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Predatory mite, Amblyseius swirskii (Acari: Phytoseiidae), for biological control of Asian citrus Psyllid, Diaphorina citri (Hemiptera: Psyllidae).

The Asian Citrus Psyllid (

), Diaphorina citri Kuwayama
(Hemiptera: Psyllidae) is an invasive pest of special concern, which has
expanded its range throughout the citrus-producing regions of Asia and
now the Americas. In the United States, the psyllid was first detected
in Florida in Palm Beach, Broward and Martin Counties in June 1998 and
quickly spread throughout the citrus-growing areas of the state (Halbert
1998; Halbert & Manjunath 2004; Michaud 2002; Tsai et al. 2002). It
has also been identified in Texas, California, Arizona, and most of the
south-eastern US (French et al. 2001, Qureshi & Stansly 2010). Host
plants of D. citri are confined to Rutaceae, in particular the genus,
Citrus and its relatives (Halbert & Manjunath 2004). The ornamental
orange jasmine, Murraya paniculata (L.) Jack, a common hedge plant in
south Florida, is considered a preferred host of ACP (Tsai et al. 2000).
Direct injury to citrus by ACP results from
 : see bark; stem.

 or bast

Plant tissues that conduct foods made in the leaves to all other parts of the plant.
 feeding on emerging
foliage (flush), which causes permanent distortion or even abscission of
new shoots with heavy
 /in·fes·ta·tion/ () parasitic attack or subsistence on the skin and/or its appendages, as by insects, mites, or ticks; sometimes used to denote parasitic invasion of the organs and tissues, as by helminths.
 (Michaud 2004; Hall & Albrigo
2007). However, most economically important damage is caused by
transmission of the bacterium ‘Candidatus Liberibacter
asiaticus’, thought to be the causal agent of
“huanglongbing” (

HLB Hotels Licensing Board  
) or citrus greening disease (Halbert
& Manjunath 2004). Symptoms indicating the presence of the bacterium

A form of chronic anemia, primarily of young women, characterized by a greenish-yellow discoloration of the skin and usually associated with deficiency in iron and protein. Also called chloremia.
 resembling zinc deficiency, a more diagnostic
1. A spot or blot; a splotch.

2. A discoloration on the skin; a blemish.

3. Any of several plant diseases caused by fungi and resulting in brown or black dead areas on leaves or fruit.

asymmetrical mottling of leaves, twig dieback with leaf and fruit drop,
uneven coloring of fruits and reduction in fruit size and quality
(Halbert & Manjunath 2004). Citrus greening disease was first
detected in Florida in 2005 (Halbert 2005) and has spread throughout the
state (http://www.freshfromflorida.com/pi/chrp/ArcReader/mi2%20Sections%20 in%20Florida%20Positive%20for%20HLB%20 15%20Mile%20Buffer.pdf)

Integrated pest management
 (IPM), planned program that coordinates economically and environmentally acceptable methods of pest control with the judicious and minimal use of toxic pesticides.
 (IPM) practices involving biological and
chemical control strategies are being developed to suppress psyllid
populations and to consequentially slow the spread of citrus greening
(Qureshi & Stansly 2007, 2009, 2010). Native or exotic biological
control agents of the psyllid include predators as diverse as
ladybeetles (Coleoptera: Coccinellidae), lacewings (Neuroptera:
Chrysopidae), spiders (Aranae), and hoverflies (Diptera: Syrphidae) that
together can greatly reduce the reproductive potential of the ACP
population by more than 90% (Michaud 2002, 2004; Pluke et al. 2005;
Qureshi & Stansly 2008, 2009). Two exotic parasitoids of ACP,
Diaphorencyrtis aligarhensis (Shafee, Alam and Agaral) (Hymenoptera:
Encyrtidae) and Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae)
were introduced in Florida in 2000 against ACP (Hoy & Nguyen 2001).
Tamarixia radiata is now widely distributed in the Florida citrus
ecosystem at variable rates of
 see parasite.


Relationship between two species in which one benefits at the expense of the other. Ectoparasites live on the body surface of the host; endoparasites live in their hosts’ organs, tissues, or cells and often rely
 (Qureshi et al. 2009) whereas
D. aligarhenis has not established. Coccinellid predator species, Olla
v-nigrum (Mulsant), Curinus coeruleus (Mulsant), Harmonia axyridis
(Pallas), and Cycloneda sanguinea (L.) (Coleoptera: Coccinellidae), are
thought to be the most important sources of
 /bi·ot·ic/ ()
1. pertaining to life or living matter.

2. pertaining to the biota.

1. Relating to life or living organisms.
 mortality on D. citri
nymphs in Florida (Michaud 2002, 2004; Michaud & Olsen 2004; Qureshi
& Stansly 2009). However, any single approach by itself is not going
to provide enough reduction of the vector psyllid and citrus greening
disease. Dormant season

 sprays of broad spectrum insecticides in
winter provide 5-27 fold reduction in ACP populations and opportunity
for biological control in spring and summer (Qureshi and Stansly 2010).
Nevertheless, the disease continues to advance, despite more frequent
sprays of insecticides during growing season in many orchards.

This situation calls for a more proactive and
1. Having the ability or tendency to augment.

2. Grammar Indicating an increase in the size, force, or intensity of the meaning of an adjacent word, as up does in eat up.

to biological control, commencing with identification of other natural
enemies of ACP. One avenue not yet investigated is biological control
using predatory phytoseiid mites (Acari: Phytoseiidae) that could feed
on the eggs and nymphs of D. citri. Phytoseiid mites are important
agents of biological control on many pests in many crops. Depending on
the species, their food may include mites,
 minute, agile insects of the order Thysanoptera. Thrips have piercing-and-sucking mouthparts and cup-shaped feet from which bladderlike adhesive organs may be extended. Some species are wingless, but many have four narrow, featherlike wings fringed with hairs.
 and Hemiptera such as
whiteflies and armored scales as well as pollen and honeydew (Dosse
1961; Putman 1962; McMurtry & Scriven 1964; Swirskii et al. 1967;
Juan-Blasco et al. 2008). These findings have heightened interest in
exploring additional possibilities for using mites to control different


 organisms in many crops (Nomikou et al. 2001; Calvo et al.
2011). Thirty-eight species of phytoseiid mites have been reported in
Florida citrus, although the biology of only a few has been studied
(Abou-Setta & Childers 1987; Abou-Setta et al. 1997; Caceres &
Childers 1991; Fouly et al. 1994; Yue et al. 1994). Much is yet to be
learned about phytoseiids that colonize Florida citrus. We are aware of
no published reports to date of any native or exotic phytoseiid
attacking eggs or the first nymphal

 D. citri.

Amblyseius swirskii Athias-Henriot (Acari: Phytoseiidae) has shown
itself to be a very efficient biological control agent of thrips [Thrips
tabaci Lindeman and Frankliniella occidentalis (Pergande)]
(Thysanoptera: Thripidae), whiteflies [Trialeurodes vaporariorum
Westwood and Bemisia tabaci (Gennadius)] (Hemiptera: Aleyrodidae) and
phytophagous mites [Tetranychus urticae Koch (Acari: Tetranychidae) and
Polyphagotarsonemus latus (Banks) (Acari: Tarsonemidae)] (Swirskii et
al. 1967; Gerling et al. 2001; Nomikou et al. 2001; Calvo et al. 2011;
Stansly & Castillo 2009, 2010). Eggs and crawlers (first instar
nymphs) of D. citri are very similar in shape and size to those of B.
tabaci. Eggs of D. citri are laid in the newly developing unfolded
leaves where they and the first instar nymphs are protected from large
predators, but easily accessible to predatory mites. For the same
reason, and due to its commercial availability and effectiveness as
biological control agent of whiteflies and other pests in greenhouse,
open field vegetable and citrus production (Stansly & Castillo 2009,
2010; Calvo et al. 2011; Juan Blasco et al. 2008), A. swirskii was
selected for a preliminary evaluation against D. citri.

Our objectives were to determine whether 1) A. swirskii mites use
D. citri eggs and nymphs as prey and 2) the mites suppress D. citri
populations on young isolated M. paniculata plants following controlled
release in a glasshouse.


Cultures and Source Material

Diaphorina citri used in the experiments was obtained from a colony
housed at the Southwest Florida Research and Education Center (SW-FREC)
in Immokalee, Florida, USA since 2005. The colony was maintained on M.
paniculata plants propagated from seed and grown in a greenhouse covered
with “Antivirus” insect netting in 15 or 20 cm pots using a
substrate consisting of processed pine bark, 60% + Canadian
  or  any species of the large and widely distributed genus Sphagnum, economically the most valuable moss.
 peat and
Any of a group of micaceous hydrated silicate minerals related to the chlorites and used in heat-expanded form as insulation and as a planting medium.
. Plants were pruned 2 to 3 wk prior to use to
induce emergence of new shoots before being moved to an air-conditioned
glasshouse maintained at 27 [+ or -] 4 [degrees]C; 70 [+ or -] 10% RH.
Six plants were placed in a wooden cage for

 twice a week (60
x 60 x 60 cm) with openings covered with fine polyester netting (40 x 10
mesh/ cm) containing 300 adult ACP. After 3 d, plants were shaken and
remaining adults vacuumed off. Plants were then transferred into an
empty cage and held until emergence of adult ACP to be used for
experimentation or to replenish the colony. Used plants were pruned,
sprayed with a 2% glycerin soap solution (Clearly Natural Essentials

 common name for some members of the Caprifoliaceae, a family comprised mostly of vines and shrubs of the Northern Hemisphere, especially abundant in E Asia and E North America.
, Pure and Natural

 Soap) and recycled for later
use. Amblyseius swirkii sustained on the dried fruit mite Carpoglyphus
lactis (L.) (Acari: Carpoglyphidae), was provided by Koppert Biological
Systems (SWIRSKI-MITE[R]; Howell, Michigan. USA). Experiments were
conducted from Nov 2007 to Feb 2008.

Laboratory Experiment

Predation by A. swirkii on eggs and nymphal instars of D. citri was
evaluated. Plants that had been pruned 8 d prior were placed in the
oviposition cage with adult psyllids to infest new shoots as described
above. A young shoot of M. paniculata infested with D. citri eggs was
removed from a plant and placed inside an experimental arena consisting
of a snap-cap polystyrene cylindrical vial (10 cm in length and 4 cm in
diam). The bottom of the vial was removed and replaced by fine polyester
organdy attached with hot glue. The polyethylene lid was perforated to
receive a smaller plastic tube (length 1.7 cm, diam 0.7 cm) filled with
water into which the stem of the shoot was inserted and sealed to the
vial lid with plasticine to prevent escape of the mites. The vial was
then inverted onto the cap to enclose the upper part of the shoot and
placed in a metal grill over a plastic tray filled with water (Fig. 1).

Five A. swirskii adult females taken directly from SWIRSKI-MITE[R]
bottles were placed in the experimental arena without alternative food.
The experiment was set up the day SWIRSKIMITE[R] bottles were received
from manufacturer. Amblyseius swirskii females used in the experiment
 /neo·nate/ () newborn infant.

A neonatal infant.


a newborn animal.
 (maximum 2 days old) (Xu & Enkegaard 2010). There were
15 replicates of 3 treatments set out in a randomized complete block
design: 1) shoots infested with D. citri eggs without A. swirskii, 2) A.
swirskii adult mites with no D. citri or any other food source and 3)
shoots infested with eggs of D. citri and A. swirskii adult mites. Vials
were held in an air-conditioned rearing room maintained at 22 [+ or -] 2
[degrees]C, 63 [+ or -] 8% RH, 16:8 (L:D) photophase. Survival of A.
swirskii adult females, the number of D. citri eggs and eventually D.
citri nymphs were tallied under a stereoscopic microscope at intervals
0, 2, 4, and 6 d without replacement. Eggs and nymphs of D. citri were
counted as dead when observed empty and desiccated.

Glasshouse experiment

The capacity of A. swirskii to suppress populations of D. citri on
isolated plants was evaluated in an air-conditioned glasshouse
maintained at 27 [+ or -] 4 [degrees]C; 70 [+ or -] 10% RH. Murraya
paniculata plants, each in a 15 cm diam pot and with 4-6 young shoots
were used for the experiment. Shoots were infested with D. citri eggs
which were counted with the aid of a magnifying head set (2.75 X). Each
plant was placed in a ventilated cylindrical cage made from a sealed
sheet of clear plastic film covered on top with coarse mesh organdy
(Qureshi et al. 2009, Fig. 2). Amblyseius swirskii adults were released
on 4 plants at ratio of 1:2 (A. swirskii adults: D. citri eggs) inside
of 30 mL plastic cups attached to the plant branch with wire. The
correct number of A. swirskii was estimated by using a mean of 10
mites/gram of substrate obtained by counting mites in 10 randomly
selected one-gram samples of substrate under a stereoscopic microscope.
The control treatment consisted of plants infested with D. citri eggs
but no mites. Emerging D. citri adults were aspirated off the plants and
counted weekly for 8 wk. Leaves were inspected on the plant using the
magnifying headset and phytoseiid eggs, larvae, nymphs and adults were
recorded for 6 to 8 wk or until no more D. citri adults had been seen on
the plants for 2 consecutive weeks. Larvae of the phytoseiids are easily
distinguished from nymphs and adults by the presence of three pairs of
legs. Nymphs and adults have four pairs of legs. Nymphs molt through the
different nymphal stages (protonymph and deutonymph) to adult and leave
the corresponding

 (Abad-Moyano et al. 2009). The counted exuviae
were removed from plants.


Data Analysis

A generalized linear mixed model (
) (Breslow & Clayton
1993) was used to analyze treatment effects on D. citri eggs and nymphs
and on survival of A. swirskii using

 Statistics version 19.0.0 (


 Inc., Chicago, Illinois, USA; www.spss.com) for the laboratory
experiment. Treatments (A. swirskii present or absent) were included as
fixed effects, and shoots observed on d 2, 4 and 6 were included as a
random effect. Selection of the best model was based on the
Information Criterion

 (AIC). This revealed that the Poisson distribution
with a logarithmic link was most appropriate to analyze for treatment

The reduction of D. citri numbers attributable to A. swirskii
predation on caged plants in the glasshouse was calculated using the
Henderson-Tilton formula (Henderson & Tilton 1955).
 Analysis of variance, see there
 was used
to compare D. citri adult emergence between the A. swirskii and control
treatments. Diaphorina citri adult emergence was square root transformed
(sqrt (x)) to correct for heterogeneity of variance.


Laboratory Experiment

Adults of A. swirskii were observed preying upon eggs and first
instar nymphs of D. citri on M. paniculata shoots. The predatory mites
were observed sucking out the body fluids of the nymphs which then
looked dried and empty. More dead D. citri eggs were observed in the
presence of A. swirskii than in its absence (GLMM: [F.sub.1,28] =
103.18, P < 0.001) (Fig. 3A).

The treatment effect on number of dead nymphs was not significant
(GLMM: [F.sub.1,28] = 0.01, P = 0.935). Honeydew excretions from nymphs
decreased over the 6-d observation period, indicating failure of the
shoots to provide complete nutrition after the first few d which may
have contributed to D. citri nymphal mortality. However, fewer live
nymphs were observed on A. swirskii treated shoots than on untreated
shoots (GLMM: [F.sub.1,28] = 46.68, P < 0.001) (Fig. 3B). This
suggests a negative effect on the development of nymphs from eggs in the
presence of A. swirskii. Indeed, fewer D. citri nymphs (dead + live)
were observed on shoots with A. swirskii adults compared with shoots
without mites after 6 days (GLMM: [F.sub.1,28] = 18.98, P < 0.001)
(Fig. 3C). However, nymphal survival was low on both A. swirskii treated
and control shoots. An average of 24 [+ or -] 9 (34%) live nymphs were
observed in the control treatment at the end of the experiment out of
the 79 [+ or -] 8 eggs at the beginning of the experiment compared with
6 [+ or -] 5 (14%) live nymphs observed in the A. swirskii treatment.

Survival of adult A. swirskii mites on shoots infested with D.
citri eggs and nymphs was not different from shoots without D. citri
eggs and nymphs up to 6 d (GLMM: [F.sub.1,28] = 2.67, P = 0.114), which
averaged 72 [+ or -] 8% on shoots infested with D. citri eggs and nymphs
and 56 [+ or -] 13% on shoots without D. citri eggs and nymphs. Mite
reproduction was limited over the 6 d period. One larva and 3 nymphs of
A. swirskii were recorded in replicates with access to D. citri, and
only 1 larva in replicates without D. citri.

Glasshouse Experiment

Initial numbers of D. citri eggs observed on A. swirskii treated
and untreated plants averaged ([+ or -] SE) 247 [+ or -] 37 and 240 [+
or -] 40 per plant, respectively. The total number of psyllid adults
collected from plants with A. swirskii averaged 42 [+ or -] 11 which was
80% less than 204 [+ or -] 31 collected from control plants without A.
swirskii (ANOVA: [F.sub.1,7] = 28.79, P = 0.002) (Fig. 4). Most psyllid
adults emerged during the second and third wk of observation. Most A.
swirskii individuals (egg to adult) were observed 2 wk after release
when a mean of 52 [+ or -] 15 per plant out of the initial 124 [+ or -]
21 per plant was found. The mean number of A. swirskii observed per
plant had decreased the third week (24 [+ or -] 14), and then during wk
4 and 5 showed the minimum number of all life stages of A. swirskii
observed, 5 [+ or -] 2 and 5 [+ or -] 3, respectively. Per plant numbers
of all life stages of A. swirskii increased during wk 6, 7 and 8,
averaging between 3 [+ or -] 0.5 – 5 [+ or -] 0.8, 9 [+ or -] 1.1 – 17
[+ or -] 1.8, 14 [+ or -] 1.5 – 33 [+ or -] 4.6, and 66 [+ or -] 5.7 –
86 [+ or -] 7.4 for eggs, larvae, nymphs and adults, respectively,
during 3 wk period.



We observed that D. citri eggs on young shoots were preyed upon by
A. swirskii mites under laboratory conditions and numbers of dead
psyllid eggs were greater in the presence of the mites. To our knowledge
this is the first time that a phytoseiid mite was recorded preying upon
eggs of D. citri. Although, predation was also observed on first instar
nymphs of D. citri (0-6 d in the laboratory experiment), there was no
statistically significant difference in the number of dead nymphs
between A. swirskii treated and control shoots,
That can be presumed or taken for granted; reasonable as a supposition:
 due to low
survivorship in untreated controls. Possibly, shoot quality declined
over time and could not provide enough nutrition for nymphs as evidenced
by reduced honeydew secretion and high control mortality. Removal of
young shoots from plants may have inhibited photoassimilate transport
from mature leaves to shoot apices where eggs were laid and newly
hatched nymphs fed. In contrast, adult emergence was 85% from nymphs
that developed on intact shoots of caged plants in the control treatment
in glasshouse.




Availability of alternative prey as well as non-prey food sources
such as pollen and insect-produced honeydew are often important for
predator establishment and persistence in the crop (Gonzalez-Fernandez
et al. 2009; Nomikou et al. 2003, 2010). Non-prey food sources not only
provide water and nutrients to complement a diet consisting of prey, but
sometimes allow for predator reproduction. Nomikou et al. (2003)
observed that
 or  any plant of the genus Typha, perennial herbs found in almost all open marshes. The cattail (also called club rush) has long narrow leaves, sometimes used for weaving chair seats, and a single tall stem bearing two
 pollen supported survival, development and
reproduction of the two phytoseiid species A. swirskii and Euseius
scutalis Athias-Henriot. Similarly, honeydew from B. tabaci greatly
increased survival of E. scutalis, and supported development to
adulthood and a high rate of oviposition. In contrast, coccid-produced
honeydew did not promote development or oviposition of E. scutalis or A.
swirskii, indicating that honeydew from B. tabaci may be of higher
quality than coccid-produced honeydew, at least for E. scutalis (Swirski
et al. 1967). Survival of adult A. swirskii was high with or without
pollen or honeydew from B. tabaci provided on cucumber leaves, but
oviposition by adults and juvenile survival was very low on a diet of
honeydew compared with pollen (Nomikou et al. 2003). However, others
(Ragusa & Swirski 1977; Momen & El-Saway 1993) observed enhanced
survival of A. swirskii on honeydew produced by B. tabaci. Therefore,
psyllid honeydew could be used by predatory mites as an alternative food
source while searching for prey in the field. Additionally, previous
studies have demonstrated that other species of phytoseiid mites are
able to feed on plant sap (Grafton-Cardwell & Ouyang 1996; McMurtry
& Croft 1997). Further investigation is needed to determinate the
suitability of psyllid honeydew for A. swirskii as well as the role that
plant sap can play in their survival.


Citrus pollen could also be a useful non-prey food source for A.
swirskii. Villanueva & Childers (2004) found a positive relationship
between the number of phytoseiids and pollen grains on grapefruit leaves
during the period of citrus flowering at
Lake Alfred, Florida

. In
addition to pollen and honeydew from psyllids, A. swirskii, a
polyphagous predator, could also benefit from other potential preys that
colonize citrus such as several species of mites and thrips. Mixed diets
are sometimes better food than any single type of food (Messelink et al.
2008). Both A. swirskii and E. scutalis had higher oviposition rates on
diets of spider mites and almond anthers than on either food alone
(Swirski et al. 1967). Amblyseius swirskii demonstrated higher
oviposition on a mixed diet of eriophyoid mites and castor bean pollen
than on pollen alone (Ragusa & Swirski 1977).

The glasshouse experiment was conducted to test if predation by A.
swirskii upon D. citri observed on individual shoots could also be
observed on isolated plants. Amblyseius swirski released at a 1:2 (A.
swirskii adult: D. citri egg, 124:247) ratio reduced the D. citri
population up to 85% compared with control plants. This result
demonstrated the potential for psyllid control using A. swirskii.
Nomikou et al. (2002) also showed suppression of B. tabaci on single
plants of cucumber using A. swirskii. Stansly & Castillo (2009,
2010) observed significant suppression of broad mite Polyphagotarsonemus
latus (Banks) in the field using A. swirskii on both pepper and
eggplant. Also, eggplant receiving A. swirskii yielded significantly
more fruit than untreated plants or even eggplants receiving sprays of
the acaracide spiromesefen. In this study, the increase in number of
immature A. swirskii 1 month after the initial release indicated that
they were probably using psyllid immatures and honeydew to support
reproduction along with the food mites present with the SWIRSKI-MITE[R]

In this study, the predator A. swirskii showed promise for
biological control against D. citri show some promise, but further tests
are necessary to determine its possible role in the management of D.
citri. Amblyseius swirskii inoculations could be used to reduce psyllid
populations on young plants in the nurseries and field and thus, be used
as a preventive measure to reduce D. citri eggs (
 /in·oc·u·lum/ () pl. inoc´ula   material used in inoculation.

n. pl.
). Future
research should be focused on appropriate host plants (e.g., varieties
of citrus or citrus relatives), reproduction of A. swirskii on D. citri
diet, its prey preference, rates and frequency of the releases, density
dependent effects and impact on D. citri populations in the field. Other
predators, particularly lady beetles and the
Any of various insects, such as the ichneumon fly, whose larvae are parasites that eventually kill their hosts.

Of or relating to a parasitic insect of this kind.
, T. radiata, are
already well established in Florida citrus and known to cause
significant mortality to psyllid populations in the citrus groves
(Michaud 2004; Qureshi & Stansly 2009; Qureshi et al. 2009). If
proved effective in the field, A. swirskii could be a useful addition to
enhance natural mortality of D. citri through its impact on eggs and
first instar nymphs which are protected in newly developing unopened
leaves and difficult to reach by most predators. These early immature
stages are not targeted by nymphal parasitoids which prefer later
instars (Chu & Chien 1991). Finally, the predatory role of native
mites on D. citri as well as their interactions with A. swirskii should
also be evaluated.

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A musical instrument of northern Africa and southwest Asia resembling a lute.

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, J. M., Wong, E., Trigo, M. M., and Montserrat, M. 2009.
Alternative food improves the combined effect of an
 see carnivore.


Animal that eats both plant and animal matter. Most omnivorous species do not have highly specialized food-processing structures or food-gathering behaviour.
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IOBC Infantry Officer Basic Course
IOBC International Online Bridge Club
IOBC Indian Ocean Biological Center


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Maria Juan-Blasco (1), Jawwad A. Qureshi (2)*, Alberto Urbaneja (1)
and Philip A. Stansly (2)

(1) Centro de Proteccion
 /veg·e·tal/ () vegetative (defs. 1, 2, and 3).

1. Of, relating to, or characteristic of plants.

 y Biotecnologia, Instituto
Valenciano de Investigaciones Agrarias (
), Ctra. Montcada-Naquera
Km. 4.5, 46113 Moncada, Valencia, Spain

(2)Department of Entomology and
The branch of zoology that deals with nematodes.

, University of
Florida/IFAS Southwest Florida Research and Education Center, 2685 SR
29N, Immokalee, FL 34142, USA

* Corresponding author; E-mail: jawwadq@ufl.edu