Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

Local and landscape constraints of adult population

of Anastrepha obliqua (Diptera: Tephritidae) in mango orchards

Nadia L. García-Olivos1; https://orcid.org/0009-0001-0102-3154

Rodrigo Lasa-Covarrubias2; https://orcid.org/0000-0003-1175-7538

Ricardo Serna-Lagunes1; https://orcid.org/0000-0003-1265-9614

Miguel A. García-Martínez1*; https://orcid.org/0000-0002-0644-4601

1. Facultad de Ciencias Biológicas y Agropecuarias Región Orizaba-Córdoba, Universidad Veracruzana. Josefa Ortiz de

Domínguez S/N, Peñuela, Amatlán de los Reyes, 94950, Veracruz, México; nadia.garciaolivos@gmail.com, rserna@

uv.mx, miguelgarcia05@uv.mx (*Correspondence)

2. Red de Manejo Biorracional de Plagas y Vectores, Instituto de Ecología, A.C. Carretera antigua a Coatepec 351, El

Haya, Xalapa, 91073, Veracruz, México; rodrigo.lasa@inecol.mx

Received 15-XII-2023. Corrected 29-III-2024. Accepted 30-VIII-2024

ABSTRACT

Introduction: The West Indian fruit fly, Anastrepha obliqua (Macquart), is one of the most economically impor-

tant pests of mangoes in Mexico and the Neotropics.

Objective: To analyze adult population patterns of A. obliqua and their relation to local and landscape factors.

Methods: We selected 11 “Manila” mango orchards along the middle part of La Antigua River, which were char-

acterized to determine their biophysical structure and landscape configuration considering their distance to six

different land uses/land covers. Anastrepha obliqua population was estimated by collecting adult flies in colorless

polyethylene plastic bottle traps baited with a protein for three weeks in June 2022.

Results: A total of 1 869 adults of A. obliqua were trapped, of which 75 % were females and 25 % were males.

Abundance varied from 68 to 490 adult flies per orchard, while capture frequency from 0.36 to 1.8 flies/trap/

day. Abundance increased in highly shaded orchards, and capture frequency decreased in orchards with highly

soil compaction. Abundance and the frequency of capture increased in orchards near isolated trees of Spondias

spp. and decreased in orchards near sugarcane plantations. Shade management in mango orchards may be used

as an effective strategy to promote biotic interactions to naturally regulate A. obliqua populations, meanwhile,

soil compaction represents a limiting ecological condition, which might severely impact fruit fly larvae survival.

Conclusions: Adult population of A. obliqua depends on several local and landscape factors because they indicate

resource availability and ecological conditions. These findings could be considered for control strategies in the

integrated management of this pest to promote protection and improve the fruit quality of mango crop.

Key words: pest ecology; control strategy; insect population; pest management.

RESUMEN

Limitantes locales y del paisaje de la población adulta de Anastrepha obliqua

(Diptera: Tephritidae) en huertos de mango

Introducción: La mosca de las indias occidentales, Anastrepha obliqua (Macquart), es una de las plagas más

importantes económicamente en el cultivo de mango de México y el neotrópico.

https://doi.org/10.15517/rev.biol.trop..v72i1.56840

TERRESTRIAL ECOLOGY

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

INTRODUCTION

The West Indian fruit fly, Anastrepha obli-

qua (Macquart), is a pest widely distributed

along tropical areas of the Americas from the

Southern United States to Southern Argentina

(Norrbom, 2008; Santos et al., 2020). How-

ever, it can potentially expand to other tropi-

cal regions worldwide (Fu et al., 2014). It is

polyphagous and infests fruits of economic

importance such as mango (Mangifera indica

L.; Sapindales: Anacardiaceae), red or yellow

mombin, Mexican apple or “jocote” plums

(Spondias purpurea L. and S. mombin L.; Sap-

indales: Anacardiaceae) and sometimes guava

(Psidium guajava L.; Myrtales: Myrtaceae)

(Aguirre-Ramírez et al., 2017; Carvalho et al.,

1996; Guillén et al., 2022; Hernández-Ortiz &

Aluja, 1993). Recently, A. obliqua is considered

a cryptic pest species (Ruiz-Arce et al., 2012)

and stands out as the most important pest of

mango crop in the Neotropics (CABI, 2021).

The monitoring of adult incidence of A.

obliqua has been performed using different

hydrolyzed proteins as baits (IAEA, 2018; Lasa

y Williams, 2022). Meanwhile, control strate-

gies for this pest in mango orchards include the

spraying of a mixture of a hydrolyzed protein

bait and an insecticide (Epsky et al., 2014). The

most common killing component of these mix-

tures is malathion, which is a broad-spectrum

synthetic organophosphate highly toxic (Díaz-

Fleischer et al., 2017). Recent studies suggest

that the use of a toxic bait formulated with

biorational insecticide spinosad (GF120, Dow

Agrosciences) represents a promising alterna-

tive to minimize impact against non-target

insects and environment (Flores et al., 2011).

However, the optimization of any control strat-

egy for this pest requires a comprehensive

analysis of population patterns of abundance

and frequency of capture at orchard level and

spatial distribution at regional scale.

Population properties play a crucial role in

the implementation of management strategies

for controlling fruit flies in mango crop (Abei-

jon et al., 2019; Dias et al., 2018; Guillén et al.,

2022; Paredes et al., 2021). For instance, spa-

tial fluctuations in local population abundance

and frequency of capture may allow growers

to identify pest infestation levels and rein-

force insecticide application at orchard level

to improve control (Pimentel, 2007). On the

other hand, pest spatial distribution is useful

for the design and development of management

programs in an “area wide pest management”

approach, which is commonly applied for con-

trolling fruit flies (López et al., 2019; Klassen et

Objetivo: Analizamos los patrones de la población adulta de A. obliqua y su relación con factores locales y del

paisaje.

Métodos: Un total de 11 huertos de mango “Manila” fueron caracterizados en estructura biofísica y configuración

del paisaje. Los adultos de A. obliqua fueron capturados con trampas (botellas de polietileno cebadas con una

proteína) durante tres semanas en junio de 2022.

Resultados: Se capturaron 1 869 adultos de A. obliqua, de los cuales 75 % fueron hembras y 25 % machos. La

abundancia varió de 68 a 490 moscas adultas por huerto, mientras que la frecuencia de captura de 0.36 a 1.8 mos-

cas/trampa/día. La abundancia incrementó en huertos altamente sombreados y decreció en huertos con suelos

compactos. La abundancia y frecuencia incrementaron en huertos cercanos a árboles de Spondias spp. y decreció

en huertos cercanos a plantaciones de caña de azúcar. El manejo de la sombra puede ser utilizado como una

estrategia efectiva para promover las interacciones que regulan naturalmente la población de A. obliqua, mientras

que la compactación del suelo representa una condición ecológica limitante que podría impactar severamente en

la supervivencia de larvas de moscas de la fruta.

Conclusión: La población adulta de A. obliqua depende de varios factores locales y del paisaje que indican dispo-

nibilidad de recursos y condiciones ecológicas. Estos hallazgos podrían considerarse en las estrategias del manejo

integrado de esta plaga para promover la protección y el mejoramiento de la calidad de frutos en el cultivo de

mango.

Palabras clave: ecología de plagas; estrategia de control; población de insectos; manejo de plagas.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

al., 2008; Pimentel et al., 2007; Silva et al., 2019).

Thus, the analysis of variations in abundance,

frequency of capture, and spatial distribution

of A. obliqua adult population, and the factors

that could potentially modulate them, may be

an effective tool to improve agricultural man-

agement planning to prevent increased damage

and economic impacts in mango crops.

Some studies have suggested that popu-

lation abundance and frequency of capture

of fruit flies are influenced by microclimatic

conditions and structural complexity of habi-

tat (i.e., local factors) (Hudiwaku et al., 2021;

Krasnov et al., 2019). Particularly, these mea-

sures are commonly influenced by several local

biotic and abiotic factors of habitat such as

ambient temperature, soil moisture, crop type,

and host abundance (Bota et al., 2018; Tiring

& Satar, 2021). For example, the incidence

and abundance of A. obliqua is also dependent

of fruit availability for oviposition in mango

crop (Aluja & Birke,1993). In fact, fruit fly

populations are more abundant and frequent

in highly shaded and humid habitats of tropical

regions because they have lower fitness costs,

meanwhile, in temperate regions they are more

abundant and frequent in warmer and sunnier

habitats (Navarro-Llopis & Vacas 2014). Due to

low-distance dispersion capacity of fruit flies,

mainly of the genus Anastrepha (250-300 m),

there are reports that their populations are

denser in small area plots than in larger ones

(Krasnov et al., 2019; Weldon et al., 2014).

Although the abundance and frequency of cap-

ture of A. obliqua population are firstly shaped

by diverse local factors, its spatial distribution

is expected to be also influenced by the type of

matrix or surrounding non-habitat uses/covers.

The relative influence of the surrounding

landscape on spatial distribution of fruit fly

populations in agricultural regions is poor-

ly understood. A few studies have suggested

that composition (i.e., covered proportion and

number of different land uses/land covers)

and the configuration (i.e., spatial arrangement

of land uses/land covers) of the surrounding

landscape presumably modulate this pattern

(Aluja & Birke 1993; Clemente & Álvarez,

2019; Krasnov et al., 2019; Ricci et al., 2009).

Adult population of A. obliqua is widely dis-

tributed within a landscape matrix in lemon

orchards (Citrus latifolia Tan.; Sapindales: Ruta-

ceae), meanwhile, it is negatively affected in a

sugarcane matrix (Saccharum officinarum L.;

Poales: Poaceae) (Pacheco-Morales, personal

communication). Moreover, López et al. (2019)

reported that shorter distances among mango

trees favors the dispersion. Indeed, this popula-

tion can use different connectivity elements of

the landscape (e.g., isolated trees, living fences,

or native vegetation remnants), which are com-

posed of plum host species (Spondias spp.),

to obtain supplementary and/or complemen-

tary resources that may compensate for limited

resource availability in mango crop (Schwarzm-

ueller et al., 2019). Therefore, landscape factors

may be an important focus for considering in

management planning to limit survival, spread

and establishment of A. obliqua population in

vast agricultural areas.

In Mexico, A. obliqua is one of the main

pests that threatens yield and profitability of

mango and other economically important crops

(Díaz-Fleischer et al., 2017) and, although

there is a pest management program in fruit

orchards, its populations have not been sup-

pressed effectively. Identification of local and

landscape factors that influence adult popula-

tion of A. obliqua may be useful in decision-

making for control strategies establishment

that allow producers to mitigate this pest.

Therefore, this study analyzes distribution pat-

terns of abundance and capture frequency of

A. obliqua adult population and their relation

to local and surrounding landscape factors in

mango orchards in central Veracruz, Mexico.

We hypothesize that variations in abundance

and frequency of capture of adult populations

depend on at least a local and landscape factor

since they are indicators of resource availability

and of environmental conditions.

MATERIALS AND METHODS

Study area: Field work was conducted in

the municipalities of Jalcomulco and Apazapan,

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

Veracruz, in the middle and lower zone of the

La Antigua River basin. The climate in this

region is markedly seasonal, with two well-

defined seasons, rainy season (June to Novem-

ber) and dry season (December to May). The

study area has a seasonally tropical dry or

semideciduous forest (Comisión Nacional para

el Conocimiento y Uso de la Biodiversidad

[CONABIO], 2023; Murphy et al., 1995; Pala-

cios-Wassenaar et al., 2014). The most common

land uses/covers are human settlements, water

bodies, remnants of native vegetation and agri-

culture. The main crops are mango, sugarcane,

lemon (Citrus limon L.) (Sapindales: Rutaceae)

and papaya (Carica papaya L.) (Brassicales:

Caricaceae). The soil type is a sandy loam

(CONABIO, 2023). Even though this region

does not stand out as one of the main mango

producers in Mexico, previous studies have

shown high population densities of A. obliqua

due to the lack of phytosanitary management

for this pest (Diaz-Fleischer et al., 2017; Lasa &

Williams, 2022).

Sampling plot selection and design: In

the study area, we selected a total of 11 per-

manent sampling plots of 100 × 100 m embed-

ded within “Manila” mango orchards, which

were regarded as experimental units in the

study (Fig. 1). The selected plots did not have

any type of agronomic management and a

minimum distance of 500 m among plots was

considered to ensure experimental unit inde-

pendence. Plots were divided in four 50 × 50

m quadrats (0.25 ha), and a mango tree placed

in the center of each quadrat was selected ran-

domly as a sampling unit.

Sampling of A. obliqua flies: Two traps

were set in each sampling unit. These traps

were constructed using 0.6 L colorless poly-

ethylene bottles. Four circular holes of 10 mm

diameter were made around the circumference

at a height of 12 cm from the base (Fig. 2). One

trap was completely colorless and other was

covered at the bottom with yellow plastic tape

(50 mm wide) as a visual stimulus. Both traps

were baited with 250 ml of CeraTrap® (Bioi-

berica, Barcelona, Spain), an enzymatic hydro-

lyzed protein attractant of animal origin highly

attractive to A. obliqua (Lasa & Williams, 2022).

CeraTrap® bait is a long-lasting attractant (Lasa

et al., 2015), so was not rebaited and only from

10 to 30 ml were added weekly to the trap to

maintain a volume of 250 ml. Traps were hung

at the upper one-third of the total tree height

Fig. 1. Location of 11 mango orchards (Mangifera indica L.) “Manila” located in the middle part of La Antigua River basin

in the municipalities of Jalcomulco and Apazapan, Veracruz. Gray circles indicate the orchard location of the 11 permanent

sampling plots.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

inside the tree canopy at a central point of lat-

eral branches hung on two different branches

(6-8 m distance) in a mango tree placed in the

center of each sample point, the trees had a

height of 10 to 26.6 m. Traps were monitored

weekly for A. obliqua and rotated to the right

of their position. Considering the 11 sampling

plots and the eight traps per plot, a total of 88

traps were revised weekly, resulting in a total

of 264 traps for the entire sampling. Adults of

A. obliqua were monitored in the study area

for three consecutive weeks, between May 14

and June 4 of 2022, matching with the main

period of A. obliqua population increase in that

region (Birke et al., 2013; Martínez-Flores et al.,

2023). During this period temperature varied

from 18.8 to 27.8 ºC and relative humidity

from 76 to 81 %.

Captured insects were transported to the

laboratory in 70 % ethanol, recording the date

and type of trap and A. obliqua adults were

identified and sorted by sex. The abundance

and frequency of capture of A. obliqua were

considered as response variables (Ativor et al.,

2012; Vayssières et al., 2013). Abundance was

considered as the total number of A. obliqua

individuals collected during the three-week

sampling period on each experimental unit.

Availability was expressed as FTD index (flies

per trap per day) which is used to know the

relative presence of fruit flies in each area and

period. This was obtained using the formula:

FTD = F ∕ (T × D)

where F is the number of flies captured, T is the

number of traps, and D is the number of days

the traps were exposed in the field.

Local scale characterization: The orchards

were structurally characterized to identify if the

changes present in each of them may directly

affect A. obliqua population. Shade cover (%)

was recorded for each sampling unit using a

convex spherical densitometer (Forestry Sup-

pliers, Jackson, Mississippi). In addition, tree

density and tree height (m) were measured in

each sampling unit to determine influence of

Fig. 2 . Frontal view of the set traps for capturing Anastrepha obliqua in 11 mango orchards in central Veracruz, Mexico. (a)

Completely colorless trap. (b) Trap covered at the bottom with yellow plastic tape.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

planting density on the levels of infestation by

A. obliqua. Also, soil compaction (kg/cm²) in

each sampling unit was recorded using a pock-

et conical penetrometer (Humboldt H-4195,

Elgin, Illinois). The proportion of soil cover

(%) was estimated by delimiting an area of 1

m2 with a square frame in each sampling unit.

All these parameters were quantified in the first

day of sampling (14 May 2022), at the begin-

ning of the rainy season.

Landscape scale characterization: To

determine landscape configuration, we deter-

mined the distance from each individual plot

to the nearest edge of other land uses/covers

such as: i) native vegetation, ii) other mango

orchards, iii) Spondias spp. remnants, iv) sug-

arcane plantations, v) human settlements and

vi) water bodies. Distance measurements were

done using the ArcMap Patch Analyst exten-

sion version 10.8 (ESRI; 2021).

Data analysis: To identify the predictor

variables that significantly influenced the popu-

lation structure of A. obliqua, single and multi-

ple regression models were performed. As these

statistical techniques are sensitive to collinear-

ity between predictor variables, Pearson’s corre-

lation coefficient was used to exclude correlated

or redundant variables because all variables are

continuous, meanwhile, Spearman’s correlation

coefficient was used for numerical discrete

variables such abundance (Cohen et al., 2009).

Of each set of significantly correlated variables,

only the most intuitive and interpretable vari-

able was retained.

We used generalized linear models (GLM)

to assess the independent effects of each land-

scape predictor on abundance and availability

of A. obliqua (i.e., a single univariate regression

between a response and a predictor variable).

We applied a Gaussian error distribution for

continuous variables (i.e., availability) and a

Poisson error distribution for count-dependent

variables (i.e., abundance). For each multi-

ple regression model, variance inflation factor

was used to exclude predictor variables that

could affect the accuracy of the estimates. We

followed an information-theoretic approach

and multi-model inference to assess the rela-

tive effect of each local and landscape factor on

abundance and availability of A. obliqua using

the package glmulti for R version 3.5.0 (Calcag-

no & de Mazancourt, 2010). This function built

a set of models representing all possible combi-

nations of landscape predictors for abundance

and availability. It also computed the Akaike’s

information criterion, corrected for small sam-

ples (AICc) for each built model. To correct for

the overdispersion associated with count data,

abundance was assessed with qAICc instead of

AICc values (Burnham & Anderson, 2002). The

goodness-of-fit of the models was estimated

as the explained deviance for each complete

model using the modEvApackage for R version

3.5.0 (Barbosa et al., 2014).

RESULTS

Adult population of A. obliqua: A total

of 2 045 individuals of Anastrepha spp. were

captured, which 1 869 were A. obliqua (Table 1).

The total abundance of this species varied sig-

nificantly between sexes (F = 54.1, p < 0.05) and

plots (F = 18.7, p < 0.05). Female abundance

varied from 46 (plot #6) to 318 (plot #5), mean-

while male abundance ranged from 16 (plot #2)

to 175 (plot #5). Availability, estimated as FTD,

varied significantly between sexes (F = 56. 4, p

< 0.05) and plots (F = 18. 5, p < 0.05). Female

availability varied from 0.3 flies/trap/day (plot

#1) to 2.5 (plot #6), while that of males ranged

from 0.11 (plot #2) to 1.15 (plot #5).

Local factors: In the sampled plots, shade

cover ranged from 62.1 % (plot #3) to 92.7 %

(plot #2) and was positively correlated with tree

availability (R = 0.74, p < 0.05) (Table 2). Tree

height ranged from 10.13 (plot #4) to 26.3 m

(plot #1) and was positively correlated with soil

cover (R = 0.70, P < 0.05) and negatively with

soil compaction (R = −0.53, p < 0.05).

Landscape factors: Distance from each

individual plot to the nearest Spondias spp.

remnant edge ranged from 10 m (plot #5) to

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

4820 m (plot #2), and it was positively corre-

lated with distance from each individual plot

to the nearest edge of other mango orchards

(R = 0.73, p < 0.05), human settlements (R =

0.83, p < 0.05) and water bodies (R = 0.60, p <

0.05) (Table 2). Meanwhile, distance from each

individual plot to the nearest edge of Spondias

spp. remnants, sugarcane plantations (R = −0.7,

p < 0.05) and native vegetation (R = −0.54, p <

0.05) were negatively correlated.

Influence of local factors on A. obliqua

population: Abundance was significantly and

negatively influenced by shade cover produced

by mango trees. When the percentage of shade

increased the number of individuals decreased

(Z = −19.4, d.f. = 10, p < 0.05) (Fig. 3A). The

remaining structural variables did not influ-

ence abundance (p > 0.05). Availability was

significantly and negatively influenced by soil

compaction (t = −3.5, d.f. = 10, p < 0.05) (Fig.

3B). Plots with compacted soils presented a

lower availability of individuals. Remaining

structural variables did not influence availabil-

ity (p > 0.05).

Influence of landscape factors on A. obli-

qua population: Abundance (Z = −17.07, d.f.

= 10, p < 0.05) and availability (t = −2.5, d.f. =

10, p < 0.05) were significantly and negatively

influenced by distance from each individual

plot to the nearest Spondias spp. remnant edge

(Fig. 3C, Fig. 3D). In addition, abundance (Z

= 26.4, d.f. = 10, p < 0.05) and availability (t =

5.6, d.f. = 10, p < 0.05) were significantly and

positively explained by distance from each indi-

vidual plot to the nearest sugarcane plantation

edge (Fig. 3E, Fig. 3F). The A. obliqua popula-

tions increased with shorter distances between

orchards and Spondias spp. trees, while popula-

tions decreased with shorter distances between

orchards and sugarcane crops.

DISCUSSION

Local and surrounding landscape char-

acteristics of diverse economically impor-

tant crops are key factors, which modulate

insect pest populations (Carrière et al., 2012).

The present study provides insights into the

Table 1

Variation in adult population measures (mean ± SE) of Anastrepha obliqua studied in mango orchards in central Veracruz,

Mexico.

Adult population measures Females Males Both sexes

Abundance (individuals) 106.36 ± 20.9 43.81 ± 13.09 150.18 ± 37.11

Availability (flies/trap/day) 0.55 ± 0.11 0.24 ± 0.06 0.72 ± 0.13

Table 2

Local and landscape characteristics of mango orchards in central Veracruz, Mexico.

a) Local scale factors Mean ± SE Range

Tree height (m) 18.82 ± 1.33 10.1-26.3

Shade cover (%) 82.63 ± 2.99 62.1-92.7

Planting density (trees/ha) 13.36 ± 0.78 9-18

Soil compaction (kg/cm²) 17.82 ± 3.94 4.8-47

Soil cover (%) 69.54 ± 6 19.2-94.8

b) Landscape scale factors

Distance to nearest native vegetation (m) 260 ± 50 50-530

Distance to nearest mango orchard (m) 150 ± 20 30-350

Distance to nearest Spondias spp. remnant (m) 1600 ± 460 10-4820

Distance to nearest sugarcane plantation (m) 690 ± 140 130-1610

Distance to nearest water body (m) 630 ± 160 20-1450

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

importance of variables, indicators of resources

and conditions, at local and landscape level and

their influence on adult population A. obliqua

associated with mango orchards. These results

also improve the understanding of the main

drivers, which commonly are not consider in

the management of A. obliqua distribution

among agricultural settings devoted to mango

production. Overall, abundance and availability

of this fruit fly in mango orchards are mainly

modulated by the shade cover, soil compaction,

distance to plum trees and distance to sugar-

cane plantations.

The shade cover is a characteristic of crop

biophysical structure, which is directly related

to physiognomy vegetation and leaf litter in

soil. This local factor generates habitat parti-

tions where a vast diversity of insects is distrib-

uted according to specific requirements. Our

results indicate that total abundance of males

and females decreased as function of shade

cover in mango orchard. This pattern is due to

in more shaded orchards there are insects that

patrol mango trees and protect or defends food

resources against competitors (Cuautle et al.,

2005). In these orchards there is a thicker leaf

litter layer, which is a microhabitat of a specious

arthropods assemblage with highly specialized

feeding habits, including predation (García-

Martínez et al., 2015). Then, A. obliqua larvae

Fig. 3. Significant regression models of local and landscape factors and the abundance and availability of adult population

of A. obliqua associated with mango orchards in central Veracruz, Mexico. The goodness-of-fit of each model is indicated in

each panel as the percentage of deviance explained by each model (D2).

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

are more susceptible to be predated by spiders,

ants and beetles when infested mango fruits fall

to ground (Cruz-Miralles et al., 2022; García et

al., 2020). Thus, although our results indicate

a significant negative relation between shade

cover in mango orchards and A. obliqua adult

population, it should be taken as a hypothesis

for testing on the ecology of this pest.

Availability and abundance of males and

females was affected by increasing soil compac-

tion. This is physical property of any type of

soil, which is associated with soil composition

and structure and is influenced by the size of

the interacting particles allowing the presence

of water and air. Compacted soils have greater

difficulty in wetting and generally have fewer

pores to allow aeration which hinders the via-

bility of organisms, such as fruit flies. This local

characteristic points to a microhabitat where A.

obliqua pupal stage takes places. For example,

when the larvae reach the last stage of develop-

ment, the mango fruits fall to the ground, sub-

sequently, they leave the fallen fruits and begin

to quickly excavate the most superficial layer

of soil to start the pupation phase. This high

compaction prevents or hinder soil penetration.

In addition, those larvae which can bury and

pupate, face the same obstacle imposed by com-

paction at the time of emerging as adults (Abei-

jon et al., 2019; Fernandes et al., 2012; Ismail

& Ahmed, 2020). Although soil compaction

is correlated with soil moisture and both are

critical for adult emergence (Ismail & Ahmed,

2020), its importance stands out as a mechani-

cal resistance to digging in that severely impacts

larvae survival in shadier orchards. (Fernandes

et al., 2012; Ismail & Ahmed, 2020; Khan et al.,

2020). Soil compaction represents an easy man-

agement practice which may severely impacts

on larvae survival and adult emergence howev-

er, its implementation should be carefully taken

due to implications on soil biodiversity associ-

ated with mango orchards (Ishak et al., 2014).

Regarding landscape factors, abundance,

and availability of A. obliqua adults increased

significantly in mango orchards near to plum

trees of S. mombin and S. purpurea. This result

is concordant with a previous study carried

out in Central Veracruz, which reports a high

level of A. obliqua infestation (435 pupae/kg)

in Spondias fruits when compared with a low

level (14.55 pupae/kg) in mango fruits (Aluja

et al., 2001). This is due to, Spondias spp. fruits

have an optimal development for being infested

about 3 to 5 weeks earlier than mango so one

fruit fly generation precedes the beginning of

mango harvest in this region (Lasa, 2016; Tino-

co-Dominguez, 2019). In addition, the increase

in plum fruit number significantly increases

infestation levels of this pest in mango orchards

(López et al., 2019). For this reason, plum trees

in surrounding landscape might be acting as

reservoirs for A. obliqua adult population asso-

ciated with mango orchards. Even though these

trees are also considered as reservoirs of fruit

fly parasitoid species (de Sousa et al., 2021),

parasitoid abundance is relatively low (usually

< 10 %) compared to adult emergence of A.

obliqua (Sivinski et al., 2000). Unfortunately,

relictual Spondias spp. trees are commonly iso-

lated, scattered and used in live fences through-

out the study area (Díaz-Fleischer et al., 2017).

Given this context, collection, and destruction

of plum fruits and/or trees in the surrounding

landscape, or even spraying insecticides, is nec-

essary to mitigate adult population growth A.

obliqua in mango production regions.

The abundance and availability of A.

obliqua decreased in mango orchards near to

sugarcane crop land cover. This result is not

surprising since there is a previous report

that a landscape matrix composed of such a

crop negatively impacts this species (Pacheco-

Morales, 2022). In Mexico, this monoculture

is intensively managed, which includes a bio-

physical structure simplification and a dispro-

portionate agrochemical use (Meza-Palacios

et al., 2019). These crop fields are deeply tilled

before planting, have a homogeneous surface

highly exposed to sun radiation and during

harvest sugarcane plants are burned to facilitate

cutting. For this conditions, A. obliqua popula-

tion associated with mango orchards near or

surrounded by this crop is adversely affected

and decreased. Therefore, sugarcane crop plays

a key role as a barrier for A. obliqua distribution

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 72: e56840, enero-diciembre 2024 (Publicado Set. 10, 2024)

and movement at regional scale (Vermunt et al.,

2019). However, broader planning instruments

of control strategies should take into consid-

eration the environmental cost of this crop in

tropical regions devoted to mango production.

This study demonstrates empirically that

adult population of A. obliqua is dependent

on factors inside (i.e., local level) and outside

(i.e., landscape level) of mango orchards. We

demonstrate the significance and intimately

relations among shade cover, soils compac-

tion and Spondias spp. and sugarcane land

covers for shaping abundance, availability, and

distribution of adult population of A. obliqua.

These findings may allow optimization for

selecting and applying agronomic management

practices against A. obliqua in mango crop.

However, to highlight the importance of local

and landscape factors to regulate A. obliqua

populations, this study should be replicated in

other mango producing regions along the Gulf

of Mexico such as Actopan or Cosamaloapan,

Veracruz. Finally, effective outcomes will only

be achieved if the scope of this study may be

included as an efficient tool in the integrated

management of this pest to promote protection

of mango crop and improve its fruit quality in

this producing region.

Ethical statement: the authors declare

that they all agree with this publication and

made significant contributions; that there is

no conflict of interest of any kind; and that we

followed all pertinent ethical and legal proce-

dures and requirements. All financial sources

are fully and clearly stated in the acknowled-

gments section. A signed document has been

filed in the journal archives.

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