lynchiella) moctezuma (Dyar and Knab).

Prior to this study, species of the genus Toxorhynchites recorded Nuevo León by a number of authors were identified as Tx. rutilus (Coquillett), Tx. theobaldi (Dyar and Knab), and Toxorhynchites sp. Since the identity of the species of Toxorhynchites in northern Mexico is still uncertain, the species recorded here as Tx. moctezuma is based on studies by Zavortink and Chaverri [12]. Toxorhynchites moctezuma is thus a new state record for Nuevo León. Adults were collected resting in vegetation in a shrubby area in Ampliación Nogales, García County, in association with Ae. aegypti and Ae. albopictus; approaching humans in the same location in association with Ae. scapularis and Ae. albopictus; and resting in vegetation in Allende County in association with Cx. coronator and Cx. quinquefasciatus. Immature stages were collected from a water tank with clear water in Rancho la Primavera, Santiago County, in association with Ae. albopictus, Cx. coronator, and Cx. quinquefasciatus; from a discarded tire with colored water in Unidad Piloto, Guadalupe County; from a variety of artificial containers, such as plastic buckets and discarded tires, in Pesquería County in association with Ae. aegypti, Ae. albopictus, Cx. coronator, and Cx. quinquefasciatus; from a tree hole in Bio-Parque Estrella, Montemorelos County, in association with Ae. epactius and Ae. albopictus; from a plastic bucket with colored water in San Juan Bautista, Santiago County, in association with Ae. albopictus, Cx. quinquefasciatus, Cx. thriambus, and Ur. coatzacoalcos; from a discarded tire in Linares County; from a tree hole in a semi-arid area in El Potrero, Villaldama County, in association with Ae. albopictus; and from a plastic bucket at the last location.

Uranotaenia (Uranotaenia) coatzacoalcos Dyar and Knab.

This species is recorded for the first time in Nuevo León state. Immature stages were collected from an artificial container (plastic bucket) filled with rain water and abundant litter that colored the water. The specimens were collected in San Juan Bautista, Santiago County; associated species were Ae. albopictus, Cx. quinquefasciatus, Cx. thriambus, and Tx. moctezuma. Adults were collected approaching humans at night in Ciénega de González, Santiago County, in oak forest in association with Ae. amateuri and Ae. trivittatus.

Aedes (Ochlerotatus) amateuri Ortega and Zavortink n. sp. 4D562DD9-C4E7-4224-9CE6-4E6E12868149.

Type specimens: Holotype: adult female (A♀) without associated larval and pupal exuviae [CC-UL, 01110805-SAN], El Manzano, Santiago, Nuevo León, Mexico (25° 22′ 24.5′′ N 100°13′11.14′′ W), elevation 1,364 m, immature stages collected from a small pond with colored water without aquatic vegetation, 11 Aug 2005, col. A.I. Ortega-Morales. Paratypes: 1A♀, (same data as holotype); 7 fourth-instar larvae [CC-UL, 01090507-CC], 5 fourth-instar larvae [Bohart Museum], 1 larval exuviae with dead pupa [Bohart Museum], 1 pupal exuviae (Pe) with A♀ [Bohart Museum] (1–102), 1 larval exuviae (Le) with Pe, and A♀ (charola 1) [Bohart Museum], Cola de Caballo, Santiago, Nuevo León, Mexico (25° 22′ 5.3′′ N 100° 9′ 29.5′′ W) elevation 556 m, immature stages collected from a small pond, 9 May 2007, col. A.E. Elizondo-Quiroga, A.I. Ortega-Morales; 10A♀ [CC-UL, 01200610-CG], 5A♀ [Bohart Museum], Cienega de González, Nuevo León, Mexico (25° 22′ 39′′ N 100° 14′ 31′′ W), elevation 1,331 m, biting collectors, 20 Jun 2010, col. J. Rodríguez-Rojas; 4A♀ [CC-UL, 02120610-EM], El Milagro, Cruillas, Tamaulipas, Mexico (24° 40′ 41.5′′ N 98° 38′ 2.4′′ W), elevation 210 m, biting collectors, 12 Jun 2010, col. A. Hernández-Hernández, A. Sánchez-Trinidad, F. Ordóñez-Sánchez, O. Mandujano-Grajales; 4A♀ [CC-UL, 07120610-SSC], 2A♀ [Bohart Museum], Sierra de San Carlos, Cruillas, Tamaulipas, Mexico (24° 39′ 32.9′′ N 98° 36′ 46.5′′ W), elevation 210 m, biting collectors, 12 Jun 2010, col. A.I. Ortega-Morales, A. Sánchez-Trinidad, O. Mandujano-Grajales.

Female. Wing: about 3.79–3.84 mm. Proboscis: about 2.0–2.30 mm. Forefemur: about 1.87–1.94 mm. Abdomen: about 3.0–3.3 mm. Integument brown to light brown, thoracic pleura sometimes lighter brown. Head: Eyes moderately separated above antennae, interocular space with several light-colored setae and narrow curved white scales. Occiput with erect scales broad, creamy-white near midline, usually becoming tan or brown laterally. Vertex with narrow curved creamy-white scales along midline and eye margin and broad flat creamy-tan to light brown scales on dorsolateral, lateral and ventral surfaces, the darkest scales usually dorsolateral. Proboscis 1.1–1.2 length of forefemur, dark-scaled. Palpus about 0.2 length of proboscis, with 3 obvious palpomeres, dark-scaled. Antenna subequal to proboscis in length; pedicel usually yellow, darker and with a few light or dark scales mesally; flagellomere 1 slightly swollen, with white scales mesally. Thorax (Fig 3A): Acrostichal setae few, on anterior area of scutum; dorsocentral setae few, on anterior and posterior areas of scutum; prescutellar and supraalar setae numerous; humeral setae 2, 3. Scutellum with 10–12 setae on each lobe. Scutal scales narrow, curved, whitish, dingy white, or cream-colored on anteriorly, around prescutellar space and in antealar area, yellowish in diffuse narrow acrostichal and broader outer dorsocentral lines, light brown to light reddish brown in diffuse inner dorsocentral line, and dark reddish brown laterally from anterior promontory to above wing base. Scutellum with numerous narrow curved cream-colored scales, those on lateral lobe sometimes yellowish to brown. Paratergite without scales (Fig 3B). Antepronotum with setae and brown and/or cream-colored narrow curved scales. Postpronotum with vertical row of setae posteriorly, narrow curved mostly brown scales dorsally, and broad flat white scales posterioventrally. Pleura with light-colored setae on postspiracular area, prealar knob, proepisternum, upper and posterior mesokatepisternum and upper mesepimeron; with broad flat mostly appressed white scales on postspiracular and subspiracular areas, the latter patch with dorsal extension to postpronotum, lower proepisternum, proepisternum, in patch on prealar knob below setae, in upper patch on mesokatepisternum that does not reach anterodorsal corner of sclerite, in lower patch along posterior margin of mesokatepisternum, and in large, somewhat divided patch on upper 0.67 of mesepimeron (Fig 3C). Wing: Completely dark-scaled. Halter: Light-scaled. Legs (Fig 3D): All coxae with broad flat white scales on anterior surface, forecoxa usually with additional dark scales. All femora with white-scaled knee spot also involving base of tibia; fore- and midfemora predominantly dark-scaled, white scales on posterior and ventral surfaces; hindfemur mostly white-scaled with dorsal dark-scaled line beginning near base and broadening distally to become an apical dark-scaled band. Tibiae each with narrow incomplete apical white band. Fore- and midtarsus with white scales in complete narrow band at base of tarsomere 1, in complete to incomplete narrow band over joint between tarsomeres 1 and 2, and usually in inconspicuous incomplete bands over remaining joints or at the base of tarsomeres 3–5; hindtarsus with white scales in conspicuous complete band at base of tarsomere 1, in bands over joints, and completely covering tarsomere 5. Abdomen (Fig 3E): Tergum I entirely or mostly dark scaled dorsally, bare or with a few scales laterally; laterotergite bare. Terga II–VII with white scales in basolateral patches and separated narrow to broad basal bands usually broadest in middle; terga V–VII or VI, VII with light scales at apex. Sterna white scaled, III–VII sometimes with dark-scales in small apicolateral patch.

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Fig 3. Adult female of Aedes (Ochlerotatus) amateuri.

A: Dorsal view of the scutum. B: Laterial view of the body. C: Lateral view of thorax. D: Lateral view of foreleg. E: Dorsal view of abdomen.

https://doi.org/10.1371/journal.pone.0217694.g003

Male. In general, as in female except for sexual characters and scutal ornamentation. Head: Proboscis about 1.15 length forefemur. Palpus long, with 5 palpomeres, 2 and 3 ankylosed and long, 4 about 0.17 and 5 about 0.15 length of palpus; apex of palpomere 3 slightly swollen, upturned and with long setae on underside; palpomeres 4 and 5 usually slightly downturned, 4 with long setae on underside, 5 with shorter setae on underside; dark-scaled except inconspicuous light-scaled patch at base of 5 dorsally. Antenna short, extending to apex of palpomere 3, flagellomeres 12 and 13 comprising 0.5 length of flagellum. Thorax: Scutal scale-pattern not as distinct as in female, light scales more extensive, sometimes without brownish scales in dorsocentral line. Legs: Fore- and midtibiae often with posterior streak of pale scales. Anterior fore- and midclaws greatly enlarged, with blunt submedian tooth and acute basal tooth; posterior fore- and midclaws moderately enlarged, with acute subbasal tooth; hindclaws small, with acute submedian tooth.

Male genitalia (Fig 4A). Segment IX: Tergum well developed, with narrow, deep emargination between prominent tergal lobes, the lobes slightly broader than long, bearing 5, 6 short, strong setae; sternum with 2 setae distally. Gonocoxite: Length about 3.6 times median width; basal tergomesal lobe well developed, bluntly conical, with 1 very strong, bent and recurved dorsal differentiated seta, about 20 weak setae, and 3 or 4 slightly longer and stronger setae along ventral margin; apical tergomesal lobe weakly developed, with short setae; tergal surface with numerous short setae and with 2, 3 long, strong setae distad; lateral surface with numerous long, strong setae and scales; sternal surface with numerous moderate to strong setae; sternomesal margin with long row of 25–30 weak to moderate setae; mesal surface membranous. Claspette: Stem elongate, extending well beyond basal tergomesal lobe, curved dorsad, with tubercle near base of ventral surface bearing single small seta; filament subequal in length to stem, expanded near middle, without retrorse process, with slender, curved, acute apex. Gonostylus: Simple, long, slightly expanded before mid-lenght, curved inward distally, with inconspicuous spicules on dorsal surface; spiniform long, slender, about 10–12 times width, and 0.25 length of gonostylus. Phallosome: Aedeagus small, more or less ovate, with slight apical emargination. Proctiger: Paraproct strongly sclerotized, with single, large, strong, apical tooth; cercal setae small, 3.

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Fig 4. Male genitalia, pupa, and larvae of Aedes (Ochlerotatus) amateuri.

A: General aspect of genitalia. B: Trumpet. C: Metanotum and seta 1-I. D: Paddle with genital lobe (male). E: Head (dorsal view). F: Pecten of siphon.

https://doi.org/10.1371/journal.pone.0217694.g004

Pupa. Abdomen: about 2–5–2.9 mm. Trumpet: about 0.45–0.47 mm. Paddle: about 0.59–0.65 mm. With typical countershading, the dorsal portions in life very light brown, the ventral areas straw-colored to nearly colorless. Cephalothorax: Seta 6-C weakly developed, remaining setae moderately developed, most setae single to few branched or few-forked; setae 8,9-C single or 2-forked; seta 10-C with 6–8 branches. Trumpet (Fig 4B): Uniformly brown, broadening gradually from base to apex, pinna small. Abdomen: Seta 1-II moderately strong, with 13–21 branches, closer to midline than seta 1-I (Fig 4C); seta 1-IV–VII weakly to moderately developed, usually double (single to triple or 2-forked). Seta 2-II inserted laterad of seta 3-II; seta 2-III–VII inserted mesad of seta 1 of corresponding segment. Seta 3-II single; seta 3-III moderately strong, single, shorter than length of tergum IV; seta 3-V–VII usually single (single or double). Seta 5-IV–VI strongly developed, longer than succeeding segment on IV, V, shorter on VI, single, double, or 2-forked; 5-VII weakly developed, shorter than 3-VII, with 2–4 branches. Seta 6-I,II moderately long, fine; seta 6-III–VI weakly to moderately developed, single; seta 6-VII with 4–6 branches. Seta 9-I–VI small, single; seta 9-VII moderately developed, double or triple; seta 9-VIII strongly developed, dendritic, with 6–9 main branches. Terminal segments (Fig 4D): Male genital lobe 1.2 times length of tergum VIII. Paddle: Very lightly pigmented, longer than broad, index 1.1–1.3, apex weakly emarginate. Midrib not extending to apex; external buttress weakly developed, indistinct, very weakly serrated. Seta 1-P long, single.

Larva. Head (Fig 4E): about 1.00–1.05 mm. Siphon: about 0.60–0.64 mm. Saddle: about 0.22–0.28 mm. Head: Broader than long, moderately pigmented, lighter in ocular area, darker posteriorly. Seta 1-C stout. Seta 4-C weakly developed, about 0.25 length of seta 6-C, 2–4 forked, inserted mesocaudad of seta 6-C. Seta 5,6-C single, nearly equidistant from midline of head, inserted laterad of 1-C. Seta 7-C with 4–6 branches. Seta 15-C short, with 2–5 branches. Dorsomentum triangular, with 12–15 teeth on each side of median tooth. Antenna: Short, less than 0.5 length of head, weakly pigmented, slightly darker beyond middle, weakly and sparsely spiculose. Seta 1-A arising near middle of shaft, not extending to apex, with 2–5 branches. Thorax: Without conspicuous pigmentation. Spicules of thoracic integument numerous, weakly pigmented, length 4–6 times basal diameter. Tracheal trunks conspicuously dilated in metathorax. Seta 1–6-P single, setae 2,3-P shorter and weaker than seta 1-P. Seta 7-P usually triple (double, triple). Seta 1-M single, less than 0.5 length seta 5-C. Abdomen: Seta 1-III with 2–4 branches; seta 1-IV,V strongly developed, long, single. Seta 6-I,II usually double (double or triple); seta 6-III–VI single. Seta 8-II single, double. Seta 13-II,VI short, multiple, subequal; seta 13-III–V strongly developed, long, single. Segment VIII: Comb scales small, evenly fringed, about 19–26 in number, in small patch of 2 or 3 irregular rows. Siphon: Moderately pigmented, with darker basal band; short, index 2.2–2.5; acus distinct, attached. Pecten spines teeth dark, 12–18 in number, progressively longer distad, those in distal half of row with 1 large and 2 or more basal denticles; evenly spaced, in straight row extending to middle of siphon (Fig 4F). Seta 1-S inserted slightly distad of and in line with pecten, with 5–8 branches, shorter than basal diameter of siphon, slightly shorter and weaker than seta 5-VIII. Seta 2-S small, inserted near apex of siphon. Accessory dorsolateral setae absent. Anal segment: Saddle completely encircles segment, moderately pigmented, with darker basal band; acus distinct, attached; without caudal marginal spicules. Seta 1-X arising on and shorter than saddle, single. Seta 2-X with 6–9 branches, less than 0.5 length seta 3-X. Seta 3-X single, long. Ventral brush (seta 4-X) strongly developed, with 14–17 seta on strongly developed grid; no precratal setae. Anal papillae pointed, dorsal papillae 2.4–3.0 length of anal saddle, ventral papillae slightly shorter.

Systematics. From its initial recognition by Dyar [13–14] until the present time, the Scapularis Group of Aedes has usually been interpreted to consist only of species with dark, unbanded tarsi. Arnell [15], noted that the male genitalia of the Mexican species Ae. shannoni Vargas and Downs and the South American species Ae. euiris Dyar, Ae. milleri Dyar, Ae. scutellalbum Boshell-Manrique, and Ae. bejaranoi Martínez, Carcavillo & Prosen were similar to those of species in the Scapularis Group. He chose not to include any of these species in the Scapularis Group, because of significant differences in the adults and larvae or because the immatures of these species were unknown to him. Although Ae. amateuri has conspicuously banded tarsi, we assign it to the Scapularis Group without hesitation because that characteristic appears to be the only significant difference from other members of the group.

Females of Ae. amateuri have the following characteristics of the Scapularis Group presented by Arnell [15]: (1) pleural scales white, not silver, (2) paratergite without scales, (3) subspiracular scale-patch present, (4) hypostigial scale-patch absent, and (5) all wing scales dark. Females of Ae. amateuri are distinguished from other species of the Scapularis Group most conspicuously by (1) the presence of white tarsal bands, the hindtarsus having white scales at both the base and apex of the tarsomeres and tarsomere 5 entirely white-scaled (Fig 3D), and also by (2) the shape of the subspiracular scale-patch, which has a prominent dorsal extension toward the postpronotum (Fig 3D). Aedes thelcter Dyar appears to be the only other species in the Scapularis Group with a dorsal extension to the subspiracular scale-patch, but in that species the extension is not as pronounced. Aedes amateuri is very similar to Ae. shannoni Vargas and Downs, the most notorious differences between these species are found in the adult female: the abdominal terga of Ae. amateuri have basal bands, whereas in Ae. shannoni they have small triangular patches on segments 5–6, and the coloration of pleural integument is light brown or golden in Ae. amateuri, whereas is dark-brown or almost black in Ae. shannoni. The color pattern of the scales on the scutum of Ae. amateuri is similar to that of the South American Ae. crinifer (Theobald), and Ae. amateuri runs to that species in Arnell’s identification key if the tarsal bands are ignored.

The male genitalia of Ae. amateuri are characterized by (1) a bluntly conical basal tergomesal lobe of the gonocoxite that bears 1 very strong, bent, and recurved differentiated seta dorsally, about 20 weaker setae, and 3 or 4 slightly longer and stronger setae along its ventral margin, (2) a weakly developed apical tergomesal lobe of the gonocoxite (Fig 4A), and (3) a simple claspette filament without an outer angle, retrorse process, or elongate spicules. The genitalia appear to be inseparable from those specimens of Ae. scapularis (Rondani) and Ae. shannoni that lack a retrorse process on the claspette filament.

Larvae of Ae. amateuri possess all the characteristics considered to be diagnostic of the Scapularis Group by Arnell [15], namely (1) spiculose thoracic and abdominal integument, (2) seta 13-VI short, multiple, and similar to 13-II, (3) comb scales more than 15, in small patch of 2 or 3 irregular rows, (4) saddle complete, and (5) anal papillae without large, darkly pigmented tracheae. Among species of the Scapularis Group, larvae of Ae. amateuri may be distinguished from those of most species by the combination of (1) head setae 5,6-C single (Fig 4E), (2) spicules of thoracic integument numerous, weakly pigmented, length 4–6 times basal diameter, (3) thoracic seta 3-P single, (4) lateral abdominal seta 6-I,II double and 6-III–VI single, (5) abdominal seta 13-III strongly developed, long, single, (6) comb scales evenly fringed, the median spinule not enlarged, and (7) pecten spines evenly spaced, in straight row to mid-lenght of siphon (Fig 4F). The larvae appear to be indistinguishable from those of Ae. euplocampus Dyar and Knab.

In keys to North American Aedes in the subgenus Ochlerotatus [16–18], females of Ae. amateuri key to Ae. canadensis (Theobald), but they differ from that species in numerous ways, perhaps most significantly by lacking complete rows of acrostichal and dorsocentral setae on the scutum and lacking scales on the paratergite and laterotergite. These species also differ in scutal ornamentation: in Ae. amateuri the lateral margins of the scutum are clothed with dark reddish-brown scales that are the darkest scales on the scutum, whereas in Ae. canadensis those scales are matte white to lustrous white, yellowish-white, pale yellow, or pale golden and are among the palest scales on the scutum. In these same publications, larvae of Ae. amateuri key to species of the Scapularis Group, either to Ae. scapularis or Ae. tortilis (Theobald) in Carpenter and LaCasse [16] depending upon how the density and pigmentation of the cuticular spicules are interpreted; to Ae. trivittatus (Coquillett), the only species of the Scapularis Group in Canada, in Wood et al. [17]; and to Ae. scapularis in Darsie and Ward [18].

Aedes amateuri appears as the undescribed species no. 1 in the previously pubished study of mosquitoes from Tamaulipas state [19].

Bionomics. Immature stages of Ae. amateuri were collected from a small pond with colored water but without aquatic vegetation in association with Ae. trivittatus and a Culex species. Adult females were collected in Nuevo León biting collectors at dusk in association with Ae. trivittatus, Ae. amabilis, Ae. brelandi/Ae. triseriatus, Ps. ferox, Cs. particeps, and Ur. coatzacoalcos; biting in Tamaulipas state during daytime in association with Ae. trivittatus, Ae. brelandi, and Hg. equinus; and biting at dusk with Ae. trivittatus, Ae. brelandi, Hg. equinus, and Ps. cyanescens. Since females of Ae. amateuri can be persistent biters of humans, the species may be involved in the transmission of pathogens.

Distribution (Fig 5). Aedes amateuri has been collected in the northeastern Mexican states of Nuevo León (El Manzano and Cienega de González, both locations in Santiago County); and Tamaulipas (El Milagro and Sierra de San Carlos, both locations in Cruillas County). All locations where the species was collected belong to the Grand Folded Sierra of the Sierra Madre Oriental in Nuevo León, and the Sierra de San Carlos in the Coastal Plain of North Gulf in Tamaulipas. Aedes amateuri may occur in the forested regions of northeastern Mexico. Since the immature stages develop in ground pools, the species may be distributed in other regions to the north and northwest, to the states of Coahuila, Chihuahua in Mexico, and in the U.S.A.

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Fig 5. Distribution of Ae. amateuri n. sp. and Ae. lewnielseni n. sp.

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Etymology. This species is named amateuri because it was collected by personnel that were inexperienced in mosquito collections. “Amateur” is a word derived from the Latin “amatum,” which means someone who does an activity for love without being professional.

Aedes (Protomacleaya) lewnielseni Ortega and Zavortink n. sp. AC1CF45C-89F0-4ACB-AC1B-31EE405DB06A.

Types: Holotype: Adult ♀ with associated pupal exuviae (Pe) [CC-UL, 243, 08191008-CC], Sierra la Camotera, Santiago, Nuevo León, Mexico (25° 20′ 4′′ N 100° 4′ 22′′ W), elevation above 1855 m, immature stages collected from large tree hole in living oak tree, 19 Oct 2008, col. A. Hernández-Velázquez, A.I. Ortega-Morales, D.A. González-Villareal, J.A. Díaz-López, J. de la Cruz-Zavala, J.J. Hernández-Rodríguez, R. Altunar-López. Paratypes (same data as holotype): 4♀, 1 adult ♂ [CC-UL], 2♀ [Bohart Museum], 1♂ [Bohart Museum], 5A♂-Pe and genitalia (G♂) (239, 244, 250, 251, 257, 277, 278, 279, 280, 281) [CC-UL], 22 ♀-Pe (240–243, 245–249, 252–256, 258–265) [CC-UL], 6 Pe (216–221) [CC-UL], 5 larval exuviae (Le) with dead pupae (DP) (223–225, 237–238) [CC-UL], 3 dead larvae (Dl) (226, 231, 234) [CC-UL], 4 DP (236) [CC-UL], 1G♂, 1Pe, 2 Dl [Bohart Museum]; 2A♀, [Bohart Museum, 07191008-CC], (same data as holotype), biting collectors.

Female. Wing about 4.30–4.60 mm; proboscis about 2.89–2.98 mm; forefemur about 2.64–2.76 mm; abdomen about 3.1–3.4 mm. Integument dark brown. Head: Erect scales slender, dingy white or tinged with tan, lateral ones sometimes brown; midline and upper eye margin with white narrow curved scales; area laterad of midline with white and black narrow curved scales; side of head with broad white and tan or brown flat scales. Proboscis and palpus black-scaled; palpus 0.20–0.24 length of proboscis. Pedicel brown medially, yellow laterally; flagellomere 1 swollen, with dark scales. Thorax (Fig 6A): Acrostichal and dorsocentral setae numerous, in complete rows, humeral, lateral prescutal and posterior fossal setae few. Scutal scales narrow curved, brown, with white scales in acrostichal line, at sides of prescutellar space, and in very broad lateral line from anterior promontory to wing base that extends mesad to or nearly to dorsocentral setae in fossal area and is continued posteriorly as outer dorsocentral line; dark-scaled areas without scattered light scales. Scutellum with long setae and narrow curved scales on all lobes, scales of midlobe white, sometimes brown laterally. Paratergite with large patch of broad white flat scales; antepronotum with setae and white scales; postpronotum with vertical row of setae posteriorly and complete covering of mostly appressed broad, flat white scales, the upper anterior ones dense, sometimes slightly narrowed and curved. Pleura with setae on postspiracular area, prealar knob, proepisternum, upper and posterior mesokatepisternum and upper mesepimeron; white broad, flat, scales in conspicuous patches on postspiracular, hypostigial and subspiracular areas, proepisternum, metameron, in 3 patches on mesokatepisternum, and in large patch on mesepimeron; lower proepisternal scales (prosternal scales of Zavortink [20]) absent or few in an oblique line (Fig 6C and 6D). Wing: Completely dark-scaled or with a few white scales near base of costa. Halter: White or white- and brown-scaled. Legs: Predominately dark-scaled, white scales on posterior surface of fore- and midfemora, on basal 0.5–0.6 of hindfemur, in large knee spots on mid- and hindleg, on underside of foretarsomere 1, and on basal 0.8–0.9 of midtarsomere 1; hindtarsus dark-scaled. Abdomen: Tergum I with long setae, a few brown and white scales dorsally, and large patch of white broad flat scales on laterotergite; terga and sterna II–VII dark-scaled with basolateral white patches, the terga sometimes with additional basal white scales.

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Fig 6. Adult of Aedes (Protomacleaya) lewnielseni.

A: Dorsal view of the scutum (female). B: Dorsal view of the scutum (male). C: Lateral view of thorax (female). D: Lateral view of body (female).

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Male. Essentially as in female except for sexual characters and scutal ornamentation. Head: Palpus slightly shorter than proboscis, apex downturned, with numerous long setae from apex of palpomere 3 distad. Thorax: Scutum entirely white-scaled (Fig 6B). Wing: Base of vein R entirely dark-scaled. Legs: Hindtarsomere 1 entirely dark-scaled.

Male genitalia (Fig 7A–7C). As described for the Kompi Group by Zavortink [20] and apparently indistinguishable from other species. Gonocoxite: Basal tergomesal area not swollen, with 10–12 slender setae; median sternomesal sclerite very weakly developed; median sternomesal tuft very weakly developed, the setae few and not strongly curved dorsad. Phallosome: Aedeagus relatively short, length 1.5–1.8 times greatest width.

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Fig 7. Male genitalia and pupa of Aedes (Protomacleaya) lewnielseni.

A: General aspect of genitalia. B: Gonostylus and gonostylar claw. C: Claspette. D: Trumpet. E: Metanotum and seta 1-I. F: Paddle with genital lobe (male).

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Larva. Head: about 1.08–1.12 mm. Siphon: about 1.07–1.10 mm. Saddle: about 0.31–0.39 mm. Head (Fig 8A): As broad as long to slightly broader than long; brown, collar darker. Labrum more or less evenly rounded in dorsal aspect. Seta 1-C stout, arising on or near front edge of labrum. Seta 4-C weakly developed, with 2, 3 short branches, much closer to seta 6-C than to midline. Seta 5-C single to triple. Seta 6-C double or triple. Seta 7-C with 2–6 branches. Seta 14-C single. Seta 15-C short, double or triple. Dorsomentum triangular, with 10, 11 teeth on each side of median tooth. Antenna: Short, 0.4 length of head; brown; shaft with a few inconspicuous spicules. Seta 1-A single, extending to or nearly to apex of antenna. Thorax: Epidermal cells bearing brownish pigment granules. Setae 1,5-P double or triple. Seta 1-M with 3–6 branches, the branches wide-spreading. Abdomen: Epidermal cells bearing brownish pigment granules. Seta 1 with branches tapering, wide-spreading; seta 1-I with 4 branches; seta 1-II with 2–4 branches; seta 1-III–VI usually double or triple (single to triple); seta 1-VII strongly developed, long, double. Seta 2-I double or triple; seta 2-II–VI strong, single. Seta 5-I–VI usually with 3–5 branches (3–6), the branches tapering, wide-spreading. Seta 6-I–II with 2–4 branches; seta 6-III–VI double. Seta 12–1 present. Seta with 13 branches tapering, wide-spreading; seta 13-II–V with 3–5 branches; seta 13-VI with 7–9 branches, the branches much shorter and finer than those of 13-V; seta 13-VII usually triple (single to triple). Segment VIII: Branches of seta 1-VIII usually finer than those of seta 1-X. Comb with 26–37 scales in patch of 2, 3 irregular rows, each scale long, tapered, evenly fringed (Fig 8B). Anal segment: Saddle incomplete, extending 0.6–0.7 down lateral surface of segment. Seta 1-X arising on and slighty shorter than saddle, with 3, 4 branches. Seta 2-X strongly developed, with 3–6 branches. Seta 3-X strongly developed, long, single. Ventral brush (seta 4-X) with 6 pairs of setae (5 pair on grid, 1 pair precratal); seta 4a-X long, at least 2.5 length of saddle; setae with 4 branches, 4c-X usually with 3, 4 branches (2–4). Anal papillae slender, tapered distally; dorsal and ventral papillae subequal in length, 2.4–3.0 length of saddle (Fig 8C). Siphon: Brown with darker basal band and lighter apex; index 3.6–4.0; acus distinct, attached. Pecten with 17–18 evenly spaced spines, each with 1 large and several small basal denticles (Fig 8D). Seta 1-S arising beyond pecten, triple.

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Fig 8. Larva of Aedes (Protomacleaya) lewnielseni.

A: Head (dorsal view). B: Comb scales. C: Segment X. D: Pecten of siphon.

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Pupa. Abdomen: about 3.4 mm. Trumpet: about 0.68 mm. Paddle: about 0.98 mm. With typical countershading, parts dorsal in resting position light brown, parts ventral in resting position straw-colored. Setae in general well pigmented and easily seen. Cephalothorax: Seta 4-C strongly developed, single or 2-forked; seta 8-C stronger than seta 9-C, usually single (single to 2-forked); seta 10-C moderately to strongly developed, subequal to or stronger and longer than 12-C. Trumpet (Fig 7D): Brown basally, becoming lighter apically. Abdomen: Seta 1-I strongly developed, longest branches extending nearly to lateral margin of segment, primary branches simple or with few secondary branches; seta 1-II moderately strong, with 4, 5 branches, the branches finer than primary branches of seta 1-I and sometimes forked (Fig 7E); seta 1-III–V weakly to moderately developed, usually double (single to 4-forked). Seta 2-IV–V inserted mesad of seta 1 and caudad of level of seta 4 of corresponding segment; seta 2-VI inserted mesad or laterad of seta 1-VI and cephalad or caudad of level of seta 4-VI. Seta 5-IV–VI strongly developed, extending to or beyond apex of following segment, 5-IV single or double, 5-V–VI single. Seta 9-VII moderately developed, double; 9-VIII strongly developed, with 3, 4 branches. Paddle: Index 1.4–1.5 (Fig 7F).

Systematics. Aedes (Protomacleaya) lewnielseni is a member of the Kompi Group as described and treated by Zavortink [20]. Females of Ae. lewnielseni are distinguished from all six previously described species of the group by having a conspicuous hypostigial patch of white scales (Fig 6C). Males are distinguished by the combination of the following characteristics: (1) scutum entirely white-scaled (Fig 6B), (2) palpus slightly shorter than proboscis, (3) base of hindtarsomere 1 entirely dark-scaled, and (4) remigium of wing entirely dark-scaled. Larvae are distinguished by the combination of (1) the double or triple head seta 6-C (Fig 8A); (2) the attenuate branches of setae 1,13-I–VI; (3) the strong, single seta 2-II–VI; and (4) the long seta 4a-X of the ventral brush (Fig 8C).

In their general habitus females of Ae. lewnielseni are most similar to Ae. burgeri Zavortink, agreeing with that species in having a very broad lateral marginal line of white scales on the scutum that extends mesad to the inner margin of the fossa and in lacking scattered light scales in the dark-scaled areas of the scutum. The six species of the Kompi Group known to Zavortink [20] appeared to belong to three groups of two species each. Aedes lewnielseni cannot be allied with any of those groups, suggesting that they have no lasting taxonomic value.

Bionomics. Immature stages of Ae. lewnielseni were collected from a large tree hole in a living oak tree (Quercus sp.). The immature stages were associated with Ae. quadrivittatus and Cx. thriambus. Adult females were collected biting collectors in association with Ae. quadrivittatus and Ae. muelleri. The possible medical importance is unknown.

Distribution (Fig 5). Aedes lewnielseni has only been found in the Sierra la Camotera, Santiago County, Nuevo León, northeastern Mexico, but may occur in the Grand Folded Sierra of the Sierra Madre Oriental of Coahuila, Nuevo León, and Tamaulipas.

Etymology. This species is dedicated to Lewis T. Nielsen (1920–2014), in recognition of his contributions to the field of mosquito taxonomy, both through the training of students and through his own research on the mosquito faunas of Utah and other western regions, and the ecology and distribution of tree hole and snow pool mosquitoes of the western United States. Lew was, additionally, a friend, colleague, and mentor of one of us (TJZ).

GROUP 2 (Fig 10).

The immature stages of the species that comprise this group inhabit in phytotelmata and artificial containers in tropical forests in the Neotropical Region and extend into northeastern Mexico where most reach their northern limit in Nuevo León and Tamaulipas states: Anopheles eiseni, Ae. quadrivittatus, Ae. amabilis, Hg. equinus, and Cx. restrictor. All members of this group occur in the Grand Folded Sierra of the Sierra Madre Oriental; these mountains comprise part of the Tropical Mesoamerican Corridor, through which all species of this group spread from farther south into northeastern Mexico. “Cumbres de Monterrey” National Park, where wooded habitats and natural regions are conserved, is located in the mountain ranges of the Grand Folded Sierra in Nuevo León. In this region, epiphytic bromeliads, where species such as Ae. quadrivittatus develop, grow on oaks and other trees; and arboricolous species such as An. eiseni, Ae. amabilis, Hg. equinus, and Cx. restrictor develop in tree holes that are filled with rain water. While most of these species reach their northernmost range of distribution in northeastern Mexico, Hg. equinus spreads into Cameron County in Texas [18, 21–22].

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Fig 10. Phytotelmatic and artificial container inhabiting species that spread from the Neotropical Region into northeastern Mexico.

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GROUP 3 (Fig 11).

Ground pool inhabiting species that extend from the Neotropical Region into northeastern Mexico: Psorophora cilipes, Cx. bidens, and Ur. coatzacoalcos. These species usually develop in brightly lit bodies of water with abundant floating and emergent aquatic vegetation. They have spread into Nuevo León from humid, wet, swampy regions southeast of Tamaulipas and Veracruz states. Psorophora cilipes was originally reported in Mexico by Dyar [23]. Vargas [24] reported the species in Mexico State, but later, in the early revision by Díaz-Nájera and Vargas [25], the species was deleted from the list of Mexican species. Considering that this species was collected only once in a single site, we suppose that Ps. cilipes is not well established in Nuevo León state or in northeastern Mexico. Culex bidens has been reported a few times by the CAIM collectors. This species could be common, developing in swamps in suburban regions of several states of Mexico [26], but the species has not been recorded as far as north as the United States. Uranotaenia coatzacoalcos extends from northern South America through southern Mexico into Nuevo León state where it reaches its northernmost limit.

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Fig 11. Pond, swamp, and marsh inhabiting species that spread from the Neotropical Region to northeastern Mexico.

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Species and names deleted from the Nuevo León mosquito fauna.

Some species reported to occur in Nuevo León state in earlier publications are not considered to occur in the state in this study. In this section, the species deleted from the Nuevo León fauna and the reasons for the deletions are given, organized by authors of the articles in which the species were reported. The reasons for deletions of Ae. atropalpus (Coquillett), Cx. peus Speiser, and Cx. reevesi Wirth [25]; Cx. pipiens L. [1]; and Cx. virgultus Theobald [24] are explained in a previous paper on the mosquitoes of Tamaulipas state [19].

Psorophora virescens Dyar and Knab [24]. This species was synonymized with Ps. howardii Coquillett by Dyar [27].

Toxorhynchites rutilus (Coquillett) [1]. Currently, the taxonomy of the Toxorhynchites species that occurs in northeastern Mexico is uncertain. Toxorhynchites rutilus occurs in southeastern Canada and the eastern United States, extending as far as Texas and possibly northeastern Mexico, where it may occur in symaptry with Tx. moctezuma in the Nearctic-Neotropical overlapping areas. Since at this point, we cannot confirm how the populations of both species are distributed, we consider this record to be based on misidentified specimens of Tx. moctezuma.

Toxorhynchites theobaldi (Dyar and Knab) [25]. This species is commonly reported in early records of Toxorhynchites species from Mexico. Since the resurrection of Tx. moctezuma and Tx. hypoptes (Knab) from synonymy with Tx. theobaldi [12], the distribution of Tx. theobaldi is uncertain, but it does not occur in Middle America or Mexico. Mexican records of Tx. theobaldi are most likely based on specimens of Tx. moctezuma.

Deinocerites cancer Theobald [28]. Voucher specimens for this record were found in the IAIM collection. Immature stages were collected from tree holes, mounted on microscope slides and misidentified as De. cancer. We reviewed those microscope slides and identified the specimens as Cx. restrictor.

Misidentifications of mosquitoes deposited in IAIM collection; the corroborated species is included in parenthesis

Anopheles perplexens Ludlow (An. punctipennis (Say)); Aedes melanimon Dyar (Culex coronator Dyar and Knab); Ae. togoi (Theobald) (Ae. epactius Dyar and Knab); Wyeomyia smithii (Coquillett). (Wyeomyia n. sp.).

Species from adjacent regions that may occur in Nuevo León.

Some species of mosquitoes that have not yet been reported from Nuevo León occur in adjacent areas and may occur within the state. Included among these are eight species that have been recorded in the Lower Rio Grande Valley of Texas and Tamaulipas state: Ae. mitchellae (Dyar), Ae. thelcter Dyar, Cx. abominator Dyar and Knab, Cx. territans Walker, Cx. corniger Theobald, Ps. discolor (Coquillett), Ps. mexicana (Bellardi), and Ur. sapphirina (Osten Sacken) [18–19, 29].

Medical importance of the mosquitoes of Nuevo León.

Some pathogens transmitted by mosquitoes have received great attention in recent years from public health authorities and society due to the arboviral diseases recently new recorded in Mexico such as chikungunya in 2014 and Zika in 2015, both of which caused epidemic outbreaks throughout the country. Currently, dengue fever and Zika are the most important diseases transmitted by mosquitoes due to the large number of cases in humans reported by the Secretariat of Health: 17,795 cases of dengue in 2016 and 9,232 cases of Zika since 2015 to July 2017 [30]. The causal agents of both diseases are transmitted by the bites of infected females of Ae. aegypti and/or Ae. albopictus, species that are well established in urban and suburban areas of Nuevo León.

In recent years, West Nile Virus (WNV) has been recognized in birds, horses, and humans in Nuevo León state [31–32]. Some vectors of WNV, such as Cx. quinquefasciatus and Cx. tarsalis, have wide distributions in Nuevo León and their populations are usually very high during the rainy season. Adults of Cx. tarsalis usually stay in dark places in houses during the winter season and become active at the beginning of spring. Other WNV vectors such as Ae. dorsalis, Cx. nigripalpus, Cx. restuans, and Cs. inornata, have restricted distributions in Nuevo León, develop only small population sizes, and are found only during the rainy season in the summer.

Some arboviruses that cause encephalitis, such as the Venezuelan Equine Encephalitis virus (VEEV), Eastern and Western Equine Encephalitis viruses (EEEV) (WEEV), St. Luis Encephalitis virus (SLEV), and LaCrosse Encephalitis virus (LCV) have not been reported recently or ever in Nuevo León. This does not mean these viruses are absent from the state because those and other arboviruses could be circulating actively in zoonotic cycles in wild areas of Nuevo León. Legislation for surveillance of diseases caused by vector-borne pathogens in Mexico is limited to the search and diagnosis of a few diseases, such as dengue fever, chikungunya, Zika, West Nile fever, and malaria [33–34], neglecting diseases caused by other pathogens that are surely circulating in Mexico, including Nuevo León state. Some vectors of encephalitis viruses occurring in Nuevo León are: (VEEV) Ae. scapularis, Ae. taeniorhynchus, Ps. ferox, Ps. columbiae, Ma. titillans; (EEEV) Ae. taeniorhynchus, Cx. nigripalpus, Cs. melanura; (WEEV) Ae. dorsalis, Cx. quinquefasciatus, Cx. tarsalis, Cs. inornata; (SLEV) Cx. nigripalpus, Cx. quinquefasciatus, Cx. restuans, Cx. tarsalis; (LCV) Ae. triseriatus.

In recent years, cases of yellow fever have been reported in some South American countries [35]. This disease occurs in two forms: urban yellow fever for which the virus is transmitted to humans by biting females of Ae. aegypti and Ae. albopictus, and sylvan yellow fever for which the virus is transmitted to humans by biting females of Sabethes and Haemagogus species. In Nuevo León, Ae. aegypti and Ae. albopictus are well established in urban and suburban areas, whereas Hg. equinus spreads through the woodlands of the Coastal Plains of North Gulf. The presence of these species represents an epidemiological risk for yellow fever, in case of this disease is reported in Mexico.

Although there have been no reports of cases of malaria in recent years in Nuevo León, vectors of Plasmodium species that cause human malaria occur in the state: An. pseudopunctipennis and An. albimanus are the main vectors of Plasmodium in the tropical regions of Mexico, whereas An. quadrimaculatus has been involved as a malaria vector in the United States. A summary of the species of mosquitoes of medical and veterinary importance in Nuevo León and the diseases caused by the pathogens that they transmit is given in Table 4.

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Table 4. Medical and veterinary importance of the mosquitoes of Nuevo León state.

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