Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

Species diversity and spatial distribution of mosquitoes (Diptera: Culicidae)

from La Isla Amazon Park, Napo Province, Ecuador

Linda Jacome1*; https://orcid.org/0000-0003-4025-3769

Jonathan Liria2; https://orcid.org/0000-0003-1611-8364

Richard C. Wilkerson3; https://orcid.org/0000-0001-6366-1357

1. Ingeniería en Ecosistemas, Facultad de Ciencias de la Vida, Universidad Regional Amazónica Ikiam, Napo, Ecuador;

linda.jacome@est.ikiam.edu.ec (*Correspondence)

2. Grupo de Investigación en Población y Ambiente, Universidad Regional Amazónica Ikiam, Napo, Ecuador;

jonathan.liria@ikiam.edu.ec

3. Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington DC, United

States of America; wilkersonr@si.edu

Received 18-V-2023. Corrected 09-VII-2023. Accepted 09-X-2023.

ABSTRACT

Introduction: Vector-borne diseases are prevalent in the Amazon and Coastal regions of Ecuador. However,

there is a scarcity of mosquito ecology studies in these areas. The most recent list of species reported for the

country comprises 8 tribes, 22 genera, and 200 species.

Objectives: To document the Culicidae species found in La Isla Amazon Park, Napo, Ecuador, including those

with epidemiological significance; and to analyze their composition, abundance, and diversity, focusing on larval

habitats during the dry and rainy periods.

Methods: We evaluated different larval habitats, considering collection duration as the primary criterion. We

used CDC and Shannon traps to collect adult mosquitoes during both rainy and dry periods. To assess sampling

effort, we used accumulation curves and non-parametric estimators of species richness, while we employed Hill

numbers to determine diversity. Additionally, we used the Berger-Parker and Pielou indices to evaluate species

dominance and evenness. We conducted cluster analysis and ANOSIM tests to assess the similarity between

habitats and the differences in taxonomic composition between periods.

Results: We collected a total of 802 individuals from 15 species and 4 taxonomic units, 5 genera, and 4 tribes.

Notably, this may be the first records of Wyeomyia felicia Dyar & Núñez Tovar and Culex derivator Dyar & Knab

from Ecuador. Additionally, the presence of Culex dunni Dyar and Psorophora ferox von Humboldt (both recog-

nized as vectors) was correlated with increased rainfall.

Conclusions: The abundance of mosquitoes, including potential vector species, increased during the rainy sea-

son, indicating a higher risk of pathogen transmission. However, the relationship between rainfall amount and

diversity patterns is not well-defined.

Key words: Phytotelmata; Sabethini; Culicini; recreational park; urban green area.

RESUMEN

Diversidad de especies y distribución espacial de mosquitos (Diptera: Culicidae)

del Parque Amazónico La Isla, provincia de Napo, Ecuador

Introducción: Las enfermedades vectoriales son prevalentes en las regiones amazónica y costera de Ecuador. Sin

embargo, hay una escasez de estudios de ecología de mosquitos en estas áreas. En el país se ha reportado 8 tribus,

22 géneros y 200 especies.

https://doi.org/10.15517/rev.biol.trop..v71i1.55184

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

INTRODUCTION

Mosquitoes (Diptera: Culicidae) are vec-

tors of pathogens that include viruses (arbovi-

ruses), filarial worms (helminths) and protozoa

that are significant causes of morbidity and

mortality worldwide, particularly in tropical

and subtropical countries (Bueno-Marí et al.,

2015). During the COVID-19 pandemic, the

significant surge in dengue cases across the

Americas emphasized the critical role of mos-

quitoes in public health (OPS/OMS, 2020).

Culicidae is diverse, currently with 3 718

species classified into two subfamilies. Anoph-

elinae comprised of three genera, Culicinae

divided into 11 tribes, encompassing 113 gen-

era (Harbach & Wilkerson, 2023). In Ecuador,

there are recorded species from both subfami-

lies, eight tribes, 22 genera, totaling 200 species

(Ponce et al., 2021; Ramón et al., 2019).

Although vector-borne diseases are preva-

lent in the Amazonian and Coastal regions of

Ecuador, there is a scarcity of ecological studies

focused on Culicidae, and the majority of the

existing research is relatively recent (Duque et

al., 2019; Navarro et al., 2015). According to

the Ministerio de Salud Pública del Ecuador

(2020), Ministerio de Salud Pública del Ecua-

dor (2021) and Ministerio de Salud Pública del

Ecuador (2022), Ecuador has repofigrted cases

of dengue, Zika, chikungunya, Mayaro, yellow

fever, and malaria. The number of dengue cases

has shown an upward trend, with 8 416 cases

reported in 2019, followed by 16 570 and 20 593

cases in 2020 and 2021, respectively. In terms of

dengue incidence, Napo province ranked third

in the Ecuadorian Amazon region in 2019.

However, there was a surge in cases in 2021,

surpassing all other provinces and recording

the highest number of dengue cases that year.

While malaria cases have also witnessed an

increase, they remain primarily concentrated in

other provinces within the Amazonian region.

Community and adventure tourism in

Napo has increased over the years, because of

the many existing attractions, high biodiversity,

and extensive protected areas. La Isla Amazon

Park (PALI, called after its Spanish initials) is an

urban and touristic green area for recreational

purposes and serves as a habitat for various

vertebrate species. PALI is essential for tourist

and ecological interests, as well as for public

health because a large flow of people and the

presence of possible vectors increases the prob-

ability and risk of disease transmission (Duque

et al., 2019).

Effective understanding and prevention of

arthropod-borne diseases hinges upon a com-

prehensive understanding of species composi-

tion, distribution, and ecology (Ceretti-Junior

Objetivos: Documentar las especies de Culicidae encontradas en el Parque Amazónico La Isla, Napo, Ecuador,

incluyendo aquellas con importancia epidemiológica; y analizar su composición, abundancia y diversidad, enfo-

cándose en los hábitats de las larvas durante los períodos seco y lluvioso.

Métodos: Evaluamos diferentes hábitats larvarios, con la duración de la recolecta como criterio. Las trampas

CDC y Shannon recolectaron mosquitos adultos durante los períodos seco y lluvioso. Evaluamos la riqueza de

especies con curvas de acumulación y estimadores no paramétricos, mientras que determinamos la diversidad

con los números de Hill. Además, utilizamos los índices de Berger-Parker y Pielou para evaluar la dominancia y la

uniformidad de las especies. Realizamos análisis de conglomerados y la prueba ANOSIM para evaluar la similitud

entre hábitats y estaciones, así como las diferencias en la composición taxonómica, respectivamente.

Resultados: Recolectamos un total de 802 individuos de 15 especies y 4 unidades taxonómicas, 5 géneros y 4

tribus. Este podría ser el primer registro de Wyeomyia felicia Dyar & Núñez Tovar y Culex derivator Dyar & Knab

en Ecuador. Además, la presencia de Culex dunni Dyar y Psorophora ferox von Humboldt (ambos potenciales

vectores) se correlacionó con el aumento de las precipitaciones.

Conclusiones: El aumento de la abundancia de mosquitos durante el periodo lluvioso indica un mayor riesgo de

transmisión de patógenos. Sin embargo, la relación entre la cantidad de precipitaciones y los patrones de diver-

sidad no está bien definida.

Palabras clave: área urbana verde; Culicini; Fitotelma; Sabethini; parque recreacional.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

et al., 2016). This study aimed to document

the Culicidae species found in La Isla Amazon

Park (PALI), Napo, Ecuador, including those

with epidemiological significance; and to ana-

lyze their composition, abundance, and diver-

sity, focusing on larval habitats during dry and

rainy periods.

MATERIALS AND METHODS

Study site: The confluence of the Pano

and Tena rivers forms a peninsula known as

PALI in Tena, Napo province (0°59’44.45’’ S &

77°49’7.31’’ W; 0°59’49.39’’ S & 77°49’2.053’’

W; 0°59’49.27’’ S & 77°49’5.92’’ W; 0°59’34.52’’

S & 77°48’55.92’’ W). The Colonso Chalupas

Biological Reserve (RBCC), created in 2014, is

the closest protected area to PALI (Fig. 1). PALI

spans 24 hectares, featuring a native species

forest and a natural wetland; eight hectares are

designated for tourism, including four kilome-

ters of trails used in this study. Despite multiple

floods over the years, the park continues to be

inhabited by many species that were donated

to it in 1995, when it was opened. The park

is in a humid, tropical zone with abundant

bimodal rainfall (peaks tend to occur in June

and November), high relative humidity (> 85

%), and 22.8 °C mean annual temperature

(Lucas-Solis et al., 2021).

Ethics Statement: The authorization

for the Collection of Specimens of Biological

Diversity for Non-Commercial Purposes No.

1176 (MAAE-ARSFC-2021-1176) issued by the

Ministerio del Ambiente, Agua y Transición

Ecológica of Ecuador was processed.

The collected specimens were deposited

in the invertebrate wet section of the Museo de

Ciencias Naturalesat the Instituto Nacional de

Biodiversidad (INABIO), from catalog number

MECN-MA-8603 to MECN-MA-8620.

Culicidae collection: Collections of mos-

quitoes were carried out for immature and

adult stages during a rainy (26-28/07/2021)

and dry (29-31/03/2021) period. Immature

sampling points were plotted over the area

Fig. 1. Location map showing PALI sampling patches and relative position in Napo, Ecuador. Empty pins indicate the dry

period, and filled pins indicate the rainy period.

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

designated for tourism development. Sampling

was standardized by using collection duration

as the main criterion (30 min per patch) and

the number of sampling points was determined

by the availability of breeding sites along the

trail. Immatures were sampled in phytotelmata,

as well as in ponds and artificial containers.

For large water bodies, the systematic scoop-

ing technique was used; the pond margin was

subdivided into subsampling points. In other

words, the 30 min were divided by 5 min, which

means there were 6 subsampling points.

Thus, distance between points depended

on the diameter of the pond (Urbinatti et al.,

2001). The specimens were collected using a

suction pipette or dipper, depending on the

type of larval habitat, and temporarily stored

in sterilized sampling bags. Some immatures

were deposited in individual breeding vials for

development, and the remainder preserved in

cryovials with a 10 % formaldehyde solution.

The collected specimens were deposited in

the invertebrate wet section of the Museo de

Ciencias Naturales at the Instituto Nacional de

Biodiversidad (INABIO), from catalog number

MECN-MA-8603 to MECN-MA-8620.

Traps baited with dry ice (CO2) and UV

light were used to sample adult mosquitoes:

CDC trap and a modified Shannon trap, which

do not use animals as bait (Silver, 2008). The

CDC trap remained active for 14 h (18:00-

08:00) and Shannon traps remained active for

4 h (18:00-22:00). Periodic trap sampling was

carried out during that time, checking the out-

side and inside of the trap every 15 min for 5

min. Mosquitoes were collected using an insect

aspirator, sacrificed through low-temperature

exposure, mounted on entomological pins, and

stored in entomological boxes treated with

naphthalene and silica gel.

An AmScope trinocular stereo microscope

with white light and 90X magnification was

used for species recognition based on available

keys and taxonomic revisions (Forattini, 1965;

González & Darsie Jr., 1996; Lane, 1953a; Lane,

1953b; Liria & Navarro, 2003; Sallum Mureb

& Forattini, 1996; Valencia, 1973; Zavortink,

1972), and the interactive identification keys

of Walter Reed Biosystematics Unit (2021).

The genera and subgenera abbreviations follow

Wilkerson et al. (2015).

Data analysis: Sampling effort was

assessed using species accumulation curves and

nonparametric estimators of species richness:

Abundance-based Coverage Estimator (ACE),

Chao1, Jackknife 1, and Bootstrap (Colwell

et al., 2004). Randomizations (1 000) without

replacement and a 95 % confidence interval

were used. The upper limit of abundance for

rare or infrequent species used in the ACE sta-

tistic was 7, based on the categories proposed

by Friebe according to Lira-Vieira et al. (2013).

It is considered rare when D < 1 %, whereas D

% = (total number of individuals of the species

/ total number of individuals captured) * 100.

Diversity was estimated through Hill num-

bers (Jost & González-Oreja, 2012), where q is

the order of diversity (qD) and determines how

influential common species or rare species are

in the measurement. The first three Hill num-

bers, q = 0 (richness), q = 1 (Shannon’s expo-

nential diversity), and q = 2 (Simpson’s inverse

diversity), were assessed to measure the effec-

tive number of species, an equivalent measure

of true diversity.

The Berger-Parker index and the Pielou

index were calculated to quantify the species

dominance and evenness component between

seasonal periods. The similarity between

habitats was assessed with a cluster analysis

(UPGMA) dendrogram, using the Bray-Curtis

distance matrix.

The differences in taxonomic composition

were assessed using the analysis of similarity

(ANOSIM) test with the Bray-Curtis distance

and 1 000 permutations, performing pairwise

comparisons using the Bonferroni correction.

ANOSIM is based on comparing distances

between groups with distances within groups,

and the resulting R-test statistic measures

whether there is a separation of community

structure, and the p-value determines signifi-

cant comparisons (Clarke, 1993).

The statistics were run in EstimateS Pro-

gram version 9.1.0 (Colwell, 2022), PAST

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

version 4.09 (Hammer, 2022), and Microsoft

Excel (Microsoft Corporation, 2016).

RESULTS

In total, 802 specimens (immatures and

adults) were collected, identified, and grouped

into 15 species and 3 taxonomic units: 10

subgenera, 5 genera, 4 tribes (Culicini: Culex,

Sabethini: Limatus and Wyeomyia, Aedini: Pso-

rophora, Toxorhynchitini: Toxorhynchites) from

7 types of larval habitats (Table 1, Table 2). Dur-

ing the dry period, 7 species and 3 taxonomic

units were recorded, compared to 13 species

and 2 taxonomic units for the rainy period.

Wyeomyia ulocoma (Theobald, 1903) was

the most abundant immature species, with 48

% of the total abundance recorded for the dry

period and 81 % in the rainy period, followed

by Wy. medioalbipes (Lutz, 1904) (28 %) during

the dry period and Culex urichii (Coquillett,

1906) (6 %) during the rainy period.

Both periods evaluated 12 breeding sites

corresponding to 7 types. Both periods have

four types of immature habitats (artificial, bro-

meliad, bamboo, and Heliconia spp.) in com-

mon. However, Calathea sp. (near Calathea

crotalifera) as a habitat was only reported dur-

ing the dry period, and tree holes and ponds

were only reported during the rainy period.

From the CDC and Shannon traps, 61

adult mosquitoes of 4 species and 3 taxonomic

units, 4 subgenera, 3 genera, and 3 tribes were

collected (Table 2). The Shannon trap was

Tabl e 1

Abundance and diversity of immature species by habitat and collection period.

Habitat SpeciesaAbundance Sbexp(H)c1/DddeJf

DRY PERIOD 253 7 4 3 0.48 0.69

Artificial Culex (Car.) urichii 24

3 2 2 0.59 0.82Limatus durhamii 13

Limatus sp. 1 4

Bamboo Culex (Car.) secundus 4

3 2 2 0.65 0.81Culex (Car.) urichii 15

Wyeomyia (Wyo.) melanopus 4

Bromeliad Wyeomyia (Wyo.) medioalbipes 67 1 1 1 1 -

Calathea Wyeomyia (Dec.) ulocoma 27 1 1 1 1 -

Heliconia Wyeomyia (Dec.) ulocoma 95 1 1 1 1 -

RAINY PERIOD 488 11 2 1 0.81 0.35

Artificial Culex (Car.) urichii 8 1 1 1 1 -

Bamboo Culex (Car.) secundus 18

4 3 2 0.48 0.69

Culex (Car.) urichii 19

Culex (Cux.) dolosus 2

Toxorhynchites (Lyn.) theobaldi 1

Bromeliad Culex (Mcx.) imitator 12 1 1 0.93 0.35

Wyeomyia (Wyo.) medioalbipes 14

Heliconia Wyeomyia (Dec.) felicia 12 2 1 1 0.97 0.19

Wyeomyia (Dec.) ulocoma 397

Pond Culex (Mel.) dunni 92 2 1 0.82 0.68

Culex (Ncx.) derivator 2

Tree hole Wyeomyia (Wyo.) celaenocephala 5 1 1 1 1 -

a: subgenera: (Car.) Carrollia Lutz, (Wyo.) Wyeomyia Theobald, (Dec.) Decamyia Dyar, (Ncx.) Neoculex Dyar, (Cux.) Culex

Linnaeus, (Mel.) Melanoconion Theobald, (Mcx.) Microculex Theobald, (Lyn.) Lynchiella Lahille. b: richness (0D); c: Shannon

diversity index (1D); d: Simpson’s diversity index (2D); e: Berger-Parker dominance index; f: Pielou’s equality index.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

more efficient than the CDC trap in recording

abundance during the dry (58 %) and rainy (69

%) periods. Psorophora dimidiata (Cerqueira,

1943) was the most abundant adult species,

with 42 % of total abundance recorded for the

dry period and 61 % in the rainy period.

The diversity of the dry period in both

immature and adult stage, when considering

all species with a weight proportional to their

abundance (Hill number 1) and giving more

weight to dominant species (Hill number 2)

showed a slight increase with four equally

common species and three effective species,

respectably (Table 1, Table 2). During the rainy

period, the Berger-Parker and Pielou indices

indicate that there was a greater dominance of

certain species and less uniformity in the abun-

dance of species, corroborating that rare species

were mainly recorded during this period.

Diversity by larval habitat showed bam-

boo as the most diverse of natural origin in

both periods. Although artificial and bamboo

exhibited the same diversity during the dry

period, bamboo showed higher diversity and

uniformity in the data (Table 1). The CDC trap

exhibited the highest diversity during the rainy

season. However, both traps displayed similar

diversity levels during the dry period (Table 2).

According to the rarefaction curves sam-

pling sufficiency was not reached since the

cumulative number of species did not reach

an asymptote at the end of both collection

periods (Fig. 2). However, ACE and Chao 1

non-parametric estimators reached 100 % cov-

erage in the dry period (Table 2), and Chao 1

reached 100 % coverage during the rainy period

(Table 3). Although species richness was higher

during the rainy period, confidence intervals

of both periods are overlapping, meaning they

do not differ significantly (Colwell et al., 2004).

Non-parametric estimators suggested that

the estimated richness was: 8.01 ± 1.31 species

for the dry period and 13.87 ± 2.57 species for

the rainy period. The sampling effort recorded

most of the richness since sampling complete-

ness exceeded 75 % (Table 3) in both periods.

Tabl e 2

Abundance and diversity of adult species by trap and collection period.

Trap SpeciesaAbundance Sbexp(H)c1/DddeJf

DRY PERIOD 12 4 4 3 0.42 0.91

CDC Culex (Cux.) sp. 1 2 3 3 3 0.4 0.96

Wyeomyia (Dec.) ulocoma 1

Psorophora (Gra.) dimidiata 2

Shannon Culex (Cux.) sp. 1 1 3 3 3 0.43 0.91

Culex (Cux.) sp. 2 3

Psorophora (Gra.) dimidiata 3

RAINY PERIOD 49 5 3 2 0.61 0.73

CDC Culex (Mel.) dunni 3 5 4 4 0.4 0.91

Culex (Cux.) sp. 2 3

Culex (Cux.) sp. 3 1

Psorophora (Gra.) dimidiata 6

Psorophora (Jan.) ferox 2

Shannon Culex (Mel.) dunni 4 4 2 2 0.71 0.66

Culex (Cux.) sp. 3 2

Psorophora (Gra.) dimidiata 24

Psorophora (Jan.) ferox 4

a: subgenera: (Dec.) Decamyia Dyar, (Cux.) Culex Linnaeus, (Mel.) Melanoconion Theobald, (Gra.) Grabhamia Theobald,

(Jan.) Janthinosoma Lynch Arribálzaga. b: richness (0D); c: Shannon diversity index (1D); d: Simpson’s diversity index (2D);

e: Berger-Parker dominance index; f: Pielou’s equality index.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

Cluster analysis produced well-supported

branches with a high cophenetic correlation of

0.99 (Fig. 3). Two groups of larval habitats were

formed based on their taxonomic composition:

1) Calathea sp. and Heliconia spp. inflorescenc-

es, 2) artificial and Bamboo cavities.

ANOSIM indicated no significant differ-

ences in taxonomic composition of the breed-

ing sites between the dry and rainy periods

(R = -0.115, P > 0.05). The negative value of R

suggested greater dissimilarity within periods

than between them (Chapman & Underwood,

1999). In other words, the taxonomic composi-

tion differs between larval habitat types rather

than between periods.

DISCUSSION

No previous studies have been conduct-

ed on mosquito diversity and distribution

Fig. 2. Accumulation curves of immatures per sample. Dashed lines show their confidence interval bounds for each period.

Tabl e 3

Observed and expected number of immature species from

both periods.

Non-parametric estimators

of richness

Periods

Dry Rainy

ACE 7 13

Chao1 7 11.33

Jack1 9.75 17.42

Bootstrap 8.3 13.73

Observed species 7 11

Meana8.01 13.87

SDb1.31 2.57

%c87.36 79.31

a: mean of the estimated richness among the 4 non-

parametric estimators; b: standard deviation; c: measure of

sampling completeness = observed species divided / mean

of the estimators * 100.

Fig. 3. Cluster analysis (UPGMA) dendrogram of larval

habitats in the dry and rainy periods, based on the Bray-

Curtis distance matrix.

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

of breeding sites in urban parks in Ecuador.

PALI shares a similar species richness to Chico

Mendes Ecological Park and Carmo Park in

São Paulo, Brazil (Bicudo de Paula et al., 2015).

The three parks are similar in size, but the

parks in Brazil had a higher abundance of

individuals due to the longer collection period

and greater availability of breeding sites. This

highlighted the richness found in PALI since a

similar number of species were found for fewer

individuals sampled.

During the rainy period almost twice as

many immature individuals and a slightly high-

er species richness were recorded, although

not enough to be significantly different from

the dry period. Nevertheless, diversity was

slightly higher during the dry period due to

the disproportionate increase of Wy. ulocoma

immatures and Ps. dimidiata adults during the

rainy period.

Although ACE and Chao1 estimators

resulted in 100 % sampling completeness, and

the mean sampling completeness exceeded 75

%, the rarefaction curves did not reach sam-

pling sufficiency. The literature states that in

invertebrate studies, such as arthropods, mainly

in tropical areas, asymptotes may never be

reached because these are taxon-rich groups

(Gotelli & Colwell, 2001).

The Calathea sp. and tree hole patch

occurred once. Consequently, Wy. celaeno-

cephala (Dyar & Knab, 1906) was only found

in that single patch. For aesthetic reasons, the

park is constantly being landscaped, cleared,

and cleaned; therefore, the Calathea sp. patch

was removed, and the bromeliad patches were

reduced after the dry period. In other words,

there is a strong human influence on the envi-

ronment, which certainly modifies the distri-

bution patterns of species. And, regarding the

availability of breeding sites between periods,

it is premature to conclude that it was the same

since anthropogenic activity played a signifi-

cant role. However, the occurrence of the tree

hole patch during the rainy season may be due

to a meteorological factor.

The species in this study were mostly

specialized in a specific larval habitat type,

except for Wy. Ulocoma and Cx. urichii. In fact,

the cluster analysis revealed two groups with

shared species: 1) inflorescences of Calathea sp.

and Heliconia spp. with Wy. ulocoma, although

the disproportionate abundance of Wy. ulo-

coma in Heliconia generated low similarity; 2)

artificial and Bamboo cavities with Cx. urichii.

However, the tree hole, ponds, and bromeliads

had distinct compositions and did not colonize

other habitat types.

Wyeomyia ulocoma, Wy. medioalbipes,

Wyeomyia felicia (Dyar & Nuñez Tovar, 1927),

Culex imitator (Theobald, 1903), Culex dolo-

sus (Lynch Arribálzaga, 1891), Culex secundus

(Bonne-Wepster & Bonne, 1920) were found

as established in the literature (Chaverri et al.,

2018; Frank & Curtis, 1981; Lane & Cerqueira,

1942; Medeiros-Sousa et al., 2015; Navarro et

al., 2007; Valencia, 1973).

Wyeomyia celaenocephala was found in a

tree hole, which has not been described as a

breeding site for this species in particular, but

for its subgenus (Frank & Curtis, 1981; Lane &

Cerqueira, 1942; Navarro et al., 2015). Likewise,

Wyeomyia melanopus (Dyar, 1919) was found

in bamboo stumps, a larval habitat not previ-

ously reported for this species, although it has

been for other members of the subgenus (Lane,

1953b; Lane & Cerqueira, 1942).

Culex urichii was found in bamboo stumps

and artificial containers. This aligns with lit-

erature describing the species in a wide vari-

ety of larval habitats (Feijó Almeida et al.,

2020; Liria & Navarro, 2003). Records of Cx.

urichii have been found in the husks of white

cacao (Theobroma grandiflorum) (Berti et al.,

2013) and bacao (Theobroma bicolor) (Booth,

2018). While both Theobroma species have

been described in Ecuador (Duque et al., 2019;

Ponce et al., 2021), there are no records linking

them to Culicidae.

Culex dunni (Dyar, 1918) and Cx. derivator

(Dyar & Knab, 1906) were only found during

the rainy period in the “moretal”, a swampy and

permanently flooded area typical of the Ama-

zon region, characterized by the predominant

growth of the morete palm tree (Mauritia flex-

uosa). Supporting the description of Cx. dunni

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

being collected from natural ponds (Bang-

her, 2020), and subgenus Neoculex inhabiting

groundwater bodies (Harbach, 2009).

Toxorhynchites theobaldi (Dyar &

Knab,1906) was found in a bamboo stump,

indicating its presence in natural larval habitats.

However, it has also been documented in tires,

plastic, and glass containers (Ceretti-Junior et

al., 2016). Toxorhynchites is a genus of great

interest in biological control studies, because

are predators of other immature species, and as

adults are not hematophagous.

Limatus durhamii (Theobald, 1901) was

found in artificial containers, but there are

records of the species in natural breeding sites

as well (Chaverri et al., 2018; Feijó Almeida et

al., 2020; Navarro et al., 2007) and has been

found with a high rate of prevalence in artificial

larval habitats along with Aedes (Talaga, 2016).

Toxorhynchites theobaldi and Li. durhamii

are ecological regulators of immatures popu-

lations and their predator efficiency in PALI

is unknown. Therefore, it is not ruled out

that both species may have contributed to

the occurrence of rare and occasional species

(low abundance).

Psorophora ferox (von Humboldt, 1819)

and Ps. dimidiata were only collected by CDC

and Shannon traps. Psorophora lay their eggs

on damp or dry mud where they withstand

long periods (months or years) of desiccation

and hatch when the habitat is inundated by

rain or flood waters (Foster & Walker, 2019).

This could potentially explain the higher abun-

dance of both Psorophora species during the

rainy period.

There is medical importance associated

with a few species in PALI. Several strains of

Ilheus and Venezuelan encephalitis viruses have

been isolated from Wy. medioalbipes (Wilker-

son et al., 2021), but little is known about its

medical importance. In Ecuador, the species

may be of interest because it was described in

Ananas comosus (Navarro et al., 2018), there-

fore crops can serve as large reservoirs. In fact,

Wyeomyia is considered to be underestimated

since several pathogens have been recovered

from or experimentally passed through species

of the genus (Walter Reed Biosystematics Unit,

2023). However, due to problematical identifi-

cation, there has been no arboviral surveillance.

Limatus durhamii is recognized as a vector of

VEE virus, Caraparu virus, and Guama virus

(Feijó Almeida et al., 2020; Navarro et al.,

2015). Psorophora ferox is a vector for Ilheus

virus and VEE virus. Also, West Nile virus

and Cache, Oriboca, and St Louis encephalitis

viruses have been isolated from this species

(Wilkerson et al., 2015). Finally, Culex dunni

has been reported as a vector of Venezuelan

Equine Encephalitis (VEE) (Berti et al., 2013).

Some mosquitoes are known to be able to

travel long distances, usually with the help of air

currents. In the spread of the Japanese encepha-

litis virus to Australia, a significant role was

played by wind dispersal (Ritchie & Rochester,

2001). Also, in parts of Africa, where surface

water is absent for months, malaria persistence

is associated with windborne long-distance

migration of mosquitoes (Huestis et al., 2019).

Mosquitoes may be able to cross the river,

but it will depend on their flight capacity. Mos-

quitoes generally have a reduced flight range as

wind velocity increases (Verdonschot & Besse-

Lototskaya, 2014). In general, the flight capacity

of Culicidae remains unknown and is likely to

vary among species. Information is only avail-

able for one of the species recorded in PALI,

Ps. ferox, which has excellent dispersal ability

(2 500 m of average maximum flight distance),

stronger than the most common urban vector

species, Aedes aegypti and Ae. albopictus (333

and 676 m, respectively) (Verdonschot & Besse-

Lototskaya, 2014).

Psorophora ferox as a vector species could

be of concern, as it is more likely to colonize the

surroundings of the park than domestic species.

But it is known that the migration process is

far more complex because it depends not only

on flight capacity but also on the prevalence of

the species, which is influenced by factors such

as the availability of breeding sites and com-

munity dynamics. For instance, although urban

vector species like Ae. aegypti and Ae. albopictus

have been observed in urban parks in Brazil

(Ceretti-Junior et al., 2016; Medeiros-Sousa et

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 71: e55184, enero-diciembre 2023 (Publicado Oct. 30, 2023)

al., 2015) it is known that under natural condi-

tions, high abundances of Li. durhamii have

been observed to lead to competitive exclusion

of Ae. aegypti in less disturbed larval habitats

(Talaga, 2016).

However, migration from the park to

the peri-urban area is not ruled out, particu-

larly for species (Tx. theobaldi, Li. durhamii,

Cx. secundus, Cx. derivator, Cx. dolosus) with

broad distributions and adaptability to dis-

turbed environments. Furthermore, proximity

and environmental degradation may accelerate

species migration and lead to epizootic cycles.

Vertebrates in PALI (monkeys, bats, rodents,

others) contribute to the epidemiological risk.

Over time, the park could host more animal

populations, potentially maintaining enzootic

transmission cycles and even leading to epizo-

otic cycles, that may introduce pathogens into

the urban ecosystem. In addition, Hendy et

al. (2020) have concluded that the community

composition between forest edge and interior

sites changes, with a gradual decrease in the

abundance of certain species further into the

forest. Studies have also concluded that the

diversity and abundance of species varies with

distance between the ground and the canopy,

and that these metrics are concentrated in the

lower stratum (Confalonieri & Costa Neto,

2012; Tantely et al., 2019). In other words, arbo-

virus exchange between humans and wildlife

can vary significantly within meters, and that

the role of species as possible bridge vectors is

likely defined around forest fragments.

Most of the species described have already

been reported in Ecuador. However, there is no

detailed record of the distribution of Wy. mela-

nopus and Cx. dolosus in the country (Angulo &

Olivares, 1993). This may be the first confirmed

report of Wy. celaenocephala, and Tx. theobaldi

in the Ecuadorian Amazon region, and the

first record of Wy. felicia and Cx. derivator for

the country, as no records were found after the

literature review.

The increase in abundance, including epi-

demiologically important species (Cx. dunni,

Ps. ferox) during the rainy period, warns about

the risk of pathogen transmission. Therefore,

since rainfall is bimodal, preventive recommen-

dations should be provided to visitors, mainly

during periods of high rainfall. Although sea-

sonality was evaluated in this study, precipita-

tion in Tena exceeds 4 000 mm/year, with no

months has less than 100 mm (Lucas-Solis et

al., 2021), meaning the dry period refers to a

period of relatively low precipitation. This may

explain why no significant differences were

found between seasons, besides abundance.

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

ACKNOWLEDGMENTS

To Marcelo Carrera, Lizeth Sabando,

Bryan Coronel, Karla Verdugo and Evelyn Oña

for field support. We thank the AECID project

PAO-002-2019 “Eco-epidemiology, citizen sci-

ence and diversity of insects of medical-vet-

erinary importance in the urban area of Tena,

Napo Province” for the funding granted. We

thank the AECID project POA-002-2019 for

partially financing this research.

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