Evaluation of date palm cultivars...
 Two South American new species...
 A new species of Uroleucon (Hemiptera:...
 Survey for potential insect biological...
 Abundance and spatial distribution...
 Do methyl eugenol-fed males of...
 Effect of nitrogen-containing dietary...
 Patterns and consequences of mating...
 Seasonality and phenology of the...
 Population explosion of the hairy...
 Technomyrmex difficilis (Hymenoptera:...
 A new species of Cecidomyia (Diptera:...
 Effect of the biological control...
 Population survey and control of...
 Reproductive biology and development...
 Suppression of whiteflies, Bemisia...
 Effectiveness of cone emergence...
 Chironomus columbiensis (Diptera:...
 Survey of the termites (Isoptera:...
 Comparative bionomics of Frankliniella...
 Update on the establishment of...
 A major pest of cotton, Oxycarenus...
 First report of Philephedra tuberculosa...
 Ability of the redbay ambrosia...
 Populations of sharpshooters in...
 Effects of temperature on the development...
 Comparison of short-term preservation...
 Larval parasitoids associated to...
 First record of Asphondylia websteri...
 Flower visitation by adult shore...
 Book reviews
 Back Matter

Group Title: Florida Entomologist
Title: The Florida entomologist
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Permanent Link: http://ufdc.ufl.edu/UF00098813/00357
 Material Information
Title: The Florida entomologist
Uniform Title: Florida entomologist (Online)
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Language: English
Creator: Florida Entomological Society
Florida Center for Library Automation
Publisher: Florida Entomological Society
Place of Publication: Gainesville Fla
Gainesville, Fla
Publication Date: September 2008
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Subject: Entomology -- Periodicals   ( lcsh )
Insects -- Periodicals -- Florida   ( lcsh )
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periodical   ( marcgt )
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 Related Items
Preceded by: Florida buggist (Online)

Table of Contents
    Evaluation of date palm cultivars for rearing the red date palm weevil, Rhynchophorus ferrugineus (Coleoptera: Curculionidae)
        Page 353
        Page 354
        Page 355
        Page 356
        Page 357
        Page 358
    Two South American new species of Neophyllaphis subgenus Chileaphis (Hemiptera: Aphididae)
        Page 359
        Page 360
        Page 361
        Page 362
        Page 363
        Page 364
        Page 365
        Page 366
    A new species of Uroleucon (Hemiptera: Aphididae) living on Adesmia (Fabaceae)
        Page 367
        Page 368
        Page 369
        Page 370
        Page 371
    Survey for potential insect biological control agents of Ligustrum sinense (Scrophulariales: Oleaceae) in China
        Page 372
        Page 373
        Page 374
        Page 375
        Page 376
        Page 377
        Page 378
        Page 379
        Page 380
        Page 381
        Page 382
    Abundance and spatial distribution of wireworms (Coleoptera: Elateridae) in Florida sugarcane fields on muck versus sandy soils
        Page 383
        Page 384
        Page 385
        Page 386
        Page 387
    Do methyl eugenol-fed males of the oriental fruit fly (Diptera: Tephritidae) induce female re-mating?
        Page 388
        Page 389
        Page 390
        Page 391
        Page 392
        Page 393
    Effect of nitrogen-containing dietary supplements on the mating success of sterile males of the Mediterranean fruit fly (Diptera: Tephritidae)
        Page 394
        Page 395
        Page 396
        Page 397
        Page 398
        Page 399
    Patterns and consequences of mating behavior of the root weevil Diaprepes abbreviatus (Coleoptera: Curculionidae)
        Page 400
        Page 401
        Page 402
        Page 403
        Page 404
        Page 405
        Page 406
    Seasonality and phenology of the butterflies (Lepidoptera: Papilionoidea and Hesperioidea) of Mexico's Calakmul Region
        Page 407
        Page 408
        Page 409
        Page 410
        Page 411
        Page 412
        Page 413
        Page 414
        Page 415
        Page 416
        Page 417
        Page 418
        Page 419
        Page 420
        Page 421
        Page 422
    Population explosion of the hairy crazy ant, Paratrechina pubens (Hymenoptera: Formicidae), on St. Croix, US Virgin Islands
        Page 423
        Page 424
        Page 425
        Page 426
        Page 427
    Technomyrmex difficilis (Hymenoptera: Formicidae) in the West Indies
        Page 428
        Page 429
        Page 430
    A new species of Cecidomyia (Diptera: Cecidomyiidae) feeding on resin of baldcypress
        Page 431
        Page 432
        Page 433
        Page 434
        Page 435
    Effect of the biological control agent Neoseiulus californicus (Acari: Phytoseiidae) on arthropod community structure in North Florida strawberry fields
        Page 436
        Page 437
        Page 438
        Page 439
        Page 440
        Page 441
        Page 442
        Page 443
        Page 444
        Page 445
    Population survey and control of Chironomidae (Diptera) in wetlands in Northeast Florida, USA
        Page 446
        Page 447
        Page 448
        Page 449
        Page 450
        Page 451
        Page 452
    Reproductive biology and development of Lixadmontia franki (Diptera: Tachinidae), a parasitoid of bromeliad-eating weevils
        Page 453
        Page 454
        Page 455
        Page 456
        Page 457
        Page 458
        Page 459
    Suppression of whiteflies, Bemisia tabaci (Hemiptera: Aleyrodidae) and incidence of cucurbit leaf crumple virus, a whitefly-transmitted virus of zucchini squash new to Florida, with mulches and imidacloprid
        Page 460
        Page 461
        Page 462
        Page 463
        Page 464
        Page 465
    Effectiveness of cone emergence traps for detecting Phyllophaga vandinei emergence over time
        Page 466
        Page 467
        Page 468
        Page 469
    Chironomus columbiensis (Diptera: Chironomidae) new to the fauna of the United States
        Page 470
        Page 471
    Survey of the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Lanyu Island, Taiwan
        Page 472
        Page 473
    Comparative bionomics of Frankliniella occidentalis and Frankliniella tritici
        Page 474
        Page 475
        Page 476
    Update on the establishment of Paragonatas divergens (Hemiptera: Rhyparochromidae) in Florida
        Page 477
        Page 478
    A major pest of cotton, Oxycarenus hyalinipennis (Heteroptera: Oxycarenidae) in the Bahamas
        Page 479
        Page 480
        Page 481
        Page 482
    First report of Philephedra tuberculosa (Hemiptera: Coccidae) in the United States Virgin Islands
        Page 483
        Page 484
    Ability of the redbay ambrosia beetle (Coleoptera: Curculionidae: Scolytinae) to bore into young avocado (Lauraceae) plants and transmit the laurel wilt pathogen (Raffaelea sp.)
        Page 485
        Page 486
        Page 487
    Populations of sharpshooters in two citrus groves in east central Florida as indicated by yellow sticky card traps
        Page 488
        Page 489
        Page 490
    Effects of temperature on the development and survival of Metriona elatior (Coleoptera: Chrysomelidae) immatures
        Page 491
        Page 492
        Page 493
    Comparison of short-term preservation and assay methods for the molecular detection of Wolbachia in the Mediterranean flour moth Ephestia kuehniella
        Page 494
        Page 495
        Page 496
        Page 497
    Larval parasitoids associated to Anastrepha distincta (Diptera: Tephritidae) in two host fruits at the Soconusco region, Chiapas, Mexico
        Page 498
        Page 499
        Page 500
    First record of Asphondylia websteri (Diptera: Cecidomyiidae) infesting Hass avocados
        Page 501
        Page 502
        Page 503
    Flower visitation by adult shore flies at an inland site in Florida (Diptera: Ephydridae)
        Page 504
        Page 505
        Page 506
        Page 507
    Book reviews
        Page 508
        Page 509
        Page 510
        Page 511
        Page 512
        Page 513
        Page 514
        Page 515
    Back Matter
        Page 516
        Page 517
Full Text

Al-Ayedh: Rearing the Red Date Palm Weevil


Natural Resources and Environmental Research Institute. King Abdulaziz City for Science and Technology,
P.O. Box 6086, Riyadh 11442, Saudi Arabia


Development of red date palm weevil Rhynchophorus ferrugineus (Olivier) was evaluated on
4 popular cultivars of date palm, viz., 'Khalas', 'Sukkary', 'Khasab', and 'Sillaj' for 2 consec-
utive generations. The weevils reared on 'Sukkary' showed significantly better growth ex-
pressed in most parameters such as length, width and weight at larval, pupal and adult
stages. Significantly greater numbers of eggs were laid on cv. 'Sukkary' as compared to the
other 3 cultivars. This might be because of higher sugar content of cv. 'Sukkary'. Adult
lifespan was significantly longer on cv. 'Khasab'. Though more cocoons were harvested from
'Khalas', frequency of adult emergence was better on 'Sukkary'. Male to female ratio was
similar on all date palm cultivars.

Key Words: red date palm weevil, Rhynchophorus ferrugineus (Olivier), host plant food qual-
ity, fecundity, adult life span, date palm cultivars, growth assessment.


El desarrollo del picudo rojo de la palmer dactilifera Rhynchophorus ferrugineus (Olivier)
fue estudiado sobre 4 variedades populares de palmer datilifera, viz., 'Khalas', 'Sukkary',
'Khasab'y 'Sillaj' por 2 generaciones consecutivas. Los picudos criados sobre la variedad
'Sukkary' expresa un crecimiento significativamente mejor en la mayoria de los parametros
como el largo, ancho y peso de los estadios de la larva, pupa y adulto. Un numero de huevos
significativamente mayor fueron ovipositados sobre la variedad 'Sukkary' en comparici6n
con las otras 3 variedades. Esto posiblemente puede ser por que el contenido mayor de azu-
car en la variedad 'Sukkary'. La duraci6n de la vida del adulto fue significativamente mas
larga en la variedad 'Khasab'. Aunque mas capullos fueron cosechados sobre la variedad
'Khalas', la frecuencia de la emergencia de adults fue mayor en la variedad 'Sukkary'. La
raz6n del numero de machos a hembras fue similar sobre todas las variedades de palmer

The red date palm weevil (Rhynchophorus fer-
rugineus Oliv., Coleoptera: Curculionidae) is the
most devastating pest of date palm (Phoenix dac-
tylifera L) with a wide geographical distribution.
In the Arabian peninsula, it was detected on date
palm in the mid 1980s (Gush 1997; Abraham et
al. 1998; Murphy & Briscoe 1999). It was reported
from Saudi Arabia for the first time in 1986 in Al-
Katif Region (Al-Abdulmohsin 1987), from United
Arab Emirates in 1986, and from the Republic of
Iran in 1992. It spread to North Africa in Egypt in
1993 (Cox 1993). This polyphagous insect is also
widely distributed in southern Asia and Melane-
sia where it feeds on a variety of palms including
coconut, sago, date, and oil palm (Murphy &
Briscoe 1999; Rajamanickam et al. 1995). Trans-
fer of date palm offshoots as a planting material
has played a major role in rapid proliferation of
the pest in the Middle East (Abraham et al. 1998).
Azam et al. (2000) noted that 88-96% infestation
resulted from off-shoot removal and leaving the
wounds without treatment.

Damage to date palm is mainly caused by the
larval stage feeding within the trunk of palms.
This concealed feeding habit of larvae makes it
more difficult to detect infestation at an early
stage. Often rotting of the internal tissues leads
to the death of the date palm tree (Abraham et al.
1998). The weevil completes several generations
per year within the same host until the tree col-
lapses (Rajamanickam et al. 1995; Avand Faghih
1996). Yield loss due to infestation can be mild to
severe (Gush 1997).
Quality of host plant tissue significantly affects
survival and development of larvae, and influ-
ences weight, reproductive ability, longevity, and
morphology of adult insects (Leather 1990; Albert
& Bauce 1994; Dodds et al. 1996; Tammaru 1998).
Egg size and quality and selection of oviposition
sites also depend on the host plant vigor (Awmack
& Leather 2002). On a poor-quality host plant, a
female insect may lay either a few good quality
eggs only or a large number of poor quality eggs
(Rossiter 1991). Yamada & Umeya (1972) con-

Florida Entomologist 91(3)

eluded that fecundity is determined by pupal size.
Williams (1963) observed that the correlation be-
tween pupal weight and fecundity was greater
than that between pupal length and fecundity in
Chilo sacchariphagus Bojer. A significant correla-
tion was noted between the fecundity of adults
and pupal weight or length of P xylostella (Wan
1970). Murphy & Briscoe (1999) documented vari-
ations in developmental and other life history pa-
rameters such as number of eggs per female, incu-
bation period, larval and pupal duration, male
and female life span in Rhynchophorus ferrug-
ineus within and between countries.
Maternal diet effect is considered an impor-
tant factor for suitability of hosts for optimal in-
sect growth. Maternal effects have been docu-
mented in many cases (Mousseau & Dingle 1991;
Rossiter 1991; Fox 1994, 1993; Bernardo 1996;
Corkum et al. 1997; Mousseau & Fox 1998a,
1998b; McIntyre & Gooding 2000; Agrawal 2001).
In many insects, maternal diet influences egg size
and offspring quality (Fox 1993; Rossiter et al.
1993; Jann & Ward 1999). Maternal effects have
also been shown to affect duration of offspring de-
velopment and adult size (Roff 1992). A sound
fundamental knowledge of the factors associated
with dynamics of the pest population is essential
for designing efficient and sustainable integrated
pest management strategies for controlling red
date palm weevil.
The objective of the present study was to eval-
uate the influence of host tissue genotype on
growth and development of red date palm weevil
for subsequent utilization in an artificial diet for-
mulation program.


Cocoons of red date palm weevils were col-
lected in Oct 2003 from infested date palm or-
chards of cv. 'Khalas', which is the predominant
cultivar in Al-Ahsa region of Saudi Arabia.
Mother colonies were established in the labora-
tory at King Abdulaziz City for Science and Tech-
nology, Riyadh. Individual cocoons were incu-
bated on moist tissue paper in covered glass con-
tainers (7.0 cm diameter x 8.5 cm height) at 25C.
After adult emergence, individual pairs of males
and females were transferred to separate contain-
ers of the same dimensions as above for mating.
Longitudinally split pieces of sugar cane were in-
troduced in the jars as food for these adults.
Subsequent rearing of the weevil was carried
out inside the trunk pieces of 4 popular cultivars
of date palm, viz., 'Khalas', 'Sukkary', 'Khashab',
and 'Sillaj' to compare suitability of host tissue of
different genotypes for colony development.
Trunk pieces were obtained from 4- to 5-year-old
plants of 1.5 m height by chopping off apical and
basal portions. These trunk pieces, measuring av-
erage 81 cm in length, were cut longitudinally

into 2 halves. On the inner side of each half, a cav-
ity (30 cm long, 12 cm wide and 10 cm deep) was
carved, and 5 mated females were introduced into
the cavity before joining the 2 halves of each
trunk piece. The halves were held together by
winding steel wire around the trunk pieces. The
pieces were kept in cages at ambient temperature
(25C 2) for egg laying. After 5 d the females
were withdrawn and the trunk pieces were re-
turned to the respective cages for hatching of the
eggs and subsequent growth and pupation of the
larvae. After 35 d, pupae form trunk pieces of
each cultivar were harvested and incubated sepa-
rately. Adults originating from the same cultivar
were allowed to mate among themselves.
Two generations of weevil were reared consec-
utively inside the trunk pieces of respective culti-
vars with the same procedures as above. For egg
laying, 10 mated females were introduces into the
cavity of each trunk piece and 3 trunk pieces of
each cultivar were maintained as replicates. After
48 h, females were withdrawn from the cavities
and eggs were allowed to hatch in-situ. On hatch-
ing, 10 larvae were picked randomly from each
trunk piece and length, width, and weight of indi-
viduals was recorded as well as length and width
of the head capsule in order to follow larval devel-
opment on host tissue of different cultivars. Data
were recorded on the same larva at 4-d intervals
until pupation.
After pupation, cocoons were incubated indi-
vidually in plastic jars lined with moist tissue pa-
per. Ten pupae were randomly picked from each
trunk piece of the 4 cultivars, and length, width,
and weight of the pupae as well as the adults that
emerged from the same were recorded for each
cultivar. Frequency of adult emergence and male
to female ratio were recorded for populations de-
rived from the different host cultivars.
Egg laying efficiency of females reared on dif-
ferent cultivars was evaluated separately. Three
individual pairs of male and female originating
from each stem piece of the 4 cultivars were al-
lowed to mate in 500-mL plastic jars. After 3 d, fe-
males were transferred to oviposition cages (26 x
16 x 9 cm) and provided with 2 perforated ovipo-
sition substrate boxes (50 mm diameter x 20 mm
high) filled with shredded sugar cane tissue. The
boxes were replaced on alternate days. The eggs
were collected by suspending the oviposition sub-
strate in 30% aqueous solution of glycerol in
which eggs floated and substrate settled down.
The average number of eggs per 3 females in each
cultivar was recorded.
The experiment was laid out in a completely
randomized block design and 3 replicates were
maintained for each treatment. For all parame-
ters, data were recorded for 2 consecutive genera-
tions. Analysis of variance was conducted with
Fisher's test (SAS 2003). Means were separated
by LSD at t5.

September 2008

Al-Ayedh: Rearing the Red Date Palm Weevil


Egg Laying Efficiency

Egg laying started 2-3 d after mating. Initially
ovipostion rate was low but progressed rapidly
and peaked after 2 weeks, remaining stable for
about a month. Thereafter, a decline was observed
in oviposition but it continued at very low rate till
death of the female. Females reared on cv. 'Suk-
kary' laid significantly greater number of eggs (F
= 2.61, df= 3, P = 0.0254) as compared to females
from other 3 cultivars (Fig. 1). Egg laying effi-
ciency of red date palm weevil is reported to be
variable on different hosts. Hopkins & Ekbom
(1999) have reported that oviposition in Cole-
optera may continue till death but the fecundity
depends on quality of the host plant. Awmack &
Leather (2002) also noted that host plant food
quality plays a major role in fecundity of herbivo-
rous insects.

Larval Growth

Larvae reared on cv. 'Sukkary' were longer (F =
9.93, df= 3, P = 0.0001) and wider (F = 14.45, df
= 3, P = 0.0001) as compared to larvae reared on
the other three cultivars (Fig. 2). There was no
significant difference (F = 0.63, df= 3, P = 0.5944)
in the length of head capsule of larvae reared on
the four cultivars, but head capsule width was
significantly greater (F = 2.96, df= 3, P = 0.0313)
for larvae reared on 'Sukkary'. Larvae reared on
'Sukkary' weighed 2.73 0.08 g and were signifi-
cantly heavier (F = 11.36, df= 3, P = 0.0001) than
those reared on 'Khalas' (2.52 g 0.08), 'Khasab'
(2.29 g 0.08) and 'Sillaj' (2.52 g 0.07) cultivars.
Quality of host plant tissue is known to have di-
rect influence on various aspects of larval develop-
ment (Leather 1990; Albert & Bauce 1994; Dodds
et al. 1996; Tammaru 1998). Salama & Abdul-
Razek (2002) reported that red date palm weevil
can be successfully reared on sugarcane and ba-
nana fruits having higher sugar contents. The
weevils also have been reared on artificial diets
where sugar is a major component (Rahalkar et al.
1978, 1985). I analyzed the stem tissue of the 4 ex-
perimental cultivars for moisture, sugar, total pro-

Fig. 1. Egg laying efficiency of red date palm weevil
females reared on different cultivars of date palm.

Fig. 2. Size of the larvae of red date palm weevil
reared on different cultivars of date palm.

tein, and major metal ion contents (data not pre-
sented). 'Sukkary' has substantially higher sugar
content as compared to the other 3 cultivars,
which might be the reason for better larval growth
on this cultivar. Survival of some of phytophagous
insects has been shown to be linked with availabil-
ity of diets rich in carbohydrates (House 1974).

Pupal Growth

Pupal length was significantly greater (F =
2.45, df = 3, P = 0.0767) when reared on cv. 'Suk-
kary' (31.50 mm 0.60) than the pupae derived
from cvs. 'Khalas' (31.10 mm 0.29), 'Sillaj' (30.29
mm 0.62), and 'Khasab' (29.67 mm 0.45).
There was no significant difference (F = 0.72, df=
3, P = 0.5453) in the width of pupae reared on the
4 cultivars. The average pupal length and width
on the different cultivars were 30.53 mm and
11.11 mm, respectively. Viado & Bigornia (1949)
recorded average pupal length and width of red
date palm weevil reared on coconut to be 33.5 mm
and 18.5 mm, respectively. In another study, Nir-
ula (1956) reported average pupal length and
width on coconut palm as 35 mm and 15 mm, re-
spectively. The variations in results between the
current and previous studies may be due to differ-
ences in food quality of the host genotypes. There
was no significant difference in pupal weight on
different cultivars (F = 1.97, df = 3, P = 0.1331)
(Fig. 3). The pupal weight and size play a signifi-
cant role in insect fecundity (Yamada & Umeya
1972; Williams 1963; Wan 1970).

Fig. 3. Weight of the red date palm weevil reared on
different cultivars of date palm.

Florida Entomologist 91(3)

Adult Growth

Male adults reared on cv. 'Sukkary' were sig-
nificantly longer (F = 4.26, df= 3, P = 0.0059) than
those reared on the other three cultivars (Fig. 4).
Females reared on 'Sukkary' and 'Khasab' were
significantly longer (F = 2.27, df = 3, P = 0.0806)
as compared to cvs. 'Khalas' and 'Sillaj'. In the
current study average length of males and fe-
males was 30.76 mm and 32.75 mm, respectively.
Viado & Bigornia (1949) recorded average body
length as 39 mm for reared males and 41.50 mm
for females. In another study, Nirula (1956) re-
ported length and width of red date palm weevil
as 35 mm and 12 mm, respectively, on coconut. In
the current study, the length and width of male
and female were slightly shorter than those re-
ported by (Viado & Bigornia 1949; Nirula 1956).
This difference may be due to variation in the
rearing environmental conditions and food
source, i.e., internal tissue of the cultivars on
which the larvae were reared.
Male width was slightly influenced by geno-
type of the cultivars (F = 3.79, df= 3, P = 0.0111).
Males reared on 'Sukkary' and 'Sillaj' were simi-
lar but significantly wider than the males devel-
oped on 'Khalas' and 'Khasab' with no significant
difference. The width of the females developed on
cvs. 'Sukkary', 'Khasab' and 'Sillaj' was greater
than those reared on 'Khalas' (F = 3.88, df = 3, P
= 0.0098) (Fig. 4). In the present study average
width of males and females reared on 4 different
cultivars were 9.87 mm and 10.25 mm, respec-
tively. Viado & Bigornia (1949) recorded average
body width as 13.40 mm and 14.60 mm for male
and female, respectively, of red date palm weevil
reared on coconut.
Significantly greater weight (F = 2.61, df = 3,
P = 0.0254) was observed for males developed on
'Sukkary' (0.81 g 0.022) as compared to those
reared on 'Sillage' (0.75g 0.018). There was no
difference in the weight gained by male adults on
'Khalas' (0.78 g 0.019), 'Khasab' (0.76 g 0.019)
and 'Sillaj'. No significant difference in weight
was observed (F = 1.29, df = 3, P = 0.2780) be-
tween female reared on the 4 cultivars. The re-
sults showed vigorous weevil growth on cv. 'Suk-
kary' as compared to those developed on other cul-

Fig. 4. Size of the adult males and females of red
date palm weevil reared on different cultivars of date

tivars of date palm. Several other authors have
also reported the effect of host plant food quality
on different aspects of adults growth and develop-
ment (Rodrigues & Moreira 2004; Calvo & Molina
2005; Awmack & Leather 2002).

Adult Lifespan

Life span of males and females depends on the
date palm cultivar (F = 6.55, df= 3, P = 0.0006) and
(F = 19.37, df = 3, P = 0.0001), respectively. Both
the sexes fed on cv. 'Khasab' presented greatest av-
erage adult life span of 176 and 172 d for males
and females, respectively. Males reared on 'Suk-
kary' showed the shortest life span while females
have the shortest life when fed on 'Sillaj' (Fig. 5).
The average male and female life span ranged
from 4-6 and 3.75-5.75 months, respectively. Our
results showed much greater longevity of red date
palm weevil as compared to previous findings (El-
Muhanna et al. 2000;Avand Faghih 1996; Viado &
Bigornia 1949; Frohlich & Rodewald 1970). Host
plant quality is considered to have significant im-
pact on longevity of adults (Leather 1990; Albert &
Bauce 1994; Dodds et al. 1996; Tammaru 1998).
Yield of cocoons per trunk piece was highest in
cv. 'Khalas' followed by 'Sillaj', 'Khasab' and 'Suk-
kary' (Table 1). Highest frequency of adult emer-
gence was noted in cv. 'Khalas' followed by 'Suk-
kary', 'Sillaj' and'Khasab'. Male to female ratio was
similar in populations reared on all 4 cultivars.
Murphy & Briscoe (1999) have noted wide
variation in developmental and other life cycle
parameters of red date palm weevil, within and
between countries, which exhibit no clear pattern
related to climate. They reported a life cycle of
48-82, 60, 60-165, 45-48, and 57-111 d from India,
Indonesia, Myanmar, The Philippines, and Iran,
Influence of host plant food quality on insect
growth and development, has been documented
by many authors (Price 1997; Speight et al. 1999;
Karowe & Martin 1989; Rossi & Strong 1991;
Yang & Joern 1994; Stockhoff 1993; Feeny 1970;
Mattson 1980; Olmstead et al. 1997; Harborne
1982, 1988; Rosenthal & Berenbaum 1991; Gange
1995; Lambert et al. 1995; Raupp 1985; Steven-
son et al. 1993; Eigenbrode et al. 1995). The
present study has shown a significant influence of

Fig. 5. Average life span of adult red date palm wee-
vil on different cultivars of date palm.

September 2008

Al-Ayedh: Rearing the Red Date Palm Weevil


Adults emergence
Variety No. of pupae % Male ratio Female ratio

'Khalas' 92.00 74.10 0.52 0.47
'Sillaj' 73.30 65.30 0.49 0.50
Khasab 66.50 59.06 0.48 0.51
Sukkary 50.16 71.91 0.51 0.48

Adult emergence % = (No. of adults emerged /Total cocoon population)* 100.
Male Ratio = Total no. of males /Total adults Population (Male + Female).
Female Ratio = Total no. of females /Total adults Population (Male + Female).

date palm tissue quality not only on food con-
sumption, survival, and development of the lar-
vae as well as weight, size, reproductive ability,
and longevity of adults. The results are in agree-
ment with the previous reports (Leather 1990;Al-
bert & Bauce 1994; Dodds et al. 1996; Tammaru
1998). In our experiments, cv. 'Sukkary' proved to
be the best overall host for rearing red date palm
weevil among the cultivars tested.


I thank Mr. Khawaja Gulam Rasool, Mr. Waleed S. Al
Waneen, and Mr. Waleed K. Al Abdulsalam for technical
assistance with this study; Dr. Norman C. Leppla from
University of Florida and Dr. Boris C. Kondratieff from
Colorado State University for reviewing and comment-
ing on the manuscript. This project was funded by
KACST through internal funding number (01-23-BM).


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Florida Entomologist 91(3)

Mier Durante et al.: New South American Neophyllaphis species


1Departamento de Biodiversidad y Gesti6n Ambiental, Universidad de Le6n, 24071 Le6n, Spain
mpmied@unileon.es; jmnien@unileon.es

2Catedra de Entomologia, Facultad de Ciencias Exactas, Fisicas y Naturales, Universidad Nacional de C6rdoba,
V61ez Sarsfield, 299, 5000 C6rdoba, C6rdoba, Argentina

'INTA EEA Junin, Casilla de Correo 78, 5570 San Martin, Mendoza, Argentina

4Laurentian Forestry Centre; Forestry Canada, 1055, rue du PEPS,
P.O. Box 10380. Sainte-Foy, Qu6bec, G1V 4C7 Canada


Two South American new species of Neophyllaphis Takahashi (Aphididae Neuquenaphidi-
nae) are described, Neophyllaphis cuschensis Nieto Nafria & Delfino sp. nov. collected on
Podocarpus glomeratus D. Don in Cusco (Peru) and Neophyllaphis iuiuyensis Mier Durante
& Ortego sp. nov. collected on Podocarpus parlatorei Pilg. in San Salvador de Jujuy (Argen-
tina). The apterous and alate viviparous females ofN. cuschensis and the apterous and alate
viviparous females, as well as the oviparous females and males of N. iuiuyensis are de-
scribed. The differences between the nominotypical subgenus and Chileaphis and also the
differences among the 4 species of subgenus Chileaphis, viz. the 2 new species and those al-
ready known and found in Chile, N. michelbacheri Essig and N. podocarpini Carrillo are

Key Words: aphids, Neophyllaphidinae, Argentina, Peru, Podocarpus


Se described dos nuevas species del g6nero Neophyllaphis Takahashi (Aphididae, Neophy-
llaphidinae) propias de la fauna de Am6rica del Sur, Neophyllaphis cuschensis Nieto Nafria
& Delfino sp. nov. recogida sobre Podocarpus glomeratus D. Don en Cusco (Peru) yNeophyl-
laphis iuiuyensis Mier Durante & Ortego sp. nov. recogida sobre Podocarpus parlatorei Pilg.
en San Salvador de Jujuy (Argentina). Se described las hembras viviparas apteras y aladas
de N. cuschensis, las hembras viviparas apteras y aladas y ademas las hembras oviparas y
los machos de N. iuiuyensis. Se exponen las diferencias entire el subg6nero nominotipico y
Chileaphis Essig y tambi6n las diferencias entire las cuatro species del subg6nero Chilea-
phis, las dos nuevas asi como las ya conocidas y propias de Chile, N. michelbacheri Essig y
N. podocarpini Carrillo.

Translation provided by the authors.

Neophyllaphidinae Takahashi, 1921, is one of
the small subfamilies of the family Aphididae La-
treille 1802 (Remaudiere & Remaudiere 1997)
and occupies a basal position in the phylogenetic
tree of the "drepanosiphids" or "drepanosiphines"
group (Quednau & Remaudiere 1994). This group
is ranked by some authors as family or subfamily,
in this last case Neophyllaphidinae is relegated to
a lower rank. It includes 1 genus, Neophyllaphis
Takahashi, 1920, currently formed by 13 species
specialized in living on Podocarpaceae or Arau-

cariaceae. N. araucariae Takahashi, 1937, feeds
on several species of Araucaria and possibly na-
tive to Australia; the others live on species of
Podocarpus L'Her. ex Pers., 1807, though N. mich-
elbacheri (Essig, 1953) was described on Pilgero-
dendron uviferum (D. Don) Florin, 1930 (Cupres-
saceae) and N. brimblecombei Carver, 1971, has
been recorded on Eucalyptus robusta Sm., 1795
(Myrtaceae) in China (Essig 1953; Remaudiere &
Remaudiere 1997; Blackman & Eastop 1994;
Quiao et al. 2001).

Florida Entomologist 91(3)

Some authors (Russell 1966, 1982; Blackman
& Eastop 1994) consider the genus to be homoge-
nous and have not divided it into subgenera. Oth-
ers (Hille Ris Lambers 1967; Carrillo 1980; Re-
maudiere & Remaudiere 1997; Quiao et al. 2001)
divided it into 2 subgenera, the nominotypical one
and Chileaphis Essig, 1953, with 11 and 2 species,
respectively. We agree with the second treatment
because the 2 known species of Chileaphis: N.
michelbacheri and N. podocarpini Carrillo, 1980,
and the 2 described below differ from the 11 spe-
cies in the nominotypical subgenus because: (1)
eyes are multifaceted in all forms, apterous vivip-
arous females included; (2) membranous distal
part of the caudal knob is differentiated from the
sclerotized proximal part in the apterous vivipa-
rous females; (3) basal width of the cauda is simi-
lar to the maximum width of the caudal knob in
the apterous viviparous females; and (4) the first
abdominal segment has a marginal process (or
small tuberculiform sclerite) in the alate vivipa-
rous females (and other alate forms).
The division into 2 subgenera coincides well
with the geographical distribution of the species,
without considering the introduction of N. arau-
cariae into the United States, Mexico and Costa
Rica, or N. podocarpi Takahashi, 1920, into North
America and Italy. The 2 known species of Chilea-
phis and those described below are found in South
America, while 2 of the species in subgenus Neo-
phyllaphis inhabit Africa and the other 9 species
are in different parts of the Asiatic-Australian
arch of the Pacific or neighboring areas (Limonta
1990; Blackman & Eastop 1994; Qiao et al. 2001).
Neophyllaphis michelbacheri was recorded on
Pilgerodendron uviferum and Podocarpus nubige-
nus Lindl., 1807; the latter may well be the habit-
ual host of the species, whereas the former may be
a stray record. In fact, Essig (1953) states "This
interesting new genus and species was collected
by Dr. A. E. Michelbacher by beating the avail-
able limbs and foliage of the Chilean cypress,
Pilgerodendron uviferum". The leaves of both
plants are relatively similar and, above all, they
share the same habitat. Podocarpus nubigenus is
included in the section Australis de Laub, 1985, in
the genus Podocarpus Labill., 1806, and is distrib-
uted throughout regions IX, X, XI, and XII in
Chile and the Argentinean provinces of Neuquen,
Rio Negro, and Chubut, this being the largest
foreseeable distribution area of the aphid-species.
The distribution area of Pilgerodendron uviferum
partly overlaps the fore-mentioned one, but is
more southern, and includes regions X, XI, and
XII in Chile and some areas of Neuquen, Rio Ne-
gro, Chubut, Santa Cruz, and Tierra del Fuego in
Argentina (Missouri Botanical Garden 1999; In-
stituto de Botanica Darwinion 2007; Stark Schill-
ing & Le-Quesne Geir 2007).
Neophyllaphis podocarpini lives on Podocar-
pus salignus D. Don, 1824. This plant-species is

included in the section Capitularis de Laub, 1985,
and has a more northern distribution area than
the other plants, in regions VII, VIII, IX, and X in
Chile, and has not been reported in Argentina
(Missouri Botanical Garden 1999; Instituto de
Botanica Darwinion 2007; Stark Schilling & Le-
Quesne Geir 2007). This could also be the maxi-
mum distribution area of the species.
During 2 visits in Cusco some of the authors
(D., M. D., and N. N.) had the opportunity to col-
lect Neophyllaphis on young cultivated specimens
of Podocarpus glomeratus D. Don, 1824. This
plant-species is included in the section Capitu-
laris and is found in Peru (Cusco area included),
Bolivia, and Ecuador (Missouri Botanical Garden
1999). Some colleagues later contributed more
specimens collected on the same plant.
Encouraged by the discovery of what could be
the third species in subgenus Chileaphis during an
expedition in northwest Argentina by some of the
authors (M. D., 0., and N. N.) in Nov 2006, a visit
to localities where Podocarpus parlatorei Pilg.,
1903, grows was planned. We collected aphids on
this plant in San Salvador de Jujuy. The plant-spe-
cies is also included in the section Capitularis and
is found in south Bolivia as well as the provinces of
Jujuy, Salta, Catamarca, and Tucuman in north-
western Argentina (Missouri Botanical Garden
1999; Instituto de Botanica Darwinion 2007).


The aphids were measured by the method nor-
mally used in our studies (Nieto Nafria & Mier
Durante 1998), only differing from Blackman &
Eastop (1994) in how the body length is mea-
sured: viz. from the front to the tip of the cauda or
the posterior edge of the anal plate, respectively.
Abbreviations used in the text, tables, and fig-
ure captions are as follows: AntI, AntII, AntIII,
AntIV, AntV, AntVIb, AntVIpt are antennal seg-
ments I to V plus base and processus terminalis of
antennal segment VI, respectively; D is the basal
diameter of AntIII; AbdI to AbdVIII are abdomi-
nal segments I to VIII. Values in parenthesis are
exceptional values.


Based on information obtained from Blackman
& Eastop (1994), Essig (1953), Carrillo (1980),
Hille Ris Lambers (1970), and Russell (1982), and
from data obtained in a study of several samples
of N. (C.) michelbacheri and N. (C.) podocarpini,
we can state that the specimens from Cusco and
San Salvador de Jujuy belong to 2 new species to
be included in the subgenus Chileaphis: Neophyl-
laphis (Chileaphis) cuschensis and Neophyllaphis
(Chileaphis) iuiuyensis, described below. The spe-
cific names are adjective in the feminine gender
because Neophyllaphis is feminine, as is Aphis,

September 2008

Mier Durante et al.: New South American Neophyllaphis species

referred to inhabitants of Cusco and Jujuy, lati-
nized as usual in the respective catholic dioceses.
These 2 species and N. podocarpini differ from
N. (C.) michelbacheri because in the apterous vi-
viparous females alatee forms are not known in
this species): (1) antennae are at least 0.5 times
body length; (2) maximum width of head, includ-
ing eyes, at most is 1.7 times AntIII; and (3) setae
on dorsum of abdomen (or at least most of them)
at least 10 pm. Their host-species belong to differ-
ent sections of the genus Podocarpus.
Neophyllaphis (C.) iuiuyensis differs from N.
podocarpini and from N. (C.) cuschensis because
(1) membranous distal part of the caudal knob in
apterous viviparous females is small, contrasting
very little with the rest of the knob; (2) the dor-
sum of head, antennae, legs, and siphunculi are
very weakly pigmented in apterous viviparous fe-
males; (3) setiferous sclerites are absent from dor-
sum of abdomen in apterous viviparous females;
and (4) they are green when alive. It differs from
N. podocarpini because the alatae do not have
dorsal setae on the first tarsal segments.
Neophyllaphis (C.) cuschensis is distinguished
from N. (C.) podocarpini because: (1) interseg-
mental sclerites are clearly visible; (2) siphunculi
relatively prominent with wide basal diameter;
(3) setae usually absent from dorsum of caudal
knob; and (4) first tarsal segments without dorsal
setae in alatae.

Neophyllaphis (Chileaphis) cuschensis Nieto Nafria and
Delfino, sp. nov.

Apterous Viviparous Females (Fig. 1). Based
on 73 specimens, 38 measured. Body length 1.47-
2.25 mm and 1.9-2.7 times hind tibia. Brown
when alive, including legs and antennae; thorax
and abdomen covered in ash-colored waxy pow-
der, more or less intensively and extensively. Se-
tae generally short, robust, and pointed; ventral
setae longer than respective dorsal ones; a few se-
tae present on dorsum of Abd8 and cauda much
longer and finer than the rest (Table 1).
Head (Fig. 1B) well-pigmented (dark brown
when mounted) bearing numerous dorsal setae
(no fewer than 15). Its maximal width, including
eyes, 1.2-1.7 times AntIII. Front gently convex.
Compound eyes complete. Antennae 0.83-1.28
mm and 0.5-0.7 times body; proximal segments as
pigmented as head becoming darker towards
apex, though proximal 1/2 AntIII can be slightly
less pigmented (Fig. 1A). AntI, AntII and approx-
imately ventral 1/4 and dorsal 1/2-3/4 AntIII
smooth; rest of antenna spinulose (Fig. 1C). An-
tIII 0.27-0.46 mm, 1.6-2.3 times AntIV (0.12-0.21
mm) and 1.6-2.2 AntVI. AntV 0.15-0.23 mm.
AntVIb 0.14-0.19 mm and 4.6-10.0 times AntVIpt
(0.02-0.03 mm). Rostrum surpassing middle
coxae, even reaching hind coxae. Apical rostral
segment pigmented (Fig. 1D) as anterior segment

Fig. 1. Neophyllaphis (Chileaphis) cuschensis Nieto
Nafria & Delfino sp. nov., apterous viviparous female.
A, habitus. B, head and prothorax, dorsal and ventral
views. C, antenna. D, apical rostral segment. E, tarsus
of hind leg. F, siphunculus. G, cauda and anal plate, dor-
sal view.

and head, 0.08-0.11 mm, 0.4-0.5 times hind sec-
ond tarsal segment; with edges almost parallel,
some scattered spinules and few complementary
setae (4 at most).
Prothorax (Fig. 1B) separated from head by
pale, fine, irregular line and normally as pig-
mented as head; marginal papillae absent. Legs
completely pigmented, like head, most proximal
part of tibia almost black. Hind femur and tibia
0.39-0.58 and 0.63-0.93 mm, respectively. First
segment of tarsi with several ventral setae, lack-
ing dorsal setae (Fig. 1E). Second segment of hind
tarsi 0.18-0.23 mm.
Mesothorax to AbdVII bearing dorsal setifer-
ous sclerites; marginal sclerites of AbdVII and
sometimes those on mesothorax may be joined to-
gether. Stigmatic sclerites well-pigmented. Si-
phunculi almost as dark as head, with one proxi-
mal part tapering, 0.06-0.14 mm basal width x
0.02-0.04 mm high, bearing 10 to 19 setae, and
one distal part cylindrical, 0.01-0.02 mm high x
0.03-0.06 mm wide at opening, with spines annu-
larly ranged (Fig. 1F). AbdVIII dorsum with wide
transversal band, almost as pigmented as head,
with anterior spinal notch and bearing spines in

Florida Entomologist 91(3)


Neophyllaphis cuschensis Neophyllaphis iuiuyensis

Longest setae on AP AL AP AL OV M

Head, dorsal [uim] 18-30 18-25 18 15-18 15-20 13-18
Head, dorsal [times D] 0.8-1.2 0.7-1.0 0.4-0.6 0.5-0.7 0.5-0.7 0.4-0.6
AntIII [Im] 10-23 13-18 15-20 10-15 10-15 13-15
AntIII [times D] 0.5-0.9 0.5-0.8 0.4-0.7 0.3-0.5 0.4-0.6 0.4-0.6
Hind femur, dorsal [uim] 18-28 18-20 15-20 15-20 15-20 15-18
Hind femur, dorsal [times D] 0.8-1.0 0.8-1.0 0.4-0.7 0.5-0.7 0.5-0.7 0.5-0.6
Hind femur, ventral [uim] (15) 25-40 20-33 30-35 23-40 28-38 30-35
Hind femur, ventral [times D] 0.6-1.8 0.8-1.5 0.7-1.0 0.7-1.3 0.9-1.3 1.0-1.2
AbdIII, dorsal [uim] 10-25 10-23 10-16 18-23 13-23 15-18
AbdIII, dorsal [times D] 0.4-1.0 0.4-1.0 0.3-0.4 0.5-0.8 0.4-0.8 0.5-0.6
AbdVIII, dorsal short [uim] 5-15 10-18 13-15 10-20 13-18 15-18
AbdVIII, dorsal short [times D] 0.2-0.6 0.4-0.9 0.3-0.4 0.3-0.7 0.4-0.6 0.5-0.6
AbdVIII, dorsal long [uim] 13-55 30-40 35-45 34-50 40-45 33-38
AbdVIII, dorsal long [times D] 0.6-2.4 1.2-1.8 0.8-1.2 1.1-1.7 1.3-1.6 1.0-1.3
Cauda, short [I'm] 8-25 20-33 20-23 20-23 13-23
Cauda, short [times D] 0.3-1.3 0.8-1.2 0.5-0.7 0.7-0.8 0.5-0.8
Cauda, long [jim] 35-65 (75) 55-60 46-60 48-58 38-53
Cauda, long [times D] 1.4-2.7 (3.0) 2.2-2.5 1.1-1.9 1.6-2.0 1.3-1.9

transversal lines and approximately not aligned
20 setae of different lengths, short and long (Table
1). Setiferous sclerites on ventral side of abdomen,
much smaller than dorsal ones. Genital and anal
plates dark. Cauda (Fig. 1G) 0.12-0.26 mm, 1.1-
1.7 times its basal width, with two easily distin-
guishable parts separated by strongly pronounced
waist; proximal part tapering, 0.08-0.14 mm and
0.4-0.5 times total length of cauda; terminal knob
subspherical (0.8-1.4 times as wide as long), 0.7-
1.3 times as wide as basal width of cauda and 2.7-
5.3 times as wide as the waist; distal part of ter-
minal knob (to 1/2 its length) much less and pro-
gressively sclerotized and pigmented than proxi-
mal part; it has (20)26-37(43) setae of various
lengths, short and long (Table 1), 2-7(10) dorsal.
Alate Viviparous Females (Fig 2). Based on 8
specimens. Body length 2.05-2.38 mm and rela-
tively similar to apterous viviparous females, so the
following includes differentiating characters only,
excluding obvious differences in the sclerotization
of the thorax and presence of wings (Fig. 2A).
Maximal width of head, including eyes, 0.7-0.8
times AntIII. Antennae 1.53-2.02 mm and 0.7-0.8
times body, as pigmented as head. Entire length
of dorsal face ofAntIII spinulate, 80-90% of length
of ventral face bearing 31-50 narrow transverse
secondary sensoria (Fig. 2C), the biggest being
semi-annular. AntIII 0.59-0.75 mm, 1.8-2.1 times
AntIV (0.30-0.43 mm) and 2.8-3.2 AntVI. AntV
0.28-0.38 mm. AntVIb (Fig. 2D) 0.18-0.23 mm and
6.0-9.5 times AntVIpt (0.02-0.04 mm). Apical ros-
tral segment 0.10 mm. Prothorax with a pig-
mented dorsal arch and without marginal papil-

lae (Fig. 2B). Legs as dark as head. Hind femur
and tibia 0.63-0.68 and 1.01-1.11 mm, respec-
tively. Second segment of hind tarsi 0.21-0.23 mm
(Fig. 2F). Stigmatic sclerites and intersegmental
sclerites present. AbdI bearing marginal plates
with a small tuberculiforme projection (Fig. 2A
detail) and trapezoid spinal plate with uneven
edges; all bearing spinules more or less set out in
lines. AbdII-AbdVII with spinal plate going from
trapezoidal to rectangular in shape, and some se-
tiferous pleural and marginal sclerites, that may
be joined together. AbdVIII with transverse band,
also spinulate. Small scattered setiferous scler-
ites present on ventral side of abdomen. Siphun-
culi 0.11-0.16 mm with at base and with 10-19 se-
tae (Fig. 2G). Cauda 0.18-0.22 mm long proximall
part 0.08-0.12 mm), 1.7-2.0 times its width at
base, maximal width of knob 0.8-1.0 times its
length and 2.0-2.7 times width the waist; mem-
branous terminal area absent or very small and
dorsally placed (Fig. 2H); with 21-30(36) setae
(Table 1), 1-4 of which at most dorsal (Fig. 21).
Bionomics and Distribution. The specimens
form compact groups on young branches of
Podocarpus glomeratus (in Peru named "romer-
illo" and "intimpa") and, to date, only are known
in Cusco (Peru). Its distribution very likely coin-
cides with that of the host plant, which also is
found in Ecuador and Bolivia. The presence of
oviparous females, which we were unable to study
due to their deteriorated state of preservation,
confirms that it is holocyclic.
Types. The holotype is the apterous viviparous
female "PER-23, ap 15" collected on Podocarpus

September 2008

Mier Durante et al.: New South American Neophyllaphis species

I ________________


Fig. 2. Neophyllaphis (Chileaphis) cuschensis Nieto Nafria & Delfino sp. nov., alate viviparous female. A, habi-
tus. B, head (dorsal and ventral views) and thorax (dorsal). C, AntIII, with detail. D, AntVI. E, apical rostral seg-
ment. F, tarsus of hind leg. G, siphunculus. H, cauda and anal plate, dorsal view. I, gonapophisis, anal plate and
cauda, ventral view.

glomeratus in Cusco, 26-VII-2006, Mier Durante
and Nieto Nafria leg., deposited in the collection
of the University of Le6n (Le6n, Spain). The
paratypes are 29 apterous viviparous females and
4 alate viviparous females collected with the holo-
type; 35 apterous viviparous females collected at
the same plant and place, 14-II-2007, A. Ro-
driguez Berrio and I. Tisoc Duenas leg.; and 8
apterous viviparous females and 4 alate vivipa-
rous females collected on same plant in Cusco at
the University Campus, 22-XI-1994, Delfino leg.;
they are deposited in the collections of the au-
thors, "Klauss Raven BMller" Museum of Ento-
mology, Universidad Nacional Agraria La Molina
(Lima, Peru), U. S. National Museum of Natural
History, Smithsonian Institution aphid collection
(Washington D.C., U.S.A.), Museum national
d'Histoire naturelle (Paris, France) and The Nat-
ural History Museum (London, U.K.).

Neophyllaphis (Chileaphis) iuiuyensis Mier Durante
and Ortego sp. nov.

Apterous Viviparous Female (Fig. 3). Based on
7 specimens. Body length: 1.72-2.50 mm and 2.0-
2.8 times hind tibia. When alive, pale green with
longitudinal bands of varying intensity, pale-yel-
lowish legs and antennae; thorax and abdomen
very lightly covered in ash-colored waxy powder.
Setae generally short (Table 1), relatively thick
and pointed; ventral ones longer than the respec-
tive dorsal ones; a few of the setae on the dorsum
of Abd8 and on the cauda much longer and finer
than the rest.
Head (Fig. 3A) delicately pigmented (pale
creamy yellow when mounted) with numerous
dorsal setae (no fewer than 15). Its maximal
width, including eyes, 1.1-1.5 times AntIII. Front
gently convex. Compound eyes complete. Anten-

If 1 4 1
%it~ .t?
I ~ t

~ t p

'~' L'

Fig. 3. Neophyllaphis (Chileaphis) iuiuyensis Mier
Durante & Ortego sp. nov., apterous viviparous female.
A, habitus. B, head and prothorax, dorsal and ventral
views, with details. C, antenna. D, apical rostral seg-
ment. E, tarsus of hind leg. F, siphunculus. G, cauda and
anal plate, dorsal view.

nae 1.22-1.41 mm and 0.5-0.7 times body, basally
pigmented as head and getting darker towards
apex. AntI, AntII and approximately ventral 1/6
and dorsal 1/2-3/4 of AntIII smooth (Fig. 3C); rest
of the antenna spinulate. AntIII 0.38-0.49 mm,
1.6-1.8 times AntIV (0.24-0.28 mm) and 1.9-2.4
AntVI. AntV 0.24-0.28 mm. AntVIb 0.15-0.19 mm
and 4.3-6.0 times AntVIpt (0.02-0.04 mm). Ros-
trum surpassing middle coxae. Apical rostral seg-
ment (Fig. 3D) 0.09-0.10 mm, 0.3-0.4 times second
segment of hind tarsi, pigmented as anterior seg-
ment and somewhat more than head, with edges
almost parallel, some scattered spinules and few
complementary setae (no more than 4).
Legs slightly more pigmented than head, tarsi
slightly darker. Hind femur and tibia 0.58-0.62
and 0.82-0.91 mm, respectively. First segment of
tarsi (Fig. 3E) with several ventral setae and
without dorsal setae. Second segment of hind
tarsi 0.24-0.27 mm.
Prothorax separated from head by fine, pale ir-
regular line, somewhat enlarged on both sides;
marginal papillae absent (Fig. 3B). Body dorsum

September 2008

completely pale, frequently no sclerites discern-
ible. Siphunculi (Fig. 3F) slightly paler than head,
but distinguishable from outline, its proximal
part wide and tapering (0.10-0.13 mm basal width
x 0.02-0.03 mm high) and with 10 to 16 setae, the
cylindrical distal part 0.01 mm high and 0.04-0.06
mm wide at the opening. AbdVIII dorsum with
about 10-15 setae, not aligned and varying in
length, short and long (Table 1). Genital and anal
plates more or less as pigmented as head. Cauda
0.08-0.19 mm, 0.9-1.1 times its basal width, with
two easily distinguishable parts separated by a
waist (Fig. 3G); proximal part tapering, 0.07-0.09
mm and 0.4-0.5 times total length of cauda; knob
subspherical (maximal width 1.2-1.7 times its
length), 0.8-1.2 times as wide as basal width of
cauda and 2.2-3.0 times width the waist. Distal
part of knob (to 1/2 its length) less sclerotized
than proximal part and even less pigmented.
Cauda with 16-30 setae of different lengths, short
and long (Table 1) with 0-4 dorsal; membranous
part lacking setae except for areas near the limit
with sclerotized part.
Alate Viviparous Females (Fig. 4 A-H). Based
on 40 specimens. Body length 1.94-2.85 mm and
relatively similar to apterous viviparous females,
so the following contains differentiating charac-
ters only, excluding the obvious differences in
sclerotization on the thorax and the presence of
wings (Fig. 4A).
When alive, head and thorax brown, abdomen
green with longitudinal brown band, both covered
in light ash-colored powder; legs and antennae
brown. Maximum width of head, including eyes,
0.7-0.8 times AntIII. Antennae 1.66-2.16 mm and
0.7-0.9 times body, as pigmented as head. Entire
length of dorsal face of AntIII spinulate, 31 to 64
narrow transverse secondary sensoria (the big-
gest being semi-annular) occupying 79-91%
length of ventral face (Fig. 4C). AntIII 0.58-0.76
mm, 1.3-1.8 times AntIV (0.33-0.44 mm) and 2.5-
3.2 AntVI. AntV 0.34-0.44 mm. AntVIb 0.17-0.24
mm and 5.2-7.7 times AntVIpt (0.03-0.04 mm)
(Fig. 4D). Apical rostral segment 0.08-0.10 mm.
Prothorax (Fig. 4B) with spinal-pleural plate and
2 marginal plates; marginal papillae absent. Legs
more or less homogenously brown. Hind femur
and tibia 0.68-0.81 and 1.04-1.26 mm, respec-
tively. Second segment of hind tarsi 0.22-0.28 mm
(Fig. 4F). Stigmatic sclerites present and inter-
segmental sclerites absent. AbdI bearing mar-
ginal plates with small tuberculiform projection
(Fig. 4A detail) and a trapezoidal spinal-pleural
plate with very uneven edges; AbdII-(AbdIV)Ab-
dVI(AbdVII) with spinal or spinal-pleural plates;
all exhibiting spinules more or less set out in
lines; if plates absent setiferous spinal sclerites
are present and sometimes partially joined to-
gether. AbdVIII with a narrow transverse band,
spinulose. In addition small scattered setiferous
sclerites also sometimes present on dorsum and

Florida Entomologist 91(3)

Mier Durante et al.: New South American Neophyllaphis species

" D'.tii i
D iftii^

Fig. 4. Neophyllaphis (Chileaphis) iuiuyensis Mier Durante & Ortego sp. nov., alate viviparous female (A-H)
and alate oviparous female (I-J). A, habitus. B, head (dorsal and ventral views) and thorax (dorsal). C, AntIII, with
detail. D, AntVI. E, apical rostral segment. F, tarsus of hind leg. G, siphunculus. H, apex of the abdomen, ventral
view. I, tibia of hind leg. J, apex of the abdomen, ventral view, with details.

ventral side of abdomen. Siphunculi 0.08-0.15
mm diameter at base and with 6-16 setae (Fig.
4G). Cauda 0.14-0.19 mm long proximall part
0.06-0.09 mm) and 0.8-1.1 times width at base,
maximal knob width 1.0-1.7 times its length and
1.3-2.4 times waist width (Fig. 4H); terminal
membranous area absent or very small; it has 10-
26 setae, 0-4 of which dorsal (Table 1).
Oviparous Alate Females (Fig. 4 I-J). Based on
11 specimens. Body length 2.20-2.53 mm and sim-
ilar to apterous viviparous females, so only the
differential characters are given below.
Antennae 1.86-2.15 mm and 0.7-0.9 times body.
AntIII 2.4-3.1 times AntVI with 21-41 secondary
sensoria not reaching distal 11-21% of segment,
AntIV and AntV 0.41-0.49 and 0.38-0.45 mm, re-
spectively. Apical rostral segment 0.07-0.10 mm.
Legs brown but less pigmented than head, espe-
cially coxae, trochanters and proximal part of fem-
ora. Hind tibiae (Fig. 41) not expanded and bear-

ing 20-55 scent plates (very exceptionally, 1 leg of
1 specimen with 1). Middle tibiae sometimes with
scent plates, 5 at most. AbdVIII with setiferous
sclerites separated or grouped together. Genital
plate very large, transversal-rectangular, pale in
median-anterior region and almost black in an
sinuate posterior part, with small club-shaped se-
tae at the edge (Fig. 4J). Cauda with semicircular
outline, with microsculpture developed at a dif-
fused reticulate pattern, a small, strongly sca-
brous protruberance on lower-side of cauda and
approximately 20 to 60 setae (Fig. 4J).
Alate Males. Based on 9 specimens. Body length
2.00-2.35 mm, relatively similar to alate vivipa-
rous females, so only differential characters are
given below, excluding the obvious differences such
as genital plate absent and male genitalia present.
Maximum width of head, including eyes, 0.6-
0.7 times AntIII. Antennae 2.10-2.30 mm and 1.0-
1.1 times body. AntIII, AntIV, AntV and AbtVIb,

Florida Entomologist 91(3)

respectively 0.73-0.84, 0.47-0.53, 0.44-0.48, 0.23-
0.26 and 0.04 mm long with 68-96, 13-25, 11-20
and 2-5 almost rectangular secondary sensoria set
out transversely. Hind femur and tibia 0.68-0.81
and 1.04-1.26 mm, respectively. Second segment
of hind tarsi 0.22-0.27 mm. Cauda 0.12-0.16 mm
long, with its proximal part 0.05-0.07 mm long.
Bionomics and Distribution. The species forms
compact groups on young branches of the conifer
Podocarpus parlatorei (named "pino del cerro" in
Argentina) and is currently known only in San
Salvador de Jujuy (Jujuy, Argentina). Its distribu-
tion may well coincide with that of the host plant,
which is present in several Argentinean provinces
and Bolivia. The species is holocyclic.
Types. The holotype is the apterous viviparous
female "ARG-1197, ap 5", collected on Podocarpus
parlatorei in San Salvador de Jujuy, Los Perales
Botanic Park, 8-XI-2006, Mier Durante, Ortego
and Nieto Nafria leg., deposited in the collection
of the University of Le6n (Le6n, Spain). The
paratypes are 6 apterous viviparous females, 40
alate viviparous females, 11 alate oviparous fe-
males and 9 alate males, collected with the holo-
type and deposited in the collections of the au-
thors and in the "Klauss Raven Btiller" Entomol-
ogy Museum, Universidad Nacional Agraria La
Molina (Lima, Peru), U.S. National Museum of
Natural History, Smithsonian Institution aphid
collection (Washington D.C., U.S.A.), Museum na-
tional d'Histoire naturelle (Paris, France) and
The Natural History Museum (London, U.K.).


We thank A. Rodriguez Berrio (Universidad Nacio-
nal Agraria La Molina, Lima, Peru), Isolina Tisoc
Duenas (Universidad Nacional San Antonio Abad,
Cusco, Peru) for help in collecting specimens; and C.
Vergara Cobidn and G. Miller for providing specimens
of the collections of the "Klauss Raven Btiller" Entomo-
logical Museum, Universidad de La Molina, and the Na-
tional Museum of Natural History of Washington,
respectively. This paper was funded by the Regional
Government of Castilla y Le6n, Spain (grants LE45/02
and LE034A05 to Prof. M. P. Mier Durante).

BLACKMAN, R. L., AND V. F. EASTOP. 1994. Aphids on
the World's Trees. An Identification and Information
Guide. CAB International, Wallingford. 8 + 988 pp.,
16 plates.
CARRILLO, R. 1980. Aphidoidea de Chile III. Agro Sur
8(1): 21-29.
ESSIG, E. 1953. Some new and noteworthy Aphididae
from Western and Southern South America. Proc.
Californian Acad. Sci. (Fourth Series) 28(3): 59-164.
Reptiblica Argentina. Catalogo de Plantas Vascu-
lares. On line: http://www.darwin.edu.ar. Consulted
April 2, 2007.
HILLE RIS LAMBERS, D. 1967. On the genus Neophylla-
phis Takahashi, 1920 (Homoptera Aphididae) with
descriptions of two new species. Zoologische Med-
edelingen 42(7): 55-66.
LIMONTA, L. 1990. Callaphididae (Aphidoidea) nuovi
per l'Italia. Bolletino di Zoologia Agraria e di Bachi-
coltura, Serie II 22(1): 93-99.
line: http://www.tropicos.org/. Consulted April 2, 2007.
Hemiptera. Aphididae. I. In M. A. Ramos et al. [eds.]
Fauna Ib6rica, vol. 11. Museo Nacional de Ciencias
Naturales (CSIC), Madrid. 424 pp.
QIAO, G., G. ZHANG, AND Y. CAO. 2001. The genus Neo-
phyllaphis Takahashi (Homoptera: Aphididae) from
China with description of one new species. Oriental
Insects 35: 91-96.
QUEDNAU, F. W., AND G. REMAUDIERE. 1994. Le genre
sud-am6ricain Neuquenaphis E. E. Blanchard, de-
scription de deux nouvelles especes et definition de
nouvelles sous-familles d'Aphididae (Homoptera).
Bull. Soc. Entomol. France 99(4): 365-384.
des Aphididae du Monde/Catalogue of the World's
Aphididae. Homoptera Aphidoidea. Institute Na-
tional de la Recherche Agronomique. Paris. 473 pp.
RUSSELL, L. M. 1966. New synonymy and new name
combination in Neophyllaphis Takahashi. Proc. En-
tomol. Soc. Washington 68(4): 340.
RUSSELL, L. M. 1982. The genus Neophyllaphis and its
species (Hemiptera: Homoptera: Aphididae). Florida
Entomol. 65(4): 538-572.
Enciclopedia de la flora chilena. On line http://
www.florachilena.cl. Consulted Abril 2, 2007.

September 2008

Mier Durante & Ortego: A New Species of Uroleucon


1Departamento de Biodiversidad y Gesti6n Ambiental, Universidad de Le6n, 24071 Le6n, Spain
2INTA EEA Junin, Casilla de Correo 78, 5570 San Martin, Mendoza, Argentina

We describe the apterous and alate viviparous females of a new species of the subgenus
Lambersius of the genus Uroleucon (Hemiptera Aphididae) from Argentina living on
Adesmia (Fabaceae). Uroleucon adesmiae sp. n. is the first South American species of this
genus exclusively living on a species of Fabaceae. A previous key for the apterous viviparous
females of the Uroleucon species recorded in South America is modified to include the new

Key Words: aphids, Aphididae, Uroleucon, Argentina, Fabaceae

Se described las hembras viviparas apteras y aladas de una nueva especie del subg6nero
Lambersius del g6nero Uroleucon (Hemiptera Aphididae) de Argentina que vive sobre
Adesmia (Fabaceae). Uroleucon adesmiae sp. n. es la primera especie del g6nero que vive ex-
clusivamente sobre una especie de Fabaceae. Se modifica una anterior clave de identificaci6n
de las hembras viviparas apteras de Uroleucon citadas de Am6rica del Sur para incluir en
ella la nueva especie.

Translation provided by the authors.

Uroleucon Mordvilko, 1914, is one of the larg-
est genera of Macrosiphini, which currently in-
cludes 224 valid species (Blackman & Eastop
2006; Nieto Nafria et al. 2007). Asteraceae species
are habitually cited as the host-plant of Uroleu-
con, but 9 species were described on Campanu-
laceae species, and another 10 species were de-
scribed on species belonging to Balsaminaceae,
Convolvulaceae, Lamiaceae, Malvaceae, Onagra-
ceae, Polygonaceae, and Scrophulariaceae. Very
few Uroleucon species have been recorded on
plant species that do not belong to Asteraceae. Re-
maudiere et al. (1985) recorded Uroleucon com-
positae Theobald, 1915, from Africa on 77 species
belonging to 15 families, including 2 species of
Fabaceae,Albizzia petersiana Oliv. andDalbergia
sp. Blackman & Eastop (2006) recorded this
polyphagous aphid also on the Fabaceae Erio-
sema psoraleoides (Lam.) G. Don and with doubts
on Lathyrus sativus L.
On an unidentified species of Adesmia (Fa-
baceae) in the Chubut province (Argentina) one of
the authors (J. Ortego) collected a sample of
aphids belonging to tribe Macrosiphini (Aphid-
idae) and to the group of Macrosiphum Passerini,
1860, and related genera, which is characterized
by the reticulated ornamentation on the distal
part of siphunculi. It is an unnamed species of
Uroleucon and is described below.

Abbreviations used in the text and figure cap-
tions are as follows: AntI, AntII, AntIII, AntIV,
AntV, AntVIb, AntVIpt are antennal segments I to
V plus base and processus terminalis of antenna
segment VI, respectively; D is the basal diameter
ofAntIII; Urs is the ultimate rostral segment; Ht2
is the second segment of hind tarsus; AbdI to Ab-
dVIII are abdominal segments I to VIII. Values in
parenthesis are exceptional values.

Uroleucon adesmiae sp. nov.

Types. Holotype: Apterous viviparous female
(ARG-1042, ap. 8), Rio Mayo, Chubut, Argentina,
13-XII-2004, on Adesmia sp. J. Ortego leg.
Paratypes: 7 apterous viviparous females and 1
alate viviparous female, same data. Holotype and
several paratypes in the collection of the Univer-
sity of Le6n, Le6n, Spain; other paratypes in col-
lections of J. Ortego, the Natural History Mu-
seum, London, United Kingdom, and Museum na-
tional d'Histoire naturelle, Paris, France.
Etymology. The specific name is the genitive of
Adesmia, the name of the plant-host genus.
Apterous Viviparous Females. Fig. 1. Based on
8 specimens. Color in life dark green to brownish
green. Body length 2.35-2.95 mm. Mounted spec-
imens more or less pale yellowish, except the most
apical part of tibiae, tarsi, 2 last rostral segments,

Florida Entomologist 91(3)

Fig. 1. Uroleucon (Lambersius) adesmiae Mier Durante and Ortego sp. nov., apterous viviparous female. A,
habitus. B, AntIII. C, AntVI. D, cauda. E, Ht2. F, siphunculus.

AntVIpt and apices ofAntIII-AntVIb. Dorso-ceph-
alic setae numerous (7-11 setae present behind
the 2 discal setae), pale, 45-50 pm, 1.1-1.7 times
D, thick with apex blurred or slightly clavate, sim-
ilar in shape to antennal, femoral, tibial, dorsal-
thoracic and dorsal-abdominal ones. Antennae
2.62-2.96 mm and 0.9-1.1 times body length;AntI-
AntIII smooth; AntIII 0.60-0.75 mm, with 4-11
small circular secondary sensoria, ventrally
aligned on proximal 30-58% of its length; setae
30-45 pm, 0.9-1.4 times D; AntIV and AntV, re-
spectively, 0.55-0.69 and 0.43-0.53 mm. AntVIpt
0.58-0.64 mm, 0.9-1.0 times AntIII and 3.8-4.1

times AntVIb, which is 0.15-0.18 mm. Rostrum al-
most reaching hind coxae. Urs 0.15-0.18 mm, 2.7-
3.4 its basal width, 0.9-1.1 times both AntVIb and
Ht2; with 10-13 secondary setae. Pale coxae. Hind
femur and tibia, respectively, 0.81-0.99 and 1.50-
1.70 mm. First tarsal segments with (4) 5 setae.
Ht2 0.15-0.17 mm. Marginal papillae and setifer-
ous sclerites absent; intersegmental and stig-
matic sclerites unpigmented. Dorsal setae on
AbdI-AbdV abundant (19-30 in all, 3-6 marginal
each side, 7-20 spinal-pleural), the spinal ones are
40-45 pm long, 1.0-1.5 times D. Ventral-abdomi-
nal setae thin and pointed, 24-50 per segment.

September 2008

Mier Durante & Ortego: A New Species of Uroleucon

Siphunculus cylindrical with enlarged base (1.6-
2.5 times the width at the beginning of the retic-
ulation) and apical flange scarcely developed;
0.66-0.85 mm, 2.1-2.4 (2.8) times cauda and 1.0-
1.2 times AntIII; with rather ill-defined reticula-
tion on 8.0-13.4 (14.3)% its length, with approxi-
mately 20-40 cells; wrinkly basal to reticulations,
but nearly smooth at base. Genital plate with 2-4
discal and 11-17 small posterior setae. AbdVIII
with 6-12 setae. Cauda scarcely pigmented; lan-
ceolate, 0.29-0.35 mm, 1.8-2.2 times its basal
width; and with 10-15 strong pointed setae, the
lateral (2-4 pairs) are longer than the dorsal ones.
Alate Viviparous Females. Fig. 2. Based on 1
specimen. Very similar to apterous ones, but with
pterothorax brown, more pigmented head, anten-
nae, prothorax, part of femora and tibiae, tarsi, si-
phunculi and cauda, and stigmatic and marginal
sclerites tenuously pigmented. Medial of fore
wings only once-branched. Metric and meristic
features like aptera in part, the following ones are
different: body length, 2.30 mm; antenna, 2.60
mm; antenna/body, 1.1; body/siphunculi, 3.9-4.0
times; femur of hind legs, 0.77-0.80 mm; AntIII,
0.57-0.58 mm; AntIII with 12-13 secondary senso-
ria distributed on 74-76% of its length; AntIV,
0.52-0.56 mm; AntVIpt, 1.2 times AntIII; AntVIpt
4.2 times AntVIb; siphunculus, 0.58-0.59 mm;
cauda, 0.23 mm and 1.5 times its basal width.
Bionomics. The only known host plant of this
aphid is an unidentified species of Adesmia (Fa-
baceae). Aphids feed on the plant stem. Sexuals
are unknown, but the species should be holocyclic,
because of cold temperate climate.
Geographical Distribution. The new species is
known from Rio Mayo in the south of Chubut
province (Argentina), 4540'S, 7016'W, 429 m. It
might live on this plant in other Argentinean lo-
calities, although it seems to be restricted to
southern Patagonia, because on many occasions

we have examined specimens ofAdesmia in sev-
eral localities situated from Rio Negro in the
south to Jujuy in the north and we have only
caught specimens of an undescribed species of
Aphis Linnaeus, 1758.
Discussion. The morphological characteristics
of the new species indicate that it should be
placed in Uroleucon, as this genus is considered at
present (Heie 1995; Carvalho et al. 1998; Black-
man & Eastop 2006), and it presents the pale
coxae of all the native South American species of
the genus, which are included in subgenus Lam-
bersius Olive, 1965 (Nieto Nafria et al. 2007). In
addition, phylogenetic analyses of mitochondrial
cytochrome oxidase II sequences performed by
C. von Dohlen (pers. comm.) found the sample of
this species, provided by one of the authors (J. 0.),
to be nested within other South American species
of Uroleucon and related species, such as Blan-
chardaphis poikila Ortego, J., J. M. Nieto Nafria,
and M. P. Mier Durante, 1997, which also has long
and apically enlarged setae but the siphunculi in
this genus are not reticulated (Ortego et al. 1997).
Uroleucon adesmiae sp. nov. is very close to
Uroleucon payuniense Ortego & Nieto Nafria,
2007. Viviparous females of both species are
slightly pigmented when mounted, they have nu-
merous setae on the posterior part of the head and
also on the dorsum of the abdomen and poor si-
phuncular reticulation; in the alatae (only 1 spec-
imen is known) the medial vein of the forewings
only has 2 branches. In spite of this it is not diffi-
cult to distinguish them because in the new spe-
cies the setae are more robust, the pigmentation
is less intense and there are other differences that
are given in the key.
It is necessary to partly modify the identifica-
tion key (disjunctives 11 to 16) key for the Uroleu-
con species recorded in South America provided
by Nieto Nafria et al. (2007), as follows.

11. AntVIpt at most 3.8 times AntVIb. Siphunculi thin, reticulated at least on 35% of their length and yellowish
brown to light brown, like cauda, which is lanceolate. [Dark-green with dark-brown head. On Gochnatia
glutinosa and Hyaloceris cinerea. Argentina: La Rioja, Mendoza, Tucuman] ............. U gochnatiae
- AntVIpt usually at least 3.8 times AntVIb; if 3.1-3.9 times then siphunculi or cauda have different features.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2
12. (without modification) AntIII in apterae with 2-4 small secondary sensoria extended to 33(44)% of the segment
length, which is short, 0.49-0.61 mm long (alatae unknown). [Papillae absent. Setiferous and postsiphun-
cular sclerites present. Coxae more pigmented than trochanters and 1/2-2/3 proximal portion of femora.
Tarsi with 5 setae. Siphunculi 1.5-1.7 times cauda, with groups of spinules homogeneously distributed to
the reticulated portion, which is 19.6-25.5% of its total length. Cauda light-brown and with 9-14 setae.
Brown to red-brown in life. On Hypochoeris. Argentina: Mendoza] .................... U malarguense
- AntIII of both apterae and alatae usually with 4 or more secondary sensoria [Marginal papillae and marginal
and postsiphuncular sclerites present or absent] . . ......... .............................. 13
13. Siphunculi usually at least 1.6 times cauda, which is 0.24-0.45 mm long and gives 7-15 setae; if 13-15 setae are
present then more than 4 setae on posterior part of cephalic dorsum............................. 14
- (without modification) Siphunculi usually at most 1.6 times cauda, which is 0.30-0.83 mm long with 9-31 setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 0

Florida Entomologist 91(3)



Fig. 2. Uroleucon (Lambersius) adesmiae Mier Durante and Ortego sp. nov., alate viviparous female. A, habitus.
B, AntIII. C, AntVI. D, cauda. E, Ht2. F, siphunculus.

September 2008

Mier Durante & Ortego: A New Species of Uroleucon

14. (without modification) Siphunculi very robust (approximately 2 times larger than hind tibiae and 0.58-0.80 mm
long), subcylindrical (somewhat inflated at beginning of reticulation), slightly curved outside, without
flange, approximately basal 1/4 as pale as coxae and rest brown to dark-brown (as dark as tibiae or anten-
nae), nearly smooth on pale proximal portion, progressively wrinkly to scaly on dark middle portion, and
reticulation extends on 21-41% of its length. On Gutierrezia iserni. [Green in life. Argentina: La Rioja]
............................................................................. U riojanum
- (without modification) Siphunculi with different feature, usually as large as hind tibiae or thinner than them
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 5
15. Posterior part of cephalic dorsum (behind the discal two setae of vertex) with 6-11 setae
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- (without modification) Posterior part of cephalic dorsum (behind the two discal setae of vertex) usually with 4
setae (the general feature in the genus) ..................................................... 17
16. AbdII-AbdIV with 3-6 marginal setae each side and 7-27 spinal-pleural setae. Siphunculi yellowish very light-
brown homogeneously or with paler proximal 1/4. Cauda lanceolate and 0.27-0.37 mm long ....... 16 bis
- (without modification) AbdII-AbdVI with 1-2 marginal setae each side and 5-13 spinal-pleural setae. Siphun-
culi brown to dark brown in general, often with a middle portion paler than both proximal and distal por-
tions. Cauda triangular or long-pentagonal in shape, 0.22-0.30 mm long. Red in life with siphunculi dark-
brown. On Mutisia spinosa. [Argentina: Chubut and Rio Negro]...................... U. patagonicum
16 bis. Siphunculi 2.4-4.3 times its width at the beginning of the reticulation, which is well sculptured and covers
12.2-18.9% the siphunculi length. Dorsal-abdominal setae 50-60 im and 1.5-1.9 times D. Ht2 0.13-0.15
mm. Cauda with 6-9 setae. AntVIpt at least 1.0 times AntIII and 4.0 times AntVIb. Urs 1.9-2.7 times its
basal width and with 7-10 secondary setae. Green in life. On Grindelia chiloensis [Argentina: Mendoza]
............................................................................ U payuniense
- Siphunculi 1.6-2.5 times its width at the beginning of the reticulation, which is poorly sculptured and covers
8.0-13.4% the siphunculi length. Dorsal-abdominal setae 40-45 im and 1.0-1.5 times D and relatively
thick. Ht2 0.15-0.17 mm. Cauda with 10-15 setae. AntVIpt at most 1.0 times AntIII and 4.1 times AntVIb.
Urs 2.7-3.4 times its basal width and with 10-13 secondary setae. Dark green to brownish green in life. On
Adesmia sp. [Argentina: Chubut] .................................................. U adesmiae

On the other hand it is useful to remake the
identification key for the aphids on Adesmia by
Blackman & Eastop (2006), with an additional

first disjunctive as follows (SIPH is siphunculus,
ABD TERG is the abdominal tergite, and MTu are
marginal tubercles):

0. SIPH with a distal zone of reticulation. ABD TERG 1 and 7 constantly without MTu .... Uroleucon adesmiae
- SIPH without polygonal reticulation. ABD TERG 1 and 7 constantly with MTu ........................ 1


We acknowledge Carol von Dohlen, Utah State Uni-
versity, for providing us with the phylogenetic analyses
of mitochondrial cytochrome oxidase II sequences of the
new species. We appreciate Luis M. Fernandez Blanco
for cooperation in preparing the illustration. This re-
search was supported by the Regional Government of
Castilla y Le6n, Spain (grants LE45/02 and LE034A05).


BLACKMAN, R., AND V. F. EASTOP. 2006. Aphids on the
World's Herbaceous Plants and Shrubs. Two volu-
mens. John Wiley & Sons, Ltd., Chichester, U.S.A.
1024 pp.
1998. The genus Uroleucon Mordvilko (Insecta,
Aphidoidea) in South America, with a key and de-
scriptions of four new species. Zoological J. Linnean
Soc. 123: 117-141.

HEIE, 0. E. 1995. The Aphidoidea (Hemiptera) of Fen-
noscandia and Denmark. VI. Family Aphididae: Part
3 of tribe Macrosiphini of subfamily Aphidinae, and
family Lachnidae. Fauna Entomologica Scandinav-
ica 31: 1-217.
AND M. V. SECO FERNANDEZ. 2007. The genus Uro-
leucon (Hemiptera: Aphididae: Macrosiphini) in Ar-
gentina, with descriptions of five new species. The
Canadian Entomol. 139(2): 154-178.
RANTE. 1997. Blanchardia poikila gen. nov., sp.
nov. (Hemiptera: Aphididae, Macrosiphini) in Men-
doza, Argentina. The Canadian Entomol. 129: 1093-
Les plants h6tes des pucerons africains, pp. 103-
139 In G. Remaudiere and A. Autrique [eds.], Contri-
bution a l'Ecologie des Aphides Africains. Organisa-
tion des Nations Unies pour l'Alimentation et
l'Agriculture (Etude FAO production v6g6tale et pro-
tection des plants, 64), Rome.

Florida Entomologist 91(3)

September 2008


1State Key Laboratory of Integrated Management of Pest Insects & Rodents, Institute of Zoology,
Chinese Academy of Sciences, Beijing 100101, P.R. China

'USDA Forest Service, Southern Research Station, 320 Green St., Athens, GA 30602-2044, USA

*Corresponding author

A systematic survey of Chinese privet foliage, stems, seeds, and roots for associated phy-
tophagous insects was conducted in China during 2005 and 2006 in order to establish basic
information about the insect communities that Chinese privet harbors and to evaluate the
abundance and damage caused by these insects. A total of 170 phytophagous insect species
in 48 families and 5 orders were collected from Chinese privet in China. The insects belong
to 4 feeding guilds: foliage, sap, stem, and root feeders. The impact of foliage feeders varied
by site and over time. The mean percent defoliation of Chinese privet over all sites and years
was 20.5 8.2%, but ranged as high as 48%.

Key Words: Chinese privet, biological control, invasive species, exotic species

Se realize un sondeo sistematico de los insects fit6fagos asociados con el follaje, ramas, se-
millas y races de Cabo chino Ligustrum sinense en China durante 2005 y 2006 para esta-
blecer informaci6n basica sobre las comunidades de insects que usan L. sinense como un
refugio y a la vez para evaluar la abundancia y dafo causado por estos mismos. Un total de
170 species de insects consumidores en 48 families y 5 6rdenes fueron recolectada de L.
sinense en China, que pertenecen a 4 grupos de consumidores: los que se alimentan sobre el
follaje, savia, tallo y races. El impact de los consumidores de follaje varia segun el sitio y
la 6poca. El promedio del porcentaje de defoliaci6n de L. sinense en todos los sitios y anos es-
tudiados fue de 20.5 8.2%, pero se observe rangos altos hasta de un 48%.

Chinese privet, Ligustrum sinense Lour., is an
invasive exotic weed in the United States where it
is a perennial, semi-evergreen shrub or small tree
that grows to 10 m in height. This species is of
great concern in the southeastern United States
(Faulkner et al. 1989; Stone 1997), but it ranges
from Texas to Florida, and as far north as the New
England states (The Nature Conservancy 2004;
University of Connecticut 2004). In addition, it
has been introduced into Puerto Rico and Oregon
(USDA-NRCS 2002).
Chinese privet is native to China, Vietnam,
and Laos (The Nature Conservancy 2004; Wu &
Raven 2003). In China it is found in the provinces
south of the Yangzi River (Qui et al. 1992). It can
be found between 200 and 2600 m in elevation
where it occurs in mixed forests, valleys, along
streams, thickets, woods, and ravines (Wu &
Raven 2003; Qui et al. 1992).
Ligustrum sinense has been studied in China
for its chemical composition and the medicinal
value of its bark and leaves (Ouyang & Zhou
2003; Ouyang 2003). It has never been recorded
as a noxious weed in either agricultural or forest

settings. Chinese privet is grown nationwide as
an ornamental hedge plant, for its berries used in
brewing, and for oils extracted from seeds and
used in soap making (Qui et al. 1992).
Chinese privet was introduced into the United
States in 1852 (Coates 1965; Dirr 1990) as an or-
namental shrub, for hedgerows (USDA-NRCS
2002), and sometimes as single specimens for its
foliage and profusion of small white flowers (Dirr
1990; Wyman 1973). It is a forage plant for deer in
the southeastern U.S. (Stromayer et al. 1998a;
Stromayer et al. 1998b; The Nature Conservancy
2004). According to Small (1933), the species was
escaping from cultivation in southern Louisiana
by the 1930s. A survey of appropriate herbaria re-
veals collection records from Georgia as early as
1900. Based on herbarium records the species be-
came naturalized and widespread in the south-
east and eastern U.S. during the 1950s, 60s, and
70s (USDA-NRCS 2002).
Chinese privet is widely believed to drastically
reduce native plant biodiversity because of its
ability to shade out native vegetations (USDI Fish
and Wildlife Service 1992; Merriam & Feil 2002)

Zhang et al.: Biological Control of Chinese Privet

and form dense, monospecific stands that domi-
nate the forest understory (Dirr 1990). Recent
surveys in the southeastern United States show
Chinese privet completely covers 0.9 million acres
and colonies of varying densities can be found on
another 17.6 million acres (Rudis et al. 2006). In
1998, the U.S. Department of Agriculture listed
privet as one of 14 species with the potential to ad-
versely affect management objectives in North
Carolina's National Forests. Similarly, the Florida
Exotic Pest Plant Council lists Chinese privet as a
Category 1 invasive species (FLEPPC 2007). More
recently, The Nature Conservancy ranked Chi-
nese privet as having high potential to disrupt the
ecological balance (NatureServe 2006).
In addition to privet's impact on natural land-
scapes, it can be directly harmful to humans. The
flower of Chinese privet is toxic to humans caus-
ing symptoms such as nausea, headache, abdom-
inal pain, vomiting, diarrhea, weakness, and low
blood pressure and body temperature (USDA-
NRCS 2002). Where Chinese privet occurs in
abundance, floral odors may cause respiratory ir-
ritation (Westbrooks & Preacher 1986).
Repeated mowing and cutting will control the
spread of L. sinense, but may not eradicate it
(Tennessee Exotic Pest Plants Council 1996). Al-
though modern herbicides, including glyphosate,
effectively kill privet (Tennessee Exotic Pest
Plants Council 1996; The Nature Conservancy
2004; Madden & Swarbrick 1990; Harrington &
Miller 2005), environmental concerns will limit
use of herbicides on public land or in sensitive ar-
eas. Faulkner et al. (1989) reported that in exper-
imental trials of prescribed burning, there was no
significant difference in the abundance of Ligus-
trum sinense in burned vs. unburned plots.
Plants unimportant in their native habitat may
reach damaging levels when released from control
by important natural enemies through introduc-
tion into new geographic areas (Van Driesche &
Bellows 1996). This suggests that exploration for
Chinese privet natural enemies in China might
detect species suitable for use in a classical biolog-
ical control program in the U.S. Chinese privet is
considered a good candidate for classical biological
control because it has no known biological control
agents capable of lowering its pest status in North
America (The Nature Conservancy 2004), and no
native Ligustrum spp. occur there. Johnson &
Lyon (1991) list at least 27 species of insects or
mites that feed on Ligustrum spp. in the United
States, however, none suppress populations of this
plant in forests. In contrast, based on published
records, China appears to have a rich complex of
natural enemies that attack Ligustrum spp.
(Zheng et al. 2004) and the potential for finding a
biological control agent is high. However, little is
known about their relative abundance and impact
on their host plant. Therefore, a cooperative pro-
gram was initiated in 2005 to survey for natural

enemies of Chinese privet in China, with the goal
of finding potential biocontrol agents. Here we re-
port results of systematic surveys conducted in
2005 and 2006, and provide a list of the phytopha-
gous insects found. We provide basic information
about the insect community on Chinese privet in
China and identify species that may have poten-
tial as biological control agents in the US. Prelim-
inary information on impacts of different feeding
guilds on Chinese privet in China is also reported.


Survey Sites

Our surveys focused primarily on 6 sites in
Huangshan city (118.16, 29.43, elevation approx-
imately 200m) in Anhui Province, China, because
the climate-matching program Climex (Hearne
Scientific Software, Melbourne, Australia) indi-
cated that this was the province most similar in
climate to the southeastern United States (Sun et
al. 2006). The seventh site was established in
Guiyang (Huaxi, 106.40, 26.25, elevation approx-
imately 1096 m), Guizhou Province, which is an-
other area where Chinese privet is prevalent in
China, but it is much further south and warmer
than the Anhui sites.
In order to collect the most natural enemies of
privet, our survey sites were selected to include
habitats varying from natural areas to semi-nat-
ural and planted sites. These were as follows:
(1) Pure natural sites included 1 site in Ling-
nan adjacent to the Jiulong Natural resource con-
servation area in Xiuning County in the most
southern region of Anhui Province; a second site
on an unnamed island in the suburb of Huangs-
han city; and a third site in Guiyang. Chinese
privet in these 3 sites grew naturally mixed with
many other plant species and most grew rela-
tively tall (over 3 m).
(2) Semi-natural sites included 1 site in Zhan-
chuan, a small town south of Huangshan city, and
a second site at the Institute of Forestry (IOF) lo-
cated in the north of Huangshan city. Chinese
privet plants in these sites grew semi-naturally
but were near agricultural lands where they were
more likely to be disturbed by local residents.
(3) Planted sites included 1 site in She County
east of Huangshan city. The other site was in the
center of Huangshan city. Chinese privets in
these sites were abundant and planted as orna-
mental shrubs along roads. All plants were small
periodically pruned shrubs less than 1.5 m tall.

Systematic Sampling

At each site, 10 Chinese privet plants were
randomly selected for sampling, marked with
stakes and surrounded by a circle of colored tape
and a sign to prevent human disturbance and for

Florida Entomologist 91(3)

ease of relocation. Privet plants in Zhanchuan,
IOF and She county in Anhui Province were sur-
veyed from May to Aug, 2005 and from Apr to Oct,
2006. Privet on the island near Huangshan city
was sample from Jun to Aug 2005 when sampling
was discontinued because it was cleared for devel-
opment. Privet in Lingnan, Huangshan city, and
Guiyang were surveyed for 1 year from Apr to Sep
2006. At all sites, surveys were conducted at 10-
day intervals during the survey periods. Collec-
tion of insects feeding on Chinese privet was ac-
complished by hand-picking, aspirating, and
sweep-netting each sample plant 30 times. In
some cases, cages were placed over branches to
capture insects as they emerged. When immature
insects were found, they were collected in a plas-
tic bag together with the plant part on which they
were feeding and returned to our lab to be reared
to the adult stage to confirm the species.
Most insect species were identified by Professor
Yang Chuncai (Anhui Agricultural University).
Some Chrysomelidae were identified by Professor
Wang Shuyong (Institute of Zoology, Chinese
Academy of Sciences [CAS]), some Lepidoptera
larvae were identified by Professor Wang Linyao
(Institute of Zoology, CAS), and some Homoptera
were identified by Professor Liang Aiping (Insti-
tute of Zoology, CAS). Pseudaulacaspis pentagon
(Targioni-Tozzetti) was identified by Professor Xie
Yingping (Shanxi University). Others were sent to
the Zoological Museum of the Institute of Zoology,
CAS, where they were distributed to appropriate
taxonomists for identification. All phytophagous
insects were evaluated based on their frequency of
occurrence on Chinese privet, stage of develop-
ment, and collecting site. Information on insect
host range was obtained from the references "Eco-
nomic Insect Fauna of China", edited by the Edito-
rial Committee of Fauna Sinica, Academia Sinica
(Chou et al. 1985; Ge 1966; Tan et al. 1985; Yu et
al. 1996; Liu 1963; Liu & Bai 1977; Wang 1980;
Zhang 1985; Zhang 1995; Zhao & Chen 1980).

Foliage Damage

Defoliation rate was estimated by averaging
defoliation of 120 leaves per plant. Samples con-
sisted of 10 leaves randomly selected from 3 layers
(high, middle, and lower layer) and 4 cardinal di-
rections (east, south, west, and north) in each
layer, for a total of 12 sampling locations on each
plant. Defoliation was estimated by placing leaves
on transparent graph paper with a 1-mm2 grid and
measuring total leaf area and leaf area removed.

Stem Damage

Altogether, 900 plants of Chinese privet were
investigated for signs of insect feeding within
stems, oviposition, and damage at all survey sites.
Stem damage was described by attributes, includ-

ing physical shape, the distance of the damage
from the ground, the diameter of stem with boring
hole and so on.

Root and Seed Damage

Fifty roots were dug from randomly selected
sample sites and examined for root damage. Adult
insects found feeding on roots were collected for
identification and larvae were returned to the lab-
oratory with pieces of root for rearing to the adult
stage. In order to detect insects feeding in seed,
500 immature seeds were collected randomly
from survey sites and half were dissected in the
laboratory. Also, 200 panicles with 25 to 58 ma-
ture seeds per panicle plus the remaining imma-
ture seeds were collected and placed in glass con-
tainers with fine gauze lids in order to collect
adult seed-feeders emerging from them.


The phytophagous insects associated with Chi-
nese privet in China are listed in Table 1. In all,
170 species in 5 orders and 48 families were col-
lected from Chinese privet in Anhui and Guizhou
Province from 2005 to 2006. Insects were found in
4 different feeding guilds: foliage, sap, stem, and
root feeders. Among them, 95.9% of insects were
collected from privet leaves, 1 species was found
feeding in stems and 6 species were root feeders.
In contrast, only 27 species of insects feed on Li-
gustrum spp. in the U.S. Table 1 also includes an
estimate of host range for each insect based on
published reports.

Foliage-feeding Insects

Among the foliage-feeding insects in China,
Argopistes tsekooni (Coleoptera: Chrysomelidae),
Leptoypha hospital (Hemiptera: Tingidae) and an
unidentified sawfly appeared to have the greatest
impact on the plant. The extent of defoliation var-
ied among sites, seasons and years (Figs. 1 and 2).
In Zhanchuan, defoliation remained relatively
constant throughout the sampling period fluctu-
ating only slightly from 20% to 28% (Fig. 1A). De-
foliation was highest at the She county site, aver-
aging over 50% in late Jul 2005 (Fig. 1B). At the
IOF site, defoliation ranged from about 15% in
early May to about 27% in mid-Aug 2005. Defoli-
ation in 2006 was generally higher at the IOF
site, averaging about 34% for the year (Fig. 1C).
Guiyang, a natural area, had the lowest defolia-
tion of all sites averaging 1.6% in 2006 (Fig. 2).
Lingnan had an average defoliation of 16% for
2006. Defoliation at this site ranged from a high
of about 16% in late Apr to a low of less than 5%
in Sep. Defoliation at the Huangshan city site, a
site consisting of planted privet, generally de-
clined over the season from a high of ca. 30% in

September 2008

Zhang et al.: Biological Control of Chinese Privet


Relative Stage Host con- Feeding Host
Order/Family Species frequency foundb formationc guild ranged

Hepialidae Phassus excrescens Butler
Pyralidae Diaphania nigropunctalis (Bremer)
Cnaphalocrocis medinalis(Giien6e)
Parapoynx diminutalis Snellen
Parapoynx vittalis (Bremer)
Ostrinia furnacalis (Gtien6e)
Diaphania indica (Saunders)
Pycnarmon cribrata (Fabricius)
Hymenia recurvalis (Fabricius)
Maruca testulais Geyer
Palpita inustata (Butler)
Endotricha theonalis (Walker)
Diaphania perspectalis (Walker)
Tryporyza incetulas (Walker)
Heliodinidae Atrijuglans hitauhei Yang
Scythridae Scythris sinensis Felder et Rogenhofer
Brahmaeidae Brahmaea ledereri Rogenhofer
Sphingidae Psilogramma menephron (Cramer)
Gelechiidae Telphusa sp.
Noctuidae Pseudaletia separate (Walker)
Pangrapta cana Leech
Helicoverpa assulta (Gtien6e)
Ericeia fraterna (Moore)
Arctiidae Nyctemera adversata (Schaller)
Spilarctia subcarnea (Walker)
Creatonotus transiens (Walker)
Amsacta lactinea (Cramer)
Miltochrista aberrans Butler
Cyana sp.
Asura sp.
Psychidae Cryptothelea variegata Snellen
Geometridae Scopula caricaria Reutti
Percnia luridaria (Leech)
Naxa (Psilonaxa) seriaria Motschulsky
Gelasma illiturata Walker
Comostola subtiliaria (Bremer)
Tortricidae Archips seminubilis (Meyrick)
Grapholitha delineana Walker
Tingidae Leptoypha hospital Drake et Poor
Pentatomidae Nezara viridula (Linnaeus)
Menida sp.
Eysarcoris sp.
Pentatoma sp.
Homoeocerus sp.
Erthesina fullo (Thunberg)
Palomena angulosa Motschulsky
Carbula obtusangula Reuter
Coreidae Riptortus pedestris (Fabricius)
Cletus punctiger Dallas
Riptortus pedestris (Fabricius)
Urostylidae Urochela distinct Distant

L, A
L, A


1 Stem
1 Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
/ Foliage
o Foliage
o Foliage
/ Foliage
\j Foliage
/ Foliage
o Foliage
o Foliage
o Foliage
/ Foliage
/ Foliage
o Foliage
o Foliage
/ Foliage
o Foliage
o Foliage
o Foliage
/ Foliage
o Foliage
/ Foliage
o Foliage
o Foliage
o Foliage
/ Foliage
/ Foliage
/ Foliage
/ Foliage
o Foliage
o Foliage
o Foliage
o Foliage

1 Sap
o Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
El Sap



aR, rare, taken at a few sites, usually in small numbers; 0, occasionally collected at sites; C, common, taken at most sites.
'L larva; A, adult; N, nymph.
"Z'observed feeding on privet, "D"collected from Chinese privet and recorded as privet feeder in literature, "O'collected from Chi-
nese privet, but not directly observed or recorded in literature as feeding on privet.
dPo, Polyphagous, feeds on plants from other families; 01, Oligophagous,feeds mainly on Oleaceae; Mo, Monophagous on Chinese

Florida Entomologist 91(3)

September 2008


Relative Stage Host con- Feeding Host
Order/Family Species frequency foundb formationc guild ranged



Megacopta cribraria (Fabricius)
Halticus minutus Reuter
Adelphocoris fasiaticollis Reuter
Hyperoncus lateritius Westwood
Nysius ericae (Schilling)

o Sap
o Sap
1 Sap
1 Sap
o Sap

o Sap
1 Sap
o Sap
o Sap
1 Sap
1 Sap
1 Sap
1 Sap
1 Sap
o Sap
o Sap
o Sap
1 Sap

Cicadellidae Cicadella viridis (Linnaeus)
Cercopidae Clovia bipunctata (Kirby)
Abidama contigua Walker
Membracidae Tricentrus sp.
Aphrophoridae Aphrophota sp.
Flatidae Geisha sp.
Salurnis marginella Gu6rin
Lawana sp.
Ricaniidae Pochazia sp.
Euricania ocellus Walker
Pochazia guttifera Walker
Issidae Sivaloka sp.
Diaspididae Pseudaulacaspis pentagon (Targioni-Tozzetti)
Chrysomelidae Argopistes tsekooni Chen
Longitarsus bimaculatus?Baly?
Psylliodes punctifrons Baly
Pseudodera xanthospila Baly
Argopus balyi Harold
Manobidia nipponica ChujjG
Altica viridicyanea Baly
Psylliodes sp.
Aphthonomorpha collaris (Baly)
Aphthona varipes Jacoby
Aphthona strigosa Baly
Longitarsus dorsopictus Chen
Longitarsus lohita Maulik
Hemipyxis plagideroides (Motschulsky)
Monolepta hieroglyphica (Motschulsky)
Cryptocephalus sp.
Basiprionota bisignata (Boheman)
Ambrostoma quadriimpressum (Motschulsky)
Paleosepharia lequidambra Gressitt & Kimoto
Acrothinium gaschkevitschii (Motschulsky)
Mimastra soreli Baly
Plagiodera versicolora Laicharting
Phaedon brassicae Baly
Euliroetis ornata (Baly)
Monolepta selmani Gressitt et Kimoto
Coenobius longicornis Chfi ja
Adiscus variabilis (Jacoby)
Adiscus exilis (Weise)
Colaspoides sp.
Oomorphoides yaosanicus (Chen)
Agetocera spp.
Gastrolinoides japonica Harold
Basilepta ruficolle Jacoby

1 Foliage
1 Foliage
o Foliage
<1 Foliage
1 Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
<1 Foliage
1 Foliage
1 Foliage
o Foliage
o Foliage
1 Foliage
o Foliage
1 Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage




aR, rare, taken at a few sites, usually in small numbers; 0, occasionally collected at sites; C, common, taken at most sites.
'L larva; A, adult; N, nymph.
observedrd feeding on privet, "D"collected from Chinese privet and recorded as privet feeder in literature, "O"collected from Chi-
nese privet, but not directly observed or recorded in literature as feeding on privet.
'Po, Polyphagous, feeds on plants from other families; 01, Oligophagous,feeds mainly on Oleaceae; Mo, Monophagous on Chinese

Zhang et al.: Biological Control of Chinese Privet


Relative Stage Host con- Feeding Host
Order/Family Species frequency foundb formationc guild ranged

Stenoluprus cyaneus Baly
Colaphellus bowringii Baly
Phaedon brassicae Baly
Plagiodera versicolora (Laicharting)
Dercefina flavocincta (Hope)
Gallerucida ornatipemmis (Duvivier)
Cleoporus variabilis (Baly)
Eutettix apricus Melichar
Adiscus grandipalpus Tan
Melixanthus pieli (Pic)
Smaragdina aurita hammarstraemi (Jacobson)
Aulacophora lewisii Baly
Paridea biplagiata (Fairmaire)
Temnaspis nankinea (Pic)
Crioceridae Lilioceris maai Gressitt and Kimoto
Lilioceris impressa (Fabricius)
Lilioceris minima (Pic)
Lema (Lema) scutellaris (Kraatz)
Lema(Lema) concinnipennis Baly
Lilioceris rugata (Baly)
Lilioceris scapularis (Baly)
Curculionidae Sympiezomias velatus (Chevrolat)
Sympiezomias spp.
Euops sp.
Apoderus geniculatus Jekel
Alcidodes sauteri (Heller)
Myllocerinus aurolineatus Voss
Anthonomus bisignifer Schenkling
Callosobruchus chinensis (Linnaeus)
Myllocerinus ochrolineatus Voss
Meloidae Zonitis japonica Pic
Lytta caraganae Pallas
Hispidae Cassida japana Baly
Laccoptera quadrimaculata (Thunberg)
Thlaspida biramosa Boheman
Cassida piperata Hope
Eumolpidae Smaragdina nigrifrons (Hope)
Coccinellidae Henosepilachna vigintioctopunctata (Fabricius)
Epilachna freyana Beilawski
Epilachna quadricollis (Dieke)
Scarabaeidae Holotrichia parallel Motschulsky
Cetoniidae Protaetia (Calo) aerata (Erichson)
Protaetia brevitarsis Lewis
Oxycetonia bealiae (Gory et Percheron)
Pseudodiceros nigrocyaneus (Bourgoin)
Rhomborrhina unicolor Motschulsky
Oxycetonia jucunda Faldermann
Rutelidae Anomala corpulenta Motschulsky
Anomala olivea Lin
Adoretus sinicus Burmeister
Hoplia communis Waterhouse
Oedemeridae Xanthochroa hilleri Harold
Tenebrionidae Gonocephalum pubiferum Reitter
Nitidulidae Librodor.japonicus Motschulsky

" Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
o Foliage
1 Foliage
o Foliage
o Foliage
1 Foliage
o Foliage
1 Foliage
1 Foliage
1 Foliage
o Foliage
o Foliage
o Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
o Foliage
1 Foliage
o Foliage
o Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
I Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Foliage
1 Root
o Foliage
o Root
^ Foliage


aR, rare, taken at a few sites, usually in small numbers; 0, occasionally collected at sites; C, common, taken at most sites.
'L larva; A, adult; N, nymph.
observedrd feeding on privet, "D"collected from Chinese privet and recorded as privet feeder in literature, "O"collected from Chi-
nese privet, but not directly observed or recorded in literature as feeding on privet.
'Po, Polyphagous, feeds on plants from other families; 01, Oligophagous,feeds mainly on Oleaceae; Mo, Monophagous on Chinese

Florida Entomologist 91(3)


Relative Stage Host con- Feeding Host
Order/Family Species frequency foundb formationc guild ranged

Elateridae Pleonomus canaliculatus Faldermann R A < Root Po
Agriotes subrittatus Motschulsky R A < Root Po
Sarpedon atratus Fleutiaux R A < Root Unknown
Languriidae Anadastus analis Fairmaire R A o Foliage Unknown
Lagriidae Lagria nigricollis Hope R A o Foliage Unknown
Cantharidae Cantharis violaceipemis Gorh 0 A o Foliage Unknown
Cantharis sp. R A o Foliage Unknown
Buprestidae Agrilus spp. R A < Foliage Po
Catantopidae Oxya intricate (Stal) 0 N,A o Foliage Po
Xenocatantops brachycerus (Will) 0 N,A o Foliage Po
Tettigoniidae Prohimerta (Anisotima)guizhouensis Gorochov 0 N,A o Foliage Po
& Kang
Mirollia formosana Shiraki R N,A o Foliage Po
Hexacentrus unicolor Serville R N,A o Foliage Po
Kuzicus (Kuzicus) suzukii Matsumura & R N,A o Foliage Po

R, rare, taken at a few sites, usually in small numbers; 0, occasionally collected at sites; C, common, taken at most sites.
'L larva; A, adult; N, nymph.
""'observed feeding on privet, "D"collected from Chinese privet and recorded as privet feeder in literature, "O'collected from Chi-
nese privet, but not directly observed or recorded in literature as feeding on privet.
'Po, Polyphagous, feeds on plants from other families; 01, Oligophagous,feeds mainly on Oleaceae; Mo, Monophagous on Chinese

mid-May to a low of ca. 8% in early Oct. A natural
site on a nearby island had a defoliation rate of
about 30% as well (Fig. 3), suggesting that defoli-
ation was similar in an area regardless of site con-
dition. The mean percent defoliation for Chinese
privet per year among all sites and all years of
study was 20.5 8.2%.

Stem Damage

Phassus excrescens (Lepidoptera: Hepialoidae)
was the only stem borer found feeding on Chinese
privet in our survey where 5.3% (48 of 900 priv-
ets) of the plants were damaged by it. Larvae of
the insect were collected from trunks of Chinese
privet where they bored in the xylem causing
galls. While feeding, they created an off white
mass consisting of silk, excrement, and wood
scraps that covered the entrance to the larval gal-
lery. Borer entrance holes were 29.771.95cm (n
= 48) from the ground and the average diameter
of attacked stems was 2.270.72cm (n = 48).

Root and Seed Damage

Six species of insects were found feeding on
roots of Chinese privet. All 6 fed on fine roots or
the root surface. Observations of the above
ground plant health gave no indication root-feed-
ers were present. No seed-feeders were found in
either immature or mature seeds.


In order to collect the most natural enemies of
privet, survey sites were selected to include di-
verse habitats varying from natural areas to
semi-natural and planted sites. Anhui province
was selected as the primary survey area, because
it was the best climatic match to the southeastern
United States. Climatic matching is important for
conventional biological control to insure the se-
lected agents are adapted to the climate where
they will be released (Andres et al. 1976; Harley
& Forno 1992). Guizhou province was another im-
portant survey area because it is near the center
of the range of Chinese privet in China.
Chinese privet is a common ornamental shrub
but not a noxious weed in China, suggesting that
natural enemies suppress populations. We found
170 phytophagous insect species on Chinese
privet in China. Most were foliage-feeding insects
despite phenolic compounds in privet leaves that
likely provide some protection against damage
from generalist herbivores (Swearingen et al.
2002). In the United States, Johnson & Lyon
(1991) list at least 27 species of insects or mites
that feed on Ligustrum spp., however, none sup-
press populations of this plant in forests. Most are
not specialist on Chinese privet so it seems likely
that the diverse and abundant insect fauna in
China is important in regulating Chinese privet
populations in its native habitat. Other factors

September 2008

Zhang et al.: Biological Control of Chinese Privet

--- 02005
....... 2006

I- -i--.

20- 50- to- 20- 50- 10- :0- so- 10- 20- 30- 10- 20- o- 20- 20- 30-
Apr Apr Aary Aa Aay Jun Jun Jun Jul Jul Jul Alu Lug lAu Sep Sep Sep

-...-. 2006

19 29:- 9- 19" 29- 9" i- 9 9" 19" 2 9" 9" 19" 29" 9" 9 9"-
Iow Air a Aany ay Jun Jun Jun Jul Jul Jul Aug Aug Aug Sep Sep Sep

T 2005
T -"I'-. T --A-- 200o

1t- 9- 8s tz- ;B- $- i9- "8- 8- fl- "s- e- i3- fs- 8- ii- 's-
Apr AWr Aay hay hat Jun Jun Jun Jul Jul Jul Aue luc lAu S ep Sep S ep

Fig. 1. Seasonal Chinese privet defoliation at: (A) Zhanchuan (a semi-natural site), (B) She County (a planted
site), and (C) IOF (a semi-natural site) in Anhui Province. Means - SME are shown.

also may be involved such as disease organisms only occasionally rested on the plants were likely
not include in this survey. Phytophagous insects included in sweep net samples. To distinguish
were collected by hand picking or net sweeping. among them, we made notes in Table 1 showing
Most were determined as feeding on Chinese which were confirmed as feeding on privet during
privet by observations made during surveys in our surveys (/), were collected during our surveys
the field and through the literature. However, and recorded as privet feeders in the literatures
other insects that were not privet feeders and ( ), or collected from Chinese privet, but were not

Florida Entomologist 91(3)

Chinese privet defoliation in 2006
- -Guiyang
- Lingnan
A Huangshan city

22- 2- 12- 22- 2- 12- 22- 2- 12- 22- 2- 12- 22- 2- 12- 22- 2-
Apr 7y Yar Kay Jun Jun Jim Jul Jul Jul Aug Aug Aug Sep Sep Sep Oct

Fig. 2. Seasonal Chinese privet defoliation in 2006 in Guiyang, a natural site in Guizhou Province, Lingnan, a
natural site in Anhui Province and Huangshan city, a planted site in Anhui Province. MeanstSME are shown.

confirmed as privet feeders by personal observa-
tion or in the literature (o). We confirmed 81 feed-
ing on privet by personal observation (Y. Z.
Zhang), 88 were collected from privet and were re-
ported in the literature as feeding on privet, and 1
species was collected but could not be confirmed
by either method. Table 1 provides the most com-
prehensive listing of phytophagous insect feeding
on Chinese privet to date.
When screening potential biological control
agents for invasive weeds, their host range is one
of the most important factors because only host
specific agents will be considered for release to
control invasive weeds. Polyphagous species were
included in Table 1 to provide a complete listing
with no attempt to differentiate good candidates
for testing.
We included all insects found on privet, not
just the most common ones, because some insects
that are rare in their native country and sup-
pressed by their own natural enemies are effec-

Chinest piveL dfohiation 2n00
40 -- a-IlnaI

o 10
20-Jxn 30-J3v to-1ul 20-J". 30-J.l Ito-aug
Fig. 3. Seasonal Chinese privet defoliation on an is-
land natural site in Huangshan city in Anhui Province.
Means SME are shown.

tive biological control agents when released from
their own population regulating fauna.
Due to host specificity and the severe damage
it caused on Chinese privet (Zhang et al., unpub-
lished data), A. tsekooni may be the most promis-
ing biological control agent. It was apparent in
our field surveys that defoliation rates were high
when populations of A. tsekooni were large. Also,
preliminary host specificity tests suggest its host
range is restricted to Ligustrum spp. (Zhang et al.
unpublished data). Examples of flea beetle as bio-
logical control agents of exotic weeds include Al-
tica carduorum Guer. (Chrysomelidae: Coleop-
tera) on Cirsium arvense (L.) Scop. (Asteraceae)
(Wan et al. 1996) and Agascicles hygrophila Sel-
man et Vogt in China for control ofAlternanthera
philoxeroides (Mart.) Griseb., a global virulent
weed from South America (Julien et al. 1995).
Argopistes tsekooni feeds on most members of the
genus Ligustrum. However, since no indigenous
Ligustrum spp. occur in the U.S. and all Ligus-
trum spp. in the U.S. are listed as invasive weeds
(Miller et al. 2004), A. tsekooni is a potential bio-
logical control that warrants further testing.
Leptoypha hospital could be another promising
biocontrol agent because it has a limited host
range in the Oleaceae (Li 2001) and often occurred
in high numbers on Chinese privet in our sample
areas. Likewise, the unidentified sawfly may also
be an important defoliator. Thus far we have been
unable to rear adults for identification so we can-
not fully evaluate its potential for biological con-
trol. However, we have not observed it attacking
other plant species during our field surveys.
Feeding by P excrescens weakened the trunk
resulting in breakage or, in some cases, the stems
died as a result of girdling by the larvae. However,

September 2008

Zhang et al.: Biological Control of Chinese Privet

it has a broad host range and is considered an im-
portant pest of many plant species. Therefore, it is
unlikely that it could be developed as a biological
control agent.
Defoliation of privet varied widely among sites.
The highest defoliation was recorded at the She
County site which was a planted site. The lowest
was on privet at the Guiyang site a natural area in
Guizhou Province. That site was selected because it
was near the center of the range of Chinese privet
and, therefore, likely to have high number of phy-
tophagous insects. We are uncertain why privet de-
foliation was low at this site but it may be the re-
sult of high numbers of defoliator natural enemies,
or the more widely scattered and shaded privet
population we sampled at that location. Defoliation
of privet at the Lingnan site in Anhui Province, an-
other natural area, was also relatively low com-
pared with the She County site, or the Zhanchuan
and IOF sites which we classified as semi-natural.
The average 20% defoliation rate of L. sinense in
China for all sites combined demonstrates that de-
foliating insects have a large impact on privet even
when their own populations are being regulated by
natural enemies. These results suggest that in the
absence of natural enemies some of these insects
may be effective biocontrol agents.


This research is part of an ongoing Sino-US Chinese
privet biological control cooperative program funded by
the USDA-Forest Service, Southern Research Station,
Research Work Unit 4552, and the Natural National
Science Foundation of China (30525009, 30621003). We
are grateful to Ding Jianqing and Wei Wei for helpful
comments on survey plan; Fang Fang, Li Li and Chen
Yuhui for field assistance. We are also grateful to the
many taxonomists who helped with identification of our


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Florida Entomologist 91(3)

Cherry & Stansly: Florida Sugarcane Wireworms


1Everglades Research and Education Center, 3200 E. Palm Beach Road, Belle Glade, FL 33430

2Southwest Florida Research and Education Center, P.O. Box 5127, Immokalee, FL 34143


Wireworms are important insect pests of Florida sugarcane. Our objective was to determine
the abundance and spatial distribution of wireworms in Florida sugarcane on muck versus
sandy soils. Fourteen commercial sugarcane fields were sampled for wireworms on farms in
southern Florida. Melanotus communis (Gyllenhal) was the most abundant wireworm found
in both soil types. Other less abundant wireworms found and discussed are Conoderus spp.,
Ischiodontus sp., and 6. i.. ...- bimarginatus Schaeffer. There were no significant differ-
ences in densities of G. bimarginatus, M. communis, or total wireworms of all species in
muck versus sand fields. Significantly more Conoderus spp. were found in sandy fields and
significantly more Ischiodontus sp. were found in muck fields. The spatial distribution of the
wireworms within fields was similar in both soil types. In muck, wireworms in 4 fields were
randomly distributed, aggregated in 3 fields, and uniformly distributed in no fields. In sand,
wireworms in 3 fields were randomly distributed, aggregated in 4 fields and uniformly dis-
tributed in no fields.

Key Words: wireworm damage, pest, Elateridae, Melanotus


Los gusanos alambres son plagas importantes de la caha de azucar en la Florida. Nuestro
objetivo fue el determinar la abundancia y distribuci6n espacial de los gusanos alambres so-
bre caha de azucar en suelos lodosos y arenosos en la Florida. Se realize un muestreo de gu-
sano alambre en catorce campos de caha de azucar de fincas comerciales en el sur de la
Florida. Melanotus communis (Gyllenhal) fue la especie mas abundante en las dos classes de
suelo. Otras species encontradas y discutidas menos abundantes fueron: Conoderus spp.,
Ischiodontus sp. y 6. ,1.....--.- bimarginatus Schaeffer. No hubo una diferencia significativa
en la densidad de G. bimarginatus, M. communis o el total de las species de gusanos alam-
bres en campos lodosos versus campos arenosos. Un numero significativamente mayor de
Conoderus spp. fueron encontrados en campos arenosos y un numero significativamente ma-
yor de Ischiodonatus sp. fueron encontrados en campos lodosos. La distribuci6n espacial de
los gusanos alambre dentro de cada campo fue similar en las dos classes de suelo. En campos
lodosos, los gusanos alambre fueron distribuidos al azar en 4 campos, agregados en 3 campos
y no distribuidos uniformemente en ninguno de los campos. En campos arenosos, los gusa-
nos alambre fueron distribuidos al azar en 3 campos, agregados en 4 campos y no distribui-
dos uniformemente en ninguno de los campos.

The Everglades Agricultural Area (EAA) is a
highly productive agricultural area located in
southern Florida. The histosol soils of the EAA,
commonly called muck soils, were formed over a
4,400 year period from partially decomposed re-
mains of hydrophytic vegetation that accumu-
lated under anaerobic wetland conditions result-
ing in highly organic soils (Rice et al. 2005). The
EAA is bordered by less organic sandy soils. Sug-
arcane, vegetables, rice, and sod are the predomi-
nant crops grown on muck soils. These same crops
are grown on sandy soils, and citrus and pastures
also occur on sandy soils.
Wireworms in Florida are primarily a pest in
newly planted sugarcane where the larvae attack

the underground portions of the plant by feeding
on the buds and root primordia during germination
and on shoots and roots after germination. Al-
though the insects are also found at higher densi-
ties in older ratoon sugarcane, they are rarely con-
sidered a pest in ratoon sugarcane because the sug-
arcane plants are large and well established. Of the
different wireworm species found in Florida sugar-
cane, Melanotus communis Gyllenhal (Coleoptera:
Elateridae) is the most important pest. Damage by
this wireworm has been thoroughly documented in
studies by Hall (1985, 1990). Cherry & Hall (1986)
reported flight activity of M. communis in Florida
sugarcane and Cherry (1988) noted distribution
and abundance of the species in Florida sugarcane.

Florida Entomologist 91(3)

Detailed studies of wireworm biology are rare
(Lefko et al. 1998). This paucity of information
has resulted largely because most wireworms are
difficult to collect in large numbers and have pro-
longed life cycles, thereby making them rather in-
tractable for study (Keaster et al. 1975). However,
understanding the role of soil type on wireworm
abundance may be important in predicting ex-
pected wireworm damage. Gui (1935) stated that
soil conditions have a marked influence on wire-
Florida sugarcane is grown in the EAA on soils
ranging from sandy to highly organic muck. Sev-
eral earlier studies (Sosa et al. 1994; Cherry &
Hall 1986; Cherry 2007) reported different as-
pects of wireworm populations in Florida sugar-
cane. However, currently no data exist on actual
wireworm densities and their spatial distribution
in the different soil types where Florida sugar-
cane is grown. Moreover, some Florida sugarcane
growers on sandy soils believe soil insecticides
may be unnecessary at planting because of re-
duced wireworm populations inherent in sandy
soils compared to muck. However, no data exist to
substantiate this belief. Hence, our objective was
to determine the abundance and spatial distribu-
tion of wireworms in Florida sugarcane fields on
muck versus sandy soils. These data should be
useful in predicting wireworm damage in these 2
soil types.


Fourteen commercial sugarcane fields were
sampled for wireworms on farms in southern
Florida. Fields were selected from different areas
and different sugarcane growers in order to ob-
tain a representative sample of wireworm popula-
tions present. Seven fields were located on muck
soils (>40% organic matter) and 7 on sandy soils
(<13% organic matter) to compare wireworm pop-
ulation differences in these 2 soil types. Newly
planted sugarcane has few wireworms present
due to discing and soil insecticide application at
planting. Hence, all fields sampled were second
ratoon (approximately 2 years old) to keep crop
age constant between fields and allow wireworm
populations to accrue since soil insecticides are
not applied after planting.
Mature sugarcane is a very difficult crop in
which to sample insects (Southwood 1969), and
Florida sugarcane may be 3 to 4 m high before
harvest. Therefore, all fields were sampled after
harvest for easy access. All sampling was con-
ducted during a 2-month period to reduce the pos-
sibility of seasonal variation in wireworm num-
bers (Cherry 2007) affecting counts between
fields. Eight fields (4 muck, 4 sand) were sampled
during Feb-Mar 2006 and six fields (3 muck, 3
sand) during Feb-Mar 2007. Sugarcane fields
ranged from 8 to 16 ha in size.

Sugarcane plants (stools) were used for sample
units since most soil dwelling pests of sugarcane
become aggregated around sugarcane plants
(Southwood 1969) as occurs with Florida sugar-
cane grubs (Cherry 1984) and wireworms (Cherry
2007). Each field was divided into 8 equal size
plots (4 x 2 configuration). Five random samples
were taken for wireworms within each plot. Each
sample consisted of a soil sample (40 x 40 x 20 cm
deep) dug around a sugarcane stool and examined
for wireworms for 10 min by 1 person. Examina-
tion time was 5 min if 2 people were present. After
collection, wireworms were brought to the labora-
tory and identified via microscope. Soil samples
were taken from plots, mixed, and the % organic
matter of soil in the field was determined by the
methods of Mylavarapu & Kennelly (2002).
The relative abundance of different wireworms
in the 14 fields was determined. To determine if
soil type influenced wireworm populations, t tests
(SAS 2007) were conducted on the total number of
wireworms found in a field for each of the differ-
ent wireworms in muck versus sandy soil. Data
were transformed before t tests by log 10 (y + 1)
transformation (Steel & Torrie 1980), Untrans-
formed data are presented in tables. A variance to
mean ratio (s2/x) was determined for wireworms
(total number of all species) per plot throughout
each sugarcane field. The ratio is a simple index
for aggregation and was tested for departure from
randomness at alpha = 0.05 with ax2 test where X2
(n -1 df) = s2 (n -1)/x (Southwood & Henderson
2000). The variance to mean ratio was used in this
study because it is the most fundamental of the
various indices of aggregation (Taylor 1984) and
has the advantage of being easy to compute and
readily understandable (Myers 1978). Aggrega-
tion analysis was restricted to total wireworms
combined for all species rather than individual
wireworm species. This was done because when
sampling sugarcane, growers do not differentiate
between wireworm species.


Melanotus communis was the most abundant
wireworm found in both soil types with more be-
ing found in muck soils (Tables 1 and 2). These
data are consistent with Cherry & Hall (1986),
who reported that more adult M. communis were
caught in light traps from Florida sugarcane
fields on muck soils than on sandy soils. However,
there was no significant difference (t = 1.1, df =
12, P > 0.05) in M. communis population densities
in muck versus sand fields in this study, which is
partially explained by the extreme variability
found in M. communis between fields in both soil
types. Hall (1988) reported that M. communis is
an important soil pest of Florida sugarcane and
that insecticides are routinely applied at planting
time for control.

September 2008

Cherry & Stansly: Florida Sugarcane Wireworms



Field # % OM Conoderus spp. G. bimarginatus M. communis Ischiodontus sp. Total

1 82 6 2 151 12 171
2 81 1 4 163 2 170
3 82 1 7 146 2 156
4 82 1 5 131 0 137
5 51 2 2 23 15 42
6 40 0 1 9 2 12
7 82 5 1 104 20 130
Total 16 22 727 53 818

aPercentage organic matter of soil.
'Total wireworms in 40 samples in each field.

Conoderus spp. were the second most abun- worm are sometimes encountered in Florida sug-
dant wireworms found in both soil types with 6 arcane which is consistent with the highly vari-
times as many being found in sandy soils. There able distribution observed among fields in this
were significantly more (t = 4.5, df = 12, P < 0.05) study. Little is known of the biology or economic
Conoderus spp. in sandy fields than muck fields. impact of Ischiodontus sp. in Florida sugarcane.
Hall (1988) reported 4 Conoderus species com- Gi pi.... bimarginatus Schaeffer were the
only associated with Florida sugarcane. How- least abundant wireworms found in the sugar-
ever, Conoderus species were not determined in cane fields with equal population densities found
this study. Conoderus have been reported to be in both soil types. Obviously, there was no signifi-
pests of sugarcane in Louisiana (Bynum et al. cant difference (t = 0.9, df = 12, P > 0.05) in G. bi-
1949) and Hawaii (Stone 1976). marginatus in muck versus sandy fields. It is
Ischiodontus sp. was the third most abundant probable that G. bimarginatus are more numer-
wireworm found in both soil types with almost all ous in Florida sugarcane than shown here be-
(95%) being found in muck soils. As expected, cause they were the smallest wireworm species
there were significantly more (t = 3.1, df= 12, P < found in this study and were probably underesti-
0.05) Ischiodontus sp. in muck fields than sandy mated in visual samples. Hall (1988) reports that
fields. Similarly, Gui (1935) reported that organic G. bimarginatus is a small wireworm often
matter content of soils had a positive relationship present in Florida sugarcane. Little is known of
with wireworm populations of Agriotes mancus the biology or economic impact of G. bimarginatus
Say in Ohio. Furthermore, Pill et al. (1976) noted in Florida sugarcane.
that the wireworm Limonius dubitans Leconte There were 37% more wireworms of all species
was a pest only in higher organic soils. Hall (1988) in muck fields than in sandy fields. However, vari-
reported that localized populations of this wire- ability in total wireworm population densities be-



Field # % OM Conoderus spp. G. bimarginatus M. communis Ischiodontus sp. Total

1 2 17 0 32 0 49
2 12 13 11 2 1 27
3 3 6 0 125 0 131
4 2 8 0 115 2 125
5 3 9 7 116 0 132
6 3 33 4 0 0 37
7 3 6 0 92 0 98
Total 92 22 482 3 599

aPercentage organic matter of soil.
'Total Wireworms in 40 samples in each field.

Florida Entomologist 91(3)


Field Variance Mean' Ratio Chi-squareb Distribution

Muck soil
1 77.4 21.4 3.61 25.3 Aggregated
2 44.9 21.3 2.11 14.7 Aggregated
3 34.8 19.5 1.78 12.5 Random
4 25.0 17.1 1.46 10.2 Random
5 14.4 5.3 2.72 20.2 Aggregated
6 1.7 1.5 1.13 7.9 Random
7 27.0 16.3 1.66 11.6 Random
Sandy Soil
1 14.4 6.1 2.36 16.6 Aggregated
2 2.9 3.4 0.85 5.9 Random
3 68.9 16.4 4.20 29.4 Aggregated
4 26.0 15.6 1.73 11.7 Random
5 54.8 16.5 3.32 23.2 Aggregated
6 3.6 4.6 0.78 5.5 Random
7 44.9 12.3 3.65 25.5 Aggregated

"Mean of all wireworms in a plot.
'Chi-square = Variance (n-1) divided by mean.

tween fields was high in both soil types and more
wireworms were found in some sandy fields than
muck fields. These data resulted in there being no
significant difference (t = 0.5, df= 12, P > 0.05) in
total wireworm densities in muck versus sandy
fields. These data show that soil type alone cannot
be used accurately to predict wireworm densities
in Florida sugarcane fields.
Overall, spatial distribution patterns of wire-
worms were similar in both soil types (Table 3). In
muck, wireworms in 4 fields were randomly dis-
tributed and aggregated in 3 fields. In sand, wire-
worms in 3 fields were randomly distributed and
aggregated in 4 fields.
Reasons for the aggregation of wireworms in
some of the fields are not known. However, we ob-
served minor soil type differences within some
fields, especially in sandy soils. Sandy soils are
subject to rapid soil transitions and also may con-
tain small areas called "muck pockets". Salt &
Hollick (1946) noted that soil type affected wire-
worm distribution in pastures and hence this may
have caused the wireworm aggregation we ob-
served in some fields. Also, we observed minor soil
moisture differences within some fields due to low
areas with wetter soil. Lefko et al. (1998) noted
that soil moisture may be important in affecting
wireworm distribution and this may have been a
factor in wireworm aggregation in some fields.
As a last note, Sosa et al. (1994) reported that
wireworm populations were greater towards field
centers in Florida sugarcane although only a
weak correlation (r = 0.16) was shown. However,
since sampling methods between the former
study and our study were quite different, it is not
possible to directly compare results of the 2 stud-

ies. Moreover, Southwood & Henderson (2000)
have noted that sampling method and sample size
may affect the apparent distribution of an organ-
ism. Since we used the same methodology in both
soil types, our main conclusions that the wire-
worm spatial distribution patterns were similar
in both soil types remains valid.
In summary, our data show that soil type alone
cannot be used to accurately predict total wire-
worm densities in Florida sugarcane fields.
Hence, sugarcane growers on both muck and sand
soils face similar wireworm pressure which may
necessitate soil insecticide application at plant-
ing. Also, our data show high inter-field variabil-
ity in total wireworm densities in both soil types.
These latter data suggest that some sugarcane
fields with low wireworm densities could be
planted without a soil insecticide if sampling
methodology existed to determine this. Currently,
we are developing sampling methods to determine
when soil insecticides are necessary for wireworm
control when planting Florida sugarcane.

trol of wireworms attacking sugarcane in Louisiana.
J. Econ. Entomol. 42: 556-557.
CHERRY, R. 1984. Spatial distribution of white grubs
(Coleoptera: Scarabaeidae) in Florida sugarcane. J.
Econ. Entomol. 77: 1341-1343.
CHERRY, R. 1988. Correlation of crop age with popula-
tions of soil insect pests in Florida sugarcane. J. Ag-
ric. Entomol. 5: 241-245.
CHERRY, R. 2007. Seasonal population dynamics of
wireworms (Coleoptera: Elateridae) in Florida sug-
arcane fields. Florida Entomol. 90: 426-430.

September 2008

Cherry & Stansly: Florida Sugarcane Wireworms

CHERRY, R., AND D. HALL. 1986. Flight activity of Mel-
anotus communis (Coleoptera: Elateridae) in Flor-
ida sugarcane fields. J. Econ. Entomol. 79: 626-628.
GUI, H. 1935. Soil types as factors in wireworm distri-
bution. Am. Potato J. 7(5): 107-113.
HALL, D. 1985. Damage by the corn wireworm, Melano-
tus communis (Gyll.) to plant cane during germina-
tion and early growth. J. American Soc. Sugar Cane
Tech. 4: 13-17.
HALL, D. 1988. Insects and mites associated with sugar-
cane in Florida. Florida Entomol. 71: 130-150.
HALL, D. 1990. Stand and yield losses in sugarcane
caused by the wireworm Melanotus communis (Co-
leoptera: Elateridae) infesting plant cane in Florida.
Florida Entomol. 73: 298-302.
ing behavior and growth of the wireworms Melano-
tus depressus and Limonius dubitans: effect of host
plants, temperature, photoperiod, and artificial di-
ets. Environ. Entomol. 4: 591-595.
1998. Spatial modeling of preferred wireworm (Co-
leoptera: Elateridae) habitat. Environ. Entomol. 27:
MYERS, J. 1978. Selecting a measure of dispersion. En-
viron. Entomol. 7: 619-621.
Extension Soil Testing Laboratory (ESTL) Analyti-
cal Procedures and Training Manual. University of
Florida Extension Circular No. 1248.

val survival and pupation of the wireworms Melano-
tus depressus and Limonius dubitans in natural
substrates. Environ. Entomol. 5: 845-848.
RICE, R., R. GILBERT, AND S. DAROUB. 2005. Application
of the Soil Taxonomy Key to the Organic Soils of the
Everglades Agricultural Area. University of Florida
document SS-AGR-246.
SALT, G., AND F. HOLLICK. 1946. Studies of wireworm
populations. II. Spatial distribution. J. Exp. Biol. 23:
SAS INSTITUTE. 2007. SAS Institute, Cary, NC.
SOSA, 0., B. GLAZ, M. ULLOA, AND V. CHEW. 1994. Sam-
pling of sugarcane fields for wireworms (Coleoptera:
Elateridae). J. American Soc. Sugar Cane Tech. 14:
SOUTHWOOD, T. 1969. Population studies of insects at-
tacking sugarcane, pp. 427-461 In J. Williams, J.
Metcalfe, R. Mungomery, and R. Mathers [eds.],
Pests of Sugar Cane. Elsevier, New York.
SOUTHWOOD, T., AND P. HENDERSON. 2000. Ecological
Methods. Blackwell Science, Oxford.
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STONE, M. 1976. Notes on the biology of the introduced
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Florida Entomologist 91(3)

September 2008


USDA-APHIS, 41-650 Ahiki Street, Waimanalo, HI 96795


Polyandry has been reported for various species of tephritid fruit flies, and efforts to explain
this phenomenon have focused primarily on the respective roles of sperm stores and male ac-
cessory gland fluids. However, recent research on the Mediterranean fruit fly, Ceratitis cap-
itata (Wiedemann), demonstrated that female re-mating propensity was influenced by the
"suitability" of available mates; preferred males were more likely to induce female re-mating
than were less attractive males. The goal of this study was to determine whether male at-
tractiveness similarly influenced female re-mating in the oriental fruit fly, Bactrocera dor-
salis (Hendel). Females of this species are known to mate preferentially with males that
previously ingested methyl eugenol (ME), a plant-borne compound whose metabolites are
used in the synthesis of the male sex pheromone. Females were mated initially (with males
denied ME), held isolated from males with access to oviposition substrate, and then offered
ME-fed or non-ME-fed males at 1, 2, 3, or 6 weeks after the initial mating. Contrary to the
Mediterranean fruit fly, there was no effect of male attractiveness (i.e., ME feeding status)
on the re-mating tendency of B. dorsalis females. Also, time elapsed since the initial mating
had no effect on the probability of female re-mating. When offered a choice of males, how-
ever, non-virgin females, like virgins, displayed a significant preference for ME-fed over non-
ME-fed males.

Key Words: Bactrocera dorsalis, polyandry, mate selection, pheromone


La poliandria ha sido reportado en varias species de moscas tefritidas de la fruta y los es-
fuerzos hechos para explicar este fen6meno se han enfocado mayormente en los papeles res-
pectivos de los almacenes de esperma y los fluidos de las glandulas accesorias del macho. Sin
embargo, investigaciones recientes sobre la mosca mediterranea (Ceratitis capitata Wiede-
mann) de la fruta, han demostrado que la propensidad de las hembras para aparearse de
nuevo fue influenciada por la idoneidad de los machos disponibles; fue mas possible para los
machos preferidos el inducir a las hembras a aparearse de nuevo que los machos menos
atractivos. La meta de este studio fue el determinar en la mosca oriental de la fruta, Bac-
trocera dorsalis (Hendel) si los machos atractivos similarmente influenciaron a las hembras
a aparearse de nuevo. Se sabe que las hembras de esta especie se aparean preferencialmente
con machos que ingerieron eugenol metilo (EM), un compuesto producido de la plant cuyos
metabolitos son usados en la sintesis de las feromonas sexuales del macho. Las hembras fue-
ron apareadas inicialmente (con machos denegados de EM), mantenidas aisladas de los ma-
chos con aceso del sustrato para ovipositar y luego ofrecidas a machos alimentados con EM
o sin EM a las 1, 2, 3 o 6 semanas despu6s del apareamiento inicial. Al contrario de la mosca
mediterranea de la fruta, no hubo efecto de los machos atractivos (alimentados con EM) so-
bre la tendencia de las hembras de B. dorsalis a aparearse de nuevo. Ademas, la cantidad de
tiempo que pasaron despu6s del apareamiento inicial no tuvo efecto sobre la probabilidad de
las hembras a aparearse de nuevo. Cuando se ofrecio una opci6n de machos, sin embargo, las
hembras no virgenes, mostraron una preferencia significativa a los machos alimentados con
EM que a los machos no alimentados con EM.

Several non-mutually exclusive hypotheses
have been proposed for the widespread occur-
rence of polyandry in insects (Ridley 1988; Sim-
mons 2001). Females may remate to (i) replace de-
pleted sperm stores (sperm replenishment), (ii)
acquire male-controlled resources (material bene-
fits), (iii) replace sperm from a previous mate with
sperm from a genetically superior partner (ge-
netic benefits), (iv) minimize male harassment
(convenience), or because (v) the propensity for
female re-mating is a genetically correlated re-

sponse to sexual selection on multiple mating by
males (correlated evolution). Empirical support
exists for all these hypotheses, with the possible
exception of the final one involving the genetic
link between multiple mating in the sexes (Sim-
mons 2001).
Polyandry has been reported for various spe-
cies of tephritid fruit flies, including representa-
tives from the following genera: Ceratitis (Boniz-
zoni et al. 2002; Vera et al. 2003), Bactrocera (Tza-
nakakis et al. 1968; Ito & Yamagishi 1989; Song

Shelly & Edu: Re-mating Propensity of Female Fruit Flies

et al. 2007),Anastrepha (Sivinski & Heath 1988),
Toxotrypana (Landolt 1994), and Rhagoletis (Opp
& Prokopy 2000). Although widespread within
the family, little work has been directed toward
identifying the factors) promoting polyandry, and
most of this has concerned only two species, the
Mediterranean fruit fly (medfly), C. capitata
(Wiedemann), and the Queensland fruit fly, B. try-
oni (Froggatt). In the medfly, sperm replenish-
ment and male accessory gland fluids both appear
to influence female re-mating (Katiyar & Ramirez
1970; Nakagawa et al. 1971; Delrio & Cavalloro
1979; Jang 1995, 2002; Miyatake et al. 1999;
Mossinson & Yuval 2003). In contrast, in the
Queensland fruit fly, male accessory gland fluids
play a major role in inhibiting female re-mating,
whereas sperm stores appear to have little influ-
ence on female propensity to remate (Harmer et
al. 2006; Radhakrishnan & Taylor 2007; see Kuba
& Ito 1993 for similar results for the melon fly,
B. cucurbitae (Coquillett)).
As evidenced by these studies, research on te-
phritids has focused exclusively on the effect of
sperm supply on female re-mating or the role of
male accessory fluids in inhibiting female re-mat-
ing. In lek-forming tephritids, including many
Ceratitis and Bactrocera species, males do not
control resources vital to females (Shelly & Whit-
tier 1997). Consequently, it appears unlikely that
females of these species mate multiply to acquire
additional material resources. Furthermore, in le-
kking species, it appears unlikely that females re-
mate to minimize male harassment, since court-
ship and copulation proceed only after female ap-
proach to stationary, signaling males (Shelly &
Whittier 1997). Alternatively, the hypothesis that
female tephritids remate to obtain sperm from a
superior mate (i.e., to 'trade up', Halliday 1983) is
conceivable but untested.
In a recent study, Shelly et al. (2004b) found
that male attractiveness influenced female re-
mating in the medfly. Previous work demon-
strated that medfly females mate preferentially
with males exposed to the aroma of certain plant-
derived oils, such as orange oil (Shelly et al.
2004a) and ginger root oil (GRO; Shelly 2001),
over non-exposed males. In assessing female re-
mating propensity, Shelly et al. (2004b) found
that non-virgin females were more likely to mate
a second time if offered GRO-exposed males as op-
posed to non-exposed males. Whether this result
indicates genetic trading up is unknown, because
the adaptive basis of female preference for GRO-
exposed males is unknown. Nonetheless, it does
indicate that female re-mating behavior is flexible
and may vary with the perceived quality of males
available for repeat matings.
The goal of the present study was to determine
whether male attractiveness similarly influenced
re-mating by females of the oriental fruit fly, B.
dorsalis (Hendel). Although field data are lacking,

Shelly (2000a) found that B. dorsalis females
have a fairly high incidence of re-mating in the
laboratory. Over an 8-week period during which
males were provided for one twilight (mating) pe-
riod per week, approximately 50% of the 128 fe-
males observed re-mated, with most of these
(87%) re-mating only once. Consistent with the
aforementioned studies on other Bactrocera spe-
cies, re-mating by B. dorsalis females was not ob-
viously related to sperm depletion, since (i) the in-
cidence of re-mating was independent of the
length of time elapsed since the initial mating
and (ii) egg production did not vary significantly
before or after re-mating for most females.
Males of the oriental fruit fly are highly at-
tracted to methyl eugenol (ME), a compound
found in over 200 families of the plants (Tan &
Nishida 1996). Males ingest this compound and
use its metabolites to produce a long-range sex
pheromone (Nishida et al. 1988). Several studies
(Shelly & Dewire 1994; Tan & Nishida 1996) have
demonstrated that ME consumption increases
male mating success, apparently owing to the in-
creased attractiveness of the male's pheromonal
signal. Thus, as in the medfly, B. dorsalis females
preferentially mate with males previously ex-
posed to a particular chemical, thus allowing ex-
perimental manipulation of male attractiveness
and subsequent evaluation of the effect of such
manipulation on female re-mating propensity.


Study Insects

All flies used in the present study were from a
laboratory colony started with 400-600 adults
reared from mangos (Mangifera indica L.) col-
lected in Waimanalo, Oahu. The colony was main-
tained in a screen cage (l:w:h, 1.2 x 0.6 x 0.6 m)
and provided a mixture (3:1, wt:wt) of sugar (su-
crose) and enzymatic yeast hydrolysate and water
ad libitum and papayas (Carica papaya L.) for
oviposition. Infested papayas were held over ver-
miculite, and the pupae were sifted from vermic-
ulite 16-18 d later. Adults used in the mating tri-
als were separated by sex within 48 h of eclosion,
well before reaching sexual maturity at 15-18 d of
age (TES, unpublished data), and held in screen-
covered, 5-L plastic buckets; 100-125 individuals
per bucket) with ample food and water. Flies were
held at 24-28C and 60-90% RH and received nat-
ural and artificial light under a 12:12 (L:D) photo-
period. When used in the study, the flies were 3-4
generations removed from the wild.

Female Re-mating

Mating activity in B. dorsalis is restricted to an
approximately 1-h long period immediately pre-
ceding sunset (Roan et al. 1954; Arakaki et al.

Florida Entomologist 91(3)

1984). Although males display vigorous wing-fan-
ning (presumably to disperse the sex pheromone,
Schultz & Bousch 1971; Kobayashi et al. 1978),
little courtship is evident, and males jump on any
approaching female. Mating pairs remain coupled
through the night and break apart at sunrise.
To obtain initial matings, we placed approxi-
mately 50 males and 50 females (all 18-23 d-old
virgins) in plexiglass cages (30 x 30 x 40 cm) in the
afternoon (1400-1600 h). On a given day, 5-10
mating cages were established. Room lights were
extinguished when flies were placed in the cages,
consequently mating occurred under natural
light. We removed unmated flies from the cages 3-
4 h after sunset (using an aspirator under dim
light) and left the mating pairs undisturbed in the
cage through the night. The following morning we
discarded the males and transferred the mated fe-
males to screen cages (30 cm cubes, 50-75 females
per cage). Papayas were provided every other day
(for 6-8 h) as an oviposition substrate (to mimic
field conditions where females likely have multi-
ple egg-laying opportunities) starting 2 d after the
initial mating and continuing until the females
were tested for re-mating either 1, 2, 3, or 6 weeks
after the initial mating. We selected these inter-
vals to monitor re-mating frequency soon after
the initial mating (in possible response to low
sperm transfer) as well as long after the initial
mating (in possible response to sperm depletion
through extensive oviposition). Ample food and
water were provided and changed regularly.
To measure re-mating, we placed 10 mated fe-
males and 10 virgin males (19-29 d old) in plexi-
glass cages and scored matings in the same man-
ner described above. In any given cage, males ei-
ther had never been given access to ME or were
fed ME the day before testing. To obtain ME-fed
males, we applied 100 pL of ME to a cotton wick
(held vertically by insertion through a hole in the
lid of a plastic cup), which was then placed in a
bucket holding 60-70 males. The chemical was in-
troduced between 1000-1200 h and removed 1 h
later. Feeding activity was not monitored, but in a
previous study (Shelly 1997) over 90% of mature
males fed on ME within a 1-h interval. For all
combinations of post (initial)-mating interval and
male treatment, re-matings were scored in 15
cages over 3-6 different days.

Female Mating Status and Mate Selection

As described below, the frequency of female re-
mating varied independently of the ME feeding
status of the available males. This finding sug-
gested that, unlike virgin females (Shelly & Dew-
ire 1994; Tan & Nishida 1996), non-virgin females
may not discriminate among potential mates
based on their ME status, and consequently we
conducted a separate experiment to investigate
this possibility.

Following the methods presented above, fe-
males were mated and held for testing 1 week
later. On test days, we placed 1 non-virgin female,
1 ME-fed male, and 1 non-ME-fed male in trans-
parent, 3.7-L plastic bottles 3-4 h before sunset
and then scored the identity of mating males 2-4
h after sunset (males were marked 1 d prior to
testing by placing a small dot of enamel paint on
the thorax of chilled individuals). Following the
same protocol, we also tested virgin females for
discrimination between ME-fed and non-ME-fed
males. On a given test day (n = 10 for mated fe-
males and n = 5 for virgin females, respectively),
we set up 31-55 bottles with individual non-virgin
females (for a total of 443 females tested) and 15-
20 bottles with individual virgin females (for a to-
tal of 94 females tested).

Statistical Analysis

As neither the raw nor (log) transformed data
was normally distributed, we assessed the impact
of male ME 'status' (fed or non-fed) using Fried-
man's test, a nonparametric equivalent of the
parametric 2-way analysis of variance (Daniel
1990), and treated time since the initial mating as
the blocking variable. The mean numbers of re-
matings observed for the different time by male
status combinations were used in the analysis. In
our analysis, weeks since the initial mating were
the blocks, and the ME status of males presented
for re-mating were the treatments. Following the
Friedman test, we compared female re-mating
frequency across time intervals with a Kruskal-
Wallis test. In the mate choice experiment, we
compared the observed mating frequencies for
virgin and non-virgin females to that expected by
chance (50% for each male type) using the normal
approximation to the binomial. Analyses were
performed with SigmaStat Statistical Software
(Version 2.0). Means 1 SE are presented.


With the effect of the blocking variable (i.e.,
weeks since initial mating) removed, we found no
significant effect of male ME feeding status on fe-
male re-mating tendency (x2 = 0.24, df = 1, P =
0.65, Fig. 1). Using data pooled from both male
types for each interval, we also found no signifi-
cant variation in female re-mating frequency
across the different time intervals (H = 4.1, df= 3,
P = 0.26). Over all time intervals, the average
number of females re-matings ranged only from
2.1-3.2 from a possible maximum of 10 when pre-
sented with ME-fed males and from 2.0-2.8 from a
possible maximum of 10 when presented with
ME-deprived males.
Consistent with previous work, virgin females
showed a significant preference for ME-fed males,
selecting them in 70% (35/50) of the total matings

September 2008

Shelly & Edu: Re-mating Propensity of Female Fruit Flies




M~ 25




1 2 3

Weeks after initiJ

Fig. 1. Re-matings by B. dors
with treated (ME-fed) or control
Values represent mean numbers
cases) of re-matings per cage (10

(Z = 2.61, P < 0.01). Non-virg
showed a significant preferen
66% (34/53) of the re-mati
ME-fed males (Z = 2.0, P < 0
mating frequency noted for n
periment (53/443 = 12%) was
that observed in the precedi
25%). This difference in ma
have reflected the differing f
the 2 experiments. Fly densi
sexes) was 3.03 flies/1000 cm
in the main experiment ex:
mating propensity but only 1
1530 cm2) in the mate choice


Based on the data present
ing frequency ofB. dorsalis fe
dent of the time elapsed sin
This result is consistent with
study (Shelly 2000a) that n
and egg production for B.
weekly intervals over an 8-wi
were supplied periodically in
the prior study, it seems likely
eggs laid by individual female
time available for ovipositi
elapsed since the initial mati
tion is valid, the time-indep
re-mating indicates that fem.
dependent of egg deposition
fly. This finding differs from
several other tephritid specie
1988; Landolt 1994). For exa
the Caribbean fruit fly, A. sus,
ski and Heath (1988) found
provided an oviposition subst
1 week of their initial matin

10% of females that were not provided an oviposi-
Treated males (ME-fed) tion substrate. However, consistent with the
- Control males (no ME) present study, Chapman et al. (1998) reported no
difference in re-mating frequency between nor-
T mal (egg-laying) and irradiated (non-egg-laying)
females of C. capitata. These differing results in-
dicate that oviposition has a variable effect on fe-
male re-mating among tephritid species.
The data presented above also reveal that fe-
male re-mating was independent of the ME-medi-
ated attractiveness of the males available for a
second mating. This result was unexpected in
light of (i) the earlier study on the Mediterranean
4 5 6 7- fruit fly (Shelly et al. 2004b) showing that GRO-
al maung exposed males induced a higher level of female re-
mating than non-exposed males, (ii) the repeated
alis females presented demonstration (Shelly & Dewire 1994; Shelly &
(ME-deprived) males. Nishida 2004; Shelly et al. 2005, 2007) of the
(1 SE, n = 15 in all strong preference of virgin B. dorsalis females for
females per cage). ME-fed over ME-deprived males and (iii) the com-
mon observation from various taxa, including
crickets (Bateman et al. 2001), spiders (Watson
,in females likewise 1991), pseudoscorpions (Zeh & Zeh 2007), newts
ce for ME-fed males: (Gabor & Halliday 1997), and guppies (Pitcher et
ig females selected al. 2003), that once-mated females display greater,
.05). The overall re- not weaker, mate selectivity than virgin females.
on-virgins in this ex- Although methyl eugenol feeding had no effect
approximately half on female re-mating frequency in no-choice tests,
ng one (301/1200 = non-virgin females given a choice between ME-
ting frequency may fed and ME-deprived males displayed a signifi-
ly densities used in cant preference for ME-fed individuals. Taken to-
ty (considering both gether, these two findings indicate that, after a
2 (20 flies/6600 cm2) certain refractory period, a certain (time-indepen-
amining female re- dent) proportion of mated females are sexually re-
.31 flies/cm2 (2 flies/ ceptive and, male type (with respect to methyl eu-
test. genol feeding) will not alter this proportion. How-
ever, when given a choice of male type, those fe-
N males that are sexually receptive will generally
select ME-fed males over ME-deprived males.
ed above, the re-mat- While virgin and non-virgin B. dorsalis females
males was indepen- show congruence in mate choice, the adaptive
ce the first mating. benefit of selecting ME-fed males remains un-
the aforementioned known. As noted previously (Shelly 2000b), fe-
ronitored re-mating males do not appear to gain direct benefits (i.e.,
dorsalis females at increased fecundity or longevity) by selecting ME-
eek period. As fruits fed mates, raising the possibility that, by select-
both the present and ing ME-fed males, females may increase the prob-
y that the number of ability of producing'sexy sons', capable of locating
*s increased with the ME and gaining copulations.

on (i.e., with time
ng). If this assump-
endent likelihood of
ale receptivity is in-
in the oriental fruit
those reported for
s (Sivinski & Heath
mple, working with
pensa (Loew), Sivin-
that 67% of females
rate remated within
ig compared to only


We thank Boaz Yuval and an anonymous reviewer
for helpful comments on an earlier draft.


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Florida Entomologist 91(3)

September 2008


USDA-APHIS, 41-650 Ahiki Street, Waimanalo, HI 96795


In Sterile Insect Technique (SIT) programs, mass-reared males of the Mediterranean fruit fly,
Ceratitis capitata (Wiedemann), are maintained on a sugar-agar diet before their release into
the environment. Several studies suggest that nitrogenous dietary supplements improve the
mating competitiveness of the sterile males, thus increasing the cost effectiveness of SIT.
Other research, however, has not supported this notion. Here, we further investigate the po-
tential usefulness of nitrogen-containing diet additives by examining the effect of 4 different
nitrogenous materials-yeast hydrolysate, urea, whey protein, and honey-on the mating
success of sterile C. capitata males. These materials were mixed directly with the sugar-agar
diet in varying concentrations (1, 5, or 10%). Trials conducted in outdoor field-cages generated
2 consistent results across all diets: (1) neither the type nor concentration of nitrogenous ma-
terial used elevated the mating success of sterile males above that observed for the standard
sugar-agar diet, and (2) wild males invariably had a significant mating advantage over sterile
males, accounting for an average of 81% of the total matings per replicate. In addition, a field
comparison of short-term dispersal from a central release point revealed no significant differ-
ence between sterile males fed (i) the sugar-agar diet or (ii) the sugar-agar diet supplemented
with yeast hydrolysate (10%) in either the number or spatial distribution of recaptured indi-
viduals. Our results do not support the proposal that pre-release nitrogenous dietary supple-
ments improve the field performance of sterile C. capitata males in SIT programs.

Key Words: Ceratitis capitata, Sterile Insect Technique, diet, sexual competitiveness, dis-


En programs con t6cnicas de insecto est6ril (TIE), los machos criados masivamente de la
mosca mediterranea (Ceratitis capitata Wiedemann) de la fruta, fueron mantenidos con una
dieta de azucar-agar antes de ser liberados al medio ambiente. Varios studios sugieren que los
suplementos con nitr6geno en la dieta mejora la habilidad de los machos est6riles para apa-
rearse, a su vez aumentando la eficiencia de los costs del program de TIE. Sin embargo, otro
studio no ha suportado este resultado. Aqui, investigamos el uso potential de los aditivos de
dieta que contienen nitr6geno con el evaluo del efecto de 4 materials de nitr6geno diferen-
tes-levadura hidrolizada, urea, protein de suero y miel-sobre el 6xito de machos est6riles
de C. capitata para aparearse. Se mezclaron estos materials directamente con la dieta de azu-
car-agar en varias concentraciones (1, 5 o 10%). Las pruebas que se realizaron en jaulas del
campo mostraron 2 resultados en todas las dietas: (1) ni la clase o la concentraci6n de material
de nitr6geno usada elevaron el 6xito de los machos est6riles para aparearse sobre el observado
con la dieta de azucar-agar y (2) los machos salvajes invariablemente expresaron una ventaja
significativa mayor en el apareamiento que los machos est6riles, representando un promedio
de 81% del total de los apareamientos por repetici6n. Ademas, una comparici6n en el campo de
la dispersion de corto plazo de los machos est6riles liberados de un punto central revel6 nin-
guna diferencia significativa entire los machos est6riles alimentados de (i) la dieta de azucar-
agar o (ii) la dieta azucar-agar suplementada con levadura hidrolizada (10%) en tanto el nu-
mero o distribuci6n espacial de los individuos capturados. Nuestros resultados no apoyan la
proposici6n que los suplementos de nitr6geno en la dieta de los machos antes de ser liberados
mejoran el desempeno en el campo de los machos est6riles de C. capitata en programs de TIE.

In a seminal paper, Yuval et al. (1998) reported contained higher levels of protein and sugar than
that the nutritional status of male Mediterranean resting males, which were perching outside the
fruit flies (medflies), Ceratitis capitata (Wiede- lek and not signaling. This result garnered con-
mann), affected their ability to participate in mat- siderable attention because it hinted that im-
ing aggregations or leks. Biochemical analysis of provements to the adult diet might effectively in-
field-captured males showed that lekking males, crease the mating competitiveness of mass-
which were actively signaling (i.e., emitting sex reared, sterile males used in the Sterile Insect
pheromone) from leaf territories when collected, Technique (SIT) against the medfly.

Shelly & Edu: Diet and Mating Success of Sterile Male Medflies

Recently, Yuval et al. (2007) reviewed the liter-
ature pertaining specifically to the effects of pro-
tein as an adult dietary supplement on the mat-
ing success of C. capitata males. Barry et al.
(2007) reported that yeast hydrolysate, the pur-
ported protein source used in these studies, actu-
ally contains very little protein and is more accu-
rately identified as a nitrogen or amino acid
source. Work on wild flies has uniformly demon-
strated that males given access to sugar (sucrose)
+ yeast hydrolysate gain an advantage in mating
competition over males fed sugar only (Kaspi et
al. 2000; Shelly & Kennelly 2002; Shelly et al.
2002). In contrast, studies on sterile males have
generated inconsistent results. For example,
Kaspi & Yuval (2000) found that sterile males fed
a sugar-agar gel (the standard pre-release, adult
diet used in SIT programs; e.g., Dantas et al.
2004; Barnes et al. 2004) supplemented with
yeast hydrolysate (9%) achieved significantly
more copulations than males fed the sugar-agar
gel alone. In contrast, in studies conducted in Ha-
waii (Shelly & Kennelly 2002; Shelly & McInnis
2003; Shelly et al. 2006), sterile C. capitata males
provided sugar only obtained similar numbers of
matings as sterile males fed sugar and yeast hy-
drolysate. Mating trials performed in Guatemala
further suggested that the effects of yeast hy-
drolysate may vary with environmental condi-
tions: supplementing sugar with yeast hydroly-
sate significantly increased the mating perfor-
mance of sterile males at a cool, high elevation
site but had no detectable effect at a warmer, low
elevation site (Shelly et al. 2004).
The present study further investigates the po-
tential effectiveness of nitrogen-containing adult
diet supplements in enhancing the mating success
of sterile C. capitata males. Unlike previous studies
(except Kaspi & Yuval 2000), we presented the ni-
trogen sources mixed in the standard sugar-agar gel
and not separately from granular sugar. Addition-
ally, whereas previous studies used yeast hydroly-
sate as a nitrogen source exclusively (excepting
Maor 2004 et al., who used dried apricots), the ex-
periments described herein were performed not
only with yeast hydrolysate, but with several alter-
nate nitrogen sources, namely urea, whey protein,
and honey. Also, these supplements were added in
different concentrations to examine possible dose-
dependent effects. Finally, in addition to mating per-
formance, we compared dispersal between sterile
males fed the sugar-agar gel and sterile males fed
the sugar-agar gel with yeast hydrolysate added.


Study Insects

Wild flies were reared from infested coffee, Cof-
fea arabica L., berries collected near Haleiwa,
Oahu. Fruits were held over vermiculite and lar-

val development proceeded in situ. Pupae were
sifted from the vermiculite 7-9 d after fruit collec-
tion, and adults used in this study were separated
by sex within 2 d of eclosion, well before reaching
sexual maturity at 6-8 d of age. Adults were held
in plastic buckets covered with nylon screening
(volume 5 L; 100-125 flies per bucket). Wild flies
were provided with a mixture (3:1 v/v) of sugar
and yeast hydrolysate and water ad libitum, held
at 24-28C and 60-90% RH, and received both
natural and artificial light in a 12:12 (L:D) photo-
period. When used in the present study, the wild
flies were 3-6 generations removed from the wild.
Mass-reared males were from a temperature
sensitive lethal (tsl) genetic sexing system (Vienna-
7/Tol-99) and were reared by the California Depart-
ment of Food and Agriculture (CDFA) Hawaii Fruit
Fly Rearing Facility, Waimanalo, Oahu In rearing
this strain, eggs are exposed to high temperature,
which selectively kills female embryos and thus al-
lows production and release of males only (Franz et
al. 1994). Pupae were obtained 2 d before eclosion
after dusting with pink fluorescent dye (particles of
which adhere to the emerging adults, thus serving
as a strain marker) and gamma irradiation at 150
Gy with a 137Cs source. On a given day, pupae were
placed in 4 screen-covered plastic buckets (approx-
imately 150 pupae per bucket), and emerging
adults were provided a prescribed diet (see below)
and water and maintained under the same condi-
tions described above.

Dietary Treatments of Mass-Reared Males

As described below, we compared the mating
success of mass-reared males that were fed the
standard sugar-agar diet or a sugar-agar diet to
which varying amounts of nitrogen-containing
material were substituted for the corresponding
amount of sugar. The sugar-agar gel was prepared
following the recipe used by the CDFA's Medfly
Preventative Release Program (95.02% sugar,
4.91% agar, and 0.07% methyl paraben; 1 L of
water per 181 g of dry matter; I. Walters, pers.
comm.). Four different nitrogenous substances
were tested: yeast hydrolysate (USB Corporation,
#216-765-5000, containing protein and amino ac-
ids, Tsiropoulos 1978; Morton & Bateman 1981;
Barry et al. 2007), urea (Mallinckrodt Baker, Inc.,
#8642-12; NH2CONH2), whey protein (BIO-
CHEM Sports), and honey (containing amino
acids plus vitamins and minerals, White et al.
1962; Hermosin et al. 2003). All 4 of these sub-
stances were added at 3 different levels, 1%, 5%,
and 10% of the total amount of sugar (with sugar
reduced by the equivalent amount), after the
sugar-agar gel had cooled below 45C to avoid pro-
tein denaturation.
Food was placed on the plastic buckets on the
day of adult emergence (i.e., 2 d after pupal place-
ment). Slabs of diet (10 x 6 x 3 cm, l:w:h) were

Florida Entomologist 91(3)

placed on the screen-covering, overlain with a
moist paper towel to reduce desiccation, and re-
placed every other day. Among the 4 buckets pre-
pared, 1 received the standard sugar-agar gel,
and the remaining 3 buckets each received the
same nitrogen-enhanced diet at 1, 5, or 10% con-
centration, respectively.

Mating Trials

Mating trials were conducted between Jul-Oct,
2006, at the USDA-ARS laboratory in Honolulu.
Groups of 75 wild females (10-14 d old), 75 wild
males (8-13 d old), and 75 mass-reared males (5-6
d old) were released between 0800-0830 h in field
cages (2.5 m in height, 3.0 m in diameter) that
contained a single artificial tree (2 m tall with
=500 leaves resembling those of Ficus benjamin
L.; Silkwood Wholesale, Honolulu, HI). Artificial
trees were used, because they provide a chemi-
cally neutral substrate on which the flies display
the entire complement of natural activities. The
cages were monitored for 4 h, mating pairs were
collected in vials, and the males identified under a
UV (black) light (tsl males-pink dye; wild
males-no dye). All unmated individuals were re-
moved from the cages after the trials; trees were
not cleaned between trials. At least 20% of the fe-
males mated in all trials, consequently all data
were included in the analysis per standard qual-
ity control guidelines (FAO/IAEA/USDA 2003).
On a given day, tests were run in 4 cages. One
cage contained tsl males maintained on the stan-
dard sugar-agar diet, and each of the remaining 3
cages contained males maintained on a particular
concentration (i.e., 1, 5, or 10%) of the same nitro-
gen source. Diet treatments were randomly as-
signed to specific tents on each test day. Weekly,
we conducted mating trials on 2-4 d and rotated
(on a random basis) the nitrogen source tested.
Ten replicates were conducted for each nitrogen
source-concentration combination. Over the
study period, air temperature ranged between 26-
32C during the trials.


The potential effect of diet on male dispersal
was examined through a mark-recapture study
conducted between Jul-Oct 2007 in a coffee field
near Haleiwa, Oahu. The mass-reared pupae from
a daily shipment were divided into 2 groups, which
were dusted with dyes of different colors. Follow-
ing irradiation, the differently colored pupae were
placed in separate storage boxes of the same type
used in CDFA's Medfly Preventative Release Pro-
gram (so-called PARC boxes, 60 x 48 x 33 cm,
l:w:h). Within each box, 100 mL of pupae were
placed in each of 6 paper bags for an approximate
total of 36,000 pupae ( 60 pupae per mL). On the
day of peak adult emergence, we placed the sugar-

agar diet on the screen-covered top of one box and
yeast hydrolysate-supplemented diet on the other
(only the 10% concentration was tested). Food
slabs (20 x 15 x 3 cm, l:w:h) were overlain with
moist paper and replaced as previously described.
Four d after peak emergence, males from both
boxes were released between 0900-1000 h at a des-
ignated point in the coffee field. At the same time
2 d later, we placed trimedlure-baited Jackson
traps at the release point, at points 25 m from the
release point along the 4 cardinal directions (N, S,
E, and W), and at points 50 m from the release
point along the 4 cardinal directions and lines off-
set 450 from the cardinal headings (i.e., NE, SE,
SW, NW). Thus, 12 traps were arranged in 2 con-
centric circles (4 and 8 traps in the inner and outer
circles, respectively) about a central trap (the re-
lease point). In baiting the traps, we applied 2 mL
oftrimedlure (a male attractant) to a cotton wick,
which was suspended in a perforated plastic bas-
ket above a sticky insert placed inside the delta-
shaped, trap body. Traps were suspended 1.5-2.0
m above ground in shaded sites within the canopy
of coffee bushes and were left in place for 24 h. The
identity of trapped males was scored under a UV
light. Seven replicates were run with the same re-
lease point and trap sites. Successive releases
were separated by 7 d, and the dye colors used to
mark males were alternated weekly.

Statistical Analysis

Multiple comparisons regarding the numbers
of matings obtained by wild and sterile males
were made among (i) different treatments for a
given nitrogen source or (ii) different nitrogen
sources with 1-way ANOVA. Data from trap cap-
tures were analyzed by a 2-way ANOVA with
male diet and distance from release point as the
main factors. Because circumference trap 'den-
sity' was equal for the 25 (4 traps/157 m) and 50 m
(8 traps/314 m) radius circles, data were com-
bined over all traps for each distance. Data for
both mating and trapping trials met parametric
assumptions. Pair wise comparisons were made
with a paired t test (proportional data were arc
sine transformed) or the nonparametric Mann-
Whitney test (test statistic T).


Mating Trials

The results of the mating trials were remark-
ably consistent over all of the adult diets tested
(Fig. 1). Over all treatments, the average of num-
ber of matings obtained per replicate varied only
between 30.5-37.5 for wild males and 6.5-9.0 for
mass-reared males. As this finding implies, the
numbers of matings obtained by wild and tsl
males, respectively, varied independently among

September 2008

Shelly & Edu: Diet and Mating Success of Sterile Male Medflies

Yeast hydrolysate









Whey protein

SA SA/1% SA/5% SA/10%



2 '

SA SA/1% SA/5% SA/10%


f i

SA SA/1% SA/5% SA/10%

Fig. 1. Mating success of wild (o) and mass-reared, sterile, tsl (o) males of Ceratitis capitata in field cage trials.
Points represent the average (+1 SE) number of matings per replicate (n = 10). Wild males were always fed sugar
and yeast hydrolysate. Sterile males were fed standard sugar-agar (SA) or the standard sugar-agar to which 1%
(SA/1%), 5% (SA/5%), or 10% (SA/10%) of a given nitrogenous material was substituted for sugar.

(i) the 4 different treatments involving the same
nitrogen source (F tests, where df = 3, 36 and P >
0.05 in all 8 cases based on 2 male types x 4 nitro-
gen sources) and (ii) the 4 different nitrogen
sources (trials involving standard sugar-agar diet
excluded; data combined across the 3 concentra-
tions tested for each nitrogen source; wild males:
F3, 116 = 0.05, P = 0.98; sterile males: F3, 116 = 1.05, P
= 0.37). In other words, neither the type nor rela-
tive amount (from 0-10%) of nitrogenous material
mixed with the standard sugar-agar diet ap-
peared to influence the mating success of tsl
males. Based on data from all trials, wild males
obtained a significantly greater number of mat-
ings per replicate than tsl males (33.2 versus 7.7,
respectively, T = 12929.5, n, = n, = 160, P < 0.001)
and, on average, accounted for 81% of the total
matings observed per replicate.
Neither male diet (F1, 36 = 0.14, P = 0.71) nor
trap distance from release point (F2 36 = 1.57, P =
0.22) had a significant effect on the number of

males captured in the trimedlure-baited traps
(Fig. 2). The interaction between these main fac-
tors was not significant (F2 36 = 0.01, P = 0.99). The
total number of males captured per replicate was
1155.1 (t 232.6) for sugar-agar fed males com-
pared to 1069.4 (t 242.2) for males fed the sugar-
agar gel supplemented with yeast hydrolysate
(paired t = 0.81, df = 6, P = 0.45). Similar propor-
tions of males fed sugar-agar gel or sugar-agar gel
plus yeast hydrolysate were captured at the re-
lease point (26% versus 28% per replicate, respec-
tively, paired t = 0.79, df = 6, P = 0.46) or 50 m
(33% versus 30%, respectively, paired t = 0.94, df
= 6, P = 0.38) from the release point.


Supplementing the adult sugar-agar diet used
in SIT programs with different nitrogenous mate-
rials had no obvious effect on the mating success
of mass-reared, sterile tsl males of the Mediterra-
nean fruit fly. Regardless of the identity or
amount of the nitrogen source used, tsl males dis-

SA SA/1% SA/5% SN10%



2 30



Florida Entomologist 91(3)

600 Sugar-ag gel

0 25 50
Distance from release point Fig. 2. Captures in trimedlure-baited traps of mass-
reared, sterile tsl males of Ceratitis capitata fed sugar-
agar gel or sugar-agar plus yeast hydrolysate (10%)
prior to release in a Hawaiian coffee field. Bar heights
represent mean (-1 SE) number of males captured over
a 24-h period 2 d after release at 0 (1 trap), 25 (4 traps),
or 50 (8 traps) m from a central release point (n = 7 rep-

played a consistent level of mating success,
achieving an average of 19% of the total matings
observed per replicate. In addition, a release-re-
capture study revealed no difference in trap cap-
tures-either number or spatial distribution-be-
tween tsl males fed sugar-agar gel or sugar-agar
gel supplemented with yeast hydrolysate.
As noted above, the addition of nitrogen-bear-
ing substances (notably yeast hydrolysate) to a
sugar-only or sugar-agar adult diet has invariably
been shown to increase the mating success of wild
males, whereas the results are mixed with mass-
reared sterile males (Yuval et al. 2007). The rea-
son(s) for this are unclear, but Yuval et al. (2007)
note several possible explanations, including (i)
genetic modifications resulting from bottlenecks
and artificial selection associated with strain col-
onization and mass-rearing, (ii) the availability of
carcasses or feces to sterile males, which may pro-
vide protein (even to males restricted to a sugar-
only diet) and consequently obscure potential ef-
fects associated with different experimental diets,
and (iii) differences in the bacterial communities
in the guts of wild versus mass-reared males,
which affect their ability to process and utilize
different experimental diets.
Regarding the latter possibility, the addition of
symbiotic gut microbes (in particular, the bacterial
species Enterobacter agglomerans and Klebsiella
pneumoniae) to the adult diet has been shown to
improve the intestinal health, and presumably the
overall physiology, of sterile male medflies (C. Lau-
zon, unpublished data cited in Niyazi et al. 2004).
However, to our knowledge, only one study (Niyazi
et al. 2004) has investigated the impact of such
bacterial supplements on male mating success,
and this examined only one dose combination of

microbial and nitrogenous (yeast hydrolysate)
supplements to the standard sugar-agar diet. The
effect of this particular combination on male mat-
ing performance was promising but not clear-cut
(a significant, positive effect was observed in the
laboratory but not in the field), and additional dose
combinations should be evaluated.
In conclusion, it appears that any decision to
use nitrogen-containing dietary supplements in
pre-release diets hinges largely on the demon-
stration of enhanced male mating success, as
there appears to be little, if any, effect of such sup-
plements on the longevity of sterile male medflies,
another parameter central to the success of SIT
programs. For example, Barry et al. (2007) found
no difference in the longevity of sterile males pro-
vided a pre-release diet (first 2 d of adult life) con-
taining sugar and hydrolyzed yeast versus a pre-
release diet of sugar only (this finding was inde-
pendent of the composition of the post-release diet
offered). Likewise, Shelly & McInnis (2003) main-
tained sterile males on a sugar plus yeast hy-
drolysate diet or a sugar-only diet, released males
in field cages (over rooted trees with or without
fruits), and found no diet-related difference in
survival over 4-d intervals. Finally, working with
large cohorts of flies in a mass-rearing facility,
Muller et al. (1997) maintained sterile males on
the same diet over their entire lifetime and found
no difference in longevity between individuals
given a sugar-only diet and those given a sugar
plus hydrolyzed yeast diet. There is evidence that
inclusion of protein in the post-release diet (>2-3 d
of age) enhances male survival (Maor et al. 2004),
but this benefit is apparently conferred to all re-
leased males independent of their pre-release diet
(Barry et al. 2007).


We thank Elaine Pahio and Jon Nishimoto for assis-
tance and Don McInnis for allowing us to work at the
USDA-ARS facility in Honolulu. We are grateful to Boaz
Yuval for reviewing an earlier draft.

BARNES, B. N., D. K. EYLES, AND G. FRANZ. 2004. South
Africa's fruit fly SIT programme-the Hex River Val-
ley pilot project and beyond, pp. 131-141 In B. N.
Barnes [ed.], Proceedings of the 6th International
Symposium on Fruit Flies of Economic Importance.
Isteg Scientific Publications, Irene, South Africa.
MORSE. 2007. Effect of adult diet on longevity of ster-
ile Mediterranean fruit flies (Diptera: Tephritidae).
Florida Entomol. 90: 650-655.
R. COSTA. 2004. The SIT control programme against
medfly on Madeira Island, pp. 127-130 In B. N. Bar-
nes [ed.], Proceedings of the 6th International Sympo-
sium on Fruit Flies of Economic Importance. Isteg
Scientific Publications, Irene, South Africa.

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Shelly & Edu: Diet and Mating Success of Sterile Male Medflies

FAO/IAEA/USDA. 2003. Manual for Product Quality
Control and Shipping Procedures for Sterile Mass-
reared Tephritid Fruit Flies. Version 5.0 IAEA, Vi-
enna, Austria.
proved stability of sex-separation strains for the
Mediterranean fruit fly, Ceratitis capitata. Genome
37: 72-82.
2003. Free amino acid composition and botanical or-
igin of honey Food Chem. 83: 263-268.
KASPI, R., AND B. YUVAL. 2000. Post-teneral protein
feeding improves sexual competitiveness but re-
duces longevity of mass reared sterile male Mediter-
ranean fruit flies. Ann. Entomol. Soc. America 25:
KASPI, R., P. W. TAYLOR, AND B. YUVAL. 2000. Diet and
size influence sexual advertisement and copulatory
success of males in Mediterranean fruit fly leks.
Ecol. Entomol. 25: 279-284.
2004. Effects of post-teneral diet on foraging success
of sterile male Mediterranean fruit flies. Entomol.
Exp. Appl. 110: 225-230.
MORTON, T. C., AND M. A. BATEMAN. 1981. Chemical
studies on proteinaceous attractants for fruit flies,
including the identification of volatile constituents.
Australian J. Agric. Res. 32: 905-916.
J. R. CAREY. 1997. Early mortality surge in protein-
deprived females causes reversal of sex differential
of life expectancy in Mediterranean fruit flies. Proc.
Nat'l. Acad. Sci. USA 94: 2762-2765.
NIYAZI, N., C. R. LAUZON, AND T. E. SHELLY. 2004. Effect
of probiotic adult diets on fitness components of ster-
ile male Mediterranean fruit flies (Diptera: Tephriti-
dae) under laboratory and field cage conditions. J.
Econ. Entomol. 97: 1570-1580.
SHELLY, T. E., AND S. S. KENNELLY. 2002 Influence of
male diet on male mating success and longevity and

female remating in the Mediterranean fruit fly
(Diptera: Tephritidae) under laboratory conditions.
Florida Entomol. 85: 572-579.
SHELLY, T. E., AND D. 0. MCINNIS. 2003. Influence of
adult diet on the mating success and survival of
male Mediterranean fruit flies (Diptera: Tephriti-
dae) from two mass-rearing strains on field-caged
host trees. Florida Entomol. 86: 340-344.
2002. Effect of adult diet on signaling activity, mate
attraction, and mating success in male Mediterra-
nean fruit flies (Diptera: Tephritidae). Florida Ento-
mol. 85: 150-155.
fects of diet, ginger root oil, and elevation on the
mating competitiveness of male Mediterranean fruit
flies (Diptera: Tephritidae) from a mass-reared, ge-
netic sexing strain in Guatemala. J. Econ. Entomol.
96: 1132-1141.
SHELLY, T. E., J. EDU, AND E. PAHIO. 2006. Dietary pro-
tein and mating competitiveness of sterile males of
the Mediterranean fruit fly (Diptera: Tephritidae):
measurements of induced egg sterility in large field
enclosures. Florida Entomol. 89: 277-278.
TSIROPOULOS, G. J. 1978. Holidic diets and nutritional
requirements for survival and reproduction of the
adult walnut husk fly J. Insect Physiol. 24: 239-242.
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124 pp. USDA Tech. Bull. #1261.
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Florida Entomologist 91(3)

September 2008


1Current Address: Department of Molecular Biology and Genetics, Cornell University,
421 Biotechnology Building, Ithaca, NY 14853

2Department of Zoology, 223 Bartram Hall, University of Florida, Gainesville, FL 32611

'USDA-ARS, U.S. Horticultural Research Laboratory, 2001 South Rock Road, Ft. Pierce, FL 34945


The tropical root weevil, Diaprepes abbreviatus (L.), is a major pest of many plants of eco-
nomic importance, including citrus and ornamentals. It was accidentally introduced from the
Caribbean into Florida in the 1960s and its range within the United States now includes
Texas and California. No safe and effective control method is available for this species. Stud-
ies of the reproductive behavior of field populations may aid the development of effective con-
trol strategies and inform laboratory experimental design. We examined mating patterns of
individually-marked D. abbreviatus in a plant nursery in Florida over 5 d. Matings occurred
throughout the day but tended to peak during the middle of the day. Mating duration ranged
from less than 3 h to over 9 h. Most males and females mated repeatedly, sometimes with the
same partner multiple times. Both same-sex and extra-pair mountings occurred at low fre-
quencies. In a follow-up study conducted in captivity, we found no short-term direct benefits
of multiple mating to females but female fertility decreased dramatically by 2 weeks after
mating if females were not allowed to re-mate. Thus, female D. abbreviatus may re-mate, in
part, to replenish sperm stores. Our results suggest that the sterile insect technique would
not be effective but support previous conclusions that pheromones may be effective attract-
ants in this species because both females and males mate multiple times in the field.

Key Words: reproduction, beetle, mating frequency, fecundity, fertility


El picudo Diaprepes abbreviatus (L.) es una plaga de plants de importancia econ6mica tales
como los citricos y plants ornamentales. Fue introducido por primera vez del Caribe a la
Florida en los anos sesenta y su rango dentro de los Estados Unidos se ha extendido hasta
Texas y California. Por el moment no se ha desarrollado un m6todo de control seguro y efec-
tivo en contra de la especie. Estudiamos el comportamiento reproductive de poblaciones
silvestres para promover el desarrollo de estrategias de control e informar disenos experi-
mentales de bioensayos. Nosotros examinamos patrons de apareamiento de individuos
marcados de D. abbreviatus en un invernadero commercial en el sur de Florida durante 5 dias.
Observamos apareamiento durante todo el dia con una tendencia a aumentar al mediodia.
La duraci6n del apareamiento vario desde <3 hasta >9 horas. La mayoria de los machos y
hembras se aparearon repetidamente, a veces con el mismo individuo. Acoplamientos entire
individuos del mismo sexo o apareamientos entire >2 individuos ocurrieron a frecuencias ba-
jas. En un studio de seguimiento conducido en cautiverio, no encontramos beneficio para las
hembras a corto plazo de los apareamientos multiples. La fertilidad disminuy6 dramatica-
mente dos semanas despu6s del apareamiento cuando se neg6 la posibilidad de re-apa-
rearse. Por lo tanto, hembras de D. abbreviatus podrian re-aparearse, en parte, para renovar
su reserve de esperma. Nuestros resultados sugieren que la t6cnica de insects est6riles no
seria efectiva en el caso de D. abbreviatus, pero apoyan conclusions previas de que las fero-
monas pueden usarse como atrayentes en esta especie ya que tanto hembras como machos
se aparean multiples veces en el campo.

Translation provided by the authors.

Diaprepes abbreviatus (L.) is a tropical root the larval feeding damage it causes to roots of
weevil thought to be native to Puerto Rico and the plants of agricultural (e.g., citrus) and horticul-
Lesser Antilles and is currently found over much of tural (e.g. ornamentals) importance (Simpson et al.
the Caribbean, and in Florida, California, and 1996). Larval damage to roots results in reduced
Texas (Lapointe et al. 2007). In the Caribbean and plant growth and exposure to pathogens (Graham
Florida, the weevil is a major economic pest due to et al. 2003). Adults emerge from the subterranean

Sirot & Lapointe: Mating Behavior ofDiaprepes abbreviatus

pupal chamber and climb onto host plants where
they feed, mate, and lay eggs. Field-caught D. ab-
breviatus females can lay over 11,000 eggs during
their lifetime (Nigg et al. 2004; Wolcott 1936). Al-
though a variety of control methods have been or
are currently being tested (Ulmer et al. 2006; Dun-
can et al. 2007), none is entirely satisfactory, and
this species is still spreading in the United States.
Understanding the reproductive biology of this
species in the field may contribute to development
of effective control method including efforts to un-
derstand the role of semiochemicals in host and
mate location (F. Otalora-Luna, pers. comm.). De-
spite several decades of research on D. abbrevia-
tus, few studies have investigated its mating be-
havior in field populations. Previous studies
showed that the weevils primarily mate diurnally
(Schroeder 1981), and are characterized by size-
assortative mating (Harari et al. 1999), with a
peak in the percentage of individuals mating in
the middle of the day (Schroeder 1981). The wee-
vils are attracted to trees that have previously
been exposed to weevils of the opposite sex
(Schroeder 1981). In captivity, females mate with
multiple males (Harari et al. 1999; Sirot et al.
2007), males compete directly for females by at-
tempting to disrupt mating pairs and displace
mating males (Harari et al., 1999), and the last
male to mate before oviposition fertilizes the ma-
jority (~70%) of eggs (Harari et al., 2003; Sirot et
al. 2007). However, several characteristics of the
mating behavior (e.g., mating frequency, number
of mates, etc.) of D. abbreviatus in the field are
currently unknown. Gaining a more complete un-
derstanding of the mating behavior of D. abbre-
viatus in the field will provide information with
which to interpret the results of mating studies
conducted in captivity and to plan more effective
reproductive control methods (Boake et al. 1996).
The goal of this study is to describe the pat-
terns and consequences of D. abbreviatus mating
behavior in the field. This is the first study of the
mating behavior of individually-marked D. abbre-
viatus in the field. We report results on D. abbre-
viatus mating patterns in a commercial plant
nursery and follow-up experiments conducted in
captivity to clarify the consequences of the pat-
terns we observed in the field.


Mating Patterns in the Field

This study was conducted from 7 to 12 May 2002
in a commercial ornamental plant nursery in Home-
stead, FL (Miami-Dade County). During the study,
daytime air temperature ranged between 26C and
32C, there was no rainfall, and cloud cover ranged
between 0% and 50%. Adult D. abbreviatus were
collected from 3 neighboring dahoon holly trees (Ilex
cassine) in the nursery in an area with a high D. ab-

breviatus concentration. On 7 May and the morn-
ings of 8 to 11 May, we collected, marked, and mea-
sured 261 individuals of D. abbreviatus (115 fe-
males, 146 males). We collected the weevils by gen-
tly removing them from leaves or branches by hand
and recorded their mating status (mating or not at
the time of collection). We also measured the length
of their right elytra with a dial caliper to the nearest
0.1 mm and marked them uniquely with a 3-dot
color code (with uniPaint Fine Line paint pens) on
their pronotum and anterior elytra before returning
them to the tree from which they came. From 8 to 12
May, we conducted censuses 4 times per day at 3-h
intervals. The 4 censuses began at approximately
0800,1100,1400, and 1700 EST and lasted 73.7 min
per census on average (SEM: 4.8 min; n = 20 cen-
suses). During the censuses, we searched the leaves,
stems, and trunk of all trees within a 5-m x 5-m area
centered on the 3 collecting trees. We recorded the
activity (standing still, feeding, walking, mating,
mounting another, or being mounted) and location
of every marked individual re-sighted during these
It was difficult to confirm actual matings with-
out disturbing the beetles because D. abbreviatus
engage in intrasexual mountings that appear sim-
ilar to mating pairs (Harari et al. 2000). For appar-
ent matings in which both individuals were
marked, we could verify their sex and whether it
was an actual mating by referring to our database
of marked individuals. For apparent matings in
which the mounting individual was marked but
the mounted individual was not marked, we cate-
gorized the mounting as mating if the mounting
individual was male (as verified by our marking
record). For apparent matings in which the mount-
ing individual was not marked but the mounted
individual was marked, we categorized the mount-
ing as mating if the mounted individual was fe-
male. This definition may have resulted in an over-
estimate of the actual number of matings, but the
overestimate should be minimal because mount-
ings by females or of males were rare during our
study (only 3% and 8% of observed mountings that
included 2 marked individuals, respectively). With
captive weevils, we assessed both short-term and
long-term effects of mating on female fitness, as
described in the following sections.

Short-term Effects of Multiple Mating on Female
Fecundity and Fertility

We measured the short-term effects of multiple
mating by comparing the fecundity and fertility of
females mated singly (n = 20) and those mated
multiply (i.e., 3 to 4 times, with the same males; n
= 20). Adults were collected from the field (Univer-
sity of Florida citrus grove at Ft. Pierce, FL) and
maintained in same-sex cages in a greenhouse
(natural light cycle; ~27C) for 2 to 7 d before the
start of the experiment. On the first day of the ex-

Florida Entomologist 91(3)

periment, we placed individually-marked males
and females in pairs (1 male and 1 female/cage) in
clear plastic containers (diameter: 10 cm; height:
8 cm; with 1 cm2 of young citrus leaf, Citrus mac-
rophylla Wester). We checked each pair for mating
every 30 min. After the pair completed mating,
the males were returned to the all-male cage. The
females were left in their containers with a piece
of wax paper added as an oviposition substrate
(Wolcott 1933). On 3 subsequent mornings, we re-
placed the oviposition substrate in each cage and
returned the male partners of females in the mul-
tiple-mating treatment to the females' containers.
We checked the pairs for mating every 30 min and
removed males from cages by 19:00 (EST), by
which time all pairs had ceased mating. On the
4th night of the experiment, we removed the
males from the mating containers, placed oviposi-
tion substrate and fresh citrus leaves in all female
containers, and allowed the females to lay eggs for
3 more nights. The wax paper strips and eggs
were placed in plastic vials coated with a mist of
distilled deionized water in an incubator at 26C
(Lapointe 2001). We counted the number of eggs
laid (fecundity) and the percentage of eggs that
were fertilized (fertility) for each female.
Our previous research suggested that females
receive direct benefits from mating (Sirot et al.
2006). Therefore, multiply-mated females were
paired with the same males repeatedly rather
than with different males to limit the effects of ex-
perimental treatments to those related to mating
itself rather than including potential genetic ben-
efits of multiple mating. We used field-caught
rather than lab-reared beetles for this experiment
because they likely differ in a number of traits
that could affect the costs or benefits of mating to
females and we were specifically interested in the
effects of multiple mating on females in the field.

Statistical Analysis

We used a generalized linear model (GLM)
with a Poisson error distribution (Agresti 2002) to
analyze the relationship between female fecun-
dity and mating treatment. We used a GLM with
a binomial error distribution to analyze the rela-
tionship between the percentage of eggs fertilized
and the mating treatment. To correct for overdis-
persion, we used the Williams (1982) method. Fe-
male body size and the night that eggs were laid
also were included in both analyses. GLMs were
performed with the R program (Version 2.2.1; Ve-
nables & Ripley 2002).

Long-term Effect of Time Since Mating on Female Fertility

We assessed the fertilization success of females
(n = 10) for thee 2-d periods to determine the long-
term ability of females to fertilize eggs after a sin-
gle mating: immediately after mating, 2 weeks

after mating, and 4 weeks after mating. We used
lab-reared unmated females that had been main-
tained in an all-female cage for 1 month after eclo-
sion and field-caught males (University of Florida
citrus grove at Ft. Pierce, FL) that had been main-
tained in an all-male cage for 1 week after collec-
tion. We used field-caught rather than lab-reared
males since these 2 groups of males might differ
in the quantity or quality of sperm produced. Fe-
males were placed individually in screened cages
with a male and with a moistened dental wick
and a piece of fresh citrus leaf. At 3-4 h after the
start of mating, the mating was interrupted by
gently separating the male and female and the
female was transferred to a 1-L cage with a wax
paper oviposition substrate, a moistened cotton
wick, and young citrus leaves (Citrus macro-
phylla Wester). The females were maintained in
their individual containers for 2 d and then were
placed together in an all-female cage. The oviposi-
tion strips and leaves containing eggs from each
female were placed in a plastic vial coated with a
mist of distilled deionized water in an incubator
at 26C. We counted the number of eggs laid (fe-
cundity) and the percentage of eggs that were fer-
tilized (fertility) for each female. At 2 and 4 weeks
after first mating, we again placed each female in-
dividually in a 1-L cage for 2 d and collected, incu-
bated, and counted the eggs as described above.


Mating Patterns in the Field

The mean SEM length of the right elytron of
D. abbreviatus collected during our field study
was 8.8 0.1 mm (n = 146, range 6.2-11.9 mm)
and 10.1 0.1 mm (n = 115, range 7.1-13.7 mm)
for males and females, respectively. At the time of
capture, 42% of females and 33% of males were
mating and there was positive size-assortative
mating (R2 = 0.15, F1,46 = 8.03, P < 0.01). Fifty-two
percent of males and females were re-sighted at
least once during the course of our censuses.
The percentage of marked individuals mating
during each census ranged from 39% to 54% for
males and 40% to 67% for females. For both males
and females, the percentage of individuals mating
peaked in the middle of the day (Fig. 1; males: X21
= 2.84; P = 0.09; females: X21 = 3.31; P = 0.07). We
observed 28 matings in which both partners were
marked and which were not disrupted by our cen-
suses. Sixty-eight percent (19 of 28) of the mat-
ings began in the morning and 84% (16 of 19) of
these matings ended before the early afternoon
(1400 h census). We could estimate mating dura-
tion for 17 matings in which both partners were
marked. The majority (14 of 17) of these matings
lasted <3 h, 1 lasted 3-6 h, and 2 lasted >9 h.
We observed high rates of re-mating by both
males and females during our study. We investi-

September 2008

Sirot & Lapointe: Mating Behavior ofDiaprepes abbreviatus

1.0 1.90 2
1.80 .90 2.00



*o U

- U
a o^. ^"



2.10 2.20 2.30 2A. 2.50-

UOI 11U0 14UU1 1iiN
S.la Timr lu Cn.l.
Fig. 1. Percentage (Mean SEM) of re-sighted D. ab-
breviatus mating throughout the day in a commercial
plant nursery at Homestead, FL (Dade County). Data
are based on censuses conducted 4 times per day on 5
consecutive days in May 2002. The percentage of wee-
vils mating tended to peak in the middle of the day for
both males and females.

gated re-mating patterns for the 21 marked fe-
males and 24 marked males that we re-sighted on
at least 3 different days. Seventy-five percent of
the females mated with multiple partners. On av-
erage, females mated slightly less than once per d
(0.98 0.12 matings/d; median = 1; range: 0-2.2).
Two of 21 females never mated whereas 1 female
participated in 11 different matings over the
course of the 5-d study. Female mating frequency
was not correlated with female size (F1,19 = 0.08; P
= 0.78; R2 = 0.004; Fig. 2).
Seventy-six percent of males mated with mul-
tiple partners. On average, males mated once ev-
ery 1.14 0.14 d (median = 1; range: 0-2.25 mat-
ings/d). Four of 24 males never mated, whereas 2
males each participated in 8 matings over the
course of the study. Male mating frequency was
positively correlated with male size (F1,22 = 6.7;P =
0.02; R2 = 0.23; Fig. 2).
For matings that were not disrupted by our cen-
suses, 50% of males and 44% of females whose mat-
ings ended before the 1400 h census mated again
later the same day. None of the 5 females whose
matings ended between the 1400 h and 1700 h cen-
suses were mating with new partners during the
1700 h census, but 3 of them were observed mating
the following day. Two of 5 males whose matings
ended between the 1400 h and 1700 h censuses
were mating with new partners during the 1700 h
census and a third male was observed mating the
following day. Of the 7 pairs still mating during the
last census of the day, 2 females and 5 males were
observed mating again the following day.
Of 33 mating pairs in which both partners
were marked, 4 (12%) of male-female pairs mated
twice during the course of our study. Two of these

Ln Right elylron length

Fig. 2. Relationship between body size and mating
frequency of D. abbreviatus in a commercial plant nurs-
ery at Homestead, FL (Dade County). Mating frequency
is the total number of matings by an individual divided
by the number of days on which it was observed. Only
individuals that were re-sighted on at least 3 d were in-
cluded in this analysis (n = 21 females; n = 24 males).
Dashed line indicates least-squares regression of the
natural log of male mating frequency and the natural
log of male elytron length. Female mating frequency
was not correlated with female size.

pairs re-mated on the subsequent d. One pair re-
mated 2 d later and 1 pair re-mated 3 d later. For
3 of the 4 pairs, the marked female mated with a
different male in between the 2 matings with the
marked male.
Of 126 observed mountings by marked individ-
uals, 4 were mountings by females and 10 were
mountings by males of mating pairs. Of 131
mountings of marked individuals, 11 were mount-
ings of males. Intra-sexual mountings did not
tend to occur at a particular time of d; such
mountings were observed at least once during
each of the census times. Mountings of mating
pairs by marked males tended to occur in the mid-
dle of the day (90% occurred during the 1100 or
1400 censuses), which coincided with the peak of
mating activity (Fig. 1). We made ad libitum ob-
servations of 26 mountings of mated pairs during
2 d of our censuses (10 and 11 May); more of these
occurred in the middle of the day than in the early
morning and late afternoon (X21 = 7.54; P = 0.006).

Short-term Effects of Multiple Mating on Female
Fecundity and Fertility

The fecundity and fertility of females allowed
to mate once or multiple times with the same
male were measured over 6 d. Females in singly-
mated and multiply-mated treatments laid a sim-
ilar number of eggs and had similar fertility over
all nights of the experiment. Singly-mated fe-
males laid a total of 146 34 eggs (range: 0 to 527;
n = 20) over nights 2-6 of the study, whereas mul-

Florida Entomologist 91(3)

tiply-mated females laid 159 37 (range: 0 to 398;
n = 20; GLM; tl09 = 0.6, P > 0.1; Table 1). The per-
centage of eggs that were fertilized on nights 2-6
was 86 7% for singly-mated females (range: 0 to
100%; n = 15) and 82 9% for multiply-mated fe-
males (range: 0 to 100%; n = 14; t54 = 0.7, P > 0.1;
Table 1).

Long-term Effect of Time Since Mating on Female

To determine whether female fertility is sperm-
limited, we measured the percentage of eggs fer-
tilized in the first 2 d (n = 10 females), at 14-15 d
(n = 7), and at 28-29 d (n = 7) after a single 3-4 h
mating. The percentage of eggs fertilized was high
in the first 2 d after mating (90 3%), but dropped
dramatically by 2 weeks after mating (42 15%
Mann-Whitney U test; U =16; P = 0.04, one-tailed
test). Only 3 out of 863 eggs laid at 28-29 d after
mating were fertilized.


Mating Patterns in the Field

Understanding mating behavior of pest species
under natural conditions provides essential infor-
mation for implementing effective control strate-
gies and for designing and interpreting experi-
ments conducted in captivity. Our study estab-
lished several characteristics of D. abbreviatus
mating behavior in a field population. (1) Mating
duration ranged from <3 to >9 h. (2) Mating activ-
ity occurred throughout the day but the percent-
age of individuals mating tended to peak during
the middle of the day, as did mounting of mated
pairs by males. (3) Both male and female D. ab-
breviatus mated multiple times and with multiple
partners. (4) Male, but not female, mating fre-
quency was positively correlated with body size.
(5) Some individuals mated multiple times on dif-
ferent days with the same partner. (6) Mounting

by females and mounting of males occurred at low
frequencies. Together, these results provide a
clearer understanding of the reproductive biology
of this important pest species.
The percentage of individuals mating tended
to peak during the middle of the day in our study.
This pattern was also found in a previous study of
D. abbreviatus in central Florida (Schroeder
1981), and was suggested to result from an accu-
mulation of mating pairs over the course of the
day. Our findings support this hypothesis: mat-
ings began throughout the day, but those that be-
gan in the morning tended to end by the late af-
ternoon resulting in maximum overlap of matings
in the middle of the day. Studies of mating fre-
quency in the field should therefore conduct cen-
suses at consistent times to avoid variation due to
diel fluctuations, and should conduct censuses
during midday if the goal is to maximize the num-
ber of observed matings.
Multiple mating was common for both male
and female D. abbreviatus at our field site. Over
70% of males and females that we observed par-
ticipated in multiple matings during the 5-d
study. Furthermore, both males and females were
observed re-mating within the same day. Re-mat-
ing during the same day was also observed in a
study of D. abbreviatus in captivity (Harari et al.
2003). In that study, mating pairs were inter-
rupted after 2, 4, or 8 h and placed separately in
cages with an excess of individuals of the opposite
sex. Almost 100% of males and females re-mated
later the same day. In contrast, in our study of
matings by marked individuals in the field, only
48% of males and 35% of females whose matings
ended before the last census were observed to re-
mate later the same day. Our data suggest that al-
though re-mating is common in D. abbreviatus in
the field, it is unlikely to occur at frequencies as
high as those reported from captivity in the
Harari et al. (2003) study.
Our study site was characterized by dense ag-
gregations of D. abbreviatus. Mating frequency


Measure Treatment Night 1 Night 2 Night 3 Nights 4-6 GLM Results'

Number Laid Singly-mated 56 12 16 5 13 5 118 29 t109 = 0.6, P > 0.1
(n = 20)
Multiply-mated 66 15 24 7 8 4 112 29
(n = 20)
Percentage Singly-mated 99 1% 83 10% 99 1% 94 5% t54 = 0.7, P > 0.1
Fertilized (n = 15) (11) (7) (12)
Multiply-mated 91 7% 94 5% 99 1% 78 11%
(14) (10) (5) (12)

Comparisons of singly-mated and multiply-mated females on nights 2-6.

September 2008

Sirot & Lapointe: Mating Behavior ofDiaprepes abbreviatus

may be positively correlated with density. Similar
studies of mating frequencies at other field sites
are necessary to determine how mating frequency
may be affected by density. It is also important to
note that we only conducted censuses once every
3 h and may have missed matings. Therefore, our
mating frequency results should be treated as
minimum estimates of the actual mating frequen-
cies at this site.
Our finding that male mating frequency was
positively correlated to male body size suggests
that pre-copulatory sexual selection for large
male size occurs in the field. The mechanisms un-
derlying pre-copulatory selection for large size
are probably competitive ability and mating
speed (Harari et al. 1999). Males often obtain
matings by competing directly with other males
(L.K.S., pers. obs.). Therefore, males who are able
to displace other males and obtain matings
quickly will have an advantage. In a captive study
of citrus grove-collected D. abbreviatus, Harari et
al. (1999) found that large males displaced
smaller males in contests over females and initi-
ated matings more rapidly after mounting.
Harari et al. (1999) attributed the latter pattern
to female mate choice because females must open
their genital aperture in order for males to insert
their genitalia. Therefore, both male-male compe-
tition and female mate choice are likely to con-
tribute to the mating advantage of large males in
the field. If this is the case, then selection may ac-
tually be acting on male size relative to other
males in the population rather than on absolute
male size.

Effects of Multiple Mating on Female Fitness

A pressing question in the field of behavioral
ecology is why females mate with multiple males.
One hypothesis to explain this pattern is that fe-
males may receive direct benefits from multiple
matings such as avoiding sperm depletion or re-
ceiving substances from the male that increase
survival or fecundity of the female or her off-
spring (Arnqvist & Nilsson 2000). Our previous
research onD. abbreviatus provided evidence con-
sistent with this hypothesis: male seminal fluid
molecules were found predominantly in the ova-
ries and eggs of mated females (Sirot et al. 2006).
In the current study, we tested for direct benefits
of multiple matings to females. Our finding that
female fecundity and fertility measured over 4 d
did not differ for females mated once to a male
compared with females mated multiple times to 1
male suggests that female D. abbreviatus do not
benefit directly from multiple matings in the
short-term, in terms of quantity of offspring pro-
duced. However, it is still possible that multiple
matings (and/or mating to multiple males) result
in higher quality offspring or greater female lon-
gevity though male-derived resources.

Our results suggest that female D. abbreviatus
may receive important fertility benefits by re-
peated matings over the course of their lifetimes.
We found that the percentage of fertilized eggs
that females lay decreased dramatically with
time since mating. This pattern was also observed
in a previous study in which females mated with
2 males in rapid succession and were then al-
lowed to lay eggs over 30 d (Harari et al. 2003).
Together, these results suggest that female D. ab-
breviatus become sperm-limited when they do not
re-mate within 2 weeks after their last mating.
Therefore, re-mating by females may function, in
part, to replenish sperm stores. Where D. abbre-
viatus are found at low densities, females may
benefit by mating with any male they encounter.
However, replenishing stored sperm does not
seem to be a sufficient explanation for the high re-
mating frequency we observed at our field site,
which is characterized by a dense aggregation of
D. abbreviatus. It is likely that females gain addi-
tional benefits from mating with multiple males.
Edvardson (2007) presents an intriguing possibil-
ity that female weevils (Callosobruchus macula-
tus) benefit from access to water provided in male
ejaculate during mating. Future research on D.
abbreviatus should consider similar effects of food
and water availability on mating behavior.
In our study, most female D. abbreviatus mated
multiple times within 5 d. These results suggest
that the sterile insect technique (SIT) would not
be effective at controlling populations of D. abbre-
viatus, at least at high population densities. Fe-
males mated to sterile males readily re-mate with
untreated males and the sperm of untreated
males are able to fertilize eggs even after a female
subsequently mates with a sterile male (Sirot et
al. 2007). Our findings that D. abbreviatus mate
repeatedly in the field and that females may need
to re-mate to replenish sperm supplies suggest
that males and females seek individuals of the op-
posite sex multiple times throughout their adult
lives and therefore support other researchers' con-
clusions that sex or aggregation pheromones may
occur in this species (Schroeder 1981; Schroeder &
Beavers 1985; Harari & Landolt 1997). Future
studies should examine whether re-mating be-
havior and attraction to conspecific odors changes
with age or condition. We also found that male
mating frequency was positively correlated with
male size, suggesting that control strategies that
depend on mating success of males should use
large males. This relationship remains to be
tested for lab-reared D. abbreviatus.


H. J. Brockmann, C. St. Mary, M. Wayne (University
of Florida, Gainesville, FL), J. Sivinski (USDA-ARS,
Gainesville, FL), and L. Harrington (Cornell University,
Ithaca, NY) provided important insights that benefited

this work. R. Duncan and J. Pena (University of Florida,
Homestead, FL) assisted in locating field infestations.
A. S. Hill and D. Grayson (USDA-ARS, Ft. Pierce, FL)
provided technical assistance. This project was sup-
ported in part by the Florida Citrus Production Re-
search Advisory Council. We also are grateful to the
University of Florida, Department of Zoology for logisti-
cal and financial support of L.K.S.


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Florida Entomologist 91(3)

Pozo et al.: Phenology of Calakmul Butterflies


1McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History,
University of Florida, SW 34th St. and Hull Rd., Gainesville, FL 32611-2710

'Present address: Depto. de Ecologia Terrestre, El Colegio de la Frontera Sur,
Av. Centenario Km 5.5, A.P. 424, Chetumal, Quintana Roo, C.P. 77014, M6xico

'Museo de Zoologia "Alfonso L. Herrera", Facultad de Ciencias, Universidad Nacional Aut6noma de M6xico,
Apdo. Postal 70-399, C.P. 04510, M6xico, D.F., M6xico


The phenology of butterflies was analyzed in the Calakmul Region (CR) in the state of
Campeche, M6xico, over the course of 3 years. Altogether, 60,662 individuals were recorded,
consisting of 359 species in 207 genera, 18 subfamilies, 5 families, and 2 superfamilies.
Greatest species diversity was recorded during Oct and Nov. Monthly fluctuation in diver-
sity was defined by rare species. Hesperiidae (135 species) and Nymphalidae (111 species)
were the most diverse families, and showed the greatest variation with respect to distribu-
tion of species richness throughout the year. Papilionidae showed the greatest species rich-
ness during the dry season. Pieridae, Nymphalidae, and Lycaenidae showed peaks of
greatest species richness and relative abundance during the rainy season. Results were
compared to faunal studies of the Sierra de Atoyac de Alvarez, in the state of Guerrero, and
of the Sierra de Manantlan, in Jalisco and Colima. Important similarities were observed
among phenological patterns in the butterfly fauna of the 3 regions, especially between CR
and Manantlan. The phenology of species with greater relative abundance was analyzed in
relation to wingspan as a parameter of adult size. The small and medium-sized groups,
taken together, showed variations in species richness. An analysis of species seasonality was
conducted with NMDS, ANOSIM and SIMPER, in the program PRIMER 4.0. Differences
among the composition of butterfly communities with respect to the seasons were found.

Key Words: Calakmul Biosphere Reserve, rare species, Yucatan Peninsula


La fenologia de las mariposas fue analizada en la region de Calakmul (CR), Campeche,
M6xico. A lo largo de tres anos se registraron 60,662 individuos, de 359 species en 207 g6-
neros, 18 subfamilias, cinco families, y dos superfamilias. La mayor diversidad fue regis-
trada durante octubre y noviembre. La fluctuaci6n mensual de la diversidad fue definida por
las species raras. Las families con mayor diversidad fueron Hesperiidae (135 species) y
Nymphalidae (111 species) y presentaron la mayor variaci6n con respect a la distribucion
de la riqueza de species a trav6s del ano. Papilionidae mostr6 la riqueza mas grande de es-
pecies durante la estaci6n seca. Pieridae, Nymphalidae y Lycaenidae presentaron sus maxi-
mos de riqueza de species y abundancia relative durante la estaci6n lluviosa. Se hizo una
comparaci6n entire el present studio y los de fauna de Atoyac de Alvarez, en el estado de
Guerrero, y de Manantlan, en Jalisco y Colima. Se observaron semejanzas entire la fenologia
de las tres regions, especialmente entire CR y Manantlan. La fenologia de las species con
mayor abundancia relative fue analizada en relaci6n a la envergadura alar como un para-
metro del tamano de las species. Los grupos pequenos y medianos, en conjunto, muestran
variaciones en la riqueza de species. Se realize un andlisis de estacionalidad de species con
las matrices de datos, aplicando NMDS, ANOSIM y SIMPER con el program PRIMER 4.0.
Se encontraron diferencias en la composici6n de las comunidades de mariposas con respect
a las estaciones.

Translation provided by the authors.

Phenology is the study of the sequence of de- tures (Williams-Linera & Meave 2002; Shapiro et
termined biological events; this type of observa- al. 2004). With respect to insects, phenology is de-
tion began with the first agrarians of various cul- fined as the process of the appearance of different

Florida Entomologist 91(3)

stages in the life cycle of a taxonomic group over
the course of a year, and across several seasons
(dry and wet or warm and cold periods). Among
butterflies, this phenomenon has generally been
described based on the appearance of the adult
stage or imago of various species. The presence
and activity of each generation has been ex-
plained as a function of diverse climatic or vege-
tation factors, such as temperature, precipitation,
seasonality of vegetation, food availability, and
food quality. Phenological patterns in butterflies
may be determined by fruiting or flowering plants
(Tanaka & Tanaka 1982; Young 1982; Scott 1986;
Wolda 1987, 1988a), annual humidity distribu-
tion (Wolda 1987), cloudiness (Luis & Llorente
1990), photoperiodic changes (Shapiro 1975), sub-
strate availability and palatability of larval food
plants (Owen et al. 1972).
As with most other insects, butterfly life cycles
are strictly linked to seasonal changes (Owen
1971), such as temperature, day length or photo-
period, and humidity, among others. Species rich-
ness is influenced by climatic factors, which deter-
mine reproduction and survival conditions, and
therefore dictate the number of individuals or bio-
mass documented. Voltinism is defined as the re-
lationship between life cycle, annual climatic be-
havior, and the number of generations per year.
Depending on the number of generations per year,
terms such as univoltinism, bivoltinism, trivolt-
inism and multivoltinism are used to characterize
the phenology of a population, species, or taxon
(Gullan & Cranston 2000). Flight in adults is fun-
damental to conducting diverse vital actions such
as feeding, reproduction, and oviposition. Given
that butterflies are heliothermic organisms, their
flight is dependent upon sunlight (Shapiro 1975).
Gilbert & Singer (1975) and Shapiro (1975) sug-
gested that interactions with hostplants and cli-
mate may explain most of the distributional pat-
terns seen among Lepidoptera. Shapiro (1975)
emphasized that this seasonal history tends to be
extremely conservative even at the family level;
an example is the stage at which diapause takes
place (e.g., various Pieridae as pupae and many
Lycaenidae as eggs).
The phenology of an insect fauna is summarized
by the pattern of species emergence throughout
the year, which shows records obtained through
several generations during several years. When
studying phenology, we can distinguish at least 3
principal components of the insect community. The
first component refers to species that are present
year after year, regardless of the particular envi-
ronmental conditions of any given year. The second
component consists of species occurring in a given
season, generally univoltine species. The third
component consists of species whose abundance is
too low to determine their seasonality.
Describing insect phenology is a complex en-
deavor. Species behavior, relative abundance, and

the sampling methods are usually biased towards
certain taxonomic groups. Often, information ob-
tained on relative abundance does not correspond
with reality. For example, greater efforts are gen-
erally employed in collecting rare or very local-
ized species, and less effort is usually made to col-
lect abundant or common, vagile, and widely dis-
tributed species (eurytopics) (Pozo et al. 2005).
The objective of this paper is to describe sea-
sonal and phenological patterns of the butterfly
community (Papilionoidea and Hesperioidea) in
the tropical forest of the Calakmul region, based
on 2 different methodologies, with data collected
during 26 months over the course of 3 years.


Study Area

The Municipality of Calakmul is located in
southeastern Campeche State, Mexico, in the cen-
ter of the Yucatan Peninsula. With an area of
16,806 km2, Mpio. Calakmul borders the State of
Quintana Roo to the east and the Department of
El Peten, Guatemala, to the south. Almost half of
the municipality was placed under protection by
decree of the Calakmul Biosphere Reserve (CR) in
1989 (Fig. 1). The reserve includes an area of
7,232 km2, with an altitude 100 to 300 m above sea
level. The Reserve has a semi-humid, warm cli-
mate. Annual precipitation ranges from 600-1200
mm, and annual average temperature is 24.6C
(Fig. 2) (INEGI 1996). The dry season extends
from Feb to May, followed by the rainy season
from Jun to Sep, which in turn is followed by the
"nortes" (or cold) season, from Oct to Jan. "Nortes"
refer to frequent fronts of cold winds that cross the
Gulf of Mexico and carry humidity from a NE to
SW direction (Williams-Linera & Meave 2002).
In the CR, 7 vegetation types are found: tall
tropical evergreen forest, medium height tropical
semi-evergreen forest, medium height tropical
semi-deciduous forest, savanna, low tropical
semi-evergreen forest, low tropical semi-decidu-
ous forest and, low tropical semi-deciduous sea-
sonally flooded forest. The medium height tropi-
cal semi-evergreen forest and low tropical semi-
deciduous forest dominate the landscape within
the CR (Martinez & Galindo-Leal 2002).


The butterfly fauna of CR was surveyed during
266 d distributed over 26 months, across 3 years
(Mar 1997-Dec 1999), representing approximately
5562 person-h and 8840 trap-h (Table 1). Sam-
pling was conducted at sites dominated by the
most representative vegetation types of the re-
gion, medium height tropical semi-evergreen for-
est (selva median, or Upland Standard Forest
sensu Schulze 1992) and low tropical semi-decid-

September 2008

Pozo et al.: Phenology of Calakmul Butterflies

0 10 20 km




'E ". A F

ROO .. ... ..



Biosphere Reserve

... C


A' H





Fig. 1. Location of the 11 study sites within and near the Calakmul Biosphere Reserve: A = Calakmul Archeo-
logical Zone; B = road to Calakmul Archeological Zone; C = Narciso Mendoza Ejido; D = Nuevo Becal Ejido; E = en-
trance to El Papagayo; F = path to Flores Mag6n; G = north of Mancolona; H = Plan de Ayala Ejido; I = west of El
Refugio Ejido, with medium tropical semi-evergreen forest; J = west of El Refugio Ejido, with low tropical semi-de-
ciduous forest. Ejido = communal property.

uous forest (selva baja or Low Forest sensu Rze-
dowski 1983), as determined by botanist Esteban
Martinez. Within each forest type, transects of
500 m were established within habitat types of
different successional stages, including primary
forest, forest disturbed more than 10 years ago,
and forest disturbed within the past 10 years. In
each habitat type, replicate transects were sam-
pled (separated by at least 300 m), for a total of 18
transects in the first year, and 12 transects dur-

ing the second and third years (Table 2), at a total
of 11 localities. Six localities are within or near
the northern portion of the CR, and 5 are within
or near the southern part of the CR (Fig. 1).
The first year, sampling was conducted 4 tines
each season at each locality. The second year,
sampling was conducted only during the months
of greatest butterfly abundance, from Jun to Nov.
The third year, all localities were sampled
monthly. Two sampling methods were utilized, di-









September 2008

Florida Entomologist 91(3)




-J F i A mJ J A SO N D


Fig. 2. Annual evaporation levels, precipitation and temperature recorded for 1997, 1998, and 1999 at the Zoh-
Laguna Research Station in the Calakmul region.

rect search (DS) and transects (T). For each
method, 2 sampling techniques were employed:
traps with bait (t) and entomological nets (n).
Transects (T) were established through the forest
by clearing a small path of 500 m length, with a
machete, and wide enough to permit the passage
of the observer without altering the canopy. In
each transect, 10 Van Someren-Rydon traps (Ry-
don 1964) baited with banana, pineapple, and
beer, were hung (from 0700 to 1800 h) at 50 m in-
tervals, on alternate sides of the path. Fresh bait
was added each morning. Two surveyors spent
120 to 150 min in each transect. Surveyors
walked each transect, spending 10 min at each
trap to record trap contents (on a pre-prepared

form) and sample nearby individuals with ento-
mological nets. Direct Search surveys (DS) were
more opportunistic. These were conducted by
walking along roads (2 observers for approxi-
mately 8 h per d) in search of sites where butter-
flies were concentrated (sunny gaps, nectar
sources, damp ground, etc.), sampling butterflies
as opportunities were encountered. In addition,
10 traps baited with fermenting fruit (as above)
were placed in "optimal" localities along the road-
sides and checked periodically.
Upon determination to species-level, trapped
butterflies were usually released. In cases where
the identity of a trapped butterfly was uncertain
or unknown, it was collected for determination at,

HOURS, FOR 1997, 1998, AND 1999.

Trap- Man-
J F M A M J J A S O N D Total hours hours

1997 12 12 12 12 14 12 12 86 3440 2322
1998 13 13 11 14 9 9 8 77 2310 1386
1999 3 9 9 10 7 7 8 10 11 9 12 8 103 3090 1854

Total 266 8840 5562




:1 . . .
e Is.

J F l A l l l A S O N D



J F n A J J A N

Pozo et al.: Phenology of Calakmul Butterflies


Primary Disturbance >10 years Disturbance <10 years
type 1997 1998 1999 1997 1998 1999 1997 1998 1999

MT 3 2 2 3 2 2 3 2 2
LTb 3 2 2 3 2 2 3 2 2
Total 6 4 4 6 4 4 6 4 4

aMedium tropical semi-evergreen forest.
'Low tropical semi-deciduous forest.

and integration into the Zoology Museum at El
Colegio de la Frontera Sur, Chetumal, Mexico
(duplicate specimens are deposited at the Zoology
Museum of the Universidad Nacional Aut6noma
de Mexico, Mexico City). To determine the extent
to which released butterflies may be re-trapped
and re-counted, all individuals released for a 3-
month period were marked before release, and re-
capture data was collected. Recapture data con-
firmed that once released from a trap, individuals
did not usually return to traps (Hughes et al.
1998). However, only long-lived and phylopatric
species, or those with a small "home range" could
cause a bias in the data through recaptures, and
few such species were encountered in the CR.
Data from fieldwork were entered into the da-
tabase of the Lepidoptera collection at the Zoology
Museum at ECOSUR, representing a total of
60,662 records of 359 species, in 207 genera, from
18 subfamilies, 5 families and 2 superfamilies
(nomenclature follows Llorente et al. 2006; A.
Warren 2000, except that for the analyses we con-
sidered the families Riodinidae and Lycaenidae


Monthly species abundance (phenology) and
seasonal distribution (seasonality) data were ana-
lyzed. In order to describe phenological patterns,
analyses were divided into 2 parts. First, the total
number of individuals and species recorded was
considered, and a family-level analysis was con-
ducted (Shapiro 1975). Second, the phenology of
the most abundant species was analyzed (species
with 100 or more records during the 3 years of
sampling. The abundant species were divided into
3 size categories (Young 1982;Vargas-Fernandez et
al. 1999). Because of the great diversity of butterfly
species within the CR, adult size showed great
variability (e.g., Pyrisitia nise nelphe) (R. Felder
1869), (forewing length = 17 mm + 0.9 mm) and
Leptotes cassius cassidula (Boisduval 1870), (12.7
mm 1.1 mm) us. Archaeoprepona demophon cen-
tralis (Fruhstorfer 1905), (54 mm 4.4) and Mor-
pho achilles montezuma Guenee 1859 (67 mm 4.0
mm). Wingspan intervals were used as a proxy of

body size (Vargas-Fernandez et al. 1999). The 3 ar-
bitrary wingspan size categories were: small (<30
mm), medium (31-50 mm), and large (>51 mm).


For each year, a data matrix was constructed,
which recorded the species and their abundance
in each season (dry, rainy, and cold or "nortes").
An NMDS analysis was applied to each matrix.
Data was log-transformed according to the Box
and Cox test (Legendre & Legendre 1998). For
these analyses we used PRIMER 4.0 software
(Carr 1996), and results were graphed with Sigma-
Plot Version 7.1, Systat. In order to statistically
evaluate differences among groups in each sea-
son, an ANOSIM test (ANalyis Of SIMilarities)
was applied, and subsequently, a SIMPER (SIMi-
larity PERcentages) analysis was applied to de-
tect species that contributed to group formation,
and those species which acted as discriminants
among groups (Herrando-Perez 2002).



In general, the phenological patterns found in
this study are similar to those described by de la
Maza and de la Maza (1985a, b) and Austin et al.
(1996), with peak butterfly diversity at the end of
the dry season and another peak during the rainy
season. Reduced species diversity was observed
from the end of the rainy season to the middle of the
dry season (Fig. 3). Nine percent of the Calakmul
butterfly species have been recorded in every
month; similarly, Austin et al. (1996) reported 10%
of the butterfly fauna from the Tikal area, Guatema-
la (about 100 km S of Calakmul), from all months.
The month with the greatest species richness
was Oct 1997, with 231 species, and the month
with the greatest relative abundance was Aug
1998, with 17,324 individuals (more than the
1997 annual total of 16,257 records). Maximum
species diversity was observed during Oct and
Nov, in agreement with previous studies in Mex-
ico (Vargas-Fernandez et al. 1992, 1999). This pe-

Florida Entomologist 91(3)


o <10 individuals
n 10 X100
m 101S X s 1000
M 1001 SXS10000
*X > 10000


Sierra de Atoyac de Alvarez

250 b)

_ 200


E 100

z 50


Sierra de ManantlAn

250 C)

Fig. 3. a) Phenological pattern of the butterfly fauna
of the Calakmul region, showing species abundance
throughout the year; b) phonological pattern of the but-
terfly fauna of the Sierra de Manantlan, Jalisco-Colima;
c) phenological pattern of the butterfly fauna of the Si-
erra de Atoyac de Alvarez, Guerrero.

riod coincides with the end of the rainy season.
The year with the greatest species richness was
1997, with 265 species recorded (74% of the total
species number), although a similar diversity was
observed in 1998 (261 species). During 1999, spe-
cies richness decreased to just 58% of the total
species diversity recorded in this study.
Large numbers of records for any given month
do not necessarily indicate great species diversity.
The greatest number of records obtained during
any month of the study was 17,324 records in Aug

1998, representing 28.56% of the total records ob-
tained during this study. However, these individ-
uals represented only 108 species (Table 3), and 3
migratory species comprised 86.5% of the sample:
Memphis pithyusa (R. Felder, 1869), Anaea trogl-
odyta aidea (Guerin-Meneville, 1844) and Eunica
tatila tatila (Herrich-Schaffer, 1855). Memphis
pithyusa was the most frequently recorded spe-
cies in this study, with 12,848 records (Table 6).
Table 4 shows that 90.8% of the species re-
corded during this study were represented by
only 19.7% of the recorded individuals. Species
which were present from 9 to 12 months equaled
28.4% of total species recorded, and 95.4% of the
total number of records. There were 140 species
present during 6 or more months of the year
(98.1% of all records), while 178 species were re-
corded only during 3 or fewer months, represent-
ing half of the fauna studied (49.58% of species
and 1.31% of records). Among species surveyed in
this study, 183 are represented by 10 or fewer
records (60 of which were only recorded once).
These are the "rare" species and represent 0.001%
of the total number of records.
Butterflies were divided into 5 groups accord-
ing to their relative abundance (Fig. 3). The first
group contains species represented by 10 or fewer
records (183 species), the second group by 11 to
100 records (110 species), the third by 101 to 1000
records (55 species), the fourth by 1001 to 10,000
records (10 species) and the fifth by more than
10,000 records. This last group only included one
species (M. p. pithyusa). Monthly variation in spe-
cies richness is clearly defined by the first 2
groups, represented by 293 species (81.6% of total
species richness and 7.5% of total relative abun-
dance), 96 of which were recorded during only 1
month (of those, 35 in Oct), and 60 of these repre-
sented by only 1 individual (21 in Oct) (Fig. 3). Ta-
ble 4 shows species frequency by month, from
those present all year (33 species), to those re-
corded only during 1 month. It is remarkable that
half of the fauna (178 species) was recorded dur-
ing 3 or fewer months, whereas 84 species were
present during 10 or more months. Thus, half of
the species are strongly seasonal (49.6%).
The 2 peaks in species richness coincide with
the greatest relative abundance (Fig. 3a and Ta-
ble 3). The largest peak was observed from Jun to
Dec, which corresponds with the rainy season, al-
though there were markedly fewer species during
Sep. The smaller peak was observed from Feb to
Apr. We estimated that a minimum of 110 species
should fly each month, if we consider that 28-35%
of species in this study were found during the en-
tire year, and every year.

Family Level Phenology

The most diverse families in the study area are
those showing the largest variation in species

September 2008

Pozo et al.: Phenology of Calakmul Butterflies


Year J F M A M J J A S O N D Total

1997 102 95 112 152 231 187 122 265
1998 50 103 108 137 101 74 60 261
1999 35 121 131 119 98 94 142 90 74 73 72 53 208
1997-1999 59 121 157 138 98 147 168 196 144 250 207 143 359
1997 800 1985 1414 2325 4277 3739 1717 16257
1998 1075 4002 17324 4621 2984 1403 680 32089
1999 320 1712 2232 1308 524 482 1906 1408 571 527 832 494 12316
1997-1999 1395 1712 3032 3293 524 1896 5908 21057 5192 7788 5974 2891 60662

richness throughout the year (Hesperiidae = 135
species, Nymphalidae = 111 species; Fig. 4). In
contrast to other families, peak diversity of Papil-
ionidae was observed during the late dry season
into the early rainy season, as reported by Austin
et al. (1996) for the Tikal area. Relative abun-
dance of Papilionidae was distributed irregularly
throughout the year, but the greatest number of
individuals (27.5%) was recorded in Apr (Table 5);
the greatest species richness was in Jul (13 spe-
cies), Apr (11 species) and Aug (11 species). Diver-
sity and relative abundance of Pieridae species
remained more or less constant throughout the
year, showing a slight increase during the rainy
months of Aug to Nov; Oct was the peak month for
species richness (with 21 species) as well as rela-
tive abundance. Hesperiidae, Lycaenidae, and
Nymphalidae showed 2 periods of increased spe-
cies richness: the first (and lesser) occurred dur-
ing the dry season, while the second (and greater)
occurred during the rainy season; these two peri-
ods are separated by the dry month of May when
a decreased species diversity was observed. Spe-

cifically, the greatest species diversity for Lycae-
nidae was Oct (44 species; a second peak in Mar
with 25 species), with the greatest relative abun-
dance in Aug; the lowest species diversity and rel-
ative abundance among Lycaenidae was observed
in Jan. The greatest species diversity for Nymph-
alidae was observed in Oct (with a smaller peak in
Mar), while the greatest relative abundance was
in Aug (with a smaller peak in April). The differ-
ence in abundance of Nymphalidae between dry
and rainy seasons was greatly accentuated: 90%
between the months of highest and lowest abun-
dance (Table 5). Similarly, over 2 times as many
species of Lycaenidae, and over 3 times as many
species of Hesperiidae were recorded in the rainy
season than in the dry season (Table 5).

Phenology of Abundant Species Related to Size

Sixty-eight species were considered abundant
(Table 6), and 75% of these were recorded from all
months. These species are distributed among 5
families, but most (51 species) are nymphalids.


Number Number Cummulative Number Cummulative
of months of species Percentage percentage of individuals Percentage percentage

1 96 26.74 26.74 231 0.38 0.38
2 60 16.71 43.45 363 0.60 0.98
3 22 6.13 49.58 203 0.33 1.31
4 20 5.57 55.15 260 0.43 1.74
5 21 5.85 61.00 378 0.62 2.37
6 11 3.06 64.07 442 0.73 3.09
7 19 5.29 69.36 556 0.92 4.01
8 8 2.23 71.59 348 0.57 4.58
9 18 5.01 76.60 1557 2.57 7.15
10 21 5.85 82.45 2770 4.57 11.72
11 30 8.36 90.81 4846 7.99 19.71
12 33 9.19 100.00 48708 80.29 100.00
Total 359 100.00 60662 100.00

Florida Entomologist 91(3)

--- Papilionidae Pieridae
- .mpteid --Lycaenklae
--- einidae

0 4-+----I------- --.-- -
Fig. 4. Phenological patterns displayed by butterfly
families present in the Calakmul region (for this figure,
Riodinidae is included within Lycaenidae).

These 68 species represent 91.6% of the records
obtained in this study, but represent only 18.9% of
the total species richness. Out of the 3 groups
formed according to size categories, 25 species are
small, 32 are medium, and 11 are large.
Some variation exists with respect to seasonal
trends in species richness of the 3 size categories
(Fig. 5). For the small-sized group, a difference of
40% exists between the months of greatest and
least species richness. For medium-sized butter-
flies, this difference is 31.25%, and for large butter-
flies, a 36% difference exists. Figure 5b shows the
largest relative abundance for the 3 categories
from Jun to Dec, with a strong decrease during Sep.


Butterfly diversity was found to be signifi-
cantly variable among dry, rainy, and cold seasons
(Fig. 6 and Fig. 7) during each year of the study
(1997: stress = 0.15; 1998: stress = 0.05; 1999:
stress = 0.18). Of the 359 species used for this
analysis, 9% (32 species) provided more than 80%
of the effective data when grouping the communi-
ties in each of the 3 seasons (Table 7), 50% of dif-
ferentiation among communities of each season
was determined by the relative abundance of 31
species (8.6% of the total). Of the 32 species com-
prising the communities in each season, 2 did not

contribute to the distinction among communities
,112 .._. ., cyaniris cyaniris Doubleday (1848) and
Siderone galanthis (Cramer 1775)), and 1 did not
contribute to the formation of groups (Marpesia
chiron marius (Cramer 1779)), although it did
when groups were analyzed separately. Of these
32 species, only 3 did not belong to Nymphalidae:
Kricogonia lyside (Godart 1819), Glutophrissa
drusilla tennis (Lamas 1981) and Polygonus sav-
igny savigny (Latreille, (1824)) (Table 7).
When species richness by family in each sea-
son is considered (Fig. 7a), it is clear that Hesperi-
idae differed in phenological behavior from the
other families. This family had the greatest spe-
cies richness in the cold season, with values more
than double of those observed during the rainy
and dry seasons. Nymphalidae and Lycaenidae
showed maximum species richness during the
rainy season, with a slight decrease in the cold
season, while Pieridae and Papilionidae showed
minimal variations. Pieridae showed a slight in-
crease during the rainy season, at a time when di-
versity of Papilionidae decreased to values lower
than those of the dry season.
Relative abundance by season was strongly in-
fluenced by Nymphalidae (Fig. 7b). Seasonal dif-
ferences among the other families were not de-
tected on this scale, due to the abundance of
nymphalid species. Nevertheless, abundance of
Hesperiidae showed an increase in the cold sea-
son (Table 5).


Phenology Considering the Relative Abundance
of Species

Wolda (1988b) concluded that there are differ-
ences in phenological patterns among insect fau-
nas typical of temperate and tropical zones. Un-
der the assumption that seasonal changes are
minimal in the tropics, adults of the majority of
species should be present throughout the year
(Owen 1971), whereas in temperate zones adults
are restricted to the most favorable seasons (usu-
ally spring and summer). Nevertheless, upon ex-
amining climatic variations for each year of our


Taxon J F M A M J J A S O N D Total

Papilionidae 0 9 73 168 69 43 83 54 22 31 26 32 610
Pieridae 9 125 256 303 38 73 235 414 264 460 347 198 2722
Nymphalidae 1363 1384 2408 2599 333 1609 5248 20127 4698 6409 5239 2508 53922
Lycaenidae 22 108 171 135 56 122 214 372 173 280 163 100 1916
Hesperiidae 1 86 127 88 28 49 128 90 35 608 199 53 1492
Total 1395 1712 3032 3293 524 1896 5908 21057 5192 7788 5974 2891 60662

September 2008


Taxon J F M A M J J A S O N D 1997 1998 1999 Total

Eunica tatila tatila (Herrich-Schaffer [1855])
Pareuptychia metaleuca metaleuca (Boisduval 1870)
Pyrisitia nise nelphe (R. Felder 1869)
Hermeuptychia hermes (Fabricius 1775)
Cissia pseudoconfusa Singer, DeVries & Ehrlich 1983
Kricogonia lyside (Godart 1819)
Mestra dorcas amymone (M6n6tri6s 1857)
Cissia pompilia (C.Felder & R. Felder 1867)
Eumaeus toxea (Godart [1824])
Juditha molpe molpe (Hiibner [1808])
Cepheuptychia glaucina (H.W. Bates 1864)
Cissia similes (Butler 1867)
Nica flavilla bachiana (R.G. Maza & J. Maza 1985)
Cissia confuse (Staudinger 1887)
Pareuptychia ocirrhoe ssp. n.
Yphtimoides renata (Stoll 1780)
Thessalia theona theona (M6n6tri6s 1855)
Tegosa frisia tulcis (H.W. Bates 1864)
Leptotes cassius cassidula (Boisduval 1870)
Pteronymia cotytto cotytto (Gu6rin-M6n6ville [1844])
Dynamine dyonis Geyer 1837
Pyrgus oileus (Linnaeus 1767)
Mesosemia tetrica Stichel 1910
Pyrisitia dina westwoodi (Boisduval 1836)
Heliopetes alana (Reakirt 1868)

Memphis pithyusa pithyusa (R. Felder 1869)
Anaea troglodyta aidea (Gu6rin-M6n6ville [1844])
Memphis forreri (Godman & Salvin 1884)
Opsiphanes cassina fabricii (Boisduval 1870)
Myscelia ethusa ethusa (Doybre [1840])
Memphis moruus boisduvali W.P. Comstock 1961
Tp- .. virgilia (Cramer 1776)
Hamadryas februa ferentina (Godart [1824])

120 456 1111 199 76
109 5 8 40 3
2 24 37 10 5
22 40 73 51 2
33 6 31 43 11
2 79 198 19
11 16 13 4
1 10 19 11 1
13 19 23 29 8
3 8 21 14
6 15 27 25 13
16 8 11 64 5
39 6 8 36 2
6 16 20
9 26 8 22
2 5 12 18 1
10 13 4 3
1 34 22 4 1
7 29 32 11
6 2
18 21 12 6
4 10 5 3
4 7 1 2
4 15 8 4

204 99
4 19
139 23
35 9
3 116
89 123
139 4
22 21

76 144 5
8 18 1
40 144 16
4 23 1
72 85 24
46 60 27
13 144 6
36 89 16

159 1640 3722 242
2 28 79 24
8 47 105 53
31 30 11 13
28 25 34 16
7 36 36 50
12 19 35 16
2 26 31 13
20 27 39 4
22 59 45 9
3 6 6 7
10 2 4 4
11 13 5
12 8 3
3 26 3
9 8 15 10
9 11 36 8
1 5 3 6
8 12 4 7
1 19 1 1
3 3 12 4
3 14 8
1 14 16
2 14 9 21
2 6 5

472 449 341
41 131 43
81 96 32
75 104 34
83 120 38
5 12 1
94 22 36
57 38 20
6 20 7
16 8 4
12 21 66
18 40 5
3 33 11
68 23 8
9 31 18
10 25 37
20 25 8
29 9 10
17 38 3
39 30 13
19 9
29 13 12
13 16 9
31 7 8

344 5261 3382 8987
328 39 146 513
308 83 109 500
315 23 148 486
239 49 180 468
20 98 327 445
169 20 89 278
135 29 65 229
88 26 101 215
54 70 85 209
51 73 83 207
125 22 40 187
107 3 57 167
156 8 164
65 3 87 155
61 43 48 152
59 15 73 147
56 10 59 125
48 26 41 115
49 3 62 114
88 13 11 112
42 2 66 110
34 46 27 107
27 29 42 98
60 1 29 90

707 6757 2154 1750 538 172 1737 10398 713 12848
742 5332 911 157 36 11 218 6873 379 7470
296 353 30 735 249 68 1408 468 334 2210
69 595 172 375 137 16 863 471 107 1441
125 333 70 19 462 1 42 996 289 1327
44 298 51 173 174 95 300 287 605 1192
28 172 41 171 220 50 776 189 88 1053
45 47 164 80 57 36 225 301 165 691


Taxon J F M A M J J A S O N D 1997 1998 1999 Total

Heliconius erato petiverana Doubleday 1847
Glutophrissa drusilla tenuis (Lamas 1981)
Hamadryas guatemalena guatemalena (H.W. Bates 1864)
Hamadryas julitta (Fruhstorfer 1914)
Polygonus manueli manueli Bell & W. P. Comstock 1948
Heliconius charithonia vazquezae Comstock & Brown 1950
Temenis laothoe hondurensis Fruhstorfer 1907
Dryas iulia moderate (Riley 1926)
Colobura dirce dirce (Linnaeus 1758)
Anartia amathea fatima (Fabricius 1793)
Fountainea eurypyle confusa (Staudinger 1887)
Marpesia chiron marius (Cramer 1779)
Biblis hyperia aganisa Boisduval 1836
Phoebis agarithe agarithe (Boisduval 1836)
Protographium philolaus philolaus (Boisduval 1836)
Pieriballia viardi viardi (Boisduval 1836)
Adelpha paraena massilia (Felder & Felder 1867)
Opsiphanes quiteria quirinus Godman & Salvin 1881
Consul electra electra (Westwood 1850)
Asterocampa idyja argus (H.W. Bates 1864)
Ganyra josephina josepha (Salvin & Godman 1868)
Myscelia cyaniris cyaniris Doubleday [1848]
Danaus gilippus thersippus (H.W. Bates 1863)
Anartia jatrophae luteipicta Fruhstorfer 1907
Archaeoprepona demophon centralis (Fruhstorfer 1905)
Archaeoprepona demophoon gulina (Fruhstorfer 1904)
Morpho achilles montezuma Guen6e 1859
Historis acheronta acheronta (Fabricius 1775)
Historis odius dious Lamas 1995
Prepona laertes octavia Fruhstorfer 1905
Anteos maerula (Fabricius 1775)
Siderone galanthis ssp. n.
Siproeta stelenes biplagiata (Fruhstorfer 1907)
Phoebis philea philea (Linnaeus 1763)
Prepona pylene philetas Fruhstorfer 1904





50 28 168
30 20 27
11 3 22
55 83 126
22 28 9
26 28 23
5 26 72
12 9 15
7 4 20
19 87 10
32 20 135
6 6
12 17 7
6 16 12
35 75
10 15 1
6 13 1
11 17
14 26 36
4 4

1 4

29 23
23 13
9 1
4 10
2 3
1 4

4 9
12 5
12 5

62 198
71 101
8 62
10 24
24 22
5 37
2 7
10 6
12 24

8 35 67
14 33 149
24 15 146
48 35 49
6 82 3
2 14 47
31 8 46
6 10 45
7 8 13
3 12 11
90 95 228
8 33 52
5 6 15
6 31 11
11 6 4
2 11
4 10 31
5 5 30
11 18
7 10
6 2 22
40 27 20
12 10 13
5 9 7

25 535 338
39 148 267
22 6 116
16 45 307
22 45 72
3 70 64
13 26 25
4 19 27
15 26
9 14 8
1 9 30

101 127 51
103 60 20
64 92 68
15 8 6
184 21 11
89 60 35
43 22 11
110 44 27
21 111 41
56 64 28
209 74 137
53 9 2
65 48 16
80 27 7

16 38 51 28
26 66 6 5
42 20 20
3 8 13 14
46 44 8 5
3 40 15 31
2 1 7
9 18 16 2
13 26 15 4

258 387 217
120 508 309
26 283 235
3 5 5
56 38 9
52 51 8
20 25 16
26 16 1
48 22 13
25 10 24
8 4 1

82 199 682
168 146 493
288 45 483
41 323 454
265 87 404
34 133 357
74 78 349
86 85 328
28 54 319
10 85 309
8 135 299
107 53 236
33 72 203
40 29 202
10 106 184
11 41 176
92 19 169
9 3 168
21 66 162
95 13 131
28 32 129
42 17 116
17 63 109
12 45 101

949 208 2088
415 237 1636
108 269 787
344 77 517
100 38 336
106 37 290
26 114 207
63 24 161
50 69 157
16 38 122
31 32 100

Pozo et al.: Phenology of Calakmul Butterflies








Fig. 5. Analysis of phenological patterns and relative
abundance according to adult size. a) Phenology repre-
sented by size groups: small (<30 mm wingspan); me-
dium (31 to 50 mm); large (>51 mm); and combined. b)
Relative abundance of individual butterflies by size

study, the expected uniformity in phenological
patterns of butterfly species at our tropical study
sites does not exist. During the 3 years of our
study, there were important differences in precip-
itation levels, and to a lesser degree, in tempera-
ture, which caused levels of evaporation to differ
between years. Our results indicate a species
turnover of more than 30% (as represented by
adults), from the dry to rainy season. This helps
explain seasonal variation in butterfly species
richness, as only 33 species (9%) fly year-round,
either as 1 long-lived generation, multiple super-
imposed generations, or several non-superim-
posed generations. The greatest species richness
was observed during 1997, the year with the larg-
est evaporation levels.
In the CR, biotic and abiotic factors are not ho-
mogeneously distributed, generally due to soil
characteristics of the region, which in turn deter-
mine the existence of a heterogeneous mosaic of
vegetation types. This mosaic is so intertwined
that it is difficult to define vegetation types, and
even more so to distinguish them (Galindo-Leal
2001; Garcia-Gil et al. 2001; Martinez & Galindo-
Leal 2002). This in turn produces a differential
spatial and temporal distribution of the lepi-
dopteran fauna. It is possible that the phenology
of each species is more closely related to these fac-

-- Large

-N- Smal
-a- Medium
-- Large

tors, in response to the different micro-habitats
where they live.
In order to investigate this relationship, our
results were compared with phenological data on
the butterflies of the Sierra de Atoyac de Alvarez,
Guerrero (Vargas-Fernandez et al. 1992), and the
Sierra de Manantlan, Jalisco-Colima (Vargas-
Fernanez et al. 1999). Both areas cover altitudi-
nal transects; the former from 300 to 2500 m, and
the latter from 250 to 1750 m. The sampling pro-
tocols used in these studies are equivalent to that
of the present study, thus we believe that compar-
isons can be made among phenological data ob-
tained for each of the 3 areas. However, the 2 Si-
erras (mountain ranges) present biotic and abi-
otic conditions that differ from those found in the
CR, and have different biogeographical histories.
While the Sierra de Atoyac de Alvarez and the Si-
erra de Manantlan are situated along Mexico's
Pacific watershed, they differ in latitude, average
precipitation levels, and average temperature.
Species richness of Papilionoidea is very similar
across the 3 areas (although Hesperiidae were not
surveyed by Vargas-Fernandez et al. 1992, 1999).
In Atoyac de Alvarez, 337 species were recorded,
with the greatest species richness in Jul (216 spe-
cies), whereas in Manantlan, 315 species were re-
corded, with a maximum diversity of 216 in No-
vember. In the CR, 359 species were recorded (250
in Oct), representing an average difference in over-
all diversity among the sites of about 12%.
Unique environmental attributes of each region
should cause phenological patterns among the
butterfly fauna to vary among them. Some reviews
of butterfly phenology suggest that climate is the
main factor controlling the activity of these organ-
isms (Brakefield & Shreeve 1992; Warren 1992;
Gutierrez & Menendez 1998). However, climatic
factors may be influenced by differences between
habitats or years, correlated with microclimatic
changes at local or regional levels. In this study,
phenological similarities were found with respect
to species richness trends throughout the year.
Two periods of maximum species richness were
distinguished: the period of greatest diversity dur-
ing the rainy season, and the other peak during
the dry season. This indicates that phenological
patterns among tropical butterfly faunas are much
more complex than some authors have recognized.
Several authors have suggested that tropical
faunas may be characterized by a large number of
species with very low densities (Lamas et al.
1991; Owen 1971; Owen et al. 1972), which form
the group of "rare" species. This group is an im-
portant part of the species diversity of each region
and is the modulating factor of phenological
changes, as demonstrated in this study. Upon ex-
amining those species represented by fewer than
10 records for each of the 3 regions we compared,
it is evident that they represent a significant pro-
portion of total species richness, and that they

Florida Entomologist 91(3)


a Cold "Nortes"




Fig. 6. Plots of the 3 axes obtained by NMDS for data from 1997, 1998 and 1999. Transformed data Log (y + 1).
1997: stress = 0.15, 1998: stress = 0.05 and 1999: stress = 0.18.

have a large impact on the overall phenological
patterns observed in each region. The group of
rare species from Manantlan is comprised of 117
species (37.1% of the total fauna), from Atoyac de
Alvarez 144 species (42.7%), and from the CR 183
species (51%). The group of rare species may
change from year to year due to seasonal and en-
vironmental variables, which have been poorly
studied. Therefore, details on the specific mecha-
nisms that lead to seasonal variation in the com-
position of rare species are needed to fully under-
stand phenological patterns in any area.

If we consider precipitation and temperature
(causes of available humidity) as principal factors
in determining the phenological patterns of vege-
tational communities, and therefore of butterflies,
the timing and severity of the dry season is likely
to be one of the most consequential factors in de-
termining regional phenological patterns. In
Atoyac de Alvarez, the difference in humidity be-
tween the two seasons is lower than that of
Manantlan, which minimizes the effects of the
dry season and leads to an earlier peak in species
richness. Furthermore, the average number of


September 2008

Pozo et al.: Phenology of Calakmul Butterflies



O Nymphalidae
* Lycaenidae

30000 -
3 25000
aS isooo .
| 10000 -
z 5000


Fig. 7. a) Diversity of Papilionidae, Pieridae, Nymph-
alidae, Lycaenidae (including Riodinidae) and Hesperi-
idae across the 3 seasons (dry, rain, cold) of the
Calakmul region. b) Relative abundance of adults of all
families combined for each season.

species present per month is greater in Atoyac de
Alvarez than in the other 2 regions. On the other
hand, in Manantlan, the process of species accu-
mulation to the point of maximum diversity is
slower, possibly because deciduous or semi-decid-
uous vegetation (drier forests than those of
Atoyac de Alvarez) is present at almost all locali-
ties below 1800 m. The same phenomenon is also
be observed in the CR, where the dry season is
very pronounced, particularly due to the soil type
characterized by calcareous rock, which causes
rapid drainage. As a consequence of this, a
greater similarity exists between the phenologi-
cal patterns of the butterfly fauna of Manantlin
and CR, than between CR and Atoyac de Alvarez.
While the "rare" species of a region comprise
an important component of overall species diver-
sity (as for CR, Fig. 8a), they do not allow for a
straightforward phenological analysis, since the
composition of rare species may change monthly
and yearly. Thus, in order to study seasonal and
annual phenological patterns, we excluded rare
species to standardize comparisons (Fig. 8b). With
rare species excluded, overall phenological pat-
terns in Manantlan and CR show great similarity,
while those of Atoyac de Alvarez are discordant.

Kricogonia lyside (Godart, 1819)
Glutophrissa drusilla tenuis (Lamas, 1981)
Heliconius erato petiveranus Doubleday, 1847
Historis odius dious Lamas, 1995
Historis acheronta acheronta (Fabricius, 1775)
Colobura dirce dirce (Linnaeus, 1758)
*Myscelia cyaniris cyaniris Doubleday, [1848]
Myscelia ethusa ethusa (Doybre, [1840])
Eunica tatila tatila (Herrich-Schaffer, [1855])
Hamadryas februa ferentina (Godart, [1824])
Hamadryas guatemalena guatemalena (H.W. Bates,
Hamadryas julitta (Fruhstorfer, 1914)
Temenis laothoe hondurensis Fruhstorfer, 1907
Nica flavilla bachiana (R.G. Maza y J. Maza, 1985)
**Marpesia chiron marius (Cramer, 1779)
Archaeoprepona demophon centralis (Fruhstorfer,
Archaeoprepona demophoon gulina (Fruhstorfer, 1904)
Prepona laertes octavia Fruhstorfer, 1905
*Siderone galanthis ssp. nov.
Anaea troglodyta aidea (Gu6rin-M6n6ville, [1844])
Consul electra electra (Westwood, 1850)
Fountainea eurypyle confusa (A. Hall, 1929)
Memphis forreri (Godman & Salvin, 1884)
Memphis phila boisduuali W.P. Comstock, 1961
Memphis pithyusa (R. Felder, 1869)
Morpho achilles montezuma Guen6e, 1859
Opsiphanes invirae fabricii (Boisduval, 1870)
Cepheuptychia glaucina (H.W. Bates, 1864)
Cissia pseudoconfusa Singer, DeVries & Ehrlich, 1983
Pareuptychia binocula metaleuca (Boisduval, 1870)
Pareuptychia ocirrhoe ssp. nov.
T. .. -. virgilia (Cramer, 1776)
Polygonus manueli manueli Bell & W. P. Comstock,

*Species that contributed to form groups.
"Species that contributed to separate the groups.

This may be due to the shorter and less severe dry
season of Atoyac de Alvarez, as compared to the
other 2 regions. However, if we displace the data
from Atoyac de Alvarez 1 month forward (Fig. 8c),
phenological patterns of the 3 regions are all re-
markably similar. The months with the greatest
species richness are Oct for CR (157 species) and
Atoyac de Alvarez (172) and Nov for Manantlin
(164). In contrast, the months of lowest diversity
are May for CR (93) and Manantlan (81), and Apr
for Atoyac de Alvarez (77). This pattern is well
known to many experienced collectors in Mexico,

Florida Entomologist 91(3)

a) -0-Maurn
-a-Atoyc do kAu

o b



SF 0



Fig. 8. Phenological patterns of the butterfly fauna of
3 regions in Mexico. RBC = Calakmul Region, Yucatan;
Manantlan = Sierra de Manantlan, Jalisco-Colima;
Atoyac de Alvarez = Sierra de Atoyac de Alvarez, Guer-
rero. a) Phenological patterns of the 3 regions consider-
ing all species; b) phenological patterns excluding rare
species; c) phenological patterns with a 1-month (for-
ward) displacement for the Sierra de Atoyac de Alvarez.

yet this is the first time this pattern has been
quantified with data from various regions.

Family Level Phenology

Papilionidae is composed of few species, most
of which are large-bodied, often with small popu-
lation sizes. Longer survivorship due to their
large adult size may allow this family to maintain
relatively constant levels of species richness
throughout the year. Larger butterflies are good
at maintaining their water balance (Janzen &
Schoener 1968), the greatest problem confronted
by insects during the dry season. The effect of hu-
midity is important with respect to insect body
size, and those organisms with small bodies des-
iccate more easily than those with medium to
large bodies (Young 1982). In addition, the rela-
tively consistent diversity of Papilionidae across
the seasons could be related to phenological pat-
terns of larval foodplants or the availability of

adult nectar or mineral resources. For species
with long reproductive lives (e.g., Heliconiinae),
availability of nectar resources may be especially
important in determining phenological patterns
(Gilbert & Singer 1975).
In contrast, the Lycaenidae and Riodinidae are
characterized mostly by small-sized species,
many of which center their reproductive activities
(and therefore adult voltinism) around the flower-
ing periods of larval foodplants (New 1993); in the
CR this is most often from Mar to Oct. Only 2 me-
dium-sized lycaenids (Eumaeus toxea (Godart,
1824) and Juditha molpe molpe (Hfibner, 1808))
are found throughout the year in the CR. The re-
maining species of Lycaenidae and Riodinidae are
seasonal, and appear only during favorable condi-
tions for reproduction.

Phenology of the Most Abundant Species with Respect
to Size

In the CR, we found that small and medium
sized species show monthly fluctuations in spe-
cies richness, contrasting with the patterns seen
in Manantlan (Vargas-Fernandez et al. 1999).
These fluctuations are not greatly pronounced,
but taken together they can help explain seasonal
variation. Small and medium-sized species show
the greatest species turnover across seasons, and
many of them are univoltine. As also found by
Young (1982), small-sized organisms tend to have
their largest populations during the most humid
season of the year.


This study shows that in the tropical CR, sea-
sonal variation in the abundance and diversity of
the butterfly fauna is small. However, through a
careful analysis, seasonal differences in the fauna
can be identified and compared to those of other
regions. We found that in the CR, variations in
phenological patterns are largely due to "rare"
species represented by fewer than 10 records,
whereas the seasonality of the butterfly commu-
nity is described by the most abundant species,
which define the faunas of the rainy, dry and cold
seasons. Of the 33 most abundant species in the
CR, the 11 most common are represented by 1000
to 10,000 records, while the remaining 22 species
are represented by 100 to 1000 records per taxon.
Of these, 15 species are present year-round.
Faunal studies on Lepidoptera of tropical re-
gions based on systematic sampling methods con-
ducted during all months of the year are scarce.
Likewise, few studies have investigated seasonal-
ity in tropical lepidopteran faunas. The informa-
tion presented herein explains phenological and
seasonal patterns of butterfly abundance and di-
versity, and should be useful in future studies on
the phenology of this immensely diverse group.

September 2008

Pozo et al.: Phenology of Calakmul Butterflies


We thank J. A. Padilla-Saldivar for the elaboration of
Fig. 1, and B. Prado-Cuellar for editing the manuscript.
We also thank El Colegio de la Frontera Sur (ECOSUR)
for support provided during the various stages of this
study. In addition, we thank 2 anonymous reviewers for
valuable suggestions that improved the manuscript.
Mexico's "Comision Nacional para el Uso y Conserva-
cion de la Biodiversidad" (CONABIO) provided finan-
cial resources through projects J112 and Q049. The
support of projects DGAPA IN 218502, DGAPA-UNAM,
PAPIME EN 202504 and CONACyT 36488 also was
very important.


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Wetterer & Keularts: Hairy Crazy Ant on St. Croix


1Wilkes Honors College, Florida Atlantic University, 5353 Parkside Dr., Jupiter, FL 33458

2Cooperative Extension Service, University of the Virgin Islands, Kingshill, St. Croix, VI 00850


The hairy crazy ant, Paratrechina pubens (Forel), is undergoing a population explosion on
St. Croix, US Virgin Islands. Here, we evaluate the status of P. pubens on St. Croix. In 2002,
residents of Calquohoun and surrounding areas in central St. Croix began reporting large
infestations of P. pubens. In 2005 and 2006, we surveyed ants at >100 sites across St. Croix.
We found 3 geographically discrete populations of P. pubens occupying ~5% of the island: a
main population centered on Calquohoun spread over ~9 km2 and 2 smaller populations oc-
cupying <1 km2 each. Locals blamed P. pubens for crop damage due to high densities of plant-
feeding Hemiptera that tended the ants. Surveys of trees in areas with and without P. pu-
bens present indicated that P. pubens has a significant negative impact on arboreal-foraging
ants. The distribution and chronology of P. pubens records on St. Croix suggest that this spe-
cies is a recently arrived exotic. It is unknown whether P. pubens will become a more serious
pest on St. Croix or whether populations will collapse to inconsequential levels.

Key Words: exotic species, Paratrechina pubens, pest ants, Virgin Islands


La poblaci6n de la hormiga peluda-loca, Paratrechina pubens (Forel) esta explotando en
St. Croix una de las Islas Virgines de los EEUU. Aqui, evaluamos el estado de P. pubens en
St. Croix. En 2002, los residents de Calquohoun y en las areas alrededores del centro de St.
Croix empezaron a reporter infestaciones grandes de P. pubens. En 2005 y 2006, realizamos
un monitoreo de las hormigas en mas de 100 sitios por toda la isla de St. Croix. Nosotros en-
contramos 3 poblaciones geograficas distintas de P. pubens ocupando ~5% de la isla: una po-
blaci6n principal con su centro en Calquohoun dispersada por ~9 km2 y 2 poblaciones mas
pequenas cada una ocupando <1 km2. La gente local culp6 a P. pubens por el dano a los cul-
tivos debido a la alta densidad de los hemipteros fit6fagos protegidos por las hormigas. Un
monitoreo de los arboles en las areas donde P. pubens estuvo present versus ausente indic6
que P. pubens tiene un impact negative significativo sobre las hormigas que buscan su ali-
mento en los arboles. La distribuci6n y cronologia de los registros de P. pubens en St. Croix
sugieran que esta es una especie ex6tica reci6n llegada. No se sabe si P. pubens llegara a ser
una plaga mas seria en St. Croix o si la poblaci6n bajara a niveles inconsecuentes.

The hairy crazy ant, Paratrechina pubens
(Forel), is known from sites scattered across the
New World tropics and subtropics. Forel (1893)
described P pubens from St. Vincent, and Smith
(1951) reported that P. pubens was known from
Mexico, Central and South America, and the West
Indies. Other West Indian records include Marti-
nique (Forel 1912), Anguilla (Trager 1984),
Guadeloupe (Trager 1984), Puerto Rico (Torres &
Selling 1997), and Cuba (Fontenla 1995). In
1990, Miller (1994) found P pubens in a barge full
of potted plants from south Florida being un-
loaded at Guana Island, British Virgin Islands.
The earliest Florida records of P. pubens come
from the Miami area in 1953 (Trager 1984), and
P pubens is now a localized exotic pest in Florida
(Deyrup et al. 2000). Wetterer (2007) recently re-
ported a previously unrecognized 1905 record of

P pubens from Bermuda, and proposed that this
species was the most likely candidate for an uni-
dentified ant that underwent a great population
explosion for 8 years in the 19th century in Ber-
muda and then disappeared (also see Wetterer
2006). Here, we report a recent population explo-
sion of P pubens on St. Croix, US Virgin Islands
and evaluate its status there.


While working for the Cooperative Extension
Service, JK received numerous complaints about
insect pests on St. Croix, including outbreaks of
P pubens. JK took notes on these complaints and
visited the sites. From 30 Oct to 5 Nov 2005 and 3-
12 Mar 2006, JW collected ants at >100 sites on
St. Croix. Collection sites included a diversity of

Florida Entomologist 91(3)

disturbed and relatively natural habitats. JW
sampled at 1-5 km intervals over most of the is-
land, but at shorter intervals in accessible areas
adjacent to sites where P pubens was found to be
From 5-10 Mar 2006, JW examined the impact
of P pubens on arboreal-foraging ants on St.
Croix. At each of 24 sites (designated sites A-X; 12
sites with P pubens present and 12 with P pubens
absent), JW surveyed ants on 10 trees with trunk
diameter >10 cm, spaced at ~10 m intervals in for-
ested areas and at more irregular intervals in ar-
eas with fewer trees. JW tacked a folded index
card holding ~1 g of canned tuna (in water) at 2 m
height on the north side of the trunk between 9:00
and 14:00h and returned in 2 h ( 15 min) to col-
lect the cards and place ants in plastic bags. The
ants were frozen, counted, and placed in 95% al-
cohol. JW initiated a parallel study of ants coming
to terrestrial baits at site A, but abandoned it be-
cause mongooses removed most of the bait cards.
Roy Snelling of the Los Angeles County Museum
of Natural History confirmed identification of
P pubens. Stefan Cover of the Museum of Com-
parative Zoology (MCZ) confirmed identification
of all specimens.


JK investigated several complaints concerning
P pubens from 3 contiguous Estates (i.e., town-
ships) in central St. Croix: Calquohoun, Little
Fountain, and Upper Love (geocoordinates given in
N, W). The earliest complaint, on 24 Aug 2002,
was from a resident in Calquohoun (site 1; 17.741,
64.793; see Fig. 1) whose house and property were
overrun by these ants. The resident reportedly
swept up a dustpan full of dead ants every day. For

more than a year, all additional complaints con-
cerning P pubens came from nearby parts of Cal-
quohoun. On 24 May 2004, a farmer in Little Foun-
tain (site 2; 17.749, 64.790; see Fig. 1) reported that
several small farm buildings as well as large areas
of pasture were overrun with P pubens. Less than
1 km from this farm, a physician in Little Fountain
(site 3; 17.752, 64.794) reported that large num-
bers of P pubens infested his house and the large
trees near the house (date not recorded). On 21 Oct
2005, a resident of Upper Love (site 4; 17.732,
64.810) complained of an infestation on his prop-
erty, and reported that he scooped up ~2 liters of
dead P pubens every day from his covered porch
and driveway area. On 6 Dec 2005, a resident of
Little Fountain (site 5; 17.755, 64.791) reported a
very large number of P pubens inside his house
and attributed the death of several rabbits kept in
cages on his property to these ants.
JW collected P. pubens at 12 sites on St. Croix
in Oct-Nov 2005 and at 19 additional sites in Mar
2006. Paratrechina pubens occurred at high den-
sities in at least part of each of these 31 sites.
We delineated 3 geographically separate popu-
lations covering ~10 km2 (~5% of the island; Fig.
1). The main population was spread across at
least 13 Estates and occupied ~9 km2, including
all sites sampled by JK plus 21 sites sampled by
JW: Bethlehem Old Works; Rte 64; 1 km S of Rte
72 (17.730, 64.796), Bethlehem Old Works, Rte
64; 0.5 km NW of Rte 70 (site E; 17.729, 64.795),
Body Slob; S of Rte 709; E of Rte 75 (17.739,
64.776), Calquohoun; Cruzan Gardens (site K;
17.739, 64.801), Calquohoun; NNE of Cruzan
Gardens (17.745, 64.793), Canaan; Rte 73; 1.5 km
NW Frangipani (17.760, 64.798), Castle Burke;
Rte 78; 2 km E of Rte 69 (17.724, 64.804), Fre-
densborg; S of Rte 707; W of Rte 73 (17.735,


0 5 1aKm
I i I 1 1 1 1 1 I IIIILLL..

Fig. 1. Ant collection sites on St. Croix. Squares = JK records of Paratrechina pubens complaints. Circles = JW
collection sites (filled = P. pubens present, open = P. pubens absent). The 3 earliest known P. pubens records are la-
beled: site 1 (Aug 2002), site K (early 2003), and site 2 (May 2004).

September 2008

Wetterer & Keularts: Hairy Crazy Ant on St. Croix

64.783), Fredensborg; turn in Rte 707 (17.733,
64.788), Hermitage; Blue Mt; NE side (17.754,
64.798), Hermitage; Blue Mt; SE side (site Z;
17.751, 64.797), Jealousy; 1 km N of Rte 72
(17.744, 64.807), Little Fountain; NE corner (site
A; 17.758, 64.792), Little Fountain; Rte 73; 0.5 km
NW Frangipani (17.753, 64.791), Little Fountain;
Rte 73; 1 km NW Frangipani (17.756, 64.794),
Lower Love; NW of gas station (site B; 17.720,
64.807), Mon Bijou; 0.2 km W of Frangipani (site
J; 17.750, 64.789), Mon Bijou; S of Rte 72; W of
Rte 73 (site I; 17.743, 64.786), Mon Bijou; NW cor-
ner of Frangipani (site Y; 17.750, 64.788), The
Glynn; Rte 72; 0.2 km W of Rte 75 (17.747,
64.774), and Upper Love; Holy Cross Church (site
D; 17.733, 64.807).
Two smaller P pubens populations occupied <1
km2 each. The Western population, sampled at 5
sites in 3 Estates, occupied ~0.2 km2: Mount Vic-
tory; 2.5 km up Rte 58 (site N; 17.744, 64.870),
Mount Victory; 3 km up Rte 58 (site P; 17.749,
64.868), Mt. Victory; 3.5 km up Rte 58 (site Q;
17.750, 64.863), North Hall; 2 km up Rte 58 (site
M; 17.745, 64.875), and Punch; 0.5 km SW of Mt.
Victory Camp (site T; 17.746, 64.871). The South-
ern population, sampled at 5 sites in 3 Estates, oc-
cupied ~0.5 km2: Anguilla; 0.5 km up landfill road
(17.7013, 64.7809), Anguilla; 1 km up landfill
road (17.697, 64.780), Bethlehem Middle Works;
Rte 64; 0.5 km S of Rte 66 (17.702, 64.785), Beth-
lehem Middle Works; Rte 64; N of Rte 66 (17.707,
64.787), and Mannings Bay; Rte 64; 1 km from
Rte 66 (17.699, 64.789).
At Cruzan Gardens (site K in Calquohoun; see
Fig. 1), a private 2.0-ha botanical garden and
nursery that imports plants from off-island (C.
Holmes, pers. comm.), a staff member who had
worked at the gardens for 30 years reported that
P pubens became apparent at the nursery in early
2003, and had an impact on many of the fruit
trees though tending enormous numbers of mea-
lybugs on the branches, leaves, and fruit. In Nov
2005, JW found that P pubens was extremely
abundant throughout most of the property, partic-
ularly in sections that were watered regularly. In
these areas, every rock and log examined housed
P pubens colonies and the trunk of every tree ex-
amined had broad trails of P pubens foragers go-
ing up and down. Nest-bound foragers were
bloated with liquid, no doubt honeydew collected
from tended Hemiptera. In Mar 2006, we ob-
served much lower densities of P pubens at Cru-
zan Garden than in Nov 2005, despite a lack of
control efforts, suggesting that this species may
already be in decline in some areas.
JW spoke about the outbreak of ants with res-
idents at several other sites. At the Frangipani
housing development (site Y in Mon Bijou), a res-
ident said that the ants had arrived in early 2005.
Along the perimeter of his house were piles of
ants killed by a white powder pesticide, along

with much-reduced trails of live P. pubens. In Her-
mitage (site Z), near the western edge of the main
population, we saw large numbers of P pubens on
a variety of fruit trees. Residents reported that
the ants appeared around Dec 2005 and that their
coconut plantation produced no coconuts this year
because high densities of ant-tended Hemiptera
covered the flowers and young fruit and caused
them to drop prematurely. At Mount Victory
Camp, an ecotourism resort in western St. Croix
(adjacent to site P), the owner reported that P pu-
bens first became evident around Dec 2005. In the
northeast corner of Little Fountain (site A), at the
northern edge of the main population, local resi-
dents first noted P pubens in Mar 2006, and the
ants were found only at the southern edge of their
property, in the direction of the main population.
In the tree surveys, we collected P. pubens plus
6 ant species we considered exotic to St. Croix
(Monomorium floricola (Jerdon), Paratrechina
longicornis (Latreille), Solenopsis geminata (Fab-
ricius), Solenopsis invicta Buren, Tapinoma mel-
anocephalum (Fabricius), and Technomyrmex dif-
ficilis Forel), and 6 we considered to be native
(Brachymyrmex heeri Forel, Crematogaster cri-
nosa Mayr, Linepithema iniquum (Mayr), Mono-
morium ebeninum Forel, Pheidole moerens
Wheeler, and Solenopsis corticalis Forel). At the
12 P pubens-occupied sites (10 trees surveyed per
site), we collected P pubens at 60 trees and made
34 additional arboreal-foraging ant records (17
records of 3 exotic species and 17 of 3 native spe-
cies; Table 1). At the 12 sites without P pubens
present, we made 75 arboreal-foraging ant spe-
cies records (37 records of 5 exotic species and 38
of 5 native species; Table 2). Thus, there were sig-
nificantly fewer records of other arboreal-foraging
ants on trees in P pubens-occupied sites versus in
sites without P pubens present (x2 = 15.4, df = 1,
P < 0.001).
The degree of dominance of P pubens varied
greatly among the P. pubens-occupied sites. At the
6 sites where P pubens occurred on 6 or more of
the 10 trees, there were only 2 other arboreal-for-
aging ant records (1 native and 1 exotic; Table 1).
In contrast, at the 6 P pubens-occupied sites
where P. pubens occurred on 5 or fewer of the 10
trees, there were 32 arboreal-foraging ant records
of species other than P pubens. Thus, there were
significantly fewer records of other arboreal-for-
aging ants at the 6 sites where P pubens were
common in the trees than at the 6 sites where P.
pubens was present, but not common in the trees
(X2 = 26.5, df= 1, P < 0.001).


We documented 3 geographically discrete pop-
ulations of P. pubens on St. Croix: a main popula-
tion occupying ~9 km2 and 2 smaller populations
occupying <1 km2 each. The chronology of com-

Florida Entomologist 91(3)



Exotic species B P T J E K A N D I M Q Total

Paratrechina pubens 10 10 10 9 8 6 5 2 60
Technomyrmex difficilis 2 10 12
Monomorium floricola 1 3 4
Paratrechina longicornis 1 1
Native species
Linepithema iniquum 5 7 12
Crematogaster crinosa 3 3
Solenopsis corticalis 1 1 2

plaints from residents and the current distribu-
tion of P pubens suggest this species is an exotic
that first arrived in the Calquohoun area circa
late 2001 or early 2002. If the main population of
P pubens on St. Croix originated at the center of
its current range in early 2002, then to reach pe-
ripheral areas 2.0-2.5 km from the center (e.g.,
sites 7 and A) by early 2006, it must have spread
at an average linear rate of 0.5-0.6 km per year.
The smaller populations of P pubens in western
and southern St. Croix may be satellite popula-
tions founded by propagules originating from the
main population as recently as 2005. Cruzan Gar-
dens, a botanical garden and nursery located near
the center of the main population, may be acting
as a hub for dispersal to other parts of St. Croix by
P pubens colonies living in potted plants.
Currently, P pubens appears to be principally a
house and garden pest on St. Croix. However, when
occurring at extremely high densities, this species
has the potential be a significant agricultural pest

by enhancing populations of the phloem-feeding
Hemiptera it tends. Hemiptera cause crop damage
both through sapping plants of nutrients and by
increasing the occurrence of diseases, including vi-
ral and fungal infections. In addition to its nega-
tive impact on arboreal-foraging ants, P pubens
may also have negative impacts on other animals,
both invertebrates and small vertebrates.

Native Versus Exotic Range

Marlatt (1928) wrote that P pubens "is believed
to be a native of Brazil, but now occurs quite abun-
dantly in Cuba and other West Indian Islands."
Records of P pubens from South and Central Amer-
ica often come from nature preserves. For example,
Leponce et al. (2004) and Theunis et al. (2005) re-
corded P pubens in Rio Pilcomayo National Park,
Argentina. Fisher et al. (1990) and Fisher (1992) re-
ported P pubens inhabiting myrmecophytic orchids
in Barro Colorado National Monument, Panama.



Exotic species C F G H L O R S U V W X Total

Paratrechina longicornis 2 1 9 12
Solenopsis geminata 2 1 3 1 2 9
Solenopsis invicta 5 5
Monomorium floricola 2 1 4 1 8
Tapinoma melanocephalum 1 2 3
Native species
Monomorium ebeninum 2 8 3 3 1 8 25
Solenopsis corticalis 1 2 1 2 3 9
Linepithema iniquum 1 1 2
Pheidole moerens 1 1
Brachymyrmex heeri 1 1

September 2008

Wetterer & Keularts: Hairy Crazy Ant on St. Croix

This pattern suggests that P pubens is native to
South and Central America. Unfortunately, the
taxonomic status of P pubens and other closely
related forms remains uncertain, and records of
P pubens from South and Central America could
represent a different species.
In the West Indies and Florida, P pubens has
been reported primarily in open, disturbed areas,
a habitat preference often indicative of exotic spe-
cies. For example, Smith (in Forel 1893) collected
P pubens from 5 locales on the island of St. Vin-
cent, four at the seashore and 1 in a sugar cane
field, writing that this ant "appears to be confined
to the seashore, or to open land not far from the
sea." Klotz et al. (1995) listed 3 records from Flor-
ida: in a Boca Raton home, in a Miami hospital,
and in a commercial building near Homestead.
Deyrup et al. (2000) reported that P pubens "is
abundant on the campus of the University of
Miami ... foraging on sidewalks and running up
and down tree trunks."
Warner and Scheffrahn (2004) recently pro-
posed the common name "Caribbean crazy ant"
for P pubens, but this name is misleading and in-
appropriate because P. pubens appears to be ex-
otic to the Caribbean islands. Therefore, we prefer
the common name used by Wetterer (2007), "hairy
crazy ant," based on its Latin name pubens which
means pubescent, i.e., covered with short soft
hairs, a prominent characteristic of this species.
It will be interesting to track the progress of
the P. pubens on St. Croix to see whether it con-
tinues to spread and becomes a major long-term
economic and ecological pest. Alternatively, P pu-
bens may decline on St. Croix and become only a
minor pest there, or disappear completely as hap-
pened in Bermuda.

We thank A. Wetterer and M. Wetterer for comments
on this manuscript; S. Cover and R. Snelling for ant
identifications; B. Wilson for information on the infesta-
tion at Mt. Victory Camp, W. O'Brien, S. Michael, and A.
Vuppuluri for GIS mapping assistance; the National
Science Foundation and Florida Atlantic University for
financial support.


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Myrmecol. Nachr. 8: 219-224.
WETTERER, J. K. 2007. Paratrechina pubens (Forel,
1893) (Hymenoptera: Formicidae), a candidate for
the plague ant of 19th century Bermuda. Myrmecol.
Nachr. 10: 39-40.

Florida Entomologist 91(3)


Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive, Jupiter, FL 33458

Technomyrmex difficilis Forel is an Old World ant often misidentified as the white-footed
ant, Technomyrmex albipes (Smith). The earliest New World records of T difficilis are from
Miami-Dade County, Florida, collected beginning in 1986. Since then, it has been found in
at least 22 Florida counties. Here, I report T difficilis from 5 West Indian islands: Antigua,
Nevis, Puerto Rico, St. Croix, and St. Thomas. Colonies were widespread only on St. Croix.
It is probable that over the next few years T difficilis will become increasingly important as
a pest in Florida and the West Indies.

Key Words: exotic species, Technomyrmex difficilis, pest ants, West Indies

El Technomyrmex difficilis Forel es una hormiga del Mundo Antiguo que a menudo es mal
identificada como la hormiga de patas blancas, Technomyrmex albipes (Smith). Los registros
mas viejos de T difficilis son del condado de Miami-Dade, Florida, recolectadas en el princi-
pio de 1986. Desde entonces, la hormiga ha sido encontrada en por lo menos 22 condados de
la Florida. Aqui, informo de la presencia de T difficilis en 5 islas del Caribe: Antigua, Nevis,
Puerto Rico, St. Croix y St. Thomas. Las colonies solamente fueron muy exparcidas en St.
Croix. Es probable que la importancia de T difficilis como plaga va a aumentar durante los
proximos anos en la Florida y el Caribe.

The white-footed ant, Technomyrmex albipes
(Smith), has long been considered a pest in many
parts of the world. Recently, however, Bolton
(2007) reported that T albipes is part of an Old
World species group that includes 43 species, 4 of
which have broad distributions: T albipes, Tech-
nomyrmex vitiensis Mann, Technomyrmex pal-
lipes (Smith), and Technomyrmex difficilis Forel.
Bolton (2007) determined that many published
reports of T albipes were misidentifications. For
example, T albipes records in Deyrup (1991) from
California and Wetterer (1997) from Hawaii were
actually T vitiensis; T albipes records in Wetterer
et al. (2006) from Madeira were actually T pal-
lipes; and T albipes records from Florida in Wet-
terer & Wetterer (2003) and Wetterer et al. (2007)
were actually T difficilis. Bolton (2007) concluded
that all published records of T albipes from the
New World were misidentifications and that all
Technomyrmex specimens he examined from Flor-
ida were actually T difficilis.
The earliest New World records of T difficilis are
from Miami-Dade County, Florida, collected in 1986
(Deyrup 1991; misidentified as T albipes). Deyrup
et al. (2000) noted that T difficilis (misidentified as
T albipes) was spreading rapidly in Florida. By
2005, T difficilis (misidentified as T albipes) was
known from 22 counties in Florida, as well as from
Georgia, South Carolina, and Louisiana (Warner &
Scheffrahn 2004; Warner et al. 2005). Here, I report
on T difficilis on 5 West Indian islands.


The ants collected were determined to be T dif-
ficilis as distinguished from T albipes according
to Bolton (2007) by "the presence of setae on the
dorsum of the head behind the level of the poste-
rior margin of the eye (never developed in albipes)
and by having the promesonotum somewhat
longer and more slender, DTI 127-135 (as opposed
to DTI 110-124 in albipes)" (DTI = Dorsal Tho-
racic Index = length from anterior pronotal mar-
gin to metanotal groove x 100, divided by pronotal


In 2005-2007, I collected T difficilis on 5 is-
lands (26 sites; geocoordinates in N & W): Pu-
erto Rico (3 sites: Old San Juan; W end; 18.468,
66.121; 16-X-2005. Isla Verde; airport; 18.439,
66.002; 16-V-2006. San Juan; park; 18.408,
66.073; 15-X-2005), St. Thomas (1 site: UVI;
parking lot; 18.344, 64.974; 7-XI-2005), St. Croix
(18 sites: Roberts Hill; Buccaneer Hotel; 17.753,
64.679; 12-III-2006. The Glynn; 0.5 km N of Rte
72; 17.751, 64.771; 5-III-2006. Mount Victory; 0.5
km NE of camp; 17.750, 64.867; 7-III-2006. Pleas-
ant Vale; Rte 58; 17.748, 64.865; 7-III-2006.
Mount Victory; Rte 58; 17.746, 64.866; 7-III-2006.
North Hall; Rte 58; 17.746, 64.876; 30-X-2005.
William; Rte 58; 17.738, 64.891; 30-X-2005. Grove

September 2008

Wetterer: Technomyrmex difficilis in the West Indies

Place; 0.6 km N of Rte 76; 17.735, 64.822; 10-III-
2006. Upper Love; Holy Cross Church; 17.733,
64.807; 4-XI-2005. Jolly Hill; Rte 76; 17.732,
64.861; 3-III-2006. La Reine; Rte 70; 17.729,
64.774; 4-III-2006. Sunny Isle; shopping center;
17.729, 64.749; 12-III-2006. Prosperity; W end
Mahogany Rd; 17.721, 64.885; 30-X-2005. St.
George; Botanical Garden; 17.716, 64.831; 1-XI-
2005. Bethlehem Middle Works; Rte 64; 17.710,
64.790; 12-III-2006. Hesselberg; Cottages by the
Sea; 17.701, 64.886; 3-XI-2005. Hesselberg; S end
Shore Dr; 17.692, 64.892; 3-XI-2005. Camporico;
Rte 66; 17.689, 64.862; 11-III-2006), Nevis (1 site:
Stuart's; Four Seasons; 17.159, 62.623; 18-V-
2007), and Antigua (3 sites: Parham; by dock;
17.113, 61.763; 26-V-2007. North Sound; pasture
tree; 17.098, 61.783; 24-V-2007. Seatons; Sting
Ray City; 17.097, 61.721; 23-V-2007). I deposited
vouchers at Harvard University's Museum of
Comparative Zoology.
On all 5 islands, I found enormous colonies of
T difficilis, but colonies were widespread only on
St. Croix, where I found this ant at 18 diverse
sites. On St. Croix, I most often found colonies in
planted trees growing in urban and residential
areas, e.g., around the grounds of Sunny Isle
shopping center, Holy Cross Church, the Bucca-
neer Hotel, and Cottages by the Sea resort. The
ants, however, also occurred at high densities in
some forested areas, such as along Route 58
(Creque Dam Road), particularly in Mt. Victory
and Pleasant Vale, where I found T difficilis colo-
nies in virtually every tree along extended
stretches of the road.
On the other 4 islands, I encountered colonies
of T difficilis at few locales. In Puerto Rico, I
found T difficilis at 3 sites: in a large tree in a
Park Luis Munoz Marnn in San Juan, in a tree in
Old San Juan, and in several trees around the
parking lot of Luis Munoz Marin International
Airport. On St. Thomas, I found swarms of T dif-
ficilis under a tree by a parking lot on the Univer-
sity of the Virgin Islands campus. On Nevis, I col-
lected T difficilis in trees and several buildings at
the Four Seasons Hotel. On Antigua, I found T
difficilis at 3 sites: nesting in branches of sea
grapes growing by the Parham dock, in branches
throughout an enormous tree in the pasture
southeast of Sir Vivian Richards Stadium, and in
several trees on the grounds of Sting Ray City.


In the 20 years since it was first collected in
Florida, Technomyrmex difficilis has quickly ex-
panded its range across a large portion of the
state, and is now spreading across the West In-
dies. Torres et al. (2001) reported the earliest
record of Technomyrmex (presumably T difficilis)
from the West Indies, males collected in 1996 and
1997 in Guanica, Puerto Rico. Warner & Schef-

frahn (2004) reported Technomyrmex collected in
2003 on Grand Cayman Island. In 2005-2007, I
collected T difficilis on 5 West Indian islands (An-
tigua, Nevis, Puerto Rico, St. Croix, and St. Tho-
mas), finding the most widespread populations on
St. Croix. Thus, the earliest and the most exten-
sive records in the West Indies come from Puerto
Rico and St. Croix, respectively. Both islands have
much commercial traffic with Florida, from which
the West Indian populations ofT difficilis proba-
bly originated.
In Florida and the West Indies, I have most of-
ten found T difficilis in disturbed urban and res-
idential areas. For example, over the past few
years, T difficilis has become a dominant arboreal
ant on our FAU campus (Jupiter, Florida) and at
local beaches (Juno Beach, Florida). I have even
experienced outbreaks of this ant in my office on
campus. Unfortunately, T difficilis also appears
to be able to invade forest habitats, where it can
more readily impact native species. It is likely
that over the next few years, T difficilis will be-
come increasingly important as a pest in Florida
and the West Indies.
Technomyrmex difficilis currently has no com-
mon name. Just as common names are useful in
distinguishing Solenopsis invicta (the red im-
ported fire ant) from Solenopsis geminata (the
tropical fire ant), a common name for T difficilis
is needed to help people appreciate that this spe-
cies spreading through our region is not the
white-footed ant, T albipes, as mistakenly
thought until recently. The common name for T
albipes, the "white-footed" ant, is a direct transla-
tion of the Latin "albipes." The species name diffi-
cilis (Latin for "difficult") apparently comes from
the taxonomic difficulties it posed. Based on spec-
imens from Madagascar, Forel (1892) described
this taxon as Technomyrmex mayri race difficilis,
writing (in French) that it was "difficult to define,"
with many characters intermediate between
Technomyrmex mayri Forel (known only from
Madagascar) and the widespread T albipes. Forel
(1892) wrote: "I do not know how otherwise to de-
fine this embarrassing form, which is perhaps a
hybrid." The name difficilis also fits well with the
reputation of this ant has acquired in Florida. For
example, Warner et al. (2005) wrote that T diffici-
lis "is an extremely difficult pest to control due to
the large size of its colonies." I propose "techno
ant" as a common name for members of the genus
Technomyrmex, a name derived from Tekhne, the
Greek goddess of art and craft, and from murmex,
Greek for ant. Further, I propose "difficult techno"
ant as a common name for T difficilis. This
straightforward common name is unlikely to be
confused with any other.
Although all Technomyrmex specimens from
Florida and the West Indies that Bolton (2007)
and I have examined are T difficilis, it would be
useful to evaluate all available Technomyrmex

specimens from this region to confirm their spe-
cies identity. It would be unfortunate if the incor-
rect assumption that all Technomyrmex in Florida
are T albipes were replaced with another incor-
rect assumption.


I thank A. Wetterer, M. Wetterer, and J. Warner for
comments on this manuscript; B. Bolton for ant identi-
fications; the National Science Foundation and Florida
Atlantic University for financial support.


BOLTON, B. 2007. Taxonomy of the dolichoderine ant ge-
nus Technomyrmex Mayr (Hymenoptera: Formi-
cidae) based on the worker cast. Contrib. American
Entomol. Inst. 35: 1-150.
DEYRUP, M. 1991. Technomyrmex albipes, a new exotic
ant in Florida (Hymenoptera: Formicidae). Florida
Entomol. 74: 147-148.
DEYRUP, M., L. DAVIS, AND S. COVER 2000.Exotic ants in
Florida. Trans. American Entomol. Soc. 126:293-326.
FOREL, A. 1892. Les Formicides (concl.), pp. 232-280 In
A. Grandidier [ed.], Histoire physique, naturelle et
politique de Madagascar. Volume XX. Histoire na-

September 2008

turelle des Hym6noptbres. Deuxieme parties. Supple-
ment au 28e fascicule. Hachette et Cie, Paris.
Seasonal and nocturnal periodities in ant nuptial
flight in the tropics (Hymenoptera: Formicidae). So-
ciobiol. 37: 601-626.
WARNER, J., AND R. H. SCHEFFRAHN. 2004. Feeding
preferences of white-footed ants, Technomyrmex al-
bipes (Hymenoptera: Formicidae), to selected liq-
uids. Florida Agric. Exper. Sta. Series R-10161.
White-footed ant, Technomyrmex albipes (Fr. Smith)
(Insecta: Hymenoptera: Formicidae: Dolichoderi-
nae). Univ. Florida IFAS Extens. Doc. EENY-273.
WETTERER, J. K. 1997. Ants on Cecropia in Hawaii. Bio-
tropica 29: 128-132.
2006. Long-term impact of exotic ants on the native
ants of Madeira. Ecol. Entomol. 31: 358-368.
WETTERER, J. K., AND A. L. WETTERER. 2003. Ants (Hy-
menoptera: Formicidae) on non-native Neotropical
ant-acacias (Fabales: Fabaceae) in Florida. Florida
Entomol. 86: 460-463.
AND S. FITCHETT. 2007. Predaceous ants, beach re-
plenishment, and nest placement by sea turtles. En-
viron. Entomol. 36: 1084-1091.

Florida Entomologist 91(3)

Gagn6 & Hibbard: New Species of Cecidomyia


1Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA,
c/o U. S. National Museum NHB 168, P.O. Box 37012, Washington, DC 20013-7012, USA
e-mail: raymond.gagne@ars.usda.gov

2Bureau of Methods Development and Biological Control, Division of Plant Industry,
Florida Department of Agriculture and Consumer Services, 3513 S, US 1, Ft. Pierce, FL 34945-3045
e-mail: hibbark@doacs.state.fl.us

Cecidomyia lamellata Gagn6, new species, is described from adults of both sexes, pupae,
and larvae taken from resin in branchlet swellings of baldcypress, Taxodium distichum (L.)
Rich. (Taxodiaceae), in Fort Pierce, Florida. Some anatomical characters of the new species
depart from the previously known limits of the genus. Baldcypress is the first known non-
Pinaceae host of Cecidomyia.

Key Words: gall midges, Taxodium distichum, Taxodiaceae, Florida, stem galls

Se describe una nueva especie, Cecidomyia lamellata Gagn6, de adults de ambos sexos, pu-
pas y larvas recolectados de la resina en ramas engrosadas de cipr6s, Taxodium distichum
(L.) Rich. (Taxodiaceae) en Fort Pierce, Florida. Algunos caracteres anat6micos de la especie
nueva son distintos de los limits conocidos para este g6nero. El cipr6s es el primero hospe-
dero conocido de Cecidomyia que no pertece a la familiar Pinaceae.

Baldcypress, Taxodium distichum (Taxodia-
ceae), is a widespread deciduous conifer of low-
land southeastern United States and northeast-
ern Mexico. Branchlet swellings 2 to 3 cm long
and 1 to 2 cm in diameter were discovered on
baldcypress in Fort Pierce, Florida by one of us
(KLH). The exterior of the swellings was similar
in texture to the rest of the branchlets, grayish-
brown and scaly, except that a part of the surface
was concave and led into a pool of resin in which
gregarious cecidomyiid larvae were suspended.
The larvae belong to an undescribed species of
Cecidomyia, one that forces a redefinition of the
genus. Cecidomyia is a mostly Holarctic genus of
18 previously known species that feed on resin of
conifers (Gagne 2004). The last pair of abdominal
larval spiracles are greatly enlarged and posteri-
orly displaced, allowing the larvae to live im-
mersed in the resin. The genus was previously
known only from Pinaceae, mainly Pinus but also
Abies, Picea, and Pseudotsuga, so this new record
from Taxodium extends the host boundary of Cec-
idomyia. Among the more conspicuous and
unique differences between the new species and
previously described congeners are the foreshort-
ened adult antennae, the bowed instead of bent
tarsal claws, the short empodia, the broad and
completely setulose gonostylus, and the lamellate
terminal segment of the larva. The new species is

described and illustrated here and placed in con-
text with its congeners and most similar genera.


Some full-grown larvae were removed from the
resin in swellings and placed directly into 70%
ethanol. Other swellings were saved intact over
the winter atop damp peat moss in a plastic con-
tainer to obtain pupae and adults in spring. These
also were saved in 70% ethanol. Samples were
mounted on microscope slides according to the
method outlined in Gagn6 (1989). Terminology for
adult morphology follows usage in McAlpine et al.
(1981) and follows Gagn6 (1989) for larval mor-
phology. Specimens and seasonal information of
the new species were obtained by KLH. The taxo-
nomic investigation in this paper was the respon-
sibility of RJG.

Cecidomyia lamellata Gagn6, New Species

Adult. Head: Eyes large, about 10 facets long at
vertex, barely separated to partly connate; facets
circular, adjacent throughout except up to 1/2 di-
ameter apart near midheight and at vertex. Frons
with 6-9 setae per side. Labella hemispherical in
frontal view, each with 5-6 blunt-tipped setae. Pal-
pus 4-segmented. Antenna: male flagellomeres

Florida Entomologist 91(3)

(Fig. 1) foreshortened, internode not developed on
proximal flagellomeres and barely so beyond fifth
flagellomere; necks shorter than wide; circumfila
with short loops not attaining next distal circum-
filum; setulae sparse. Female flagellomeres (Fig.
2) cylindrical with short necks, some circumfilar
segments with loops on distal part of node.
Thorax: Anepisternum with a few scales dor-
sally; anepimeron with 26-34 setae; pleura other-
wise bare. Wing length, male 1.9-2.2 mm (n = 5),
female 2.1-2.5 mm (n = 4). Acropods (Figs. 3-4):
claws falciform, untoothed; empodia less than 1/2
length of claws; pulvilli about 1/4 length of claws.
Male abdomen: First through seventh tergites
entire, rectangular, each with mostly single, unin-
terrupted, posterior row of setae, a few lateral se-
tae, a pair of trichoid sensilla on anterior margin,
and otherwise evenly covered with scales. Eighth
tergite rectangular, smaller than seventh, with
anterior pair of trichoid sensilla and bare or with
a few setae posteriorly. Second through seventh
sternites with single row of posterior setae, addi-
tional setae along lateral margins and medially at
midlength, a pair of closely set anterior trichoid
papillae, and elsewhere covered with long, thin
scales. Eighth sternite with posterolateral setae,
a pair of widely set anterior trichoid sensilla, and
scales covering distal third. Genitalia (Figs. 5-6):
cerci elliptical; hypoproct wider than aedeagus,
deeply bilobed, each lobe with 1 apical and 2-3
subapical setae, entire surface setulose; aedeagus
cylindrical, slightly recurved dorsally, rounded
apically, subequal in length to hypoproct and
cerci, with several sensilla; gonocoxite cylindrical,
unlobed; gonostylus terete, wide throughout
length to apical tooth, with setae evenly dis-
persed, longest laterally, entire surface setulose.
Female abdomen: First through seventh terg-
ites as for male. Eighth tergite rectangular,
smaller than seventh, with anterior pair of tri-
choid sensilla and covered with setiform scales.
Second through seventh sternites as for male.
Eighth sternite weakly sclerotized with postero-
lateral setae, a pair of widely set anterior trichoid
sensilla, and scales covering posterior third. Ovi-
positor (Figs. 7-8) elongate-protrusible; extended
eighth segment (beyond tergite and sternite) with
evenly scattered, short setae; ninth segment
about twice length of seventh tergite, with numer-
ous short setae throughout; tenth tergite asetose;
cerci (Fig. 8) discrete with setae dispersed over en-
tire surface, each with pair of apical peglike setae.
Pupa (Fig. 9). Cephalic sclerite with 2 papillae
atop a protuberance on each side, one papilla with
short, one with long seta. Base of antenna with
short, wide, pigmented ridge ventrally. Frons
smooth with ventrolateral triplet of papillae, 2
with setae, and 8 frontal papillae, 4 with setae.
Prothoracic spiracle fairly short. Abdomen uni-
formly covered with short verrucae, pointed on
dorsum and venter, rounded on pleura.

Larva. Third instar (Figs. 10-12): Body elon-
gate, spindleform, without conspicuous lobes ex-
cept posteriorly. Integument smooth except for
anterior horizontal rows of spinose verrucae on
second thoracic segment through seventh abdom-
inal segment of both dorsal and ventral surfaces
and additionally on eighth dorsal abdominal seg-
ment and surrounding anus. Spatula (Fig. 10)
with short, convex anterior tooth and long shaft
thinning gradually from both ends toward
midlength. Papillae typical of supertribe Cecid-
omyiidi (Gagne 1989), all present and setose or
not setose as in other Cecidomyia spp. (Gagne
1978), the setae where present fairly short except
for the long dorsal and pleural setae on the eighth
segment (Fig. 12). Eighth segment exceeding ter-
minal (ninth) segment in length, hind spiracles of
eighth abdominal segment greatly enlarged, set
at end of posteriorly directed lobes, tapering to
closed apex, opening laterally. Terminal segment
(Fig. 11) modified posteriorly as a dorsoventrally
flattened lamella ending in 2 elongate append-
ages each subtending a corniform terminal pa-
pilla; 2 of remaining terminal papillae without se-
tae, situated dorsally at base of lamella, 2 with
moderately long setae situated laterally at
midlength, the remaining 2 with short setae, as
wide as long, situated ventrally on lamella.
Holotype: Male, from stem swelling of Taxo-
dium distichum, Ft. Pierce, Florida, collected X-6-
2006, K.L. Hibbard, emerged III-19-2007, depos-
ited in the National Museum of Natural History
in Washington, DC (USNM). Other material ex-
amined: 7 males, 6 females, 6 pupae, 12 larvae,
same pertinent data as holotype except adults
emerged III-5 to 23-2007. All deposited in USNM.
Etymology. The specific name is an adjective
that refers to the lamellate terminal larval abdom-
inal segment, a structure unique to this species.
Biological Note. Larvae of this species spend
their lives immersed in resin. The full-grown
third instars spin a cylindrical cocoon within the
resin mass, effectively separating them from the
liquid. They then turn end to end inside the co-
coon with their heads facing the exterior and
eventually pupate in that position. When adults
are fully developed inside the pupae, they force
the still closed pupal skin through the end of the
cocoon and any dried resin and become lodged
part way out of the cocoon. Adults then emerge
through sutures in the pupal thorax.
Remarks. The chief distinguishing characters
of Cecidomyia are as follows: Eyes are more or
less confluent at vertex, their facets either hexag-
onal and closely juxtaposed, or circular and not
touching. The occipital process is absent. The an-
tenna has 12 flagellomeres, the first and second
not connate; male flagellomeres are binodal and
tricircumfilar, except foreshortened and cylindri-
cal in C. lamellata; female flagellomeres are cylin-
drical with short necks. The palpus is usually

September 2008

Gagn6 & Hibbard: New Species of Cecidomyia




-3~-w ~
-v ~


Figs. 1-8. Cecidomyia lamellata n. sp. 1, Male third flagellomere (ventral). 2, Female third flagellomere (ventral).
3, Acropod (lateral). 4, Same (dorsal). 5, Male genitalia (dorsal). 6, Hypoproct and aedeagus (dorsal). 7, Female ab-
domen, eighth segment to cerci (lateral). 8, Detail of female cerci (lateral).

four-segmented, three-segmented in one species.
The R, wingvein is curved distally to join C behind
the wing apex, the costa is broken at its juncture
with Rs, the M 3+4 fold is present, and the cubitus
is forked. The tarsal claws are simple, curved be-
yond midlength or (in C. lamellata only) bowed,
empodia are slightly longer than claws or (in

C. lamellata only) but half as long, and pulvilli
are diminutive, about 1/4 length of the claws.
Male and female first through seventh tergites
have 1 to 2 rows of posterior setae, several lateral
setae, and a pair of anterior trichoid sensilla. The
gonostylus is setulose basally, asetulose and
ridged apically, or, in C. lamellata, completely set-

Florida Entomologist 91(3)



d e

Figs. 9-12. Cecidomyia lamellata n. sp. 9, Pupal head (dorsal, but cervical sclerite anterior). 10, Larval head and
cervical and first thoracic segments (ventral). 11, Larval eighth and terminal segments (ventral): a = anus; b =
lamellate terminal segment; c = hind spiracle. 12, Larval eighth segment (dorsal): d = pair of pleural papillae; e =
dorsal papilla.

ulose. The ovipositor is usually short-protrusible, Cecidomyia lamellata is a remarkable addition
but long-protrusible in C. lamellata. Female cerci to Cecidomyia because of several anomalies.
are discrete, evenly covered with setae, 1 apicolat- These are the foreshortened adult antennae, the
eral pair thick and blunt-tipped. The pupal abdo- bowed instead of bent tarsal claws, the short em-
men is covered both ventrally and dorsally with podia, the broad and completely setulose gonosty-
uniformly short, acute verrucae, and laterally lus, the longer than usual ovipositor, and on the
with rounded verrucae. Larvae have elongate larva, the lamellate terminal segment and the
cephalic apodemes and greatly enlarged and pos- merged 4 apical points of the hind spiracle. Bald-
teriorly displaced hind spiracles. cypress further extends the known host range of

September 2008

Gagn6 & Hibbard: New Species of Cecidomyia

Cecidomyia beyond Pinaceae from which the ge-
nus was previously known.
In the key to Nearctic genera in Gagne (1981),
the bowed instead of bent claws of C. lamellata
will prevent this species from running to the
same place as other species of Cecidomyia. It will
instead run to couplet 126 with Sequoiomyia and
Taxodiomyia. Interestingly, both these genera are
restricted to baldcypress. Unlike either Sequoi-
omyia and Taxodiomyia, the first and second
flagellomeres are not connate in Cecidomyia. Se-
quoiomyia is further distinguished from Cecid-
omyia lamellata by its fully connate eyes with all
facets hexagonal and the presence of at least 2
complete rows of posterior setae on the abdominal
terga. In C. lamellata the eyes are not fully con-
nate and the facets are circular and not all adja-
cent, and the posterior setae of the abdominal
terga form only a single row. Taxodiomyia female
cerci are fused instead of discrete and the gono-
stylus is widest basally and is asetulose and lined
with ridges apically. In C. lamellata the female
has discrete cerci and the male gonostylus is
broad throughout its length, completely setulose
and without ridges. In addition, Sequoiomyia and
Taxodiomyia larvae live, respectively, in seeds
and complex leaf galls and their larvae are ovoid,
while Cecidomyia larvae live in resin and are
spindleform. The short empodia of these 3 genera
is a departure from the general rule that conifer
feeding gall midges have empodia that are appre-
ciably longer than the tarsal claws. It is a striking
coincidence that the short empodia and bowed

tarsal claws occur on all 3 genera of gall midges
on baldcypress.


We are grateful to Nit Malikul, Systematic Entomology
Laboratory (SEL), for preparing the microscopic slides; Di-
ana Marques for arranging the drawings and photos onto
plates; and, for their helpful comments on drafts of the
manuscript: David A. Nickle and Allen L. Norrbom, Sys-
tematic Entomology Laboratory; Keith M. Harris, Ripley,
Surrey, UK; and two anonymous reviewers.


GAGNE, R. J. 1978. A systematic analysis of the pine
pitch midges, Cecidomyia spp. (Diptera: Cecidomyi-
idae). U.S. Department of Agriculture Tech. Bull.
1575: 1-18.
GAGNE, R. J. 1981. Cecidomyiidae, pp. 257-292 In J. F.
McAlpine, B. V. Peterson, G. E. Shewell, H. J. Tes-
key, J. R. Vockeroth, and D. M. Wood [eds.], Manual
of Nearctic Diptera. Vol. 1. Research Branch, Agri-
culture. Canada Monograph No. 27.
GAGNE, R. J. 1989. The Plant-Feeding Gall Midges of
North America. Cornell University Press, Ithaca,
New York. xi and 356 pp. and 4 p1s.
GAGNE, R. J. J. 2004. A catalog of the Cecidomyiidae
(Diptera) of the world. Mem. Entomol. Soc. Washing-
ton 23: 1-408.
1981. Manual of Nearctic Diptera. Vol. 1. Research
Branch, Agriculture. Canada Monograph No 27. vi +
674 pp.

Florida Entomologist 91(3)

September 2008


1MacArthurAgro-Ecology Research Center, 300 Buck Island Ranch Rd., Lake Placid, FL 33852

'Entomology & Nematology Department, University of Florida, Gainesville, FL 33852


Field experiments were conducted during the 2006-2007 growing season to determine the ef-
fect of the predatory mite, Neoseiulus californicus (McGregor) on arthropod community
structure when released as a biological control agent for the twospotted spider mite, Tet-
ranychus urticae Koch, in north Florida strawberries (Fragaria x ananassa Duchesne). Re-
leases of N. californicus were conducted at approximately 1-month intervals from Dec 2006
to Feb 2007 to compare effects of predator release times on arthropod community structure.
Evaluations of community structure were conducted 3 times during the growing season. The
Shannon-Weaver index of diversity was used to quantify differences among release and non-
release plots. Our results indicate that the release of N. californicus does not affect the ar-
thropod diversity in the strawberry system studied. The generalist feeding behavior of N.
californicus, coupled with a high level of richness and diversity in the strawberry ecosystem,
may diffuse the measurable effect of N. californicus releases on the arthropod community
structure. This makes N. californicus a desirable biological control agent for management of
twospotted spider mite in strawberries while preserving arthropod diversity.

Key Words: biological control, arthropod assemblage, strawberry ecosystem, twospotted spi-
der mite, pest management


Experimentos de campo fueron establecidos para determinar el efecto del acaro predador,
Neoseiulus californicus (McGregor), en la estructura de las comunidad de artr6podos cuando
es liberado como control biol6gico de la aranita roja, Tetranychus urticae Koch, en Fresas
(Fragaria x ananassa Duchesne) en el norte de la Florida. Liberaciones de N. californicus se
realizaron en intervalos de un mes desde Diciembre 2006 hasta Febrero 2007, para compa-
rar los efectos del tiempo de liberaci6n de los predadores en la estructura de la comunidad
de artr6podos presents. Evaluaciones de la estructura de las comunidades se realize tres
veces durante la temporada de producci6n. El indice de Shannon-Weaver fue usado para
cuantificar las diferencias entire los lotes donde se hicieron las liberaciones y los lotes de con-
trol. Nuestros resultados demuestran que la liberaci6n de N. californicus no afecta significa-
tivamente la diversidad de artr6podos en el sistema de fresas estudiado. El comportamiento
alimentario generalista de N. californicus y el alto nivel de riqueza y diversidad del sistema
de producci6n de fresas, puede dispersar el efecto cuantificable de N. californicus en la es-
tructura de la comunidad. Esta caracteristica hace que N. californicus sea un controlador
biol6gico important para el manejo de la aranita roja en fresas, al mismo tiempo que se con-
serva la diversidad del sistema.

Translation provided by the authors.

Twospotted spider mite, Tetranychus urticae
Koch (TSSM), is a key pest of strawberries
(Fragaria x ananassa Duchesne) in north Florida.
High populations of TSSM can reduce foliar and
floral development thereby decreasing the quality
and quantity of mature fruit (Rhodes et al. 2006).
Twospotted spider mite populations have become
resistant to most acaricides due their short life cy-
cle and high fecundity (Huffaker et al. 1969; Wil-
liams 2000; Cross et al. 2001; Stumpf & Nauen
2001; Sato et al. 2004). Outbreaks of TSSM have
become more frequent over the last few decades

due to increased use of pesticides in modern cul-
tural practices. As a result, more growers are uti-
lizing biological control as an alternative to chem-
ical management (Huffaker et al. 1969; Escudero
& Ferragut 2005; Rhodes et al. 2006). However,
little is known about the non-target effects of bio-
logical control releases on beneficial arthropods in
the strawberry ecosystem.
Phytoseiid mites have been found to be highly
effective predators in controlling TSSM (Zhi-Qui-
ang & Sanderson 1995). Two of the most com-
monly used phytoseiids are Phytoseiulus persimi-

Fraulo et a.: N. californicus and Arthropod Community Structure

lis Athias-Henriot and Neoseiulus californicus
(McGregor) (McMurtry & Croft 1997; Cloyd et al.
2006). Oatman et al. (1972) found that although
P persimilis was effective in controlling TSSM, it
is a type I specialist predator of Tetranychus spe-
cies and tends to decimate TSSM populations, al-
tering the arthropod complex (McMurtry & Croft
1997). However, Colfer et al. (2004) found that re-
leases of generalist species of phytoseiid mites
such as N. californicus do not affect the diversity
or abundance of arthropod populations.
As a type II generalist, N. californicus has a
broad diet range that includes not only various ar-
thropods, but also plant sap, honeydew, and pollen
(McMurtry & Croft 1997).Neoseiulus californicus
can adapt to fluctuations in prey populations, pro-
viding stable pest suppression over time (Croft et
al. 1998; Castagnoli et al. 1999; Escudero & Fer-
ragut 2005; Greco et al. 2005). Rhodes et al. (2006)
observed that N. californicus was able to maintain
more consistent control of TSSM populations com-
pared with P persimilis throughout the season in
north Florida strawberry fields. The ability of N.
californicus to survive on a broad array of food
sources contributes to its stability and may miti-
gate its effect on community structure and other
beneficial arthropods (Jones 1976; Powers & Mc-
Sorley 2000; Cross et al. 2001; Rhodes et al. 2006).
There are many natural enemies capable of
suppressing TSSM populations (Oatman et al.
1985). However, their use in biological control has
been limited due to their generalist feeding pref-
erences. Field studies conducted between 1964-
1980 in southern California identified 9 phyto-
seiid mite species and several species of insects
within the families Thripidae, Cecidomyiidae,
Coccinellidae, Staphylinidae, Anthocoridae, Lyga-
edae, Chrysopidae, and Hemerobiidae as natural
enemies of TSSM (Oatman et al. 1985). Rondon et
al. (2004) conducted laboratory studies evaluating
the big-eyed bug, Geocoris punctipes Say, minute
pirate bug, Orius insidiosus (Say), and the pink
spotted lady beetle, Coleomegilla maculata De-
Geer, as predators for TSSM. They found that
while they feed on TSSM, they preferred other
phytophagous insects, thereby limiting their util-
ity as successful biological control agents.
Conserving a robust ecosystem is essential to
sustain the myriad of natural enemies that con-
tribute to a sustainable integrated pest manage-
ment (IPM) program. In this study, field experi-
ments were conducted to determine the effect of
releasing N. californicus as a biological control
agent for TSSM on the arthropod complex in
strawberry fields, and evaluated with the Shan-
non-Weaver index (IT) (Shannon 1948). Our hy-
pothesis was that inundative releases of N. cali-
fornicus to control TSSM in strawberries will not
negatively impact other key natural enemies and
arthropod diversity in the strawberry system. If
so, N. californicus could be released as part of an

IPM or biological control program in north Flor-
ida strawberry while preserving non-target bene-
ficial arthropods.


A preliminary study was conducted during
2005-2006 (Jan to Mar) to assess the accuracy of
sample methods and level of diversity in the
strawberry system. The main study was con-
ducted during the 2006-2007 (Oct to Mar) grow-
ing season to evaluate the effect of predatory re-
leases at 3 phenological periods: foliar, floral, and
fruit development.

Field Preparation

The field experiment was located at the Uni-
versity of Florida Plant Science Research and Ed-
ucation Unit in Citra, Florida (82.17W, 29.410N).
Sixteen research plots of 53.29 m2 with a bare
ground buffer of 11 m between plots were planted
with strawberries, variety 'Festival', during the
first week of Oct 2005 and 2006 in raised beds cov-
ered with 6-mil black plastic mulch. Strawberry
plants were fertilized through the drip irrigation
system once per week with 18.5 kg of ammonium
nitrate (Southern States Cooperative, Inc., Rich-
mond, VA) and 32.7 kg of muriate of potash
(Southern States Cooperative, Inc., Richmond,
VA) per ha. Nitrogen was increased in Feb to 27.1
Kg/ha of ammonium nitrate to accommodate in-
creased nutrient demand during fruit develop-
ment. Fungicides were applied 3 times per week
in rotation to all experimental plots throughout
the season to combat Botrytis fruit rot (Botrytis ci-
nerea) and anthracnose fruit rot (Colletotrichum
acutatum). The fungicides used were Abound
(azoxystrobin) (Syngenta Crop Protection,
Greensboro, NC), Topsin (thiophanate) (Cerexa-
gri, Inc., King of Prussia, PA), Aliette (alumi-
num tris) (Bayer Crop Science, Research Triangle
Park, NC), and Serenade (Bacillus subtilis)
(Agraquest, Davis, CA). No insecticides or acari-
cides were applied to the research plots. Prepara-
tion and management procedures are described in
detail in Fraulo & Liburd (2007).
The experimental design used was a random-
ized complete block with 4 treatments and 4 rep-
lications. Neoseiulus californicus (Koppert Biolog-
ical Systems, Romulus, MI) was released in all
treated plots at the recommended rate of 1-2
predators per square meter. Viability was as-
sessed by observing 20-30 predatory mites in a
Petri dish with a dissecting binocular microscope
(10-20x) (Leica MZ12.5, McBain Instruments,
Chatsworth, CA) for 15 min before each release to
ensure that the mites were vigorously active. The
treatments included releases ofN. californicus as
follows: (1) in the "early" season at foliar develop-
ment, 4 weeks after planting (WAP), (2) during

Florida Entomologist 91(3)

the "mid" season at floral development, 8 WAP, (3)
"late" in the season at fruit development, 12-16
WAP, and (4) "no release" untreated control.

Preliminary Study

Field preparations for the preliminary study
are described above and in Fraulo and Liburd
(2007). The goal of the preliminary study was to
determine an appropriate sample size and to eval-
uate diversity with the Shannon-Weaver diver-
sity index. Data were collected at 12 WAP, during
the "late" season of the 2005-2006 field season to
assess the overall effect of N. californicus on the
arthropod assemblage in the strawberry field.
One yellow sticky Pherocon AM Trap (YST)
(Tr4ce, Inc., Adair, OK), 28 cm x 23 cm surface
area with 56 squares of 6.45 cm2 (1 in2) forming a
grid on the board was hung on a garden stake 30
cm above plants. Each trap was placed in the cen-
ter row of each of the 24 treatment plots. Traps
were collected weekly for 5 weeks and placed into
Zipper Seal Storage Bags@ (American Value, Dol-
gencorp, Inc., Goodlettsville, TN) and transported
to the Small Fruits and Vegetable IPM Labora-
tory at the University of Florida, Gainesville, FL
to be examined under a dissecting microscope (10-
20x) (Leica MZ12.5, McBain Instruments, Chat-
sworth, CA). To develop a sub-sample protocol, 3
YSTs were randomly chosen from all samples and
each of the 56 one-inch squares on each trap was
examined to determine the arthropod families
found per square. The families observed on each
square of the YST were counted and compiled into
a comprehensive list, and recorded. A cumulative
frequency distribution was plotted for each
square to determine the optimal sub-sample size.
These data were used to create a sub-sampling
scheme for data analysis in the main study. The
frequency tables of the families with their key
taxonomic characteristics were recorded and used
for primary identification during the main study.

Main Study

We evaluated the effect of N. californicus by
sampling at 1-month intervals 2 weeks after each
release date, based on the homogeneity of the pre-
liminary results, field observations, and previous
research by Garcia-Mari & Gonzalez-Samora
(1999). Arthropod sampling was conducted during
the 2006-2007 field season at (a) 2 months after
planting ("early" season), (b) 3 months after
planting ("mid" season), and (c) 4 months after
planting ("late" season). Individuals were identi-
fied to family or genus depending upon the level of
functional variation within the taxon, based on
notes from the preliminary study described above
and previous research (Jones 1976; Cross et al.
2001; Arevalo et al. 2006; Klein et al. 2006). The
number of individuals present from each taxon

was collected by the method described in the pre-
liminary sampling, identified, and recorded. Un-
known taxa were identified at the Department of
Plant Industry, Gainesville, FL. Data were col-
lected throughout the season and compared to de-
termine the effect of N. californicus releases on
arthropod assemblages in the field within each
distinct period.
We employed 4 sampling methods in order to
increase the probability of encountering a higher
diversity of taxa and to avoid bias, as follows: (1)
In situ (visual inspection), (2) foliar sampling, (3)
pitfall traps, and (4) yellow sticky traps.
In situ Sampling. Twenty-four strawberry
plants from the interior rows of each plot were vi-
sually inspected once weekly for 2 weeks during
each sample period. The visual inspection con-
sisted of a scan for 30 s for each plant. This en-
abled us to sample the larger arthropods occur-
ring in the field including macro-hymenopterans,
hemipterans, and coleopterans.
Foliar Sampling. Four young trifoliate leaves
from the inside of the plant crown and 4 old trifo-
liates from the outer-crown were taken randomly
from each treatment plot. Samples were con-
ducted weekly for 2 weeks after each of the 3 pred-
atory releases. The leaves were placed in a ziplock
bag and transported to the laboratory where they
were visually inspected under the dissecting bin-
ocular microscope for leaf-dwelling and minute
Pitfall Traps. Traps were constructed of white
polypropylene deli containers 14 cm deep and
10.5 cm in diameter (Fabri-Kal Corp., Kalamazoo,
MI) filled with 0.15 L of 10% dish soap and water
solution. The traps were placed in the soil under
the black plastic mulch in one of the 2 center rows
of each treatment plot to capture cursorial soil ar-
thropods and soil dwellers (Southwood 1966). The
traps were left in the field for 48 h each week for
a 2-week period after each of the 3 predatory re-
Yellow Sticky Traps. One trap per plot was
placed in one of the 2 center rows at foliar height,
approximately 30 cm above ground to capture
winged arthropods. The YST were left in the field
for 48 h each week for a 2-week period after each
of the 3 predatory releases. In the laboratory, a
proportion of each trap surface area was observed
for analysis based on results of the preliminary

Statistical Analysis

The arthropod assemblages among treatments
in both the YST and pitfall traps throughout the
season were analyzed with a Non-Metric Multidi-
mensional Scaling (NMS) ordination with PC-
ORD 4 (Kruskal 1964). This method is well-suited
for describing patterns in community data and
does not assume normality (McCune & Grace 2002).

September 2008

Fraulo et a.: N. californicus and Arthropod Community Structure

The ordination was conducted by the S0rensen
distance measure (McCune & Grace 2002) with a
random starting configuration. The autopilot set-
ting on medium thoroughness was used to deter-
mine the dimensions of the ordination (McCune &
Grace 2002). Fifteen runs with real data with 200
iterations were used. To assess the strength of the
ordination, the Monte Carlo option with 30 runs
of randomized data was selected. Data were
square-root transformed if the correlation of vari-
ation among rows and columns were greater than
50% (McCune & Grace 2002) and outliers were re-
moved to strengthen the structure of the data.
Rare families that were present in numbers too
low to be included in the ordination were recorded
in a comprehensive list for each treatment and
were included in diversity calculations. All taxa
found were included in the calculation of the Sh-
annon-Weaver diversity index and results among
treatments were compared by analysis of vari-
ance (ANOVA) followed by mean separation with
LSD test (SPSS 2004). To convert the results of
the Shannon-Weaver index into a true diversity
measure, the exponential of the entropy value
was calculated (Jost 2006), as follows:

D = exp pilnpi = exp(H')

where pi = the proportional abundance of taxon.
D = true diversity measure.


Preliminary Study

Sub-sampling procedures demonstrated that
28 squares on the YST consistently included at
least 90% of the arthropod families recorded (Fig.
1). Therefore, 28 squares (47% of the trap area,
excluding borders) were observed for analysis in
the main study. The Shannon-Weaver index indi-
cated no significant differences in level of diver-
sity among any of the treatments in the prelimi-
nary study (F = 0.5; df = 3, 12; P = 0.69).

Main Study

Arthropod community structure following re-
leases of N. californicus for biological control of
TSSM was analyzed during various periods
throughout the growing season. The NMS ordina-
tion conducted following releases of N. californi-
cus during "mid" season sampling showed a weak
community structure and did not provide any
groupings of the data related to the treatments
imposed (Fig. 2). A similar lack of structure was
observed in ordinations for the "early" and "late"
sampling dates (data not shown). In the "early"

o, i

0 5 10 a6 20
N',ite ofS%--s

26 M0 a.

Fig. 1. Distribution frequency used in the prelimi-
nary trial to calculate optimal sub-sample size for
squares on yellow sticky card trap (YST).

season (Dec) we compared the plots treated with
N. californicus releases and untreated plots
(without N. californicus releases). The arthropod
assemblages found in pitfall traps showed a weak
structure (axis 1, P = 0.13; axis 2, P = 0.13; axis 3,
P = 0.32) indicating that N. californicus releases
did not have a significant effect on the arthropod
assemblages. When data were square-root trans-
formed and outliers were removed, the NMS ordi-
nation remained non-significant (axis 1, P = 0.13;
axis 2, P = 0.29; axis 3, P = 0.48). Results from
NMS ordination of the YST also were not signifi-
cant (axis 1, P = 0.52; axis 2, P = 0.68; axis 3, P =
0.48). The coefficient of variation was low
(39.17%) indicating that transformation of data
has no effect on data structure.
During the "mid" season (Jan) no significant
differences were recorded in assemblages col-
lected in the pitfall traps (axis 1, P = 0.29; axis 2,
P = 0.26; axis 3, P = 0.26) by the NMS ordination.
Yellow sticky traps also showed no significant dif-
ferences among treatments (axis 1, P = 0.29; axis
2, P = 0.13; axis 3, P = 0.23). Transformations
were not recommended as it did not affect data
In the final NMS ordination, no significant dif-
ferences among treatments were found in the late
season (Feb), for pitfall trap data (axis 1,P = 0.13;
axis 2, P = 0.16; axis 3, P = 0.32). When outliers
were removed from the data set results remained
non-significant (axis 1, P = 0.13; axis 2, P = 0.16;
axis 3, P = 0.13). Yellow sticky traps showed no
differences among arthropod assemblages with
introduction of N. californicus (axis 1, P = 0.52;
axis 2, P = 0.71; axis 3, P = 0.71). Removing outli-
ers did not affect data structure (axis 1, P = 0.29;
axis 2, P = 0.32; axis 3, P = 0.58).
The true diversity index based on the Shan-
non-Weaver index ("D") showed a significant ef-
fect on arthropod diversity in the YST in the
"early" season (F = 5.35; df = 3,12; P =0.01), but
not in the pitfall traps (F = 0.3; df = 3,12; P =
0.83). "Mid" season YST and pitfall traps showed
no significant effect on diversity (F = 1.3; df =

.......... .

Florida Entomologist 91(3)

Mid-Season Pitfall Traps

X 0
02- S
.02- X x


*1 0 ..
-4 -12 -1.0 -0.8 -0.46 -02 00 0.2 0.4 06 0.
Axis I
Mid-Season Yellow Slidcy Traps

OS. *
1 0 x

-12 -10 -00 -06 -04 -02 00 02 04 06 0.B

September 2008

Mid-Season Pitfall Traps

D 5


-14 -12 -1.0 -0 -06 -04 -02 00 02 04 0-6 0-8


-0X2 X
00 a


-12 -12 0 -0B -06 -04 -02 00 02 04 06 08

Axi 1

Fig. 2. Non-Metric Multidimensional Scaling (NMS) ordinations of arthropod communities in "mid" season pit-
fall traps (a, b) and yellow sticky traps (c, d) showing weak data structure along axis 1 and 2 (a, b), and axis 1 and
3 (c, d) with no significant differences (P < 0.05) among treatments. A = Early release; 0 = Mid-season release; =
Late release; x = No Release (control).

3,12; P = 0.7; and F = 0.3; df = 3,12; P =0.57, re-
spectively). Yellow sticky traps (F = 2.8; df= 3,12;
P = 0.08) and pitfall traps (F = 6.5; df = 3,12; P =
0.6) also showed no significant difference among
treatments late in the season (Tables 1 and 2).
Overall, the visual and foliar samples did not
produce sufficient numbers of arthropods to con-
duct robust statistical analysis. However, they
should not be dismissed because they revealed in-
teresting phenological trends and important nat-
ural predators of strawberry pests. Early in the
season thrips (Frankliniella spp.) and Chalci-
doidea populations were significantly higher in
the treated plots compared with the untreated
plots (F = 4.81; df= 1, 30; P = 0.04, F = 8.44; df=
1, 30; P = 0.01, respectively) During the "mid" sea-
son high numbers of Pachybrachius spp. and a
dramatic decline in aphid population directly fol-
lowing an increase in syrphid abundance were ob-
served (Tables 3, 4, 5, and 6). Foliar sampling in-
dicated that as the season progressed, the abun-
dance of Coccinellids increased in all treatment

plots (personal observation), as did six-spotted
thrips Scolothrips sexmaculatus (Pergande) and
Geocorid bugs Geocoris spp. Late in the season,
numbers of taxa decreased, and an increase of
Bradysia spp. (Sciaridae) was observed.


Ordination results confirmed the lack of treat-
ment effects observed through the analysis of
variance. If the treatments had affected commu-
nity structure, points representing the same
treatment would have clustered together in the
ordination figures (Klein et al. 2006), but this was
never observed. Ordination used to describe the
community patterns and the measures of diver-
sity both indicated that presence ofN. californicus
did not disrupt the natural ecology of the system.
Neoseiulus californicus was released at several
times during the season and persisted through
the experiment, not disrupting arthropod assem-
blages while significantly reducing TSSM popula-

Fraulo et a.: N. californicus and Arthropod Community Structure


Sample date

Treatment Dec Jan Feb Preliminary

Early 2.9 0.16 c 9.4 1.00 7.1 0.68 8.6 0.41
Middle 2.5 0.20 b 8.6 0.30 8.8 0.91 8.7 1.40
Late 3.9 + 0.34 a 7.5 0.66 5.7 1.00 8.2 0.14
No-release 3.1 0.28 b 8.2 0.53 6.3 0.47 7.0 0.95

Means with the same letter within columns are not significantly different (P < 0.05) based on LSD test. Dates with no letters
represent dates with no significant differences.

tions as observed in a related study (Fraulo &
Liburd 2007). The Shannon-Weaver index on YST
indicated a significant difference on community
structure between treated and untreated plots in
the "early" season. However, this effect was short
lived and was non-significant for the remainder of
the season. This early disruption may be due to
sparse and clumped populations within arthropod
communities at the beginning of the season and
did not have a significant overall effect.
The majority of predators in the strawberry
system are generalist feeders and N. californicus
itself is a generalist predator. Feeding preference
likely moderated the impact on the arthropod as-
semblage. Our results indicate that the release of
N. californicus does not have a statistically signif-
icant effect on the assemblage and community
functioning of arthropods in the strawberry sys-
tem. Its release did not affect the balance and reg-
ulation of the natural ecosystem in our study.
Generalist feeding behavior coupled with a high
level of richness and insect diversity in the straw-
berry system may be key factors in reducing the
effect of N. californicus releases on the structure
of the strawberry system.
There were no overall non-target effects on
natural enemies. Major insect families (Thripi-
dae, Cecidomyiidae, Coccinellidae, Staphylinidae,
and Lygaedae) as cited by Oatman et al. (1985)
and Rondon et al. (2004), were present through-
out our study. Thrips (Frankliniella spp.) and

Chalcidoidea populations were present through-
out the field early in the season. Neoseiulus cali-
fornicus is a known predator of thrips. However,
adult thrips migrate into flowering strawberry
plants and take shelter within the styles of the
strawberry flower where N. californicus cannot
access them (Cross et al. 2001). Increased levels of
the superfamily Chalcidoidea, which are parasi-
toids of thrips, were also observed in higher num-
bers in the plots with high numbers of thrips.
Aphids were abundant in all treatments dur-
ing the "early" season and decreased throughout
the season. Decreasing numbers of aphids in con-
junction with increased numbers of Syrphidae are
consistent with studies conducted in north Wales
showing that Syrphidae can cause considerable
reduction in aphid numbers (Cross et al. 2001).
This function was not affected by the presence of
N. californicus. Foliar sampling indicated that as
the season progressed, the abundance of sixspot-
ted thrips (S. sexmaculatus) and Geocoris spp. in-
creased in all treatment plots (both are predators
of TSSM).
Our findings from this and previous release
studies demonstrate that releasing N. californi-
cus in the field at the recommended rate of 1-2 N.
californicus per m2 when TSSM populations are
low (<70-80 TSSM per trifoliate) provides season-
long control of TSSM and does not disrupt other
natural enemies of seasonal pests (Jones 1976;
Fraulo & Liburd 2007). A one-time application of


Sample date

Treatment Dec Jan Feb

Early 5.4 0.76 4.7 0.77 3.8 0.42
Middle 5.0 0.44 3.7 0.44 3.7 0.09
Late 5.2 0.62 3.7 0.54 4.4 0.37
No-release 5.8 0.63 3.8 0.46 3.9 0.47

No significant differences (P < 0.05) among treatments on any sampling date.


Dec Jan Feb


Acanalonidae 2 1 2 1 Acanalonidae 4 1 3 3 Aleyrodidae 4 -
Aleyrodidae 2 16 26 14 Aleyrodidae 2 17 3 1 Aphididae 6 6 1 3
Aphididae 383 643 579 539 Aphididae 12 10 5 11 Cecidomyiidae 4 6 8 1
Bethylidae 1 Tachinidae 5 8 2 3 Chalcidoidea 33 26 33 31
Bibionidae 5 5 Bibionidae 3 1 1 Chrysomella 2 2 2 -
Cecidomyiidae 11 10 6 9 Braconidae 2 1 3 5 Cicadellidae 7 2 1 7
Chalcidoidea 34 59 83 91 Cecidomyiidae 10 8 14 4 Coccinella (Hippodamia spp.) 3 1 6
Chrysomelidae 1 Chalcidoidea 61 66 78 53 Drosophilidae 1 -
Cicadellidae 2 11 17 5 Chrysomelidae (Alticinae spp.) 1 1 1 Ichneumonidae 3 8 4 4
Staphylinidae 2 Cicadellidae 3 3 4 2 Lepidoptera 1 -
Cucujidae 1 11 Coccinellidae (Hippodamia spp.) 4 3 2 Muscidae 17 14 14 19
Dolichopodidae 7 17 4 3 Drosophilidae 4 1 2 1 Nitulidae 2 1 2 -
Drosophilidae 3 Ichneumonidae 4 3 Lygaeidae (Pachybrachius spp.) 1 4 5 2
Ichneumonidae 4 4 Leiodidae 1 1 1 Phoridae 2 6 1 2
Lepidoptera 1 1 1 Dolichopodidae 2 4 4 5 Psychodidae 2 1
Muscidae 8 7 9 30 Lygaeidae (Pachybrachius spp.) 3 6 1 Sciaridae (Bradysia spp.) 52 48 94 48
Nitulidae 2 Muscidae 23 13 15 15 Staphylinidae 2 1 1 1
Phloeothripidae 3 2 4 3 Nitulidae 1 1 Syrphidae (Sphaerophoria spp.) 3 2 1 1
Phoridae 3 16 7 7 Phloeothripidae 1 Thripidae (Frankliniella spp.) 2 8 5 2
Platygatroidae 1 Phoridae 10 14 40 9 Araneida 1 1 1 3
Psychodidae 1 3 10 Psychodidae 5 2 1 Dolichopodidae 10 7 3 4
Sciaridae (Bradydia spp.) 5 5 50 10 Sciaridae (Bradysia sp.) 23 17 24 8 Tachinidae 2 4 4 2
Thripidae (Frankliniella sp.) 39 39 23 Staphylinidae 2 1 -
Syrphidae (Sphaerophoria spp.) 1 1
Thripidae (Frankliniella spp.) 16 17 9 10

E = early treatment, M = middle treatment, L = late treatment, and NR = no release.


Dec Jan Feb


Chrysomelidae (Alticinae spp.) 1
Aphididae 19 9
Apoidea -
Bibionidae 2 -
Cecidomyiidae 8 7
Chalcidoidea 4
Chironomidae 2
Cicadellidae 1 -
Collembola 41 21
Cucujidae 1 -
Drosophilidea -
Elateridae -
Formicidae 35 26
Ichnueumonidae 1
Lepidoptera -
Miridae 1 -
Muscidae 2 2
Mutillidae 1 -
Nematode 2
Phloeothripidae 1
Phoridae 2 3
Sciaridae (Bradysia spp.) 1
Staphylinidae 3 1
Tettigoniidae 2
Thripidae (Frankliniella spp.) 1 5
Araneida 1 2

1 Aphididae 11 2 6 5 Aphididae
5 9 Cecidomyiidae 4 6 2 Braconidae
1 1 Chalcidoidea 1 2 1 Cecidomyiidae
- Cicadelidae 1 Chalcidoidea
7 17 Collembola 50 85 132 65 Chrysomellidae
3 2 Gryllidae 1 Collembola
- Cucujidae 1 1 Formicidae
1 Drosophilidae 2 1 Lygaeidae (Geocoris spp.)
23 31 Formicidae 44 9 2 2 Sciaridae (Bradysia spp.)
- Lygaeidae (Pachybrachius spp.) 18 21 47 29 Staphylinidae
- 2 Miridae 1 Thripidae (Frankliniella spp.)
1 Muscidae 1 Vespidae
6 42 Nitulidae 1 Araneida



2 2
1 2
1 1

3 1
5 2

E = "early" treatment, M = "mid" treatment, L = "late" treatment, and NR = no release.

1 1
- 1 -

- -
1 1 -
- -- 1
10 24 13
1 1 11
4 1 -
9 7 14
- 1

- 1 -
3 1 1
- 2 -


- &


- Phoridae 1 -
2 Sciaridae (Bradysia spp.) 4 2 1 2
- Araneida 4 3 2 2
- Staphylinidae 1 -
- Tettigoniidae 1 -
- Thripidae (Frankliniella spp.) 2 2 2 4
- Scollidae 1 -


Florida Entomologist 91(3)


Dec Jan Feb


Tettigoniidae 3 1 3 3 4 7 2 4
Apoidea 6 9 5 0 1 3 2
Syrphidae (Sphaerophoria spp.) 3 3 1 4 16 13 6 8 2 1 1
Araneida 1 2 1 8 4 2 -
Lepidoptera 1 1 1 1 1 -
Muscidae 1 2 2 2 1 1 2
Chrysomelidae 1 1 1
Coccinelidae (Hippodamia spp.) 1 1 2 1 -
Lygaeidae (Tristicolor spp.) 1
(Pachybrachius spp.) 8 2 6 12 1 1
Sciaridae (Bradysia spp.) 52 28 39 14
Cicadellidae 1 -

E = "early" treatment, M = "mid" treatment, L = "late" treatment, and NR = no release.


Dec Jan Feb


Aphididae 43 64 28 40 1 4 4 1 1 -
Aleyrodidae 29 22 22 2 2 2 2 2 3
Thripidae (Frankliniella spp.) 7 2 1 1 14 4 14 14 3 3
(Scolothrips sexmaculatus) 1 2 2 6 3 10 10 4 1 3 2
Chalcidoidea 1 -
Lepidoptera 1 6 6 2 9
Syrphidae (Sphaerophoria spp.) 9 8 9 9 4 5 10 15
Lygaeidae (Geocoris spp.) 3 9

E = "early" treatment, M = "mid" treatment, L = "late" treatment, and NR = no release.

N. californicus at the recommended rate would
cost one-third the price of chemical treatments
while having a non-significant effect on benefi-
cials in strawberry. Although N. californicus is
tolerant to many insecticides and fungicides
(Easterbrook 1992; Croft et al. 1998; Escudero &
Ferragut 2005; and Liburd et al. 2007), we do not
recommend additional chemical applications as
they may adversely affect other beneficial. We
found that when released in the field, N. californi-
cus is able to maintain consistent control of TSSM
populations while maintaining an array of benefi-
cials throughout the season in north Florida
strawberry fields.


We thank Elizabeth Boughton, Pedro Quintana-As-
cencio, and H. Alejandro Ar6valo for providing assis-
tance with statistical analysis and critical reviews of
the project.


Key to the most common species of thrips found in
early-season blueberry fields in Florida and south-
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CLOYD, R. A., C. L. GALLE, AND S. R. KEITH. 2006. Com-
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S. L. HSU. 2004. Evaluation of large-scale releases of
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Comparative life histories and predation types: are
Neoseiulus californicus and N. fallacis (Acari: Phy-

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Florida Entomologist 91(3)

September 2008


1Mid-Florida Research and Education Center and Department of Entomology and Nematology,
University of Florida, IFAS, 2725 Binion Road, Apopka, FL 32703-8504

2Leon County Hazardous Waste Management, 7550 Apalachee Parkway, Tallahassee, FL 32311

'Department of Zoology, University of Burdwan, West Bengal, India


A qualitative and quantitative population survey of immature and adult Chironomidae was
conducted for 1 year in a country club wetlands in northeast Florida, USA. (. i."' ...."hl"
paripes and Goeldichironomus cars were the 2 predominant chironomid species in the wet-
lands. Adults of these 2 species emerged at nuisance levels from Apr through Jun, and in
Aug and Sep. Polypddiliume' Cryptochironomus, Tanytarsini, and Tanypodinae collected in
low numbers during the survey were not identified to species. In laboratory bioassays, LCg0
values of technical grade temephos against G. paripes and G. cars were 0.01 and 0.009 ppm,
respectively. For s-methoprene the LCg0 values were 0.082 and 0.055 ppm, and for Bacillus
thuringiensis serovariety israelensis (Bti) 1.056 and 0.467 ppm, respectively. In experimen-
tal field plots in the wetlands, 5% AI Skeeter Abate@ (temephos) pellets at 0.1 kg AI/ha re-
duced midge larvae by 52-86% and at 0.2 kg/ha by 74-92% during 4 weeks posttreatment.
Sand formulated technical powder of Bti at 1,000,000 and 2,000,000 ITU (International
Toxic Units) Bti/m2 reduced midge larvae by 47-52% and 82-88%, respectively, during 6 to 20
days posttreatment. STRIKE pellets (4.25% AI s-methoprene) at 0.14 kg AI/ha suppressed
a maximum of 80% total chironomid adult emergence at 7 days posttreatment; this IGR at
0.28 kg AI/ha reduced adult emergence up to 92% during 15 days posttreatment. Temephos
and Bti were more cost-effective and provided midge control for relatively longer period than
s-methoprene in the field evaluations.

Key Words: nuisance midges, Glyptotendipes paripes, Goeldichironomus carus, population
management, temephos, Bacillus thuringiensis serovar. israelensis, s-methoprene


Se realize un monitoreo cualitativo y cuantitativo por 1 ano de los inmaduros y adults de la
familiar Chironomidae en los humedales de un club en el noreste de la Florida, EEUU. Glyp-
totendipes paripes y Goeldichironomus carus fueron las species de la familiar Chironomidae
mas predominantes en los humedales. Los adults de estas species emergieron en niveles
fastidiosos desde el mes de abril hasta junio, y en agosto y septiembre. Los Polypedilum,
Cryptochironomus, Tanytarsini y Tanypodinae recolectados en numerous bajos durante el mo-
nitoreo no fueron identificados al nivel de especie. En los bioensayos del laboratorio, el valor
CLg0 de 'temephos' al grado tecnico contra G. paripes y G. carus fue 0.01 y 0.009 ppm, respec-
tivemente. Los valores de CL,0 para 'S-methoprene' fueron 0.082 y 0.055 ppm y para 'Bacillus
thuringiensis' serovariedad israelensis (Bti) 1.056 y 0.467 ppm, respectivemente. En ensayos
de parcelas en el campo en los humedales, pelotillas al 5% de AI Skeeter Abate@ (temephos)
al 0.1 kg IA/ha reducieron 52-86% de las larvas y al 0.2 kg/ha 74-92% de las larva durante
las 4 semanas despues del tratamiento. El polvo tecnico de Bti formulado con arena al
1,000,000 y 2,000,000 UTI (Unidades Toxicos Internacionales) Bti/m2 reducieron las larvas
de chironomidos por 47-52% y 82-88%, respectivemente, durante los 6 a 20 dias despues del
tratamiento. Las pelotillas de STRIKE (4.25% de IA S-methoprene) al 0.14 kg =IA/ha su-
primieron la emergencia de los adults de chironomidos a un maximo de 80% a los 7 dias des-
pues del tratamiento; este RCI (Regulador del Crecimiento de Insectos) aplicado al 0.28 kg
IA/ha redujo la emergencia de adults hasta 92% durante los 15 dias despues del trata-
miento. Temephos y Bti fueron los mas efectivos en cuanto al costo y controlar los chironomi-
dos por un period de tiempo mas largo que S-methoprene en las evaluaciones del campo.

Adult emergence of chironomid midges can oc- review). At Ponte Vedra Beach, northeast Florida,
cur at nuisance levels in areas surrounding urban USA, a "labyrinth" of shallow wetlands developed
and suburban aquatic habitats (see Ali 1996 for for residential and recreational purposes, support

Ali et al.: Control of Chironomidae

chironomid populations at nuisance levels for sev-
eral months each year that interfere with human
activities, necessitating control measures. We ex-
amined the composition of the midge larval com-
munity in these wetlands, as well as the preva-
lence of adults in adjacent residential and recre-
ational land areas. Laboratory and field evalua-
tions of the organophosphorus larvicide
temephos, the insect growth regulator (IGR), s-
methoprene, and the biological insecticide, Bacil-
lus thuringiensis serovar. israelensis (Bti) for
midge control also were conducted. The relative
cost of controlling midge larvae with each mate-
rial was compared.

The habitat in northeast Florida (3011'N,
8122.5'W) includes a series of man-made shallow
lakes (<1-3 m deep, surface area 65 ha) with ca. 19
km of irregular shoreline (Fig. 1). Prior to routine
larval and adult sampling, a survey of the entire
habitat was conducted for midge fauna by collect-
ing benthic samples with a 15 x 15 cm Ekman
dredge from a boat at ca. 0.05-min latitude/longi-
tude intervals. Spatial coordinates were recorded
with a Global Positioning System receiver. All
benthic samples were washed in the field through
a 350-pm pore sieve, and retained material trans-
ferred to labeled bottles for transport to the labo-
ratory to identify (Epler 1995) and count the im-
mature (larvae and pupae) Chironomidae. Based
on these data which identified areas supporting
nuisance or near-nuisance (>100 larvae/m2) levels
of midge larvae, a routine spatio-temporal strati-
fied larval sampling plan was established. This
plan facilitated the efficient use of sampling re-
sources (Lobinske et al. 2002). From Jul 2003 to
Jul 2004, 45 Ekman dredge samples were col-
lected at monthly basis, processed as above, and
the midge larvae identified and counted.
Adult midge populations were assessed based
on 9 permanently placed New Jersey (NJ) light
traps (Fig. 1). Trap jars were replaced at least
weekly and collected chironomids were identified
(Weiderholm 1989) and counted.
A procedure similar to that of Ali (1981) was
used for laboratory evaluation of technical teme-
phos (90% active ingredient, Al) and the biologi-
cal insecticide Bti (potency: 7,468 International
Toxic Units (ITU)/mg) against field-collected lar-
vae of the predominant midge species. The serial
dilutions for temephos were made in acetone
while those for Bti were made in deionized water.
Larval mortality in the bioassay cups was noted
24 h (temephos) and 48 h (Bti) after treatments
and corrected for any mortality in corresponding
control cups.
Technical grade s-methoprene (96% Al) was
tested in small rearing units with continuous air
supply (Ali & Lord 1980). Larval and/or pupal




Basin C'


Fig. 1. Map of study wetlands for Chironomidae
showing Basins A to D (larval sampling) and location of
New Jersey light traps 1 to 9 (adult sampling), Ponte
Vedra Beach, Florida, USA.


Florida Entomologist 91(3)

mortality and adult emergence was assessed
daily until all immatures had died or emerged as
adults in the controls. Bioassays for all materials
were replicated at least 3 times. The corrected
mortality data were analyzed by Probit analysis
to estimate dose response of larvae of each species
to the test materials.
Temephos (5% Skeeter Abate@ pellets), s-me-
thoprene (4.25% STRIKE pellets), and Bti (Vec-
toBac technical powder, containing 5,000 ITU/
mg) were evaluated for chironomid control in the
wetlands. For this, 21 open field plots (each 20 x 5
m) in wetland areas supporting consistent larval
densities of >1,000/m2 were permanently marked
by driving a stake in each corner of a plot. Each
test material was evaluated at 2 rates in separate
evaluations that followed at the termination of
the prior field evaluation. Each treatment rate
was applied to 3 randomly selected plots (repli-
cates) while 3 plots served as controls, utilizing a
total of 9 plots per evaluation. Treatments were
made by hand from a boat to ensure uniform dis-
tribution. The first evaluation started on 20 Apr
2004, when temephos was applied at 0.1 and 0.2
kg AI/ha. On 2 Jun 2004, Bti formulated on sand
grains at 1,000,000 and 2,000,000 ITU/m2 was ap-
plied. On 4 Oct 2004, s-methoprene was applied at
0.14 and 0.28 kg AI/ha for the final evaluation.
For temephos and Bti evaluations, 3 Ekman
dredge samples were randomly collected from each
plot immediately prior to treatment and at post-
treatment d 3, 7, 14, 21, and 28 (temephos), and 2,
6, 13, 20, and 30 d (Bti). All benthic samples were
washed and processed in the laboratory as above.
For evaluation of s-methoprene, a single night's
adult emergence at pretreatment and at 7, 15, 22,
and 29 d posttreatment was sampled utilizing 3
randomly placed 30-cm high (0.25 m2 base area)
metal-cone submerged emergence traps (Ali 1980)
per plot; thus 27 total traps were utilized during
each sampling. A removable glass jar at the apex of
the trap containing the trapped adult chironomids
was collected and transported to the laboratory for
midge identification and counting. The degree of
reduction in posttreatment larvae and adults was
calculated according to Mulla et al. (1971).


Two chironomid species, Glyptotendipes
paripes and Goeldichironomus cars predomi-
nated in Ponte Vedra Beach wetland and were the
primary nuisance in the vicinity. In the prelimi-
nary survey of the entire wetland, G. paripes
formed 99% and G. cars <1% of the total chirono-
mid larvae. Midge larvae of the taxa, Polypedi-
lum, Cryptochironomus, Tanytarsini and Tanypo-
dinae were found in very low numbers and were
not identified to species. Spatially, mean midge
densities in different basins (Fig. 1) of the wetland
varied considerably; Basin A supported the high-

est (1,714 larvae/m2) and Basin B the lowest (57
larvae/m2) densities of midge larvae. Basins C and
D, respectively, supported 982 and 344 midge lar-
vae/m2. The overall mean density of total midge
larvae in all basins amounted to 923 larvae/m2.
Monthly mean larval densities (Fig. 2) varied
from 152 larvae/m2 (Aug 2003) to 1,666 larvae/m2
(May 2004) for G. paripes; 20 larvae/m2 (Jul 2003)
to 581 larvae/m2 (Aug 2003) for G. cars; and 363
larvae/m2 (Nov 2003) to 2,809 larvae/m2 (May
2004) for total Chironomidae. Glyptotendipes
paripes and G. cars, respectively, formed 56 and
14% of total Chironomidae larvae collected during
the study period. The larval maxima for G.
paripes occurred in Apr-May 2004 and minima in
Aug and Nov 2003. The highest density of G.
cars occurred in Aug 2003 and lowest in Jun
2003 and Jun 2004. It was interesting to note that
total chironomid densities (predominately G.
paripes) during winter (Dec 2003 and Jan and Feb
2004) exceeded 1,200 larvae/m2.
Glyptotendipes paripes formed 86% and G.
cars 7% of the total adult chironomids collected
during the study period (Fig. 2). The daily mean
number of G. paripes occurring in NJ traps was
highest in Apr 2004 (3,096 adults/trap/day) and
lowest in Jan 2004 (<5 adults/trap/day). Goeldi-
chironomus cars populations were <1 adult/trap/
day in Dec 2003 and Jan 2004, with maxima (363
adults/trap/day) occurring in Apr 2004. In gen-
eral, peak adult activity was somewhat bimodal,
with major peaks occurring during Apr to Jun
and relatively smaller peaks from Aug to Nov.
There were appreciable numbers of adults in the
study area for almost 9 months of the year.
Susceptibility of G. paripes and G. cars larvae
in the laboratory to temephos was very similar;
both species were highly susceptible to temephos
as indicated by LC90 values of 0.01 ppm (G.
paripes) and 0.009 ppm (G. cars) (Table 1). The
Bti technical powder was twice as effective
against G. cars (LC90 = 0.467 ppm) compared to
G. paripes (LC90 = 1.056 ppm) (Table 1).
The IGR s-methoprene was more effective
against G. cars (LC90 = 0.055 ppm) compared to
G. paripes (LC90 = 0.082 ppm) (Table 1). The LC90
data indicate that G. paripes and G. cars, respec-
tively, were 8.2 and 6.1 times more susceptible to
temephos compared to s-methoprene.
In field tests, temephos at 0.1 kg AI/ha gave ap-
preciable larval control with 38-78% reduction in
the population of G. paripes and G. cars, and 52-
86% reduction in the total number of chironomid
larvae for 21 d posttreatment. At the application
rate of 0.2 kg AI/ha, temephos produced between
58 and 94% reduction in G. paripes and G. cars,
and 74-92% reduction of total midge larvae dur-
ing 28 d posttreatment (Fig. 3).
Posttreatment larval reduction with Bti was
lower compared to the temephos treatments. For
example, at 2 d posttreatment, only a 12% (low

September 2008

Ali et al.: Control of Chironomidae

U Geoldichironomus cars
S2500 Glyptotendipes paripes
W TotalChironomidae
2000 -

E 1500 -

+1 1000 J |
500 -

0 I
2003 I 2004

3500 -

3000 -
2500 -

X 2000 -

E 1500 -

') 1000 -

z 500 -

2003 2004

Fig. 2. Monthly mean densities of chironomid larvae prevailing in wetlands and corresponding population
trends of adult Chironomidae in 9 New Jersey (NJ) light traps permanently placed around wetlands at the Ponte
Vedra Beach, Florida (Jul 2003 to Jul 2004).

Florida Entomologist 91(3)

Geoldichironomus cars & Glyptotendipes paripes
10010.1 kg AI/ha T 0e

20 120
80 o80

G60 60

040 40

0 D

3 7 14 21 28

Bacillus thuringiensis se rovar. israelensis
1l 000000lA ITU/Hm2 2000000Afl ITIUrm2


S" 40


2 6 13 20 30 o
2 8 13 20 30

F_10.14 kg AI/ha

7 15 22 29

September 2008

Total Chironomidae

3 7 14 21 28

2 v 13 20 30r

2 6 13 20 30


1 0.28 kg Al/ha

Days Posttreatment

7 15 22 29
7 15 22 29

Fig. 3. Percent reduction of chironomid larvae after application of temephos (Skeeter Abate 5% pellets) utilized
at two rates (0.1 and 0.2 kg AI/ha), and a Bacillus thuringiensis serovar. israelensis powder containing 5,000 ITU/
mg, formulated on sand utilized at 2 rates (1,000,000 and 2,000,000 ITU/m2), and percent reduction of chironomid
adult emergence after application of s-methoprene (STRIKE 4.25% pellets) at 2 rates (0.14 and 0.28 kg AI/ha) in
field plots established in wetlands at the Ponte Vedra Beach, Florida (Apr-Nov 2004).








+H 40

0 0


Ali et al.: Control of Chironomidae


Lethal concentration (ppm)

Chi square for heterogeneity

Tabular value
Midge species LC50 95% CL LCg, 95% CL Slope Calculated at 0.05 level

G. paripes 0.005 0.004-0.005 0.010 0.008-0.012 4.30 6.797 7.815
G. carus 0.005 0.004-0.006 0.009 0.007-0.011 5.17 1.672 7.815
Bacillus thuringiensis serovar. israelensisb
G. paripes 0.175 0.127-0.224 1.056 0.778-1.624 1.64 4.046 11.070
G. carus 0.089 0.059-0.121 0.467 0.335-0.749 1.77 2.040 9.488
G. paripes 0.022 0.016-0.028 0.082 0.061-0.123 2.23 5.476 7.815
G. carus 0.013 0.010-0.017 0.055 0.040-0.086 2.04 4.551 7.815

aLarval mortality checked after 24 h.
'Larval mortality checked after 48 h.
'Continuously exposed in laboratory aquaria.

rate of application) and a 23% (high rate of appli-
cation) reduction in numbers of total larvae were
noted. However, at 6, 13, and 20 d posttreatment,
the low rate of Bti gave 38-48% reduction of G.
paripes and G. carus and 47-52% reduction in the
total larval chironomid population. The high rate
of Bti produced 76-86% reduction in G. carus and
G. paripes and 82-88% reduction in total Chirono-
midae during the 6-20 d posttreatment (Fig. 3).
The low rate of s-methoprene application re-
sulted in 80% reduction of adult Chironomidae at
7 d posttreatment, but thereafter this IGR was
generally ineffective. At 0.28 kg AI/ha, s-metho-
prene induced 71-92% emergence suppression of
adult Chironomidae for up to 15 d posttreatment
(Fig. 3).


Two species of chironomids, G. paripes and G.
carus vastly predominated the Ponte Vedra Beach
wetlands with other chironomid taxa found only
in very small numbers. Adults of these species
emerged at nuisance levels from Mar to Nov, with
large peaks of emergence occurring during Apr to
Jun, coinciding with highest larval densities dur-
ing this period. Larvae of the 2 midge species
were susceptible to the larvicides, temephos and
Bti, and the IGR, s-methoprene. In field trials, te-
mephos applied as 5% Skeeter Abate@ pellets at
0.2 kg AI/ha gave good control of larvae for up to
28 d; Bti at 2,000,000 ITU/m2 gave control for 20
d; and s-methoprene STRIKE@ pellets at 0.28 kg
AI/ha suppressed significant adult emergence for

15 d posttreatment. Considering the current mar-
ket price in relation to field control of these chi-
ronomids, temephos gave control for the longest
period and is 3-4 and 4-5 times more economical
than Bti and s-methoprene, respectively. How-
ever, due to the possible development of resis-
tance (Ali & Mulla 1978a), we recommend that
Bti and s-methoprene be used in rotation with te-
mephos as alternate options for midge control as
part of a resistance management program. Each
control material has a separate mode of action
and thus with alternation, the risk of resistance
to any one compound will be reduced. Although
temephos is environmentally more hazardous
than Bti and s-methoprene, the temephos use
rate of 0.2 kg AI/ha would probably have tempo-
rary and reversible impact on non-target biota co-
existing with chironomids in the aquatic ecosys-
tem, as described by Ali & Mulla (1978b).


Gratitude is expressed to the Sawgrass Association,
Incorporated, for a grant-in-aid to the University of
Florida to undertake this study.


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