Influence of methoprene on the...
 State records, confirmations, and...
 Flight phenology of male Cactoblastis...
 Two new species of Dryinidae (Hymenoptera:...
 High population density and egg...
 Ground-dwelling arthropod association...
 Biological control of Fenusa pusilla...
 Expression of feeding symptoms...
 A new species of Dacnusini from...
 Two new genera of Tropidocephalini...
 A survey of hymenopterous larval-pupal...
 Feeding, reproduction, and development...
 Natural enemies of Brazilian peppertree...
 Occurrence of Camponotus pennsylvanicus...
 A new host association of Commoptera...
 Reovirus-like sequences isolated...
 Studies on the genus Onesia (Diptera:...
 Taxonomic notes on the genus Haphsa...
 Descriptions of two new species...
 Kneallhazia (=Thelohania) solenopsae...
 Fecundity and mating propensity...
 Pupal development, longevity and...
 Sympatry of Orius insidiosus and...
 Description of late-instars of...
 The effect of larval diet and sex...
 Distribution of Verodes aequalis...
 Marking blueberry maggot flies...
 Detection surveys and population...
 The red palm weevil, Rhynchophorus...
 Sampling methods for Myllocerus...
 Patterns of ant activity on Opuntia...
 First record of Erythmelus klopomor...
 Greenidea psidii (Hemiptera: Aphididae:...
 Corrected species identification...
 Building roads and bridges for...
 Book reviews
 Back Matter

Group Title: Florida Entomologist
Title: The Florida entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00360
 Material Information
Title: The Florida entomologist
Uniform Title: Florida entomologist (Online)
Abbreviated Title: Fla. entomol. (Online)
Physical Description: Serial
Language: English
Creator: Florida Entomological Society
Florida Center for Library Automation
Publisher: Florida Entomological Society
Place of Publication: Gainesville Fla
Gainesville, Fla
Publication Date: June 2009
Frequency: quarterly
Subject: Entomology -- Periodicals   ( lcsh )
Insects -- Periodicals -- Florida   ( lcsh )
Genre: review   ( marcgt )
periodical   ( marcgt )
Additional Physical Form: Also issued in print.
System Details: Mode of access: World Wide Web.
Language: In English; summaries in Spanish.
Dates or Sequential Designation: Vol. 4, no. 1 (July 1920)-
Issuing Body: Official organ of the Florida Entomological Society; online publication a joint project of the Florida Entomological Society and the Florida Center for Library Automation.
General Note: Title from caption (JSTOR, viewed Sept. 13, 2006).
General Note: Place of publication varies.
General Note: Latest issue consulted: Vol. 87, no. 4 (Dec. 2004) (JSTOR, viewed Sept. 13, 2006).
 Record Information
Bibliographic ID: UF00098813
Volume ID: VID00360
Source Institution: University of Florida
Holding Location: University of Florida
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Resource Identifier: isbn - 0015-4040
issn - 1938-5102
oclc - 33223434
lccn - sn 95026670
 Related Items
Preceded by: Florida buggist (Online)

Table of Contents
    Influence of methoprene on the age-related mating propensity of males of the oriental fruit fly and the Mediterranean fruit fly
        Page 193
        Page 194
        Page 195
        Page 196
        Page 197
        Page 198
    State records, confirmations, and habitats of Aradidae (Hemiptera: Heteroptera) from Louisiana, U.S.A.
        Page 199
        Page 200
        Page 201
        Page 202
        Page 203
        Page 204
        Page 205
        Page 206
        Page 207
    Flight phenology of male Cactoblastis cactorum (Lepidoptera: Pyralidae) at different latitudes in the southeastern United States
        Page 208
        Page 209
        Page 210
        Page 211
        Page 212
        Page 213
        Page 214
        Page 215
        Page 216
    Two new species of Dryinidae (Hymenoptera: Chrysidoidea) from China
        Page 217
        Page 218
        Page 219
        Page 220
    High population density and egg cannibalism reduces the efficiency of mass-rearing in Euscepes postfasciatus (Coleoptera: Curculionidae)
        Page 221
        Page 222
        Page 223
        Page 224
        Page 225
        Page 226
        Page 227
        Page 228
    Ground-dwelling arthropod association with coarse woody debris following long-term dormant season prescribed burning in the longleaf pine flatwoods of north Florida
        Page 229
        Page 230
        Page 231
        Page 232
        Page 233
        Page 234
        Page 235
        Page 236
        Page 237
        Page 238
        Page 239
        Page 240
        Page 241
        Page 242
    Biological control of Fenusa pusilla (Hymenoptera: Tenthredinidae) in the northeastern United States: a thirty-four year perspective on efficacy
        Page 243
        Page 244
        Page 245
        Page 246
        Page 247
    Expression of feeding symptoms from pink hibiscus mealybug (Hemiptera: Pseudococcidae) by commercially important cultivars of hibiscus
        Page 248
        Page 249
        Page 250
        Page 251
        Page 252
        Page 253
        Page 254
    A new species of Dacnusini from Montecristo Island, with description of the preimaginal phases and venom apparatus of Antrusa curtitempus (Hymenoptera, Braconidae, Alysiinae)
        Page 255
        Page 256
        Page 257
        Page 258
        Page 259
        Page 260
    Two new genera of Tropidocephalini (Hemiptera: Fulgoroidea: Delphacidae) from Hainan Province, China
        Page 261
        Page 262
        Page 263
        Page 264
        Page 265
        Page 266
        Page 267
        Page 268
    A survey of hymenopterous larval-pupal parasitoids associated with Anastrepha fraterculus and Ceratitis capitata (Diptera: Tephritidae) infesting wild guava (Psidium guajava) and peach (Prunus persica) in the southernmost section of the Bolivian Yungas forest
        Page 269
        Page 270
        Page 271
        Page 272
        Page 273
        Page 274
        Page 275
    Feeding, reproduction, and development of the red palm mite (Acari: Tenuipalpidae) on selected palms and banana cultivars in quarantine
        Page 276
        Page 277
        Page 278
        Page 279
        Page 280
        Page 281
        Page 282
        Page 283
        Page 284
        Page 285
        Page 286
        Page 287
        Page 288
        Page 289
        Page 290
        Page 291
    Natural enemies of Brazilian peppertree (Sapindales: Anacardiaceae) from Argentina: their possible use for biological control in the USA
        Page 292
        Page 293
        Page 294
        Page 295
        Page 296
        Page 297
        Page 298
        Page 299
        Page 300
        Page 301
        Page 302
        Page 303
    Occurrence of Camponotus pennsylvanicus (Hymenoptera: Formicidae) in trees previously infested with Enaphalodes rufulus (Coleoptera: Cerambycidae) in the Ozark Mountains of Arkansas
        Page 304
        Page 305
        Page 306
        Page 307
        Page 308
    A new host association of Commoptera solenopsidis (Diptera: Phoridae) with the ant Pheidole dentata (Hymenoptera: Formicidae) and behavioral observations
        Page 309
        Page 310
        Page 311
        Page 312
        Page 313
    Reovirus-like sequences isolated from adult Asian citrus psyllid, (Hemiptera: Psyllidae: Diaphorina citri)
        Page 314
        Page 315
        Page 316
        Page 317
        Page 318
        Page 319
        Page 320
    Studies on the genus Onesia (Diptera: Calliphoridae) from China, with the descriptions of two new species
        Page 321
        Page 322
        Page 323
        Page 324
        Page 325
        Page 326
        Page 327
        Page 328
        Page 329
    Taxonomic notes on the genus Haphsa (Hemiptera: Cicadidae) with descriptions of two new species
        Page 330
        Page 331
        Page 332
        Page 333
        Page 334
        Page 335
        Page 336
        Page 337
    Descriptions of two new species of the Platylomia spinosa species group (Hemiptera: Cicadidae)
        Page 338
        Page 339
        Page 340
        Page 341
        Page 342
        Page 343
    Kneallhazia (=Thelohania) solenopsae infection rate of Pseudacteon curvatus flies determined by multiplex PCR
        Page 344
        Page 345
        Page 346
        Page 347
        Page 348
        Page 349
    Fecundity and mating propensity of Toxotrypana curvicauda (Diptera: Tephritidae) on an alternative host, Jacaratia mexicana (Caricaceae)
        Page 350
        Page 351
        Page 352
        Page 353
        Page 354
    Pupal development, longevity and behavior of Carmenta theobromae (Lepidoptera: Sesiidae)
        Page 355
        Page 356
        Page 357
        Page 358
        Page 359
        Page 360
        Page 361
    Sympatry of Orius insidiosus and O. pumilio (Hemiptera: Anthocoridae) in North Central Florida
        Page 362
        Page 363
        Page 364
        Page 365
        Page 366
    Description of late-instars of Bryothinusa koreana Ahn and Jeon (Coleoptera: Staphylinidae: Aleocharinae) by association of life stage based on DNA sequence data
        Page 367
        Page 368
        Page 369
        Page 370
        Page 371
        Page 372
        Page 373
    The effect of larval diet and sex on nectar nicotine feeding preferences in Manduca sexta (Lepidoptera: Sphingidae)
        Page 374
        Page 375
        Page 376
    Distribution of Verodes aequalis (Coleoptera: Zopheridae), a poorly known species from cloud forests of Mexico
        Page 377
        Page 378
    Marking blueberry maggot flies (Diptera: Tephritidae) with fluorescent diet for recapture studies
        Page 379
        Page 380
        Page 381
    Detection surveys and population monitoring for Pseudacysta perseae on avocados in Southern California
        Page 382
        Page 383
        Page 384
        Page 385
    The red palm weevil, Rhynchophorus ferrugineus (Coleoptera: Curculionidae), newly reported from Zhejiang, China and update of geographical distribution
        Page 386
        Page 387
    Sampling methods for Myllocerus undecimpustulatus undatus (Coleoptera: Curculionidae) adults
        Page 388
        Page 389
        Page 390
    Patterns of ant activity on Opuntia stricta (Cactaceae), a native host-plant of the invasive cactus moth, Cactoblastis cactorum (Lepidoptera: Pyralidae)
        Page 391
        Page 392
        Page 393
    First record of Erythmelus klopomor (Hymenoptera: Mymaridae) as a parasitoid of the avocado lace bug, Pseudacysta perseae (Heteroptera: Tingidae)
        Page 394
        Page 395
    Greenidea psidii (Hemiptera: Aphididae: Greenideinae) new invasive aphid in Costa Rica
        Page 396
        Page 397
        Page 398
    Corrected species identification of the predator Orius pumilio (Heteroptera: Anthocoridae) in a research colony
        Page 399
    Building roads and bridges for integrated pest management in Florida: a tribute to Madeline and Charles Mellinger of Glades Crop Care
        Page 400
        Page 401
        Page 402
        Page 403
    Book reviews
        Page 404
        Page 405
        Page 406
        Page 407
        Page 408
        Page 409
        Page 410
        Page 411
    Back Matter
        Page 412
        Page 413
Full Text

Shelly et al.: Methoprene and Mating Activity of Male Fruit Flies


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


The application of juvenile hormone (or chemical analogs, such as methoprene) to newly
emerged adult male fruit flies (Diptera: Tephritidae) represents a promising method to im-
prove the efficiency of the Sterile Insect Technique against economically important species.
This procedure has been shown to accelerate male sexual maturity in species with a long
pre-copulatory period, and could allow for release of sterile males at younger ages and a
greater release rate of sterile males overall. Topical application of methoprene has been
shown to enhance male mating competitiveness. The present study investigated the effect of
methoprene on maturation speed in males of the oriental fruit fly, Bactrocera dorsalis (Hen-
del) (a 'slow' maturing species) and the Mediterranean fruit fly, Ceratitis capitata (Wiede-
mann) (a 'fast' maturing species). For both species, newly emerged males were treated with
acetone containing methoprene (treated) or acetone alone (control). The mating propensity
of males was then monitored in non-competitive environments with mature females. Con-
trary to other studies, we found no evidence that methoprene accelerated male sexual activ-
ity in either a wild-like or mass-reared strain ofB. dorsalis or a mass-reared (genetic sexing
strain) of C. capitata. Possible explanations for these results are discussed.

Key Words: Bactrocera dorsalis, Ceratitis capitata, Sterile Insect Technique, hormone ther-
apy, Tephritidae, methoprene


La aplicaci6n de hormones juveniles (o andlogos quimicos, como metoprena) a machos adul-
tos de moscas de la fruta (Diptera: Tephritidae) reci6n emergidos represent un m6todo pro-
metedor para mejorar la eficiencia de la t6cnica del insecto est6ril (TIE) contra species de
importancia econ6mica. Este procedimiento ha de mostrado que se puede acelerar el tiempo
de la madurez sexual de machos en species con un period largo de pre-copulaci6n, y puede
permitir la liberaci6n de machos est6riles mas j6venes e incrementar el numero de machos
est6riles liberados. Se ha mostrado que aplicaciones de contact de metoprena mejora la
competividad reproductive de machos. Este studio investiga el efecto de metoprena sobre
la velocidad de madurez de los machos de la mosca oriental de la fruta, Bactrocera dorsalis
(Hendel) (una especie que madura lentamente) y la mosca mediterranea de la fruta (Cerati-
tis capitata Wiedemann), (una especie que madura rdpidamente). Por ambas species, ma-
chos reci6n emergidos fueron tratados con acetona que contiene metoprena (el grupo
tratado) o solo acetona (el grupo control). Se monitored la propensidad de los machos para
aparearse en ambientes no-competidos con hembras maduras. Al contrario de otros studios,
no encontramos ningun evidencia que metropena acelera la actividad sexual de machos en
cepas de B. dorsalis salvajes o criados masivamente o en una cepa de C. capitata que sepa-
raron los sexos gen6ticamente criados masivamente. Se discuten las posibles causes de estos

The Sterile Insect Technique (SIT) is widely
used to suppress or eradicate fruit fly pests, in
particular the Mediterranean fruit fly, Ceratitis
capitata (Wiedemann) (Hendrichs et al. 2002;
Klassen 2005; Enkerlin 2005). The process in-
volves the mass production of the target species
and the subsequent release of irradiated (sterile)
individuals in the environment to achieve mat-
ings between sterile males and wild females,
which yield infertile eggs and thus depress the re-
productive potential of the pest population. Ulti-
mately, the chief goal of the SIT is to produce max-
imally competitive sterile males at the lowest cost

over the shortest time interval. Because the
large-scale production and release of insects are
inherently expensive, there is a persistent need to
evaluate these processes and reduce associated
In recent years, many studies have been con-
ducted to assess the impact of adult diet (Aluja et
al. 2001; Shelly et. al 2005; Barry et al. 2007;
Yuval et al. 2007) and adult olfactory environ-
ment (Shelly et al. 2007) on the field perfor-
mance of sterile fruit fly males. Another promis-
ing avenue of investigation involves the applica-
tion of juvenile hormone (or synthetic analogs) to

Florida Entomologist 92(2)

newly emerged males. This procedure may result
in 2 significant benefits to SIT programs. First,
Teal et al. (2000) demonstrated that topical ap-
plication of juvenile hormone (or the synthetic
mimics methoprene and fenoxycarb) to young
adult males of the Caribbean fruit fly, Anas-
trepha suspense (Loew), dramatically acceler-
ated sexual maturation. For example, the mean
age at which all males in a replicate mated was
7 d for control males compared to 4 d for hor-
mone-treated males. Similar trends have also
been reported for males of the West Indian fruit
fly, A. obliqua (Macquart) and the Mexican fruit
fly, A. ludens (Loew) (Teal et al. 2007). Particu-
larly for species, like Anastrepha, with relatively
long pre-maturation intervals of 1-3 weeks de-
pending on strain and species (Aluja 1994), in-
creased rate of sexual development allows for
the earlier release of sterile males, which, in
turn, promotes a greater release rate of sterile
males into the environment. Methoprene treat-
ment may be less effective on fruit fly species
with shorter pre-maturation intervals, and Faria
et al. (2008) found no methoprene-mediated ef-
fect on sexual development in mass-reared, male
Mediterranean fruit flies, whose normal pre-cop-
ulatory period was only 2-4 d.
In addition, treating young males with metho-
prene may enhance their copulatory success.
Working with A. suspense, Pereira (2005) found
that protein-fed males treated with methoprene
achieved significantly more matings than sexu-
ally mature, protein-fed males that had not re-
ceived methoprene. However, a comparable study
on C. capitata yielded inconsistent results. Com-
paring mating success among 4 treatment groups
(methoprene treated or untreated and protein-fed
or deprived), Faria et al. (2008) found no effect of
methoprene in field cage trials (regardless of diet
regime) and in laboratory trials found a signifi-
cant effect of methoprene among protein-deprived
males but not protein-fed males.
The present study investigated the effect of
methoprene application on sexual maturation in
males of the oriental fruit fly, Bactrocera dorsalis
(Hendel) and the Mediterranean fruit fly. In Bac-
trocera species, males of wild populations have
lengthy pre-copulatory periods of 2-4 weeks
(Yang et al. 1994; Wee & Tan 2000), while males
of mass-reared strains mature in approximately
6-8 d (Vargas et al. 1984) Thus, the pre-matura-
tion interval in Bactrocera species is comparable
to that of many Anastrepha species, and we an-
ticipated that methoprene would effectively ac-
celerate sexual maturation in B. dorsalis, as re-
ported for the melon fly, B. cucurbitae (Coquil-
lett) (Haq et al. 2008). The work presented here
on C. capitata was undertaken to confirm the re-
sults of Faria et al. (2008), i.e., the absence in
this species of a methoprene effect on male sex-
ual maturation.


Study Insects

The majority of tests with B. dorsalis involved
flies from a laboratory ("wild-like") colony started
with 300-500 adults reared from mangos
(Mangifera indica L.) collected in Waimanalo,
Oahu. The colony was maintained in a screen
cage (l:w:h, 1.2 x 0.6 x 0.6 m) and provided a mix-
ture (3:1 wt:wt) of sugar (sucrose) and enzymatic
yeast hydrolysate. Water was supplied ad libi-
tum, and papayas (Carica papaya L.) were intro-
duced for oviposition. Infested papayas were held
over vermiculite, and the pupae were sifted from
vermiculite 16-18 d later. Adults used in the mat-
ing trials were separated by sex within 24 h of
eclosion and held in screen-covered, plastic buck-
ets (5-L volume; 100-125 individuals per bucket)
with ample food (the sugar-yeast hydrolysate
mixture) and water. Flies were held at 24-28C
and 60-90% RH and received natural and artifi-
cial light under a 12:12 (L:D) photoperiod. When
used in the study, these wild-like flies were 4-5
generations removed from the wild.
As shown below, tests with the wild-like strain
of B. dorsalis failed to show an effect of methop-
rene. Consequently, we performed a second set of
tests using B. dorsalis from a mass-reared strain
that had been maintained by USDA-ARS for over
20 years (D. McInnis, personal communication).
Females of this strain oviposited in perforated
tubes, and eggs were placed on an artificial diet
(Tanaka et al. 1969) for larval development.
Adults were handled in the same manner de-
scribed above and held under the same environ-
mental conditions.
For medfly, experiments were performed with
mass-reared males and wild-like females. Mass-
reared males were from a tsl (temperature sensi-
tive lethal) genetic sexing system (Vienna-7/Tol-
99) reared by the California Department of Food
and Agriculture's (CDFA) Hawaii Fruit Fly Rear-
ing Facility, Waimanalo, Oahu. In rearing this
strain, eggs are exposed to high temperature,
which selectively kills female embryos and allows
production and release of males only (Franz et al.
1994). Undyed pupae were obtained 2 d before
eclosion after irradiation in air at 150 Gy of
gamma radiation with a 137Cs source.
Female medflies were from a laboratory colony
started with 400-600 adults reared from infested
coffee berries, Coffea arabica L., collected near
Haleiwa, Oahu. The colony was maintained in
screen cages and provided with ample food
(sugar-yeast hydrolysate mixture), water, and ovi-
position substrate (perforated plastic vials con-
taining small sponges soaked in lemon juice).
Eggs were placed on standard larval medium
(Tanaka et al. 1969) in plastic containers over ver-
miculite for pupation. Adults were separated by

June 2009

Shelly et al.: Methoprene and Mating Activity of Male Fruit Flies

sex within 2 d of eclosion, well before reaching
sexual maturity at 7-9 d of age. Prior to testing,
the adult mass-reared male and wild-like female
medflies were maintained under the conditions
described above for B. dorsalis. When used in the
present study, the wild-like C. capitata females
were 5-6 generations removed from the wild.

Chemical Treatments

For both B. dorsalis and C. capitata, we followed
the methods of Teal et al. (2000) and Pereira (2005),
with 1 exception noted below. For treated males, me-
thoprene was applied topically at a dose of 5 pg in 1
pL of acetone per male. For control males, 1 pL of ac-
etone alone was applied per male. For both male
groups, the procedure was conducted within 24 h of
adult emergence. Males were immobilized by chill-
ing (5-7C for 5-10 min), and the chemical was ap-
plied via pipette onto the dorsal surface of the tho-
rax. Because the aforementioned authors did not
anesthetize the males in their work, we assessed
the possible effect of chilling on the chemical treat-
ment by performing additional mating trials for B.
dorsalis (wild-like strain) in which males were im-
mobilized in a net bag (i.e., without chilling) and the
chemicals were applied directly through the net-
ting. Based on data obtained with chilled males (see
below), non-chilled males were tested only at ages of
10 and 12 d. The effect of chilling was not investi-
gated in the mass-reared males ofB. dorsalis or C.

Mating Propensity

Mating propensity was monitored in labora-
tory cages for both study species. For the wild-like
strain ofB. dorsalis, we placed 15 males of a given
age and treatment and 20 mature females (18-25
d old) in plexiglas cages (30 x 30 x 40 cm, with a
screened opening on the top and a sleeve-covered
opening on 1 side). Males were tested at 2-d age
intervals between the ages of 2-18 d (i.e., when 2,
4, 6 ... 18 d old). Because mating occurs at dusk
in this species (Arakaki et al. 1984), the cages
were set up in mid-afternoon (1400-1600 h), and
the room lights were extinguished (natural light
entered through west-facing windows). Cages
were checked 3-4 h after sunset, and the number
of mating pairs was recorded for individual cages.
On a given test day, we set up equal numbers of
cages with treated or control males of a particular
age, with 6, 8, or 10 total cages established per
age group per day and 1-3 different age groups
tested on any given day. Fifteen cages (replicates)
were run in total for treated and control males in
each age group. In tests involving non-chilled
males ofB. dorsalis, 10 cages (5 treated, 5 control)
were established per age group per day on each of
2 d, yielding 10 total cages per treatment group
per age category.

Tests involving the mass-reared strain of B.
dorsalis were conducted in the same manner, ex-
cept that only 1 age group of males (5 d old) was
tested. Initially, to describe the age-dependent
mating activity of the mass-reared strain (inde-
pendent of any chemical treatment), we recorded
mating pairs in cages containing 15 untreated
males of a given age (i.e., when 2, 3, 4... 12 d old,
respectively) and 20 sexually mature females
(>14 d old). Fifteen cages were run in total for
each 1 d age group, with 1-5 age groups tested per
day and 3-5 cages tested per age group per day. As
shown below, untreated males from the mass-
reared strain first showed noticeable mating ac-
tivity when 5 d old, and consequently we pre-
dicted that an effect of methoprene treatment
would be detectable at this age. We tested 7 cages
for both treated and control 5 d old males on each
of 3 different days, yielding 21 replicates in total
for each treatment group. Females used in these
tests were all sexually mature (>14 d old).
For C. capitata, we placed 20 males of a given
age and treatment and 30 females (8-12 d old) in
screen cages (30 cm cubes). Males were tested at
1 d age intervals between the ages of 2-6 d. Mat-
ing occurs primarily in the morning in natural
populations in Hawaii (Shelly et al. 1994), conse-
quently the cages were set up in the morning
(0700-0800 h). Cages were checked periodically
for 4 h, and mating pairs were removed by gently
coaxing them into vials. On a given day, we set up
5 cages with treated males and 5 cages with con-
trol males of the same age. Males of the same age
were tested over 5 days, yielding a total of 25
cages (replicates) per treatment group.

Statistical Analysis

For both the wild-like strain ofB. dorsalis and
C. capitata, the effects of male age and chemical
treatment were analyzed by a Friedman's test, a
2-way ANOVA by ranks, because assumptions re-
garding normality and homoscedasticity of the
data were not met. In our analyses, male ages
were blocks, and the presence or absence of meth-
oprene in the acetone applied to the males were
treatments. For each species, the test was per-
formed with mean numbers of matings for the dif-
ferent male age-chemical treatment combinations
(following Daniel 1990; test statistic X2 with df =
1). Thus, in our case, the Friedman test (i) ranked
the (mean) number of matings obtained by
treated and control males, respectively, within
each age group and then (ii) compared the
summed ranks across all age intervals (thus re-
moving the effect of the blocking variable, i.e.,
male age).
For trials involving non-chilled B. dorsalis
males from the wild-like strain, pair wise compar-
isons between the number ofmatings observed for
treated and control males were made with the

Florida Entomologist 92(2)

Mann-Whitney test (test statistic T) for each of
the 2 age groups considered (i.e., 10 and 12 d old),
where n, = n = 10 in both cases. For each of these
age groups, data gathered on all 4 sets of males
(chilled or non-chilled and treated or control)
were compared by a Kruskal-Wallis test, where df
= 3 in both cases (test statistic H).
For trials involving the 5 d old males from the
mass-reared strain of B. dorsalis, pair wise com-
parisons between the number of matings ob-
served for treated and control males were made
by the Mann-Whitney test, where n = n = 21.


For the wild-like strain ofB. dorsalis, we found
no significant effect of methoprene treatment on
mating propensity (X2 = 1.70, P = 0.21, Fig. 1).
Over both treatment groups, male mating fre-
quency increased dramatically between 10-16 d of
age and then appeared to level off between 16-18
d of age (Fig. 1).
Data gathered for non-chilled males from the
wild-like strain of B. dorsalis similarly revealed
no effect of methoprene on male mating activity.
For 10-d-old, non-chilled males, individuals
treated with methoprene obtained an average of
1.9 (+0.2) matings per replicate (13% of males)
compared to 1.7 (+0.2) (11% of males) for control
males (T = 98.5, P = 0.65). Likewise, for 12 d, old
non-chilled males, individuals treated with meth-
oprene obtained an average of 2.9 (+0.3) matings
per replicate (19% of males) compared to 2.6
(0.4) (17% of males) for control males (T = 114.5,
P = 0.49). Within each of these age groups, there
were no significant differences in mating propen-

--- Melhoprene
- No methopene

I -

E 20

sity among chilled/treated, chilled/control, non-
chilled/treated, and non-chilled/control males (10
d: H = 0.53, P = 0.91; 12 d: H = 0.25, P = 0.97).
The mating activity of untreated males from
the mass-reared B. dorsalis strain was nil at 2 or
3 d, very low at 4 d, old, but then increased rap-
idly between 5-9 d, after which it appeared to
level off (Fig. 2). Based on these data, we selected
5 d old males to test for a possible effect of meth-
oprene treatment on mating frequency. Among 5-
d-old males, individuals treated with methoprene
(when < 1 d old) obtained an average of 2.5 (+0.3)
matings per replicate (17% of males) compared to
2.1 (+0.3) (14% of males) for control males (T =
476.5, P = 0.54).
As with B. dorsalis, methoprene had no appar-
ent effect on the mating activity of C. capitata
males (X2 = 1.80, P = 0.19, Fig. 3). For both treat-
ment groups, male mating frequency increased
noticeably between the ages of 2-4 d but was sim-
ilar for the ages of 4-6 d old.


In the present study, topical application of the
juvenile hormone mimic methoprene to newly
emerged adult males produced no significant ac-
celeration of sexual maturation in either B. dorsa-
lis or C. capitata. This result was obtained for 2
strains of B. dorsalis, 1 wild-like and the other
mass-reared, and for 1 mass-reared strain of C.
The absence of a methoprene effect on matura-
tion in mass-reared C. capitata was consistent
with the study ofFaria et al. (2008) and the notion
that the already short pre-copulatory period (2-4

0 2 4 5 8 10 12 14 16 1t 20 0 2 4 6 8 10 12 14
Male age (d) Male age (d)

Fig. 1. Mating frequency of Bactrocera dorsalis
males from a wild-like strain treated with acetone con-
taining or lacking methoprene. Values along ordinate
represent the mean (+1 SE) proportion of males mating
per cage (replicate) at a given age interval. Fifteen
males were observed per cage, and 15 cages were run
per treatment group per age group.

Fig. 2. Mating frequency of untreated (no chemical
treatment) Bactrocera dorsalis males from a mass-
reared strain. Values along ordinate represent the mean
(1 SE) proportion of males mating per cage (replicate)
at a given age interval. Fifteen males were observed per
cage, and 15 cages were run per treatment group per
age group.

June 2009

Shelly et al.: Methoprene and Mating Activity of Male Fruit Flies

0- species resulted exclusively in horn-less adults
-- Nometne (Moczek & Nijhout 2002), but see Zera (2007) for
a an alternate explanation. Thus, the demonstra-
tion of a methoprene effect on male sexual matu-
o. ration in one species of Bactrocera, namely B. cu-
curbitae (Haq et al. 2008), does not imply that
. / such an effect exists in all Bactrocera species.
Second, the absence of a methoprene effect in
0 B. dorsalis might have been an inadvertent conse-
quence of our experimental protocol. As noted, un-
like previous studies, we anaesthetized B. dorsa-
0- lis males via chilling. However, this procedural
difference appears inconsequential, because sup-
S 2 3 4 5 elementary tests with non-chilled males likewise
ale age (d) failed to demonstrate a methoprene effect. Alter-
natively, although a methoprene effect has been
Fig. 3. Mating frequency of Ceratitis capitata males demonstrated inAnastrepha species in small lab-
n a genetic sexing strain treated with acetone con- oratory cages (Teal et al. 2000; Pereira 2005), it is
ing or lacking methoprene. Values along ordinate possible that the use of such cages for B. dorsalis
resent the mean (+1 SE) proportion of males mating somehow obscured this effect. For example, if
replicate at a given age interval. Twenty males were mating activities (e.g., pheromone release) of sex-
erved per cage, and 25 cages were run per age group, ually mature males are inhibited in laboratory
cages, then differences in mating "drive" between
methoprene-treated and control males may be
essens the opportunity for additional develop- minimized and undetectable. However, the obser-
ntal acceleration via methoprene application. vation that age-related mating frequency curves
ever, our failure to detect a methoprene effect were nearly identical between methoprene-
B. dorsalis was unexpected, because males of treated and control B. dorsalis males suggests the
Species have a relatively long maturation in- unlikely scenario that a cage effect, if existing,
val and all previous studies (Teal et al. 2000, was so severe that it completely eliminated be-
7; Pereira 2005) ofAnastrepha species with a havioral differences between treated and control
ilar life history have demonstrated a pro- males. Still, as Haq's (2008) study ofB. cucurbitae
nced increase in male sexual development fol- was performed in field cages, the possibility exists
ing methoprene treatment. In addition, Haq that a significant methoprene effect might yet be
08) reported that methoprene significantly detected for B. dorsalis if tested under more nat-
rtrned the male nrp-cpnnlnlatrv interval in an- ural conditions.

other Bactrocera species, B. cucurbitae.
There may be 2 main reasons to account for the
absence of a methoprene effect in B. dorsalis.
First, the effect of methoprene treatment of 5 pg
methoprene in 1 pL of acetone per male (Teal et
al. 2000) on the rate of male sexual maturation
may vary among tephritid species and exogenous
application of juvenile hormone or chemical mim-
ics may have different effects in different species.
For example, among aphids, application of juve-
nile hormone has been shown to promote brac-
hyptery in some species but not in others (Zera &
Denno 1997). Likewise, the application of juvenile
hormone (JH III) to diapausing females was
found to promote the development of vitellogenic
(yolky) oocytes in one species of heteropteran (Ad-
ams et al. 2002) but not in another (Shinoda et al.
1996). A variable effect of juvenile hormone treat-
ment has even been reported among populations
within a single species: application of an interme-
diate dose of juvenile hormone to male larvae of a
US population of the beetle Onthophagus taurus
Schreber resulted in the production of horned
adults, whereas the same dose applied to male
larvae of an Australian population of the same


We thank Peter Teal for providing the chemicals
used to treat the insects.


ADAMS, T. S., FILIPI, P. A., AND YI, S.-X. 2002. Effect of
age, diet, diapause and juvenile hormone on oogene-
sis and the amount of vitellogenin and vitellin in the
twospotted stink bug, Perillus bioculatus (Het-
eroptera: Pentatomidae). J. Insect Physiol. 48: 477-
ALUJA, M. 1994. Bionomics and management ofAnas-
trepha. Annu. Rev. Entomol. 39: 155-178.
Effect of adult nutrition on male sexual performance
in four neotropical fruit fly species of the genus
Anastrepha (Diptera: Tephritidae). J. Insect Behav.
14: 759-775.
Arakaki, N., Kuba, H., AND Soemori, H. 1984. Mating
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J. G. 2007. Effect of adult diet on longevity of sterile


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Mediterranean fruit flies (Diptera: Tephritidae).
Florida Entomol. 90: 650-655.
DANIEL, W. W. 1990. Applied Nonparametric Statistics.
2nd ed. PWS-Kent, Boston.
ENKERLIN, W. R. 2005. Impact of fruit fly control pro-
grammes using the sterile insect technique, pp. 651-
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[eds.], Sterile Insect Technique: Principles and Prac-
tice in Area-Wide Integrated Pest Management.
Springer, Dordrecht, The Netherlands.
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survival, maturation and sexual performance of
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Improved stability of sex-separation strains for the
Mediterranean fruit fly, Ceratitis capitata. Genome
37: 72-82.
PEREIRA, R. M. C., AND STAUFFER, C. 2008. Effect of
juvenile hormone analog and dietary protein on sex-
ual competitiveness of male melon fly, Bactrocrea cu-
curbitae. Oral presentation, 3rd Research Coordina-
tion Meeting of the International Atomic Energy
Agency: Improving Sterile male Performance in
Fruit Fly Sit Programs. Valencia, Spain.
KERLIN, W. 2002. Medfly area-wide sterile insect
technique programmes for prevention, suppression
or eradication: the importance of mating behavior
studies. Florida Entomol. 85: 1-13.
KLASSEN, W. 2005. Area-wide integrated pest manage-
ment and the sterile insect technique, pp. 39-68 In V.
A. Dyck, J. Hendrichs, and A. S. Robinson [eds.],
Sterile Insect Technique: Principles and Practice in
Area-Wide Integrated Pest Management. Springer,
Dordrecht, The Netherlands.
MOCZEK, A. P., AND NIJHOUT, H. F. 2002. Developmen-
tal mechanisms of threshold evolution in a poly-
phonic beetle. Evol. Devel. 4: 252-264.
PEREIRA, R. 2005. Influence of a juvenile hormone ana-
log and dietary protein on male Caribbean fruit fly,
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system of the Mediterranean fruit fly, Ceratitis capi-
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87: 470-481.
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diet and methyl eugenol on the mating success of
males of the oriental fruit fly, Bactrocera dorsalis.
Florida Entomol. 88: 307-313.

B., AND MCINNIS, D. 2007. Aromatherapy on a large
scale: exposing entire holding rooms to ginger root
oil increases the mating competitiveness of sterile
males of the Mediterranean fruit fly. Entomol. Exp.
Appl. 123: 193-201.
1996. Vitellogenins and vitellins in the bean bug,
Riptortus clavatus (Hemiptera: Alydidae): purifica-
tion, immunological identification, and induction by
juvenile hormone. Arch. Insect Biochem. Physiol. 31:
TO, R. 1969. Low-cost larval rearing medium for
mass-production of oriental and Mediterranean fruit
flies. J. Econ. Entomol. 62: 967-968.
2000. Mating experience and juvenile hormone en-
hance sexual signaling and mating in male Caribbe-
an fruit flies. Proc. Nat. Acad. Sci. USA 97: 3708-
LER, T. C., AND OLSON, S. 2007. Improving the effica-
cy of the sterile insect technique for fruit flies by in-
corporation of hormone and dietary supplements
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Insect Technique: Principles and Practice in Area-
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Ann. Entomol. Soc. American 77: 651-656.
WEE, S. L., AND TAN, K. H. 2000. Sexual maturity and
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YANG, P., CAREY, J. R., AND DOWELL, R. V. 1994. Com-
parative demography of two cucurbit-attacking fruit
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phritidae). Ann. Entomol. Soc. American 87: 538-545.
AND SHELLY, T. 2007. Breakfast of champions or kiss
of death? Survival and sexual performance of pro-
tein-fed, sterile Mediterranean fruit flies (Diptera:
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ZERA, A. J. 2007. Endocrine analysis in evolutionary-de-
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June 2009

Taylor & Gil: Aradidae from Louisiana


'Illinois Natural History Survey, 1816 South Oak Street, Champaign, Illinois 61820
E-mail: sjtaylor@illinois.edu

2Department of Entomology, Louisiana State University, Baton Rouge, Louisiana 70803-1710
E-mail: sgill@lsu.edu

The Aradidae of Louisiana are poorly known, with only 5 species reported from the state. We
examined 251 adult flat bugs from Louisiana in the Louisiana State Arthropod Museum,
confirming the presence of 4 species (Aradus fallen Stal, Acaricoris ignotus Harris and
Drake, Notapictinus aurivilli (Bergroth), and Mezira sayi Kormilev) and adding 14 more in
4 subfamilies (Aneurinae: Aneurus fiskei Heidemann, Aneurus pygmaeus Kormilev; Aradi-
nae:Aradus acutus Say, Aradus aequalis Say,Aradus kormilevi Heiss,Aradus ornatus Say,
Aradus robustus Uhler; Carventinae: Neoproxius gypsatus (Bergroth); Mezirinae: Neurocte-
nus pseudonymus Bergroth, Neuroctenus simplex (Uhler), Mezira emarginata (Say), Mezira
froeschneri Davidova-Vilimova et al., Mezira granulata (Say), and Mezira lobata (Say)) to
the state's fauna. Habitats recorded for these species are discussed.
Key Words: flat bug, distribution, habitat, Aneurus, Neuroctenus, Neoproxius

Los Aradidae de Louisiana han sido poco estudiados y solo cinco species son conocidas para
en el estado. Al examiner 251 chinches adults de Louisiana pertecientes al Louisiana State
Arthropod Museum, confirmamos la presencia de cuatro species (Aradus fallen Stal,Aca-
ricoris ignotus Harris y Drake, Notapictinus aurivilli (Bergroth), y Mezira sayi Kormilev) y
aiadimos catorce mas, representando cuatro subfamilias (Aneurinae: Aneurus fiskei Heide-
mann, Aneurus pygmaeus Kormilev; Aradinae: Aradus acutus Say, Aradus aequalis Say,
Aradus kormilevi Heiss, Aradus ornatus Say, Aradus robustus Uhler; Carventinae:
Neoproxius gypsatus (Bergroth); Mezirinae: Neuroctenus pseudonymus Bergroth, Neurocte-
nus simplex (Uhler), Mezira emarginata (Say), Mezira froeschneri Davidova-Vilimova et al.,
Mezira granulata (Say), y Mezira lobata (Say)), a la fauna del estado. Los habitat registrados
para estas species son discutidos.

Translation provided by the authors.

In America North of Mexico, 127 species of
Aradidae are known (Davidova-Vilimova et al.
1996; Froeschner 1988; Kormilev & Froeschner
1987; Vaisrhelyi 1994). In contrast, the flat bug
fauna of Louisiana is poorly known, with only 5
species (Aradus fallen Stal, Aradus quadrilinea-
tus Say, Acaricoris ignotus Harris and Drake, No-
tapictinus aurivilli (Bergroth), and Mezira sayi
Kormilev) reported before the present study
(Davidova-Vilimova et al. 1996; Froeschner
1988). Herein, we report new state records and
confirmations of flat bugs from Louisiana on the
basis of an examination of material collected from
21 parishes (Fig. 1).


We examined 251 adult specimens housed in
the Louisiana State Arthropod Museum (LSAM),
all of which were identified by the senior author.
Abbreviated terms in specimen data are indicated
in brackets at first usage.


Subfamily Aneurinae Douglas and Scott, 1865
Aneurus Curtis, 1825

The review of north and central American
Aneurus species by Kormilev (1968) has been su-
perseded by that of Picchi (1977), which stands as
the definitive work for this genus in North Amer-
ica, and provides keys to species.

Aneurus fiskei Heidemann

Aneurus fiskei occurs throughout much of east-
ern North America from Michigan and Connecti-
cut south to Georgia, Texas and Mexico (Froe-
schner 1988; Picchi 1977). Its occurrence in Loui-
siana is not unexpected.
EAST BATON ROUGE Parish: Baton Rouge
[BR], Place Du Plantier Apts., Yellow sticky board
trap in Cypress tree,18-IV-1986, E. G. Riley (1 ;);
BR, at mercury vapor and black light [MV & BL],

Florida Entomologist 92(2)

Fig. 1. Map of Louisiana, showing 21 parishes from
which we report Aradidae in the present study. Parish:
A-Calcasieu, B-Catahoula, C-Caddo, D- East Baton
Rouge, E-East Feliciana, F-Franklin, G-Grant, H-Iber-
ville, I-Jackson, J-Lafayette, K-Lincoln L-Livingston,
M-Natchitoches, N-Orleans, O-Pointe Coupee, P-
Rapides, Q-St. James, R-St. Landry, S-St. Martin, T-
West Feliciana, U-Webster. U. S. EPA level III ecore-
gions adapted from Daigle et al. (2006).

8-V-1987, D. A. Rider (1 9); BR, under bark of
dead limb, 26-1-1991, M. S. Strother (1 9); BR, on
pecan trunk, 14-X-1979, Andeab Berhane (2 9, 1

Aneurus pygmaeus Kormilev

Aneurus pygmaeus has a southern distribu-
tion, having been reported from Georgia and Flor-
ida to Texas, from Arkansas, and also from Cali-
fornia (Froeschner 1988; Picchi 1977; Taylor &
McPherson 1989a), but it has not previously been
reported from Louisiana.
NATCHITOCHES Parish: Kisatchie National
Forest [KNF], 3128'59"N 9304'28"W, barking
tree, 5-IV-2003, A. Cline (1 9, 1 6).

Subfamily Aradinae Amyot and Serville, 1843
Aradus Fabricius, 1803

This large genus is represented in North
America north of Mexico by 84 species (Froe-
schner 1988; Vaisrhelyi 1994). Parshley's (1921)
monograph is still the definitive work and useful
for identifications, although more than 20 new
species have been described from the region since
that time. An adult male collected from a pitfall
trap at Palustris Experiment Station (Evangeline
Parish) is not included here as it requires further
study, and may represent a new species nearAra-
dus inornatus Uhler and Aradus blaisdelli Van

Aradus acutus Say

Aradus acutus is a common, widespread spe-
cies reported from Alaska and Maine south to Cal-
ifornia, Texas, and Florida (Froeschner 1988;
Parshley 1921; Matsuda 1977), but it has not been
previously reported from Louisiana.
CADDO Parish: 14-III-1944, R. L. Stone (2 Y Y
,2 66); LINCOLN Parish: Ruston, 26-IV-1941,
W. W. Wirth (1 Y); WEBSTER Parish: Shongaloo,
23-XII-1939, Newsom (5 Y 2 66 ).

Aradus aequalis Say

Aradus aequalis is relatively infrequently col-
lected, but occurs from Maine, Quebec, and On-
tario south to Illinois and Virginia. It also has
been reported from Texas (Froeschner 1988;
Parshley 1921; Matsuda 1977). Its occurrence in
Louisiana constitutes a significant range exten-
sion into the southeastern United States, and the
coloration of this specimen deviates slightly from
the descriptions given by Say (1832) and Parshley
(1921). Parshley (1921) gives the coloration of this
species as "Brown; broad lateral margins and pos-
terior region of pronotum, margins of scutellum
at middle, basal expansions of corium and neigh-
boring veins, posterior margins of connexival seg-
ments, and inner margins of genital lobes, yellow-
ish; membrane with irregularly reticulate pale
markings; legs yellow, femora and tibiae with
broad brown band at middle." The Louisiana spec-
imen has little if any yellow on the pronotum, and
only the apical third of the scutellum is yellowish,
not the margins of the scutellum at the middle.
The brown bands on the tibiae and femora indi-
cated by Parshley (1921) are quite faint.
WEST FELICIANA Parish: Feliciana Pre-
serve 6 mi. ESE of St. Francisville, 3047'N
9115'W, flight intercept trap, 10-30-III-1996,
D. Pashley(1 9).

Aradus fallen Stil

Aradus fallen is the most widespread aradid
species in the Western Hemisphere. It occurs from
British Columbia and Connecticut, throughout
much of the United States and into Mexico and
Brazil (Froeschner 1988; Parshley 1921). We con-
firm its presence in Louisiana with these records.
Wildlife Management Area [WMA], MV & BL, 23-
VI-1990, V. L. Moseley (1 9); KNF, 2 mi. E Red
Bluff Cgd, MV & BL, 17-18-VI-1988, C. B. Barr
(1 9).

Aradus kormilevi Heiss

Aradus kormilevi is widespread in North
America, ranging from British Columbia and
New Brunswick south to California, Texas, and

June 2009

Taylor & Gil: Aradidae from Louisiana

Florida (Froeschner 1988; Heiss 1980), but has
not previously been reported from Louisiana.
Three Louisiana specimens, 2 brachypterous fe-
males and 1 macropterous male, reside in the
Louisiana State Arthropod Museum. This species,
along with Aradus antennalis Parshley which oc-
curs in the western United States and Canada, is
unusual among the North American Aradus spe-
cies in that it is thought to feed on living pines,
whereas the remaining 82 species occur on dead
trees and are thought to be mycetophagous
(Usinger & Matsuda 1959).
EAST BATON ROUGE Parish: BR, under
bark, 24-IX-1976, M. T. Smith (1 2); BR, LSU
Campus, 3024'49"N 9110'47"W, sweeping vege-
tation, 24-III-2005, D. Bustamante (1 6); NAT-
CHITOCHES Parish: KNF, Red Dirt WMA, 19-
20-IV-1985, E. G. Riley (1 2).

Aradus ornatus Say

Aradus ornatus is reported from Indiana to
Pennsylvania, south to Virginia (Parshley 1921),
as well as New York (Drake 1923) and Georgia
(Froeschner 1988). Emergence traps captured 24
adults (11 2 2, 13 6 6) of this distinctive but un-
commonly collected species in 2 areas of West Fe-
liciana Parish from 17 samples of dead woody de-
bris (limbs) with obvious signs of decay, collected
from the ground, with samples comprised either
of 1-2.5 inch diameter limbs probably dead for
more than 1 year, or of 2.5- to 10-inch diameter
limbs with loose bark and fungal staining but still
structurally sound (Mike Ferro, personal commu-
nication, Jan 2008). Blatchley (1926) reports 1
male of this species "taken from beneath bark of
an oak log" in Indiana. This is the first record ofA.
ornatus from Louisiana, and the second from the
southeastern United States.
WEST FELICIANA Parish: Feliciana Pre-
serve: 3047.607'N 9115.272'W, 25-III to 22-IV-
2007, LN14A-CWD Rear 1, M. Ferro (1 9);
30046.647'N 9115.210'W, 25-III to 22-IV-2007,
LN22A-CWD Rear 1, M. Ferro (1 9, 1 6);
30046.647'N 9115.210'W, 25-III to 22-IV-2007,
LN22C-CWD Rear 1, M. Ferro (1 6);
30046.647'N 9115.210'W, 25-III to 22-IV-2007,
LN23A-CWD Rear 1, M. Ferro (1 9);
30047.760'N 9115.356'W, 25-III to 22-IV-2007,
LN32A-CWD Rear 1, M. Ferro (2 66);
30047.760'N W91015.356'W, 25-III to 22-IV-2007,
LN32C-CWD Rear 1, M. Ferro (1 9); Tunica
Hills WMA: 30056.261'N 91030.522'W, 25-III to
22-IV-2007, LP11A-CWD Rear 1, M. Ferro (1 9);
30056.261'N 91030.522'W, 25-III to 22-IV-2007,
LP12A-CWD Rear 1, M. Ferro (1 6); 30056.261'N
91030.522'W, 25-III to 22-IV-2007, LP14B-CWD
Rear 1, M. Ferro (1 9); 30'55.826'N 91'31.260'W,
25-III to 22-IV-2007, LP22A-CWD Rear 1, M.
Ferro (1 6); 30055.826'N 91031.260'W, 25-III to
22-IV-2007, LP22B-CWD Rear 1, M. Ferro (1 6);

30055.826'N 91031.260'W, 25-III to 22-IV-2007,
LP24A-CWD Rear 1, M. Ferro (1 6); 30055.826'N
91031.260'W, 25-III to 22-IV-2007, LP24C-CWD
Rear 1, M. Ferro (4 9 2 66); 30059.702'N
91032.716'W, 25-III to 22-IV-2007, LP31B-CWD
Rear 1, M. Ferro (1 6); 3059.702'N 9132.716'W,
25-III to 22-IV-2007, LP32B-CWD Rear 1, M.
Ferro (1 6); 30059.702'N 91032.716'W, 25-III to
22-IV-2007, LP33C-CWD Rear 1, M. Ferro (1 6);
30059.702'N 91032.716'W, 25-III to 22-IV-2007,
LP34B-CWD Rear 1, M. Ferro (1 2).

Aradus robustus Uhler

Aradus robustus occurs from Maine, Ontario,
and Nebraska south to Texas and Florida
(Froeshner 1988; Leschen & Taylor 1987; Parsh-
ley 1921), and is one of the most commonly col-
lected Aradus species in North America. Parsh-
ley (1921) designated a new variety of this spe-
cies, which is treated by Kormilev & Froeschner
(1987) and Froeschner (1988) as a subspecies,
Aradus robustus insignis Parshley, 1921, in ac-
cordance with the 1985 International Code of
Zoological Nomenclature. However, Parshley
(1921) noted that A. r. insignis was a color vari-
ant of "no geographical significance." Although
some of the Louisiana material has the reddish-
brown abdomen characteristic of A. r. insignis,
we have refrained from using the subspecific
designation because these animals need further
study to clarify the taxonomic relevance of the
observed variation. This is the first published
record for A. robustus in Louisiana, but it's pres-
ence is not unexpected.
EAST BATON ROUGE Parish: BR, 29-1-1977,
D. W Brown (1 2); EAST FELICIANA Parish: 3-
VI-1979, C. B. Barr (1 6); FRANKLIN Parish:
Macon Ridge Res. Sta., Flight intercept trap, 21-
III-2003, R. H. Gable, (1 2); IBERVILLE Parish:
St. Gabriel, in dead log, 11-IV-1991, M. S.
Strother, (1 2,2 66).

Subfamily Carventinae Usinger, 1950

Of the 39 genera (Kormilev & Froeschner
1987; Grillo Ravelo 1988) of Carventinae only 2
are known to occur in North America north of
Mexico (Acaricoris, Neoproxius). In contrast to
the other subfamilies, for which most genera are
typically encountered under bark or in flight or
light traps, nearly all of Carventinae in the
United States for which habitat data are re-
corded have been collected from Berlese or litter
samples (Taylor & Lewis 1989; Taylor & McPher-
son 1989b, and see below). A key to the apterous
Aradidae of North and South America is pro-
vided by Drake & Kormilev (1958), and to the
genera of neotropical Carventinae is provided by
Kormilev & Doesburg (1977) and by Grillo Rav-
elo (1988).

Florida Entomologist 92(2)

Acaricoris Harris and Drake, 1944

This is an entirely wingless genus whose mem-
bers occur primarily in the tropical and subtropi-
cal Americas, with only 2 species, Acaricoris flori-
dus Drake and A. ignotus occurring north of Mex-
ico (Harris & Drake 1944; Drake 1957).

Acaricoris ignotus Harris and Drake

Acaricoris ignotus occurs from Texas and Ar-
kansas to Georgia and Florida (Froeschner 1988;
Taylor & Lewis 1989; Taylor & McPherson
1989b). It previously has been reported from Lou-
isiana (Drake & Kormilev 1958), and additional
records are provided here.
WEST FELICIANA Parish: 3047'N 9115'W,
mixed pine hdwd Berlese, [no date], J. L. Fass-
bender (1 2); Tunica Hills WMA, 3056'32"N
9129'75"W, Berlese, 1-II-2003, S. A. Gil, B. W.
Collier, A. M. Sanchez (1 Y).

Neoproxius Usinger and Matsuda, 1959

Usinger & Matsuda's (1959) subgenus
Neoproxius was elevated to generic status by
Kormilev & Froeschner (1987). This genus is dis-
tributed across South and Central America, with
only 2 species in America north of Mexico:
Neoproxius schwarzii (Heidemann), which is
known from Cuba (Grillo Ravelo 1988) and
Tampa, Florida, and Neoproxius gypsatus (Ber-
groth). The 2 North American species are keyed
by Blatchley (1926), while Usinger & Matsuda
(1959) key 4, and Kormilev (1982b) keys 11 of the
13 Neoproxius species listed by Kormilev & Froe-
schner (1987).

Neoproxius gypsatus (Bergroth)

Neoproxius gypsatus is known from Florida,
Panama, and Venezuela (Blatchley 1926; Froe-
schner 1988; Taylor & McPherson 1989b). The oc-
currence ofN. gypsatus in Louisiana is a notable
range extension to the west, and suggests that the
species ought to be encountered in Mississippi
and Alabama.
EAST BATON ROUGE Parish: 5-IV-1979, C.
B. Barr (1 9); WEST FELICIANA Parish: Felici-
ana Preserve 6 mi. ESE of St. Francisville,
3047'N 9115'W, litter sample from ravine, 18-
XI-1995, C. Carlton (4 9 9, 3 66).

Subfamily Mezirinae Oshanin, 1908

Five genera of Mezirinae occur in America
north of Mexico, 3 of which are reported here.
The remaining 2 genera are each represented by
a single species in the United States: Aphle-
boderrhis pubescens (Walker) 1873 is reported

from Texas and South America (Froeschner
1988), and Nannium pusio Heidemann 1909 is
known only from Ohio. A specimen from Felici-
ana Preserve, West Feliciana Parish, may repre-
sent an undescribed species, and is not included

Neuroctenus Feiber, 1860

Six species of Neuroctenus occur in America
north of Mexico. Four of them are keyed by Blatch-
ley (1926) and Torre-Beuno (1939), and a key to 5
species is provided by Kormilev (1982a).
Vasarhelyi (1994) was apparently unaware of Kor-
milev's (1982a) key when he described Neurocte-
nus unistellatus Vasarhelyi from central Texas.

Neuroctenus pseudonymus Bergroth

Neuroctenus pseudonymus Bergroth is re-
ported from Indiana and the District of Columbia
southwest to Texas and Tennessee (Froeschner
1988). Its occurrence in Louisiana represents a
southeastern range extension. It is distinguished
by the large, rounded paratergites of the female
(Kormilev 1982a) and the distinct transverse
ridge on connexivum VII of the males.
WEST FELICIANA Parish: Tunica, 8-IV-1977,
M. L. Israel (1 9).; Tunica Hills W of Weyanoke,
MV & BL, 18-V-1985, C. B. Barr (1 6); Tunica
Hills W of Weyanoke TIS R4W, on dead log Quer-
cus nigra, 8-V-1986, J. E. Barr (1 6); 3055'N
9130'W, hardwood Berlese, 18-V-1999, J. L. Fass-
bender (1 ).

Neuroctenus simplex (Uhler)

Neuroctenus simplex is the most widespread
and common of the North American Neuroctenus
species, occurring from Montana and Maine south
to Texas, Florida, and Cuba (Froeschner 1988).
Neuroctenus unistellatus is closely related to N.
simplex (Vasarhelyi 1994), and the ranges of
these 2 species in North America need reevalua-
tion. Neuroctenus simplex appears to be common
and widespread in Louisiana.
CADDO Parish: 14-III-1948, R. L. Stone (1 6);
Shreveport, on rotting log, 12-II-1991, V. L. Mose-
ley (1 9); CALCASIEU Parish: Charles City
[=Lake Charles], 14-IX-1993, W. W. Wirth (1 6);
EAST BATON ROUGE Parish: BR, 23-1-1983, D.
A. Rider (5 9 9, 3 66); BR, 22-11-1976, D. W.
Ragsdale (1 9, 16); BR, Oak Log, 30-XI-1978, W.
E. Blust (1 9, 2 66); GRANT Parish: Gray
Branch, SW of Breezy Hill off Hwy. 123, under
bark, 28-VII-1982, C. B. Barr (1 6); WEBSTER
Parish: Shongaloo, 23-XII-1939, Newsom (1 6);
WEST FELICIANA Parish: Tunica Hills W of
Weyanoke, TIS R4W, at night on dead log Quer-
cus nigra, 8-V-1986, J. E. Barr (1 9, 1 6).

June 2009

Taylor & Gil: Aradidae from Louisiana

Notapictinus Usinger and Matsuda, 1959

This genus comprises 40 species, occurring pri-
marily in South and Central America (Kormilev
& Froeschner 1987). Only N. aurivilli occurs in
the United States.

Notapictinus aurivilli (Bergroth)

Notapictinus aurivilli has been reported from
Florida, Georgia, and Bayou Sara (West Feliciana
Parish), Louisiana (Blatchley 1926; Froeschner
1988; Taylor & McPherson 1989b). Here we pro-
vide 1 new parish record for Louisiana.
ST. LANDRY Parish: 2-XI-1971, D. C. Herzog
(1 2, 1 6); WEST FELICIANA Parish: Feliciana
Preserve 6 mi. ESE of St. Francisville, 3045'N
9116'W, Berlese, 6-III-1996, C. Carlton (1 6).

Mezira Amyot and Serville, 1843

Ten species of this large genus (more than
160 species) are known from America north of
Mexico. Blatchley (1926) keys 4 species from
the eastern United States. Seven of the United
States species of Mezira are keyed by Usinger
(1936) and Torre Bueno (1939). Kormilev's
(1971) key to 89 New World Mezira species in-
cludes 6 from America north of Mexico. Kormi-
lev (1982c) keyed 3 and Davidova-Vilimova et
al. (1996) keyed 4 small Mezira species. How-
ever, the 3 relatively recent descriptions
(Mezira smith Kormilev, Mezira froescheneri
Davidova-Vilimova et al. 1996, and M. sayi)
make previous distributional information ques-
tionable, especially for the 4 smaller Mezira

Mezira emarginata (Say)

Mezira emarginata has been recorded from 9
states, from North Carolina and Florida west to
Nevada and California, as well as Mexico (Froe-
schner 1988), but it has not previously been re-
ported from Louisiana. It's occurrence in Louisi-
ana is not unexpected.
CADDO Parish: 4-1-1948, R. L. Stone (1 9, 2
66); EAST BATON ROUGE Parish: Baker, 30-
VIII-1934, H. M. Harris (4 2); BR, 29-1-1977,
D. W. Brown (5 9 9, 5 66); BR, 5-XII-1982, D. A.
Rider (14 2 2, 6 66); BR, in house, 10-III-1927,
O. W. Rosewall (1 2); IBERVILLE Parish: St.
Gabriel Res. Farm, under bark of dead sugar-
berry Celtis laevigata Wild., 11-IV-1991, V.
Moseley and M. S. Strother (1 2, 2 66); OR-
LEANS Parish: New Orleans, LA Nature Cen-
ter, Blk.lt.trap, 25-IV to 4-V-1984, C. B. Barr (1
2); ST. JAMES Parish: 18-IV-1972, (1 2); ST.
LANDRY Parish: duff, 25-III-1980, L. D. New-
som(1 2).

Mezira froeschneri Davidova-Vilimova, Taylor and

Mezira froeschneri was described from speci-
mens collected via Berlese sampling of hardwood
litter in northern Florida (Davidova-Vilimova et
al. 1996), the single specimen recorded here adds
a second state to the species distribution, and its
occurrence in a pitfall traps is also suggestive of a
ground-dwelling, probably leaf litter, habitat.
Based on this new record, it also should be found
in Mississippi and Alabama.
WEST FELICIANA Parish: 15 mi. E. St. Fran-
cisville, Feliciana Preserve, pitfall trap, 2-VI-
1995, D. Colby and D. Landau (1 2).

Mezira granulata (Say)

Mezira granulata has been widely reported
from 15 primarily eastern states and Mexico, and
occurs from Maryland and Florida west to Illinois
and Texas, as well as Arizona and Mexico (Davi-
dova-Vilimova et al. 1996; Froeschner 1988; Tay-
lor & McPherson 1989a). Further evaluation of
M. granulata's range since the description of M.
froeschneri is warranted. Although its occurrence
in Louisiana is not unexpected, it is somewhat
surprising that only 1 specimen from the state is
present in the LSAM, given that the species oc-
curs in Florida and Arkansas (Davidova-Vilimova
et al. 1996; Taylor & McPherson 1989a).
CATAHOULA Parish: Sicily Island Hills
WMA, 7-V-1987, C. B. Barr (1 2).

Mezira lobata (Say)

Mezira lobata occurs primarily in eastern
North America, from New York and Florida west
to Illinois, Missouri, and Texas, as well as Califor-
nia, Canada, and Mexico (Froeschner 1988). This
is the first record of M. lobata from Louisiana.
EAST BATON ROUGE Parish: BR, 29-1-1977,
D. W Brown (3 92 1 6); BR, oak log, 30-XI-1978,
W. E. Blust (1 2); BR, Mississippi R. levee, under
bark, 31-VIII-1996, T Clarke (2 66); 1.2 mi S of
Central on LA Hwy. 3034, C. B. Barr (1 6); Ben
Hur Res. Farm, dead log, 12-III-1999, R. Costello (1
S); Bluebonnet Swamp, under bark, 20-IV-2005, J.
Hamm & S. Tewari (2 9 9); Zachary, Sunshine Rd.,
under bark, 13-XI-1992, J. T McBride (1 9, 5 6 6);
IBERVILLE Parish: St. Gabriel, in dead log, 11-IV-
1991, M. S. Strother (3 9 9, 2 66); St. Gabriel,
sweeping vegetation, 16-IV-2005, L. Eisenberg (1
9); LAFAYETTE Parish: Lafayette, 28-III-1967, D.
A. Neal (1 9); LIVINGSTON Parish: Hogbranch
Crk at US 190, 30-1-1983, D. A. Rider (3 9 9, 3
6 6); ORLEANS Parish: 12-III-1982, G. L. Stone (1
9); POINTE COUPEE Parish: 18-IV-1975, D. F.
Lower (1 9); RAPIDES Parish: Camp Beaure-
gard, "@ large," 12-IX-2004, E. Ott (1 6); WEST
FELICIANA Parish: Elm Park, 24-IV-1966 (1 9);

Florida Entomologist 92(2)

Feliciana Preserve 6 mi. ESE of St. Francisville,
3047'N 9116'W, flight intercept trap, 1-10-IV-
1996, C. Carlton (1 ); Feliciana Preserve, 3047'N
9116'W, 9-10-IV-2005, ENTM 4005 Class (1 2).

Mezira sayi Kormilev

Mezira sayi has been reported from 10 states,
from North Carolina and Florida west to Illinois
and Texas (Davidova-Vilimova et al. 1996; Froe-
schner 1988; Taylor & McPherson 1989a), and was
reported from Louisiana by Davidova-Vilimova et
al. (1996) without specific locality data. Here, we
reexamine the same, and additional, material,
providing distribution data. This is the most com-
monly encountered of the small Mezira species in
the state. It is most often found under the bark.
CALCASIEU Parish: Sam Houston St. Pk.,
3019'N 9316'W, sifting, 11-II-2003, A. R. Cline
and S. T. Dash (1 2, 1 6); CATAHOULA Parish:
Sicily Island Hills WMA, 7-V-1987, C. B. Barr (7 Y
2, 8 66); EAST BATON ROUGE Parish: Place
Du Plantier Apts., flight trap, 15-III-1984, E. G.
Riley (1 Y); BR, 5-III-1991, M. Sean Strother (2 9
9, 1 6); Bluebonnet Swamp, under bark, 4-II-
2001, J. E. Light (5 66); Bluebonnet Swamp,
sweeping, 12-IV-2005, P Jayasimha (1 6); Green-
well Springs, 3029'44"N, 9059'34"W, under bark,
4-II-2001, S. A. Gil (4 9 9, 4 66); Zachary, Sun-
shine Rd., under bark, 13-XI-1992, J. T. McBride
(2 9 9, 2 66); FRANKLIN Parish: N.E. Research
Sta. Winnsboro, sweeping soybean, 23-IX-1994, C.
F. Grymes and D. E. Fairbanks (1 ); IBERVILLE
Parish: St. Gabriel, near river road, under bark,
13-IX-1996, J. Adamczyk (1 6); JACKSON Parish:
Schoolhouse Spring, R1W T17 NNE Sec.12, 6 mi.
N Eros 14-15-IV-1988, C. B. Barr (1 6); ST. LAN-
DRY Parish: 5 mi E. of Washington on Hwy 103,
on dead log, 3-XI-1990, V. L. Moseley (4 9 9, 3
6 ); under bark, 3-XI-1990, M. S. Hicks (1 6); ST.
MARTIN Parish: 4 mi. S. of Belle River, sticky
trap in swamp, 13-X-1994, D. A. Duerr II (1 6);
WEST FELICIANA Parish: Feliciana Preserve 6
mi. ESE of St. Francisville, 3047'N 9115'W,
flight intercept trap, 10-30-III-1996, D. Pashley (2
6 6); Feliciana Preserve, 15 mi. East of St. Fran-
cisville, under bark, 28-IX-1996, G. Turano (1 9);
Feliciana Preserve, 3047'N 9115'W, 9-10-IV-
2005, ENTM 4005 Class (1 9); Tunica Hills .5 mi
W of Weyanoke, MV & BL, 15-VIII-1986, C. B.
Barr (1 6); Tunica Hills W ofWeyanoke, TIS R4W,
collected under bark, 3-V-1985, C. B. Barr (2 6 6);
Tunica Hills W of Weyanoke, TIS R4W, on dead
log Quercus nigra, 8-V-1986, J. E. Barr (6 9 9, 5
66); Tunica Hills area, under bark, 28-IX-1996,
G. Turano (1 ).

Habitat Associations

Usinger & Matsuda (1959, pp. 35-42) summa-
rized the literature on habitat data for Aradidae,

including host tree records and flight records.
Subsequent work on the North American aradid
fauna has resulted in very modest additions to
this information (Deyrup & Mosley 2004; Taylor
& Lewis 1989; Taylor & McPherson 1989a,b;
McPherson & Weber 1981). Some additional in-
formation can be gleaned from the material exam-
ined in this study. For all taxa combined, nearly
half of the specimens examined (47% or 119 indi-
viduals) had no habitat data whatsoever on the
specimen labels, while the remaining 52% (132
specimens) had some minimal habitat data. For
specimens with habitat data, 59% (78 specimens)
were associated with trees. Half (39) of these were
found under bark: A. fiskei (1), A. pygmaeus (2),
M. emarginata (3), M. lobata (10), and M. sayi
(23), and this is where Aradidae are most fre-
quently encountered in temperate North Amer-
ica. The remaining specimens associated with
trees were mostly recorded from in, on, or under
dead wood-descriptions that likely include sub-
cortical collections. These include A. robustus (3),
M. lobata (7), M. sayi (18), N. pseudonymus (1),
and N. simplex (6). With the exception ofM. emar-
ginata, all of these species previously have been
recorded from under bark or in association with
dead wood (Leschen & Taylor (1987):A. robustus;
Picchi (1977): A. pygmaeus; Taylor & McPherson
(1989a): M. lobata, M. sayi, N. simplex; Taylor &
McPherson (1989b): M. sayi; summarized in
Usinger & Matsuda (1959):A. fiskei, M. lobata, N.
pseudonymous, and N. simplex)).
Of the remaining records with habitat data, 4
specimens were obtained by sweeping: A. kormi-
levi (sweeping vegetation [1]), M. lobata (sweep-
ing vegetation [1]), and M. sayi (sweeping [1],
sweeping soybean [1]). Two specimens were col-
lected from sticky traps: A. fiskei (yellow sticky
board trap in cypress tree [1]),M. sayi (sticky trap
in swamp [1]). Davidova-Vilimova et al. (1996)
also report M. sayi from a sticky trap in Georgia,
and Picchi (1977) recordedA. fiskei in flight in the
District of Columbia. Three specimens were taken
in pitfall traps, including 2, anAradus sp. and an
apterous Carventinae, which may be undescribed
species, while the third, M. froeschneri, is other-
wise known only from Berlese litter samples in
northern Florida (Davidova-Vilimova et al. 1996).
Specimens from habitats given as "Berlese,"
"duff," "sifting," and "litter sample" includedA. ig-
notus [2], N. gypsatus [7], N. pseudonymus [1], N.
aurivilli [1], M. emarginata [1], and M. sayi [2],
and these litter habitats are probably very impor-
tant at least for A. ignotus, which was also re-
corded from similar habitat by Taylor & Lewis
(1989) and for N. aurivilli and N. gypsatus, which
were both reported from Berlese litter samples by
Taylor & McPherson (1989b). Aradus ornatus is
notable in that all 24 specimens (18% of the 132
specimens with habitat data) were recovered by
emergence traps to collect them from fallen, de-

June 2009



S(1) -i -i -- --1

Subfamily / Species
Subfamily / Species ^


Aneurus fiskei
Aneurus pygmaeus

Aradus acutus
Aradus aequalis
Aradus fallen
Aradus kormilevi
Aradus ornatus
Aradus robustus

Acaricoris ignotus
Neoproxius gypsatus

Neuroctenus pseudonymus
Neuroctenus simplex
Notapictinus aurivilli
Mezira emarginata
Mezira froescheneri
Mezira granulata
Mezira lobata
Mezira sayi

Total Number of Species



+ * *

+ + +

x + + + + *

+ . 3
+ + 1
S + 1 -"

1 0
+ 1

+ + 2
+ + 6

* *

+ +

x +
x + + + + + + +

+ + + + + + + + +

14 2 2 3 8 1 2 1 4 1 1 1 1 3 2 1 1 1 3 1 10

+ 2
+ 1




Florida Entomologist 92(2)

caying twigs and limbs. Two species were col-
lected from flight traps, including A. robustus
(flight intercept trap [1]), and M. sayi (flight trap
[1], flight intercept trap [2]). Finally, 2 species
were captured at lights: A. fiskei (at mercury va-
por and black light [1]) and M. emarginata (black
light trap [1]).

Distribution of Collections

We have documented 14 new state records
and confirmed 4 of the 5 species of Aradidae pre-
viously reported from Louisiana (Table 1). This
material includes the first record of the subfam-
ily Aneurinae, and first records for the genera
Aneurus, Neuroctenus, and Neoproxius from the
state. Specimens are recorded from 21 parishes,
only about one third of the 64 parishes in Louisi-
ana. Sampling in parishes in direct contact with
the Gulf of Mexico largely has not been done.
Coverage of ecoregions (Daigle et al. 2006) is
fairly good in the upper two thirds of the Missis-
sippi Alluvial Plain, the South Central Plains,
and the Mississippi Valley Loess Plains (Fig. 1).
Few records are reported from the Western Gulf
Coastal Plain, and none come from either the
South Central or Southeastern plains. We sus-
pect that additional species of Aradidae should
be found in the state, especially with focus on un-
der-sampled regions and habits, especially leaf


We thank Victoria M. Bayless and Christopher E.
Carlton (Department of Entomology, Louisiana State
University, Baton Rouge) for facilitating SJT's visit to
the Louisiana State Arthropod Museum and for the loan
of material. Mike Ferro graciously provided specific
habitat data forA. ornatus.


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MATSUDA, R. 1977. The Insects and Arachnids of Cana-
da. Part 3. The Aradidae of Canada. Hemiptera: Ara-
didae. Canadian Dept. Agric. Publ. 1634: 1-116.
MCPHERSON, J. E., AND WEBER, B. C. 1981. Seasonal
flight patterns of Hemiptera in a North Carolina
black walnut plantation. 6. Tingidae and Aradidae.
Gt. Lakes Entomol. 14: 137-140.
PARSHLEY, H. M. 1921. Essay on the American species
of Aradus (Hemiptera). Trans. American Entomol.
Soc. 47: 1-106.
PICCHI, V. D. 1977. A systematic review of the genus
Aneurus of North and Middle America and the West
Indies (Hemiptera: Aradidae). Quaest. Entomol. 13:
TAYLOR, S. J., AND LEWIS, S. P. 1989. Notes on the hab-
itat and distribution ofAcaricoris ignotus Harris and
Drake (Heteroptera: Aradidae). Southwest. Nat. 34:
TAYLOR, S. J., AND MCPHERSON, J. E. 1989a. State
records and confirmations of Arkansas flat bugs

June 2009

Taylor & Gil: Aradidae from Louisiana

(Heteroptera: Aradidae). Gt. Lakes Entomol. 22: 19-
TAYLOR, S. J., AND MCPHERSON, J. E. 1989b. Distribu-
tional records of some Florida Aradidae (Het-
eroptera). Florida Entomol. 72: 265-267.
TORRE-BUENO, J. R. DE LA. 1939. A synopsis of the
Hemiptera-Heteroptera of America north of Mexico.
Entomol. Americana 19: 141-304.

USINGER, R. L. 1936. Studies in the American Aradidae
with descriptions of new species (Hemiptera). Ann.
Entomol. Soc. America 29: 490-516.
USINGER, R. L., AND MATSUDA, R. 1959. Classification of
the Aradidae. British Museum, London. 410 pp.
VASARHELYI, T. 1994. Two new flat bug species from
North America (Heteroptera, Aradidae). J. New York
Entomol. Soc. 102: 86-90.

Florida Entomologist 92(2)

June 2009


'USDA-ARS-CMAVE at Center for Biological Control, FAMU, Tallahassee, FL 32308

2USDA-ARS-CPMRU, Tifton, GA 31794


Long term trapping studies of the invasive moth Cactoblastis cactorum (Berg) were con-
ducted at various latitudes from Puerto Rico to South Carolina. Three flight periods per year
were identified at the 5 temperate sites studied, which covered the majority of the infested
range on mainland United States. In general, the 3 flight periods across a latitudinal gradi-
ent from south Florida to central, coastal South Carolina were a spring flight during Feb-
May, a summer flight during Jun-Aug, and a fall flight during Sep-Nov. At any 1 site, each
flight period lasted about 2 months. In the tropical areas of the Florida Keys and a Carib-
bean Island, the insect exhibited overlapping generations. Previous studies of this insect (as
a biological control agent) report 2 flight periods per year in its native range of Argentina
and its introduced range of Australia and South Africa. A synthetic pheromone-baited trap
was a good indicator of generational time, and we suggest that trapping assays in these ar-
eas will likely identify 3 generations rather than 2. Initiation and timing of the 3 genera-
tional flights has importance in the current United States and Mexico monitoring program
for presence and expansion of this invasive pest, development of mapping programs to iden-
tify monitoring windows and management efforts with the Sterile Insect Technique.

Key Words: flight phenology, Cactoblastis cactorum, invasive pest, pheromone trapping


Se realizaron studios sobre la capture a largo plazo de la polilla invasora Cactoblastis cac-
torum (Berg) en varias latitudes de Puerto Rico hasta el estado de Carolina del Sur (EEUU).
Se identificaron tres periods de vuelo por ano en los 5 sitios estudiados de clima templado,
que abarco la mayoria del rango infestado en la area continental de los Estados Unidos. En
general, los 3 periods de vuelo que cruzan el gradiente latitudinal desde el sur de Florida
hasta la costa central del Carolina del Sur fueron los vuelos de primavera de febrero a mayo,
de verano de junio hasta agosto, y de otono de septiembre a noviembre. En todos los sitios,
el period de vuelo duro como 2 meses. En las areas tropicales de los Cayos de Florida y en
una Isla de Caribe, el insecto present generaciones que se superponen. Studios hechos an-
teriormente sobre este insecto (como un agent de control biol6gico) informan de 2 periods
de vuelo por ano en su rango native de Argentina y en su rango introducido de Australia y
Sudafrica. Una trampa cebada con una feromona sint6tica fue un buen indicador del period
de generaci6n, y sugerimos que los ensayos usando trampas en estas areas probablemente
identificaran 3 generaciones en vez de 2. El inicio y la coordinaci6n del tiempo de los 3 vuelos
de los generaciones tiene importancia en los programs actuales de monitoreo para la pre-
sencia y expansion de esta plaga invasora en los Estados Unidos y M6xico, asi como en el de-
sarrollo de programs de mapas para identificar ventanas de monitoreo, y esfuerzos de
manejo de esta plaga con la t6cnica del insecto est6ril.

The cactus feeding pyralid Cactoblastis cac-
torum (Berg) has the notoriety as both a beneficial
introduced insect and a harmful alien insect pest.
In the 1920s the moth was first used as a classical
biological control agent against non-native
prickly pear cactus (Opuntia spp.) in Australia
(Dodd 1940; Mann 1970). Because of the success
of this insect to help reclaim some 24 million ha of
prickly pear infested lands in Australia (Raghu &
Walton 2007), the insect was used as a biological
control agent in other parts of the world (Pettey
1948; Fullaway 1954; Zimmermann et al. 2004).

Initially, C. cactorum was used to manage non-na-
tive species of prickly pear. However, with the in-
troduction of C. cactorum into the Caribbean Is-
land of Nevis in 1957 (Simmonds & Bennett
1966), the moth was used to attack native Opun-
tia species. At the time, concern for native species
and biodiversity was superseded by the need to
help subsistence farmers reclaim pastures over-
run with prickly pears (Zimmermann et al. 2000).
Through natural and/or human-aided dispersal,
the moth made its way throughout most of the
Caribbean and, unfortunately, was found in the

Hight & Carpenter: Flight Phenology of Cactoblastis

Florida Keys in 1989 (Dickle 1991). Introduction
and establishment of C. cactorum into Florida
from the Caribbean occurred at least twice (Simo-
nsen et al. 2008) with human intervention (Pem-
berton 1995; Simonsen et al. 2008). The moth's
spread and attack of native Opuntia spp. in the
United States threatens rare and endangered
species (Stiling et al. 2000), ornamental interests
(Mahr 2001), and agricultural systems in the
United States and Mexico (Sober6n et al. 2001;
Vigueras & Portillo 2001; Garrett 2004).
Studies have been conducted to identify the
distribution (Stiling 2002) and dispersal (Hight et
al. 2002; Solis et al. 2004) ofC. cactorum on North
American prickly pear species. Since the initial
find of C. cactorum in 1989, the moth has spread
rapidly along the Gulf and Atlantic Coasts and
now occurs as far west as Petit Bois and Horn Is-
lands off the coast of Mississippi (unpublished
data), and as far north as Bull Island, South Caro-
lina (R. Westbrooks, USGS, personnel observa-
tion, 2004). The rate of spread away from coastal
areas into the Florida Peninsula has been slower,
but the occurrence of the insect in the interior is
relatively common as far north as central Florida
(Orlando) (unpublished data).
Early monitoring techniques consisted of vi-
sual surveys for infested plants. A more precise
monitoring system for C. cactorum presence and
dispersal relies on an insect trap baited with a
synthetic 3-component synthetic sex pheromone
(Heath et al. 2006). Traps are unpainted "wing-
type" traps positioned 1.5 m above the ground
(Bloem et al. 2005a). This pheromone trap has be-
come incorporated into the operational program
to monitor and detect C. cactorum in the United
States and Mexico.
South African growers have practiced insecti-
cide applications and insect removal, either egg-
sticks or pads infested with larvae, since the
1950s (Zimmermann et al. 2004). Recent evalua-
tions of insecticides have revealed promising com-
pounds for targeted applications (Bloem et al.
2005b; Zimmermann 2008), but in areas with ex-
tensive native prickly pear this technique can
damage native species of Lepidoptera and be cost
prohibitive. Biological control of C. cactorum has
been reviewed by Pemberton & Cordo (2001a,
2001b), and additional studies are being con-
ducted in Argentina at the USDA's South Ameri-
can Biological Control Laboratory (SABCL) (G.
Logarzo, personal communication, 2008). How-
ever, biological control will not prevent the insect
from expanding its range in North America. The
primary aim of managing C. cactorum in the
United States is to stop the westward spread of
this insect into the rich prickly pear areas of the
Southwestern United States and Mexico (Stiling
2002). The Sterile Insect Technique (SIT) is an
area-wide approach that might prevent the
spread of C. cactorum and eradicate isolated pop-

ulations beyond the leading edge (Bloem et al.
2007). Radiation biology for this insect was exam-
ined by Carpenter et al. (2001a, 2001b), overflood-
ing ratios were assessed by Hight et al. (2005),
and a field validation of the technology was re-
viewed in Bloem et al. (2007).
One contradictory aspect of the biology of C. cac-
torum has been the number of annual generations
for the insect. Early studies in South America iden-
tified 2 generations in Argentina (Dodd 1940). In
Australia (Dodd 1940), South Africa (Pettey 1948),
and generally worldwide (Mann 1969), C. cactorum
is considered to have 2 generations per year. How-
ever, Mann (1969) reported a "partial third genera-
tion" in warmer parts of Australia. Distribution of C.
cactorum in the United States spans a wide latitu-
dinal range from the semi-tropical Florida Keys to
temperate South Carolina. This study was con-
ducted to determine the number and timing of C.
cactorum generations along a latitudinal gradient
in the eastern United States from semi-tropical to
temperate regions. Knowledge gained from this
study has implications for timing of various man-
agement techniques and monitoring efforts, and
predictions ofC. cactorum population dynamics.


Seven study sites where prickly pear cactus
plants were infested with C. cactorum were se-
lected. Two sites (Edisto Beach, SC and Lower
Sugarloaf Key, FL) were near the latitudinal ex-
tremes of known C. cactorum infestations in
North America, and 4 were intermediate sites
(Fig. 1). The seventh site was located in Guanica,
Puerto Rico. All study sites were located less than
8 km from the coast except at Sebring, FL, which
is about 100 km from either coast. Trapping stud-
ies were conducted from 2005-2008 (Tables 1 and
2, Figs. 2-8).

Sebring, FL

Lower Sugarloaf Key, FL .-

Gunca, PR

Fig. 1. Map of the southeastern United States and
Puerto Rico identifying the locations of Cactoblastis cac-
torum trapping study sites (map not to scale).

Florida Entomologist 92(2)

June 2009


Study Site and Years of Trapping

Sebring, FL St. Marks, FL Pensacola Beach, FL Jekyll Island, GAEdisto Beach, SC
Flight Parameter 2005-2007 2005-2008* 2005-2008* 2005-2006 2005-2006

Spring start flight 6.7 (0.62) 10.5 (0.72) 9.5 (1.00) 10.0 (1.19) 15.0 (0.84)
mid-Feb early-Mar early-Mar early-Mar mid-Apr
Spring end flight 20.0 (0.44) 23.0 (0.45) 21.5 (0.38) 23.5 (0.59) 21.5 (1.03)
mid-May early-Jun late-May early-Jun late-May
Spring duration of flight 13.7 (0.71) 12.5 (0.69) 12.0 (1.03) 13.5 (2.06) 6.5 (0.59)
Between flight duration 2.7 (0.44) 3.2 (0.61) 3.8 (0.35) 2.0 (0.00) 6.5 (1.03)
Summer start flight 23.0 (0.00) 26.2 (0.49) 25.2 (0.49) 25.5 (0.59) 28.0 (0.00)
early-Jun late-Jun late-Jun late-Jun mid-Jul
Summer end flight 33.3 (0.44) 34.0 (0.45) 33.8 (0.35) 35.0 (0.84) 35.0 (0.84)
mid-Aug late-Aug mid-Aug late-Aug late-Aug
Summer duration of flight 10.7(0.44) 7.8 (0.56) 8.5 (0.38) 9.5 (1.03) 5.0 (0.84)
Between flight duration 1.0 (0.00) 3.0 (0.45) 2.2 (0.35) 1.5 (0.59) 6.5 (1.03)
Fall start flight 34.7 (0.44) 37.0 (0.45) 36.0 (0.45) 36.5 (1.03) 39.5 (0.59)
late-Aug mid-Sep early-Sep mid-Sep late-Sep
Fall end flight 46.3 (0.44) 46.3 (0.71) 47.7 (0.85) 47.0 (1.19) 47.5 (1.03)
mid-Nov mid-Nov late-Nov late-Nov late-Nov
Fall duration of flight 11.7 (0.44) 9.3 (0.44) 12.0 (0.81) 10.5 (0.59) 8.0 (0.84)
Between flight duration 12.5 (0.59) 17.0 (1.00) 12.7 (1.30) 15** 17.0 (0.00)

*Last data collected for 2008 trap collections ended with "fall start flight".
**Only a single year estimate.

Except for Pensacola Beach, FL, 2 Pherocon 1-
C Wing traps (Trece Incorporated, Salinas, CA)
were run at all sites, each trap separated by 5 m
and placed within 10 m of C. cactorum-infested
Opuntia spp. The Pensacola Beach site was sam-
pled with 70 pherocon 1-C wing traps; each trap
positioned near an infested C. cactorum, about
250 m apart, distributed over an area approxi-
mately 9 x 0.3 km. Traps were baited with a red
rubber septa lure impregnated with a synthetic
female sex pheromone (Suterra, LLC, Bend, OR).
Male moths entering the wing traps were cap-
tured when they contacted the sticky bottom.
Collaborators assisted with the trap service ef-
forts at all sites except the St. Marks, FL and
Pensacola Beach sites, which were serviced by one
of the authors (SDH). Traps were serviced by col-
laborators or 1 of the authors (SDH) weekly at all
sites. However, the exact day of the week for trap
service varied among sites. If a moth was present,
trap bottoms were removed and sent to one of the
authors (SDH) for identification. Lures were re-
placed every 2 weeks.
Four flight period parameters were measured:
beginning of flight, end of flight, duration of flight,

and duration of gap between flights. Beginning of
a flight period was defined as the week traps be-
gan catching male C. cactorum after the traps had
no weekly captures. End of a flight period was de-
fined as the week that traps caught males before
the weekly trap catch dropped to zero. On occa-
sion, the weekly catch between flight periods did
not drop to zero, but an obvious low point in trap
catch (ending week of flight) was followed the
next week by a sustained weekly increase in trap
catch (beginning week of flight). In this instance,
the duration of the gap between flights was iden-
tified as 1 week. Cumulative degree-day (CDD)
values for the calendar years of 2005, 2006, and
2007 were calculated from National Weather Ser-
vice data from weather stations nearest each
trapping site. The degree-day model was set with
an upper threshold of 45C and a lower threshold
of 10C. Statistical analyses were conducted on
data collected during 2005-2007 for 5 sites that
exhibited 3 discrete generations: Sebring, St.
Marks, Pensacola Beach, Jekyll Island, and
Edisto Beach. The relationships between CDD of
these sample sites and parameters associated
with the temporal occurrence of the 3 generations

Hight & Carpenter: Flight Phenology of Cactoblastis

S4 -e-2006 CatchI


,1 5 9 13 17 21 25 29 33 37 41 45 49
Week of Year (200)
X 3


the independent variable (PROC GLM) (SAS In-C
H 1

-stitt sttisti i t

.following dependent variables: week of the year
i.. that marked the beginning of a flight; week of the
z X X X 1 5 9 13 17 21 25 29 33 37 41 45 49
year that marked the end of a flight; dur(2)ation

o flights. Flight phenology of Cactoblastis cactorum at
the Gunica, Puerto Rico weekly t rapping sites was examdurined by PROC
S20 Cl2006.

1 5(SAS 17ttt 219 c29
y '1 ~ di (dependent variables) on those sample sites were
a " M examined by regression analysis with CDD set as
m ^the independent variable (PROC GLM) (SAS In-
5 [stitute 1989). The statistical model included the
r following dependent variables: week of the year
^F 'S ^ X XX X that marked the beginning of a flight; week of the
cc m C CO year that marked the end of a flight; duration
(weeks) of a flight; and duration (weeks) between
flights. The relationship between latitude and
Sthe CDD of the trapping sites was examined by PROC
oo CORR (SAS Institute 1989).


Y) ~ 200 C6.h
& 0
C stitute 1989). They sttsical mekly incl d sthe

Sm oya -th- 2005Caroh
I^ ^^ ~~6----t --


fis Th a t ap

CDD o e p site wa e xaminedby*

C COR (A Ii 19 8 9)

c i S Z!"a Fig. 3A Flight phenology of Cactoblastis cactorum at
Y disin the Lower Sugarloaf Key, Florida weekly trapping site
t during-2005.

Florida Entomologist 92(2)

-4-2005 Catch
---2006 Catch
A 2007 Date


1 5 9 13 17 21 25 29 33 37 41 45 49
Week of Year (2005 2007)

Fig. 4. Flight phenology of Cactoblastis cactorum at
the Sebring, Florida weekly trapping site during 2005-

nated spring, summer, and fall), except at the
Florida Keys and Guanica sites (Figs. 2-8). The
Lower Sugarloaf Key site was too remote for con-
sistent weekly service and the population level of
C. cactorum too low to identify a consistent pat-
tern. In addition, this site was damaged in 2005
by Hurricanes Katrina (26 Aug), Rita (20 Sep),
and Wilma (24 Oct) and traps were not visited
for several weeks. The Guanica site was substi-
tuted for the Lower Sugarloaf Key site in 2006.
Traps were serviced weekly at Guo nica, but
again, the population was low. No distinct gener-
ational flight pattern emerged for either tropical
site. Both sites exhibited overlapping genera-
tions since adult males were caught in traps vir-


1 5 0 13 17 21 25 29 33 37 41 45 49
Week of Year (2005 2008)

Fig. 5. Flight phenology of Cactoblastis cactorum at
the St. Marks, Florida weekly trapping site during 2005

1000 2005 Catch
--2006 Catch
SA 2007 Catch
800 2008 Catch

= 400


1 5 9 13 17 21 25 29 33 37 41 45 49
Week of Year (2005 2008)

Fig. 6. Flight phenology of Cactoblastis cactorum at
the Pensacola Beach, Florida weekly trapping site dur-
ing 2005-2008.

tually throughout the year (Figs. 2 and 3). Trop-
ical sites were dropped from the study after 1
year of trapping, and were not included in phe-
nological analysis of male C. cactorum flight
times or duration.
Sites outside Puerto Rico and Lower Sugar-
loaf Key had large populations of C. cactorum.
The highest yearly average of male C. cactorum
captured was at the Pensacola Beach site (4,437
25) (mean SEM), followed by Jekyll Island
(498 4.8), Sebring (441 2.7), St. Marks (324
4.8), and finally Edisto Beach (241 2.0). At
different years across the latitudinal gradient,
male C. cactorum were flying at generally the
same periods within each study site (Table 1).

--2005 Catch
-- 200] Calch

Week of Year (2005 2006)

Fig. 7. Flight phenology of Cactoblastis cactorum at
the Jekyll Island, Georgia weekly trapping site during

June 2009

Hight & Carpenter: Flight Phenology of Cactoblastis

* 20


1 5 9 13 17 21 25 29 33 37 41 45 49
Week of Year (2005 2006)
Fig. 8. Flight phenology of Cactoblastis cactorum at
the Edisto Beach, South Carolina weekly trapping site
during 2005-2006.

In general, the start and end of each of the 3
flight periods was most similar for the St.
Marks, Pensacola Beach, and Jekyll Island
sites. This was not unexpected since the lati-
tudes of these 3 sites were more similar to one
another than either of the other 2 sites (Table
2). The site with the shortest duration of male
flight period and the longest number of weeks
between flight periods was the most northern
site (Edisto Beach), while the site with the long-
est duration of male flight was generally the
most southern site (Sebring) (Table 1). The
shortest duration between flight periods was at
the southern site, except between the Spring
and Summer flights when Jekyll Island was the
shortest duration (Table 1).
Latitude and CDD were significantly related
to each other (R2= 0.98). There were significant
relationships between the CDD of the sample
site and the initiation of the spring, summer,
and fall flights (Fig. 9). However, no significant
relationships were observed between the CDD
of the sample site and the termination of the
spring, summer, or fall flight. The durations of
the spring and summer flights were not signifi-
cantly influenced by the CDD of the sample
sites, but the duration of the fall flight de-
creased significantly as the CDD of the sample
site decreased (F = 10.240, df = 1, 11; P <
0.0085; Y = 0.00215x 7.435; R2= 0.4821). Con-
versely, there was a significant increase in the
duration of the period between the summer and
fall flights (F = 11.198, df = 1, 11; P < 0.0652; Y
= -0.00216x + 18.449; R= 0.5045) with decreas-
ing CDD. No significant relationships were ob-
served between the CDD of the sample site and
the duration of the period between the fall and
spring flights or the duration of the period be-
tween the spring and summer flights.

a Spring
* Summer
* Fall


6000 7000 8000 9000
Cumulative annual degree day

Fig. 9. Linear regression of the influence of annual
cumulative degree-days on the week of the year that
Cactoblastis cactorum males begin their spring (F =
6.841, df= 1, 11;P <0.02402;Y= -0.00317x + 32.789;R2
=0.3834), summer (F = 41.746, df = 1, 11; P < 0.0001; Y
= -0.00242x + 43.193; R2= 0.7914), and fall (F = 32.698,
df = 1, 11; P < 0.0001; Y = -0.00247x + 54.451; R2=
0.7483) flights.


Cactoblastis cactorum had 3 flight periods per
year along a latitudinal gradient that extended
over the majority of the United States range in-
fested by the insect and that had an annual CDD
range from 6430 to 8486. In the most tropical ar-
eas of South Florida and the Caribbean Islands,
the insect has overlapping generations through-
out the year. The tendency for shorter duration
between flights with increasing southern latitude
is consistent with the tropical sites not experienc-
ing a gap at all between flights and loosing any
sign of distinct generations. Flight duration and
number of weeks between flights was generally
similar for the 3 middle latitude sites and inter-
mediate to the 2 extreme latitude sites. Variation
in the pattern of flight period and flight duration
along the latitudinal gradient was likely caused
by differences in the microclimates at the sites
that differed from the surrounding area. For ex-
ample, the Jekyll Island site was situated at the
edge of a saltwater marsh and protected by small
trees and a large earthen berm. These physical
factors influenced the ambient wind, solar irradi-
ation, and temperature at the study site and
likely influenced the development rate of C. cac-
Identification of 3 flight periods for C. cac-
torum throughout its adventive range in the
United States is useful for current monitoring
and management efforts. Various United States
and Mexican Federal and State Departments of
Agriculture and natural area managers have es-

Florida Entomologist 92(2)

tablished trapping sites to monitor for the pres-
ence of C. cactorum from South Carolina to Cali-
fornia and throughout the Yucatan Peninsula of
Mexico. Knowing the general dates when C. cac-
torum moths will be flying focuses the timing
when traps need to be set up and serviced, saving
expenses and work effort. Including flight infor-
mation from a wide latitudinal array is useful in
the production of mapping models for C. cactorum
based on development and phenology data. Pre-
liminary flight phenology based on virgin C. cac-
torum female baited traps (unpublished data) and
life history information (McLean 2004) was used
by USDA, Animal and Plant Health Inspection
Service (APHIS), to develop a risk zone mapping
program to visually identify adult activity for pre-
dicting the most appropriate times to monitor es-
tablished populations or survey new areas for the
spread of this insect (Bloem et al. 2007). Inclusion
of our multi-year and multi-latitude flight data
along with recent development studies and de-
gree-day models (McLean 2004; McLean et al.
2006; Legaspi & Legaspi 2007; Legaspi et al.
2009) will improve the predictive power of the
APHIS and other models, both in the timing of
various C. cactorum life stages and the expanded
geographical usefulness. Initiation of the spring,
summer, and fall flight periods was the best over-
all regression model that significantly explained
the relationship between latitude and timing of
the flight periods, and duration of the flight or
time between flights. The significant relationship
between the CDD of sites at different latitudes
and the week of year that the 3 flights began field-
validates the use of the APHIS model.
To conduct the SIT program for C. cactorum,
knowledge of the timing, duration, and number of
adult flights is crucial. The latitude of the Missis-
sippi barrier islands on which occurs the current
western leading edge of C. cactorum is between
the latitudes of St. Marks and Pensacola Beach.
Therefore, the flight period information obtained
at St. Marks and Pensacola Beach will be useful
in determining the flight periods and durations at
the current leading edge. Sanitation efforts
against C. cactorum currently rely on the search
and destruction of eggsticks on prickly pear
plants and larvae feeding inside prickly pear
pads. Oviposition and larval development is
timed by the 3 flight periods and sanitation ef-
forts against immature stages would be appropri-
ate between the end of the flight period and the
beginning of the next flight period. For infested
plants at the St. Marks and Pensacola Beach lat-
itudes, sanitation efforts for larvae should be tar-
geted during the months of Jun, Sep, and the
overwintering months of Dec-Feb. In addition,
mitigation of the reproductive capacity of the
adult stage of C. cactorum could be targeted
through mating disruption, a compatible control
tactic with the SIT (Bloem et al. 2001; Carpenter

2000). Such an activity could be conducted during
the 3 adult flight periods in the months of Mar-
May, Jul-Aug, and late Sep-Nov. In the SIT pro-
gram, releases of sterile insects are timed to occur
when wild C. cactorum are present in the field.
Based on our trapping surveys, sterile insects are
effectively released in the months of Mar-May,
Jul-Aug, and late Sep-Nov.
Previous studies conducted of this insect as a
biological control agent indicated 2 flight periods
per year. In terms of southern hemisphere sea-
sons, a summer flight occurs in Argentina, Aus-
tralia, and South Africa generally during Jan-
Mar and a spring flight during Sep-Nov (Dodd
1940; Pettey 1948). Based on northern hemi-
sphere seasons our trapping efforts from south-
ern Florida to central coastal South Carolina re-
vealed a summer flight during Jun-Aug, a fall
flight during Sep-Nov, and a spring flight during
Feb-May. Difference between the northern and
southern hemispheres is the addition of a fall
generation in the United States. While immature
stages of C. cactorum can be easy to identify with
visual searches for infested plants, observing the
cryptic and reticent flying adults is extremely dif-
ficult. Like many insects, identifying the occur-
rence of C. cactorum can be challenging: host
plants are difficult to find, infestation can be con-
fused with other sources of damage (i.e., native
cactus moth), or the population occurs at low
level (Lalone 1980; Hanula et al. 1984; Walters et
al. 2000). Traps baited with C. cactorum syn-
thetic pheromone attract male C. cactorum
(Heath et al. 2006) and have detected the occur-
rence of this insect where no larval damage was
evident (Hight et al. 2002; Bloem et al. 2005a).
Traps were a powerful tool to identify flight times
of male C. cactorum. It would not be surprising
that a third generation of C. cactorum would be
identified throughout much of the northern lati-
tudes of Argentina, Australia, and South Africa
in areas with a higher accumulation of degree-
days. In these warmer climates, we predict a
third flight in the southern hemisphere range of
C. cactorum, and the 3 flight periods would gen-
erally occur during Aug-Oct, Nov-Jan, and Feb-
Apr. The increased accumulation of degree-days
will shorten the time between the spring and fall
flight period, allowing enough time to complete a
third flight period in the fall before the onset of
cooler winter temperatures. However, where C.
cactorum in the southern hemisphere is truly a 2-
generation per year insect, then the additional
generation in the United States may help explain
the rapid spread and buildup of insects in its in-
troduced range. Estimates of spread in the
United States range from 50 to 160 km/yr (Stiling
2002; Solis et al. 2004), while earlier studies lim-
ited the natural spread of C. cactorum to 16-24
km over 2.5 year in Australia (Dodd 1940) and 3-
6 km over 2.5 year in South Africa (Pettey 1948).

June 2009

Hight & Carpenter: Flight Phenology of Cactoblastis


We thank John Mass, Chris Albanese, Michael Get-
man, Sofia Baez, Nathan Herrick, Carla Evans,
Stephen McLean, and Melany Coombs (USDA-ARS-
CMAVE, Tallahassee, FL), Isabel Perez (APHIS, Puerto
Rico), Chuck Byrd (TNC, Florida Keys), MonaLisa
Payne, Bear Moore, and Cecilia Carreo-Turnbull
(APHIS-PPQ, Sebring, FL), Robi Gray and Glynn
County 4-H Club, Brunswick, GA, and Elaine Freeman,
Bess Kellett, and Kate Hall (Edisto Beach, SC) for their
technical assistance. We thank Stuart Reitz and Susie
Legaspi (USDA-ARS-CMAVE, Tallahassee, FL), David
Held (Auburn Univ.), and 2 anonymous reviewers for
comments on earlier drafts of this manuscript. Mention
of trade names or commercial products in this publica-
tion is solely for the purpose of providing specific infor-
mation and does not imply recommendation or
endorsement by the U.S. Department of Agriculture.


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June 2009

Xu et al.: New Species of Dryinidae from China


'College of Natural Resources and Environment, South China Agricultural University,
Guangzhou, Guangdong 510642, P. R. China

2Department of Plant Protection, University of Tuscia, 1-01100 Viterbo, Italy

'Institute of Insect Sciences, Zhejiang University, Hangzhou, Zhejiang 310029, P. R. China

Two new species, Fiorianteon isoneuron sp. nov andAnteon gaoligongense sp. nov, are de-
scribed, respectively, from Xiaowutaishan Provincial Nature Reserve (Hebei Province,
China) and Gaoligongshan National Nature Reserve (Yunnan Province, China). Fiorianteon
isoneuron can be recognized from the related Palaearctic species F junonium Olmi by the
different shape of parameres (without a notch in dorsal view in F junonium, with a deep
notch in F isoneuron) and stigmal vein (distal part much shorter than proximal part in F
junonium, about as long as in F isoneuron). Anteon gaoligongense can be recognized from
the related Oriental species A. indicum Olmi by the different shape of segment 5 of fore tar-
sus (with basal part much longer than distal part inA. indicum, slightly shorter than distal
part inAnteon gaoligongense). Keys are provided for the determinations of the above species
with illustrations of male genitalia and female chelae.

Key Words: taxonomy, Hymenoptera, Dryinidae, Fiorianteon,Anteon, new species, China

Se described las nuevas species Fiorianteon isoneuron sp. nov., proveniente de la Reserva
Natural Provincial Xiaowutaishan (Provincia de Hebei, China), y Anteon gaoligongense sp.
nov. de la Reserva Natural Nacional Gaoligongshan (Provincia de Yunnan, China). Fiorian-
teon isoneuron se separa de la especie paleartica mas relacionada, F junonium Olmi, por la
morfologia de los parameros (con una muesca, en vista dorsal, en F junonium, y una mar-
cada escotadura en F isoneuron), asi como la de la vena estigmal (con la parte distal much
mas corta que la proximal en F junonium, siendo casi iguales en F isoneuron). Anteon gao-
ligongense puede ser reconocida de la especie Oriental mas pr6xima, A. indicum Olmi, por
la forma del quinto segment del tarso anterior (que cuenta con una parte basal much mas
larga que la distal enA. indicum, siendo ligeramente mas corta enAnteon gaoligongense). Se
aportan claves para la identificaci6n de las species antes mencionadas e ilustraciones de la
genitalia masculina y la quela de la hembra.

Translation provided by the authors.

Dryinidae (Hymenoptera: Chrysidoidea) are
parasitoids of Hemiptera Auchenorrhyncha
(Guglielmino & Olmi 1997, 2006, 2007). Fiorian-
teon Olmi, 1984, is a genus present in the Palae-
arctic and Oriental region. Only 3 species have
been described from Japan and China (including
Taiwan). Anteon Jurine, 1807, is a genus present
in all zoogeographical regions. About 326 species
have been described from all continents (Olmi
1999) and the genus was revised by Olmi (1984,
The species of Fiorianteon and Anteon inhab-
iting China were studied in the last 20 years
mainly by Olmi (1991, 1995), He & Xu (2002), Xu
et al. (1998, 2001), and Xu et al. (2003, 2006a,
In 2008 we examined additional specimens of
Fiorianteon andAnteon from P. R. China and have
found 2 new species described herein.


The descriptions follow the terminology used by
He & Xu (2002) and Olmi (1984, 1994, 1999). The
measurements reported are relative, except for the
total length (head to abdominal tip, without the an-
tennae), which is expressed in millimeters. In the
descriptions, POL is the distance between the inner
edges of the two lateral ocelli; OL is the distance be-
tween the inner edges of a lateral ocellus and the
median ocellus; OOL is the distance from the outer
edge of a lateral ocellus to the compound eye; OPL is
the distance from the posterior edge of a lateral ocel-
lus to the occipital carina; TL is the distance from
the posterior edge of an eye to the occipital carina.
All material studied in this paper is deposited
in the Hymenoptera collection of South China Ag-
ricultural University, Department of Entomology,
Guangzhou, Guangdong, P. R. China (SCAU).

Florida Entomologist 92(2)

Fiorianteon isoneuron sp. nov. (Fig. 1)

Description. Holotype Male. Fully winged.
Length 2.8 mm. Head black, except mandibles,
clypeus and part of genae near antennal toruli
testaceous-whitish; antennae brown; mesosoma
black; gaster brown; legs testaceous, except mid
coxae basally black and hind coxae almost totally
black. Antennae filiform; antennal segments in the
following proportions: 8:5:12:12:11:10:9:8:8:10.
Head shiny, smooth, punctate, without sculpture
among punctures; frontal line complete, very nar-
row in the posterior half of the face; occipital car-
ina complete; POL = 5; OL = 2; OOL = 10; OPL =
6; TL = 11; greatest breadth of posterior ocelli
longer than OL (4:2). Pronotum short, transverse,
punctate, without sculpture among punctures.
Scutum, scutellum and metanotum shiny,
smooth, punctate, without sculpture among punc-
tures. Notauli incomplete, reaching approxi-
mately 0.5 x length of scutum. Propodeum dull,
reticulate rugose, without transverse or longitu-


dinal keels. Forewing hyaline, without dark
transverse bands; distal part of stigmal vein
about as long as proximal part. Parameres with a
deep notch in dorsal view (Fig. 1). Tibial spurs 1,
Female. Unknown.
Holotype: Male, P R. CHINA, Hebei Prov.,
Zhangjiakou, Xiaowutaishan Provincial Nature
Reserve, 20-22.VIII.2005, Jingxian Liu & Liqiong
Weng (SCAU).
Etymology: This species is named isoneuron
because the distal part of stigmal vein is very long
and about as long as proximal part.
Remarks. Fiorianteon isoneuron is similar to
the only known Palaearctic species F junonium
Olmi, 1984 (known from Japan). The above spe-
cies can be recognized by the different shape of
parameres (without a notch in dorsal view in F
junonium, with a deep notch in F isoneuron) and
stigmal vein (distal part much shorter than
proximal part in F junonium, about as long as in
F isoneuron), as follows:

Figs. 1 and 2. Male genital armatures (in dorsal view) of Fiorianteon isoneuron (1; holotype; scale bar = 0.06 mm)
and Fiorianteonjunonium (2; specimen from Japan, Uradani; scale bar = 0.11 mm). Right half removed.

June 2009

Xu et al.: New Species of Dryinidae from China

1. Parameres without a deep notch in dorsal view (Fig. 2); distal part of stigmal vein much shorter than proximal
part .......... ... ...................... ................... ........... F junonium Olmi
- Parameres with a deep notch in dorsal view (Fig. 1); distal part of stigmal vein about as long as proximal
part.................................................. ........... F. isoneuron sp nov.

Anteon gaoligongense sp. nov. (Fig. 3)

Description. Holotype Female. Fully winged.
Length 3.3 mm. Head black, except mandibles
testaceous; antennae testaceous; mesosoma
black; gaster brown; legs testaceous, except
basal part of hind coxae blackish. Antennae
clavate; antennal segments in the following
proportions: 12:8:13:13:11:10:9:8.5:8:12. Head
shiny, smooth, punctate, without sculpture
among punctures; frontal line absent; face with-
out 2 lateral keels near orbits directed towards
the antennal toruli; POL = 6; OL = 3; OOL = 7;
OPL = 7; TL = 5; greatest breadth of posterior
ocelli shorter than OPL (3:7); occipital carina
complete. Pronotum shiny, with anterior sur-
face rugose; posterior surface shiny, punctate,

without sculpture among punctures, shorter
than scutum (7:21), more than twice as broad as
long (27:7). Scutum, scutellum and metanotum
shiny, smooth, finely punctate, without sculp-
ture among punctures. Notauli incomplete,
reaching approximately 0.6 x length of scutum.
Propodeum with a strong transverse keel be-
tween dorsal and posterior surface; dorsal sur-
face reticulate rugose; posterior surface with 2
complete longitudinal keels and median area
almost completely smooth and shiny, rugose
only along the lateral margins. Forewing hya-
line, without dark transverse bands; distal part
of stigmal vein slightly shorter than proximal
part (9:11). Fore tarsal segments in the follow-
ing proportions: 10:3:3.5:5:18. Enlarged claw
(Fig. 3) with a proximal prominence bearing a

Figs. 3 and 4. Chelae ofAnteongaoligongense (3; holotype; scale bar = 0.08 mm) andAnteon indicum (4; holotype;
scale bar = 0.08 mm).

Florida Entomologist 92(2)

long bristle. Segment 5 of fore tarsus (Fig. 3)
with basal part slightly shorter than distal part
(8:11), with 2 rows of 19 + 19 lamellae; distal
apex with a group of about 9 lamellae. Tibial
spurs 1, 1, 2.
Male. Unknown.
Holotype: Female, P. R. CHINA, Yunnan
Prov., Gaoligongshan National Nature Reserve,
1-18.VIII.2005, Juanjuan Ma & Yali Cai

Etymology: The specific name derives from the
Latin adjective gaoligongense (inhabiting Mt. Ga-
Remarks. Anteon gaoligongense is similar to
A. indicum Olmi, 1984 (known from India and
Taiwan). The above species can be recognized by
the different shape of segment 5 of fore tarsus
(with basal part much longer than distal part in
A. indicum, slightly shorter than distal part inA.
gaoligongense), as follows:

1. Segment 5 of fore tarsus with basal part much longer than distal part (Fig. 4) ............. A. indicum Olmi

Segment 5 of fore tarsus with basal part slightly shorter than distal part (Fig. 30).... A. gaoligongense sp. nov.

We are grateful to Mr. Jingxian Liu (Zhejiang Uni-
versity), Miss Liqiong Weng, Miss Juanjuan Ma, and
Miss Yali Cai (South China Agricultural University) for
help in collecting dryinids in the field. This study was
supported by the Ministry of Science and Technology of
P. R. China (MOST grant N. 2006FY110500).


GUGLIELMINO, A., AND OLMI, M. 1997. A host-parasite
catalog of world Dryinidae (Hymenoptera: Chrysi-
doidea). Contrib. Entomol. Internat. 2: 165-298.
GUGLIELMINO, A., AND OLMI, M. 2006. A host-parasite
catalog of world Dryinidae (Hymenoptera: Chrysi-
doidea): first supplement. Zootaxa 1139: 35-62.
GUGLIELMINO, A., AND OLMI, M. 2007. A host-parasite
catalog of world Dryinidae (Hymenoptera: Chrysi-
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(Ser. ii) 39: 121-129.
HE, J., AND XU, Z. 2002. Hymenoptera Dryinidae. Fau-
na Sinica 29. Science Press, Beijing: xii + 464 pp.
OLMI, M. 1984. A revision of the Dryinidae (Hy-
menoptera). Mem. American Ent. Inst. 37: xii + 1913
OLMI, M. 1991. Supplement to the revision of the world
Dryinidae (Hymenoptera Chrysidoidea). Frustula
entomol. (1989)(N.S.) 12(25): 109-395.

OLMI, M. 1994. The Dryinidae and Embolemidae (Hy-
menoptera: Chrysidoidea) of Fennoscandia and Den-
mark. Fauna Entomologica Scandinavica 30. Brill,
Leiden. 100 pp.
OLMI, M. 1995. A contribution to the knowledge of the
Dryinidae of Taiwan (Hymenoptera Chrysidoidea).
Boll. Zool. Agr. Bachic. (Ser. ii) 27: 19-34.
OLMI, M. 1999. Hymenoptera Dryinidae-Embolemi-
dae. Fauna d'Italia 37. Edizioni Calderini, Bologna:
xvi + 425 pp.
Xu, Z., HE, J., AND OLMI, M. 1998. New species of Dry-
inidae from China (Hymenoptera, Chrysidoidea).
Phytophaga 8: 21-37.
Xu, Z., HE, J., AND OLMI, M. 2001. Descriptions of new
species of Dryinidae from China (Hymenoptera
Chrysidoidea). Frustula entomol. (2000)(N.S.)
231(36): 1-22.
XU, Z., OLMI, M., AND HE, J. 2003. Three new species of
Dryinidae from China (Hym., Chrysidoidea). Ento-
mol. Monthly Mag. 139: 233-236.
Xu, Z., OLMI, M., AND HE, J. 2006a. Description of a
new species ofAnteon Jurine from the People's Re-
public of China and of the male of Anteon fidum
Olmi (Hymenoptera: Dryinidae). Zootaxa 1164:
Xu, Z., OLMI, M., AND HE, J. 2006b. Descriptions of five
new species of Anteon Jurine from China (Hy-
menoptera: Chrysidoidea: Dryinidae). J. Kansas En-
tomol. Soc. 79: 92-99.

June 2009

Kuriwada et al.: Mass-rearing in Euscepes postfasciatus


1Okinawa Prefectural Plant Protection Center, Maji 123, Naha, Okinawa, 902-0072, Japan

2Ryukyu Sankei Co. Ltd, Maji 123, Naha, Okinawa, 902-0072, Japan


The West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curcu-
lionidae) is a major pest of sweetpotato Ipomoea batatas (L.) Lam (Solanales: Convolvu-
laceae) in some countries. In order to improve mass-rearing for an eradication program
employing the sterile insect technique (SIT), optimal population density of E. postfasciatus
in an artificial diet was examined. Six population densities (1000, 4000, 7000, 10000, 13000,
and 16000 individuals per container with 200 g of artificial diet) were compared for effect on
the number of eggs collected and hatchability. The total number of eggs collected after 24 d
increased with an increase in population density and reached a saturation level at 13,000 in-
dividuals, whereas hatchability was not affected by population density. The results indicated
that optimal population density in mass rearing was 13,000 individuals on 200 g of artificial
diet. Furthermore, we examined cannibalism by adult weevils in the presence of other diets.
The result suggested that egg cannibalism may be a major reason for the low rate of egg col-
lection in the mass rearing of E. postfasciatus.

Key Words: West Indian sweetpotato weevil, artificial diet, sterile insect technique, density
effect, mass rearing


El picudo de camote de las Antillas Occidentales, Euscepes postfasciatus (Fairmaire) (Co-
le6ptera: Curculionidae), es una plaga mayor del camote, Ipomoea batatas (L.) Lam (Solana-
les: Convolvulaceae) en algunos pauses. Para mejorar la cria en masa en un program de
erradicaci6n empleando la t6cnica del insecto est6ril (TIE), se examine la densidad de pobla-
ci6n optima de E. postfasciatus usando una dieta artificial. Se compararon seis densidades
de poblaci6n (1000, 4000, 7000, 10000, 13000 y 16000 individuos por recipient con 200 g de
dieta artificial) para su efecto sobre el numero de huevos recolectados y su habilidad para
eclosionarse. El numero total de los huevos recolectados despu6s de 24 dias incremento con
el aumento de la densidad de la poblaci6n y llego al nivel de saturaci6n a los 13,000 indivi-
duos, mientras que la habilidad para eclosionarse no fue afectada por la densidad de la po-
blaci6n. Los resultados indicaron que la densidad 6ptima de la poblaci6n para la cria masiva
fue de 13,000 individuos con la dieta artificial de 200 g. Ademas, examinamos el canibalismo
por los adults de los picudos en la presencia de otras dietas. El resultado indica que el ca-
nibalismo de los huevos puede ser una de las razones mayores para la baja tasa de la colec-
ci6n de huevos en la cria masiva de E. postfasciatus.

Sweetpotato Ipomoea batatas (L.) Lam
(Solanales: Convolvulaceae) is one of the world's
major food crops, especially in developing coun-
tries (Jansson & Raman 1991). The presence of
the West Indian sweetpotato weevil Euscepes
postfasciatus (Fairmaire) (Coleoptera: Curculion-
idae) and the sweetpotato weevil Cylas formicar-
ius (Fabricius) (Coleoptera: Brentidae) together
have prevented growers from expanding produc-
tion of sweetpotatoes into tropical and subtropical
regions (Jansson & Raman 1991). Lower levels of
pre-and postharvest infestations can reduce both
quality and marketable yield and render sweetpo-
tato storage roots unfit for consumption because
of toxic sesquiterpenes that are produced by roots

in response to weevil feeding (Uritani et al. 1975).
Much of the research has been conducted on con-
trol of C. formicarius (Jansson & Raman 1991;
Miyatake et al. 2000; Kumano et al. 2008), but
few studies have been done on E. postfasciatus.
Euscepes postfasciatus is the major pest of
sweetpotato in the South Pacific, Caribbean ba-
sin, islands of southwestern Japan, and some
countries of Central and South America (Sher-
man & Tamashiro 1954; Kohama 1990; Raman &
Alleyne 1991; Yasuda 1993). The presence of this
weevil on South Pacific islands and southwestern
Japan (Okinawa) has resulted in strict quaran-
tine regulations, including banning the export of
sweetpotato from these islands (Raman & Alleyne

Florida Entomologist 92(2)

1991; Kuba et al. 2003). Because the adults of E.
postfasciatus do not seem to produce sex phero-
mones, pheromone traps cannot be used to sup-
press the weevils (Kuba et al. 2003). However,
sterile insect technique (SIT) is an effective
method for suppression or eradication of weevils
(Kuba et al. 2003).
Establishing an insect mass-rearing technique
is a basic requirement for implementing SIT.
Sterile insect technique involves introducing a
high proportion of sterile matings into a natural
population in order to reduce reproduction to a
level below population maintenance (Knipling
1979). In past successful SIT projects, millions of
insects per week were produced in mass- rearing
facilities (e.g., 280 million per week of the te-
phritid melon fly Bactrocera cucubitaes (Conquil-
lett); 6 million per week of the Old World screw-
worm C'.i .' ... bezziana Villeneuve; and 14
million per week of the codling moth Cydia
pomonella (L.) (cf, Dyck et al. 2005). InE. postfas-
ciatus, a large number of weevils can be reared on
sweetpotato roots (one million per week; Yamag-
ishi & Shimoji 2000), but rearing involves some
challenges. It is very expensive to procure large
quantities and maintain stable quality of the
sweetpotato roots throughout the year. Further-
more, weevils that are reared on sweetpotato
roots face a high risk of infection with the proto-
zoan Farinocystis sp. (undescribed species) at the
Okinawa Prefectural Plant Protection Center
(OPPPC) (Morita et al. 2007). The weevils were
infected with the protozoan orally through the
sweet potato root. Infection remarkably reduces
the fecundity and longevity of adult E. postfascia-
tus (N. K. & D. H., unpublished data). Therefore,
the development of an artificial diet for mass-
rearing E. postfasciatus is needed.
Shimoji & Kohama (1996a) and Shimoji &
Yamagishi (2004) developed an artificial larval
diet, and Sakakibara (2003) developed an artifi-
cial diet for adult E. postfasciatus to feed and ovi-
posit. The larval diet contains mainly sweetpotato
powder, agar, cellulose, protein, sugar, dried
yeast, and water. The adult artificial diet consists
of the larval diet in addition to an antibiotic
(chloramfenicol) and KOH. However, optimal pop-
ulation density for mass-rearing has not been ex-
amined. Generally, in mass-rearing the insects
are reared at higher than optimal density for eco-
nomical reasons. For example, a mass-rearing fa-
cility may have limited space and budget, and in-
sects may be reared with a limited number of con-
tainers and diets. The high density is expected to
have negative effects on the insects' biological
performance because of resource shortages and
accumulation of weevil waste (Pearl & Parker
1922; Ito et al. 1969). Furthermore, high density
may promote egg cannibalism by adult insects.
Females of E. postfasciatus excavate the surface
of either the sweetpotato root or the artificial diet,

oviposit in the cavity, and then seal the eggs in the
cavity with a plug (Raman & Alleyne 1991; Shi-
moji & Kohama 1994). The buried eggs near the
surface of the artificial diet are at a high risk of
being cannibalized by adult weevils intentionally
or accidentally at high density. Egg cannibalism
by adult insects is expected to occur frequently
under mass-rearing conditions, and results in a
reduction in the number of eggs collected from the
mass-rearing procedure. In this study, we exam-
ined the optimal population density for egg collec-
tion in large-scale mass-rearing experiments, and
the number of eggs that were cannibalized in
small-scale laboratory experiments. Our results
provide useful information for establishing an ar-
tificial mass-rearing system for use in an SIT
strategy against this weevil.



The West Indian sweetpotato weevils used in
our study were from a colony that was originally
established from insects collected in Yaese Town,
Okinawa, in Nov 2004. They were cultured on
sweetpotato roots at 25 + 1C, 50-90% Relative
humidity (RH), and a photoperiod of 14: 10 (L: D)
for 18 to 20 generations at OPPPC in Naha, Oki-
nawa. The experiments were conducted from Mar
to Aug 2008. The artificial diet for adult weevils
was modified from Sakakibara (2003) by replac-
ing sweetpotato powder with the powder of rail-
road vine, Ipomoea pes-caprae (L.) (Solanales:
Convolvulaceae). Other components of the artifi-
cial diet (agar, cellulose, protein, sugar, and anti-
biotic, etc.) were the same as Sakakibara (2003).

Effect of Population Density on Fecundity and Hatch-
Population Density
Subsequent to emergence from the sweetpo-
tato roots, adult weevils (ages 2-9 d old) were
placed in a mesh container and fecundity of the
weevils was measured for 24 d. Most adults
stayed in the sweetpotato roots until sexual mat-
uration, which was about 10 d after emergence
(Kohama & Shimoji 1998). The weevils were
maintained on sweetpotato roots during the pe-
riod between emergence and the setting up of our
experiments. To determine the effect of popula-
tion density on fecundity, 1,000 (5 replications),
4,000 (5 replications), 7,000 (5 replications),
10,000 (5 replications), 13,000 (5 replications),
and 16,000 (4 replications) adults were intro-
duced into each meshed container (357 x 287 x
120 mm) with 200 g of artificial diet cut in the
shape of a square pillar (5 x 5 x 40 mm). When
greater amounts of the artificial diet was supplied
to each container, it became damp and then

June 2009

Kuriwada et al.: Mass-rearing in Euscepes postfasciatus

spoiled (T. K. unpublished data). Numbers of wee-
vils per container were measured by using the
mean body weight from samples of 50 weevils.
The sex ratio was assumed to be approximately
1:1 on the basis of Raman & Alleyne (1991). The
containers were kept at 25 + 1C, 50-80% RH, and
a photoperiod of 14:10 (L: D).

Egg Collection

Egg collection procedure in the experiment
was based on the technique of egg extraction for
the boll weevil, Anthonomus grandis Boheman
(Coleoptera: Curculionidae) (Roberson 1984),
which exhibits similar oviposition behavior. After
the setup of the mass-rearing containers, eggs
were collected 9 times in 24 d (Table 1). We rinsed
the container, weevils, and diet all together with
running tap water to remove the eggs that were
oviposited on the artificial diet as well as at the
bottom of the container. First, the weevils were
scooped out of the suspension with a 1.0-mm
mesh net (see weevil collection section). The large
pieces of diet were rubbed vigorously by hand in
water for 15 min in order to free the eggs buried in
the diet. The remaining suspension was filtered
through a 0.5-mm meshed sieve to separate the
eggs from the diet and waste particles. The mix-
ture of eggs and remaining waste particles were
mixed with 14.6% salt-water in a glass (100 mL),
and freshwater was gently added to the glass to
separate the eggs from the waste (Ohno et al.
2005). After a few min, we collected the eggs from
the boundary between the salt-water and the
freshwater by pipette. The number of eggs per
container was estimated by the method from
Ohno et al. (2006). When there were very few eggs
and the number could not be estimated by this
method, we counted the eggs directly. For surface
disinfection, the eggs were dipped in 70% ethanol
solution for 40 min, and then rinsed with sterile
water (modified from Ohno et al. 2004). The
hatchability of the eggs was then determined (see
hatchability section). From the first egg collec-

tion, the collected eggs were discarded because
some fungi and mites adhered to the weevils and
containers. After this treatment, few fungi and
mites were observed on the weevils and contain-

Weevil Collection

After being removed from the suspension, the
weevils were rinsed in water, and dried with an
electric fan for 3.5 h. They were then re-intro-
duced to containers with 200 g of fresh artificial
diet to continue the egg-collection experiment. We
did not record the status of individuals (dead or
alive) for logistic reasons. This procedure was con-
ducted 9 times during a 24-day period for each


To examine the effects of population density on
hatchability, we randomly chose 200 eggs that
had been oviposited 7 d after the initial setup of
each treatment. One hundred disinfected eggs
were then placed on a nylon cloth (block) on wet
filter paper in Petri dishes (9 cm in diameter x 2
cm in height) (2 blocks per density treatment).
The Petri dishes were kept at 25 + 1C, 50-80%
RH, and a photoperiod of 14:10 (L:D). Previous
studies showed that the mean egg-hatching pe-
riod was about 7-9 d (Sherman & Tamashiro 1954;
Raman & Alleyne 1991). We recorded the number
of hatched eggs at 6, 7, 10, 12, and 14 d after the
day of egg collection. The experiment was repli-
cated 3 times.

Egg Cannibalism

To examine the incidence of egg cannibalism,
we conducted 2 small-scale laboratory experi-
ments. In one we examined whether food shortage
caused egg cannibalism, and in the other experi-
ment whether adult weevils cannibalized eggs
even in food-rich conditions. To examine whether


Monday Wednesday Friday

Week 1 Set up mass rearing
Week 2 Discard collected eggs First egg collection
due to contamination. (5 d after setting up). Second egg collection.
Week 3 Third egg collection. Fourth egg collection. Fifth egg collection.
Week 4 Sixth egg collection. Seventh egg collection. Eighth egg collection.
Week 5 Ninth egg collection
(24 d after setting up).

Florida Entomologist 92(2)

egg cannibalism was influenced by the artificial
diet, we used both sweetpotato root and artificial
diet for these experiments. We used weevils aged
1 d after emerging from the sweetpotato, and we
selected only male weevils because of the poten-
tial problem of identifying original eggs from the
newly oviposited ones during the experimental

Experiment 1. Effect of Starvation on Egg Cannibalism

This experiment was intended to clarify the re-
lationship between food shortage and egg canni-
balism by adult male weevils. The number of eggs
cannibalized by either starved or unstarved
adults was measured in the absence of other food
sources. In the starved condition, each male was
kept individually in a well of a 24-well multiplate
(AS ONE Corporation, Osaka, Japan). The males
were given neither food nor water for 7 d. Each
unstarved control male was set in a small plastic
container (200 mL) with either a small piece of
sweetpotato root (60 mg) or artificial diet (60 mg)
for 7 d as well. The diet was changed daily. After
the starvation procedure, each male and 5 eggs
were placed at the bottom of a well of a 24-well
multiplate, and we counted the number of re-
maining eggs 5 d after setting up the treatment.

Experiment 2. Effect of the Presence of Other Diets on
Egg Cannibalism

This experiment examined whether male E.
postfasciatus cannibalized eggs when other food
sources were available. The numbers of cannibal-
ized eggs were compared in the presence or ab-
sence of a small piece of sweetpotato root or arti-
ficial diet. Each male was kept individually in a
well of a 24-well multiplate with no diet or water
for 3 d. We then divided the individual weevils
into 3 treatments: (1) 5 eggs and no diet, (2) 5 eggs
and a small piece of sweetpotato root (5 mg), and
(3) 5 eggs and a small piece of artificial diet (5 mg)
at the bottom of a well of a 24-well multiplate. For
each group, we counted the number of lost eggs 2
d after setting up the experiment.

Statistical Analyses

Generalized linear mixed models (GLMMs)
were performed by using R 2.60 (R Development
Core Team 2007) in our analyses (Schall 1991;
Crawley 2005). When female fecundity was a de-
pendent variable, we used GLMM with Poisson
error structure and a log link function. When the
hatching rate or the number of cannibalized eggs
was a dependent variable, we used GLMM with
binomial error structures and logit link functions.
When the effect of population density on female
fecundity and hatching rate was examined, the
dependent variable was fecundity or hatching

rate, fixed independent variables were population
density and quadratic term (population density2);
and the random effects were replication and
blocks (hatching rate only). In experiment 1, the
dependent variable was the number of cannibal-
ized eggs, fixed independent variables were the
diet treatment (i.e., no diet vs. sweetpotato root),
and the random effects were the identities of the
focal individuals. In experiment 2, the dependent
variable was the number of cannibalized eggs, the
independent variable was the diet treatment and
the random effects were identities of the focal in-
dividuals. A Wald test was used to test the statis-
tical significance of each coefficient in the model.
To examine the relationship between fecundity
per female and population density, we used the
non-parametric Jonckheere-Terpstra test for or-
dered alternatives (Hollander & Wolfe 1999) be-
cause the error structure of fecundity per female
was not able to be determined. In egg cannibalism
experiments, the P-values were corrected by
Tukey's all-pairwise comparisons (R add-on pack-
age "multcomp," Hothorn et al. 2008). We used the
two-tailed test for statistical significance.


Effect of Population density on Fecundity and Hatch-

The peak of oviposition was 10 d after setting
up the experiment; the number of eggs then de-
creased over time (Fig. 1). The fitted GLMM equa-
tion indicated the saturation curve (quadratic
term (population density)2 = -8.39e-9 1.7e-9 (SE),
z = 71.45, P < 0.001; linear term (population den-
sity) = 2.83e-04 2.34e 06, z = 120.85, P < 0.001) .

-- 16000
-- 3ooM
/ --- 41000


, x --... o

5 7 10 12 14 17 19 21 24
Days After Selling

Fig. 1. Oviposition pattern of Euscepes postfasciatus
at different population densities.

June 2009

Kuriwada et al.: Mass-rearing in Euscepes postfasciatus

The result suggested that the total number of
eggs collected per 24 d increased with the in-
crease of population density, and reached satura-
tion at 13,000 individuals (Fig. 2a). No significant
relationship was observed between fecundity per
female and population density (Fig. 2b, J* = 249,
k = 6, P = 0.084).
Hatchability was 57.5% with the treatment of
10,000 to 13,000 individuals, and no significant
differences in hatchability among the treat-
ments were observed (Fig. 2c, the quadratic
term; -4.88e 09 + 5.20e 02, z = 9.38e-8, P = 1.0, lin-
ear term; 1.07e-"4 9.33e02, z = 1.14e7, P = 1.0).

Egg Cannibalism
Experiment 1. Effect of Starvation on Egg Cannibalism

Starved males cannibalized a significantly
larger number of eggs than unstarved males did
during 5 d (Fig. 3 and 4, starved vs. artificial diet;
b = 1.76 + 0.50 (SE), z = 3.55, P = 0.0011, starved
vs. sweetpotato root; b = 1.07 + 0.46, z = 2.35, P =
0.050). No significant differences were observed
between treatments containing artificial diet or
sweetpotato roots (b = 0.69 + 0.53, z = 1.32, P =
0.38). The b values indicated contrast coefficients
(Crawley 2005).

Experiment 2. Effect of the Presence of Other Diets on
Egg Cannibalism

No significant differences in the number of
eggs cannibalized were apparent among the diet
treatments (Fig. 4, no diet vs. artificial diet; b =
0.28 + 0.44, z = 0.64, P = 0.80, no diet vs. sweetpo-
tato root; b = -0.11 0.43, z = 0.25, P = 0.97, and
artificial diet vs. sweetpotato root; b = -0.39 +
0.45, z = 0.87, P = 0.66).


The total number of eggs collected over 24 d in-
creased with population density, but leveled off
above 13,000 individuals, although we did not in-
vestigate the treatment of more than 16,000 indi-
viduals. However, hatchability and fecundity per
female were not affected by population density.
These results implied that optimal population
density in mass rearing lay around 13,000 adult
individuals with 200 g of artificial diet. Few eggs
were collected after 19 d. In the treatment of
13,000 individuals, a female laid 0.08 eggs per
day. Raman & Alleyne (1991) reported that wee-
vils reared on sweetpotato roots oviposited 1.0 egg
per day per female. This large difference in the
number of eggs collected was not likely due to
only damage to weevils by the rinsing procedure
or to the quality of artificial diet.
Egg cannibalism was considered to be a major
reason for the low rate of egg collection. With
some insects, eggs or immature larvae are canni-


IOW 4000 l000 10000 i30 1W0



100 4 Pp 7l00 in D 13it 1y00
Pqmtatn Dwuuty

Fig. 2. (a) Relationship between population density
of Euscepes postfasciatus adults in cages with 200 g of
artificial diet and number of eggs collected. The horizon-
tal bar in the middle of each box denotes the median
number of eggs. The top of the box (above the median)
denotes the 75th percentile, and the bottom of the box
(below the median) denotes the 25th percentile. The
vertical dashed lines indicate the maximum and mini-
mum values of the y axis except for the outliers. Open
circles represent the outliers. Outliers were values more
than 1.5 times the interquartile range above the 75th
quartile or below the 25th quartile. (b) Relationship be-
tween population density of E. postfasciatus adults and
number of eggs collected per female. Vertical bars repre-
sent SD. (c) Relationship between population density
of E. postfasciatus adults and hatched larvae observed
on the nylon cloth in Petri dishes. Vertical bars repre-
sent SD.

Florida Entomologist 92(2)

of egg collection in the mass-rearing of this spe-
n=55 n= 52 n = cies. Based on our data that each weevil cannibal-
S= ized an average of 0.4 eggs during a 2-day period,
1/ we estimated that 13,000 weevils cannibalized
2,600 eggs per day. We assumed that if egg canni-
a *balism were fully prevented, a total of 74,400 eggs
would be collected in 24 d with the treatment of
w 13,000 individuals (in the present state, 12,000
eggs were collected). Therefore, if a method to pre-
Svent egg cannibalism is developed, the rate of egg
b collection would change dramatically. For exam-
ple, Shimoji & Kohama (1996b) and Shimoji &
b Yamagishi (2002) used an aluminum meshed con-
tainer that was set in a plastic cup to collect eggs
easily. Eggs laid into the container fell through
the mesh and into the bottom of the plastic cups.
Artificial Diet Sweet Potato Because a mesh container separates the eggs
.3 Starved Unstrved from the adult weevils, this method would also be
useful in preventing egg cannibalism. Further-
Fig. 3. Egg cannibalism among groups of starved and more, hatchability would probably be improved
unstarved E. postfasciatus adults in the absence of because this method does not involve rinsing the
other food sources. Different letters indicate a signifi- eggs, which can damage them.
cant difference in the number of cannibalized eggs. Ver- Food shortages and an accumulation of weevil
tical bars represent SD. waste also may cause a reduction in female per-
formance. Therefore, a greater frequency of food
replacement and cleansing of containers might
balized by other individuals (Banks 1956; Fox increase the number of eggs collected. If eggs had
1975; Kinoshita 1998). In the present study, the been collected daily instead of every 2-3 d, num-
starved male weevils cannibalized eggs more fre- bers of eggs collected may have increased. How-
quently than the well-fed ones did. This result im- ever, this approach would not be practical because
plied that cannibalism due to food shortage con- labor reduction is essential in mass-rearing
tribute to the reduction in the number of eggs (Parker 2004). More fruitful approaches to in-
collected. Furthermore, adult weevils cannibal- creasing egg output may involve improvement of
ized some of the eggs even in the presence of shape and water content of the artificial diet. The
available diet. This finding suggested that egg artificial diet sometimes became moldy during
cannibalism may be a major cause of the low rate mass-rearing. If water content in the diet can be
decreased, growth of the mold may be suppressed.
Furthermore, small changes in diet composition
can often lead to large changes in insect survival
n=48 n=43 n 48 and reproduction (Cohen 2004). For example,
Fisher & Bruck (2004) found that larval black
S vine weevil Otiorhynchus sulcatus survival was
I 23% higher and body weight was 1.6 times higher
I on an improved diet with 0.8 g decrease in methyl
I paraben and replacement of sorbic acid with po-
a tassium sorbate. Therefore, experiments should
S0 a be conducted to determine whether alteration of
SI_ artificial diet components increases the reproduc-
Sa ive performance ofE. postfasciatus. For example,
S0 reduction in antibiotic content may increase the
Sl performance of the weevil because microorgan-
Sisms are symbiotically associated with E. postfas-
I ciatus (T. Hosokawa, personal communication).
In conclusion, we demonstrated an optimal
Population density around 13,000 individuals of
No diet Artificial Diet Sweet Potato adult E. postfasciatus with 200 g of artificial diet.
Fig. 4. Egg cannibalism by E. postfasciatus adults in Under mass-rearing conditions, we conclude that
the presence or absence of other food sources. No signif- egg cannibalism was a major cause of the low rate
icant differences among the treatments were observed, of egg collection in the artificial diet treatment
as indicated by the same letters for each treatment. Ver- compared with the treatment of sweetpotato
tical bars represent SD. roots. Improvement of egg collection techniques

June 2009

Kuriwada et al.: Mass-rearing in Euscepes postfasciatus

and artificial diet is urgently needed for the mass-
rearing ofE. postfasciatus.


We thank the staffs of OPPPC and Ryukyu-Sankei
Co. Ltd. for helpful assistance, and A. Oyafuso and Dr.
M. Yamagishi, who provided valuable comments for this
study. The manuscript was further greatly improved by
4 anonymous reviewers, who provided valuable com-
ments on the manuscript.


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lease, pp. 209-232 In V.A. Dyck, J. Hendrichs, and A.
S. Robinson [eds.], Sterile Insect Technique, Princi-
ples and Practice in Area-Wide Integrated Pest Man-
agement. Springer, Dordrecht.
PEARL, R., AND PARKER, S. L. 1922. On the influence of
density of population upon the rate of reproduction
in Drosophila. Proc. Natl. Acad. Sci. USA 8: 212-219.
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management of the West Indian sweet potato weevil,
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ment: A Global Perspective. Westview Press, Boul-
der, CO, USA.
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June 2009

Hanula et al: Arthropods and Coarse Woody Debris


'USDA Forest Service, Forestry Sciences Laboratory, 320 Green St., Athens, GA 30602-2044

2USDA Forest Service, Department of Forest Resources, Clemson University, Clemson, SC 29634



A 5-year study of long-term (40 years) study plots was conducted on the Osceola National
Forest in northern Florida to determine how dormant-season fire frequency (annual, bien-
nial, quadrennial, or unburned) affects ground-dwelling macroarthropod use of coarse
woody debris in longleaf pine (Pinus palustris Mill.) forests. Pitfall traps were used to sam-
ple arthropods near logs or metal drift fences of equal length. Samples were identified to ge-
nus or the lowest practical taxonomic level. Overall, significantly more arthropods and more
arthropod biomass were captured near drift fences than near logs. Similarity of arthropods
captured near logs or drift fences ranged from 64.4% in annually burned plots to 69.2% in
quadrennially burned plots, with no significant differences noted. Likewise, Shannon diver-
sity, evenness, richness, and number of rare genera were the same for traps regardless of the
trap location. Interaction between fires and trap location were observed in 31 of 932 arthro-
pod taxa. Of those, 10 taxa had significantly higher numbers captured in traps near logs in
some burn treatments but there was no consistent pattern between log use and fire fre-
quency. In most cases, more were captured in log pitfalls in frequently burned plots but that
was not the case for at least 4 taxa. Where interactions between trap location and fire fre-
quency were not significant, arthropods in an additional 101 taxa were captured in higher
numbers at 1 trap location or the other. Of those, 73 were captured in higher numbers in pit-
falls near drift fences and 28 were captured in higher numbers near logs. Results showed no
increase in log usage by general or more mobile ground-dwelling arthropods as more fre-
quent burning reduced the herbaceous and woody under story. However, logs were clearly
important to a wide variety of arthropods regardless of burn frequency.

Key Words: prescribed burning, coarse woody debris, habitat heterogeneity


Una investigaci6n de 5 aios usando parcelas de studio de largo plazo (40 aios) fue reali-
zado en el Bosque Nacional de Osceola en el norte del estado de la Florida para determinar
como la frecuencia de los incendios en la estaci6n latente annual bienal, cuadrienal o no que-
mada) afecta el uso de los escombros leiosos en bosques del pino, Pinus palustris Mill. por
los macroartropodos que viven en el suelo. Se usaron trampas de caida para muestrear los
artr6podos cerca de troncos o de postes metalicos de retensi6n de una misma longitud. Las
muestras fueron identificadas al nivel de g6nero o al nivel taxon6mico mas practice. Sobre-
todo, se capturaron significativamente mas artr6podos con un mayor cantidad de biomasa
cerca de los postes metalicos que cerca de los troncos. La similitud de los artr6podos captu-
rados cerca de los troncos y los postes metalicos varia de 64.4% en las par6celas quemadas
anualmente a 69.2% en las parcelas quemadas cuadrienales (cada 4 aios) sin diferencias
significativas anotadas. Asimismo, la diversidad Shannon, la igualdad, riqueza y el numero
de g6neros raros fueron los mismos en las trampas a pesar de la ubicaci6n de la trampa. Se
observaron la interacci6n entire los incendios y la ubicaci6n de las trampas en 31 de los 932
taxa de artr6podos. De estos, 10 taxa tuvieron un mayor numero capturados en trampas
cerca de los troncos en algunos de los tratamientos quemados pero no hubo un patron con-
sistente entire el uso de los troncos y la frecuencia de los incendios. En la mayoria de los ca-
sos, se capturaron mas en las trampas de caida puestas cerca de los troncos en las parcelas
quemadas frecuentemente pero esto no fue el caso en por lo menos 4 taxa. Donde las inte-
racciones entire la ubicaci6n de la trampa y la frecuencia del incendio no fueron significati-
vas, los artr6podos en 101 taxa adicionales fueron capturados en numeros mas altos en la 1
ubicaci6n de trampa o la otra. De estos, 73 fueron capturados en numerous mas altos en tram-
pas de caida cerca de los postes metalicos y 28 fueron capturados en numerous mas altos cerca
de los troncos. Los resultados mostraron ningun aumento en el uso de los troncos por los ar-
tr6podos generals o los mas m6viles que viven en el suelo mientras que la quemada mas fre-

Florida Entomologist 92(2)

cuente redujo la vegetaci6n herbacea y lenosa en las plants bajeras. Sin embargo, los
troncos fueron claramente importantes a una amplia variedad de artr6podos a pesar de la
frecuencia de la quema.

Large dead wood in terrestrial forest habitats
is an important resource for a number of arthro-
pods and other animals that use it for food, ovipo-
sition sites, protection from environmental ex-
tremes, and foraging habitat (Elton 1966; Har-
mon et al. 1986; Grove 2002). Within this habitat
saproxylic arthropod communities vary with spe-
cies of tree, stage of decomposition, types of fungal
colonists, and location (aquatic to xeric) of dead
wood in the landscape (Harmon et al. 1986;
Speight 1989; Okland et al. 1996; Grove 2002). In
addition to direct contribution to forest diversity,
saproxylic arthropods are an important part of
the food web supporting a variety of predators
and parasites (Harmon et al. 1986). Considerable
research on coarse woody debris and its function
in forests has been done in North America but
most of the emphasis has been in the Pacific
Northwest and Canada (Harmon et al. 1986; Wol-
dendorp et al. 2002). Within North America there
is a large body of literature on the Scolytinae
(Curculionidae: Coleoptera) and their associates
but relatively little work on other saproxylic spe-
cies or those that occur in later stages of decay
(Savely 1939; Howden & Vogt 1951; Hammond et
al. 2001, 2004), and less is known about how dead
wood affects the distribution of arthropods that
may be associated with it but not totally depen-
dent on it (Irmler et al. 1996; Marra & Edmonds
1998; Andrew et al. 2000; Buddle 2001). Despite
the growing interest in this topic, interactions be-
tween woody debris and arthropod communities
in the Southeastern United States have received
little attention (McMinn & Crossley 1996) and
this is particularly true for longleaf pine (Pinus
palustris) forests.
Longleaf pine once occupied about 30 million
hectares in the Southeastern region of North
America (Frost 1993). Today <1.3 million hectares
remain as small, isolated parcels (Outcalt & Shef-
field 1996) resulting in longleaf pine and associ-
ated communities being classified as the third
most endangered ecosystem in the United States
(Noss et al. 1995). Conservation and restoration
of these communities is a priority for forest man-
agement and conservation groups throughout the
region (Van Lear et al. 2005).
Prior to European arrival in North America,
longleaf pine communities were maintained by
frequent fires (1-3 year intervals) started by light-
ning or Native Americans (Landers 1991) but this
is no longer the norm. An unanticipated result of
reduced fire frequency is the increased accumula-
tion of fuels which foster fires that are signifi-
cantly more likely to damage both the environ-
ment and stand improvements, and pose a much

greater threat to human health and safety (Oult-
calt & Wade 2004). A number of remedies are
available and, although prescribed burning is the
only alternative that addresses the full suite of ec-
osystem components, other alternatives are com-
monly chosen to achieve short-term results (Van
Lear et al. 2005). Even when fire is used, these re-
maining longleaf pine communities are often not
managed under the appropriate fire regime
(Brose & Wade 2002).
A number of studies and reviews have looked
at effects of fire on arthropods but no clear gen-
eral trends have emerged (Rice 1932; Heyward &
Tissot 1936; Pearse 1943; Buffington 1967; Ahl-
gren 1974; Hurst 1971;Warren et al. 1987; Muona
& Rutanen 1994; Buddle et al. 2000; Niwa & Peck
2002; Hanula & Wade 2003). However, despite the
lack of a general trend, fire has a significant effect
on many epigaeic arthropods and this is particu-
larly true for the longleaf pine flatwoods ecosys-
tem where it reduces diversity, community simi-
larity and populations of many taxa (Hanula &
Wade 2003). In addition to direct mortality, fire al-
ters the forest habitat and community structure
resulting in either positive or negative effects on a
given species. For example, by reducing predators
(spiders, centipedes, ground beetles, etc.) some
species may experience population increases or,
conversely, reductions in prey may reduce preda-
tor populations. Fire may also reduce food for de-
tritivores (e.g., termites) and cover for other spe-
cies. By removing refuges such as litter or low
growing shrubs, fire may increase susceptibility
to predation, temperature extremes or desicca-
Elton (1966) recognized the importance of
woody debris as a source of forest diversity and he
noted that as wood decomposes it is increasingly
colonized by generalists that do not require spe-
cific tree species or even depend on woody debris
as their sole habitat. For example, Irmler et al.
(1996) found that the increasing variety of wood-
dwelling species as dead wood aged was due
largely to immigration of litter dwelling species to
dead wood and to species using dead wood for
overwintering. Likewise, Carcamo & Parkinson
(1999) reported that decomposed coarse woody
debris was a major factor in shaping ground bee-
tle assemblages, and Evans et al. (2003) and
Jabin et al. (2004) reported that proximity to logs
was an important factor in shaping litter-dwell-
ing invertebrate communities. These studies
demonstrate the role dead wood can play in shap-
ing epigaeic arthropod communities, which can in
turn affect a variety of ecological processes
(Evans et al. 2003; Jabin et al. 2004).

June 2009

Hanula et al: Arthropods and Coarse Woody Debris

While planning a study on the long-term ef-
fects of frequent dormant-season burning on epi-
gaeic arthropods (Hanula & Wade 2003) we hy-
pothesized that the presence of coarse woody de-
bris would be more important to these arthropods
on annually burned plots where much of the leaf
litter, understory vegetation and structure is
sparse and has little time to recover between
burns. To address this hypothesis, we conducted a
study over a 5-year period to examine whether
the presence of woody debris influenced pitfall
trap captures under varying burn frequencies in a
longleaf pine forest. In addition, we measured the
amount of coarse woody debris and other habitat
variables to determine how long-term frequent
burning interacted with these variables to shape
the epigaeic arthropod community.


Study Site and Prescribed Burn Treatments

The study was conducted in the Osceola Na-
tional Forest in Baker County, Florida. Study
plots were established in 1958 to examine effects
of burning frequency on fuel reduction for wildfire
prevention. At that time the overstory trees were
45-year-old longleaf pines that were 20 m tall and
29 cm diameter (DBH) with a few slash pine (P el-
liottii) of similar size. The presence of remnant
"boxed" longleaf pine trees from past turpentine
operations suggest that the area was never
cleared for agriculture. The understory consists of
typical flatwoods vegetation dominated by saw
palmetto (Serenoa repens), gallberry (Ilex glabra),
Vaccinium sp. and wiregrass (Aristida beyri-
chiana). The study was a randomized complete
block design consisting of 24, 0.8-ha plots ar-
ranged in 6 blocks of 4 treatments. Initial treat-
ments were winter burns applied every 2, 4 or 6
years and unburned controls. However, in 1964
the 6-year interval treatment was replaced with
annual winter burns. The entire site was burned
in 1958 to initialize the study area, and treat-
ments have been applied as scheduled since then
between Dec and early Mar. Fire intensity varied
from year to year depending on weather condi-
tions, dead fine-fuel moisture content (which typ-
ically ranged from 7-20%), and firing techniques.
Our study, superimposed on this long-term winter
burn study, started in fall, 1994 just before the ap-
plication of annual burns in the winter of 1994-
1995. The study continued for the full cycle of
burn treatments (annual, biennial, and quadren-
nial) ending after the quadrennial plots were
again burned the winter of 1999-2000. Control
plots had been unburned for 42 years at the close
of our study.
We sampled the naturally occurring large dead
wood on the plots in Dec 2003, three years after
the study. Small dead wood was not affected by

fire frequency on our plots (Hanula & Wade 2003),
so it is unlikely that large dead wood was, and
there was no evidence of increased tree mortality
following our study so the delay in measuring this
attribute did not result in significant changes in
dead wood volumes between the time insects and
dead wood were sampled. We estimated large
dead wood volume by measuring the end diame-
ters of all down woody debris in five 10-m wide
transects which resulted in sampling approxi-
mately 50% of the plot surface. In addition, we
conducted a 100% survey of standing dead wood.
We used Huber's equation (volume = m x 1; where,
m = mid-point cross-sectional area and 1 = length)
to estimate volume of dead wood (Avery 1975).
The mid-point diameter of downed wood was esti-
mated by taking the average of the end diameters.
The mid-point diameter of standing dead wood
was estimated using taper equations for coastal
plain longleaf pine (Clark et al. 1991).
Pre-burn live and dead plant biomass (dried)
was estimated on each plot from eight 1-m2 sub-
plots. Details of sampling procedures for live and
dead plant material can be found in Hanula &
Wade (2003). Sampling methods were adapted
from Shea et al. (1996). Two clusters of 8 transect
lines were established in each plot. We ran 15 m
long transects from the center of each cluster in
the 4 cardinal directions and 4 additional 15-m
long transects were established at 90 from the
ends of the original transects. Sampling frames (1
m2) were placed on the ground 4 m from the start-
ing point of each transect. All stems <1.9 cm basal
diameter were collected by category from each
sample point, bagged, oven-dried at 42C for a
minimum of 48 h, and weighed. Plant biomass
was separated into 11 categories consisting of (1)
live palmetto, (2) dead palmetto, (3) live gallberry,
(4) live pine needles (seedlings), (5) live grasses
and forbs, (6) litter layer (01 or L layer), (7) hu-
mus (02 and 03 or F and H layers), (8) pine cones,
(9) 0-0.6 cm dead branches, (10) 0.6-2.5 cm dead
branches, and (11) other dead woody material.

Arthropod Sampling

We hypothesized that logs could function in
two ways to increase trap captures of arthropods:
(1) they could be a preferred habitat resulting in
concentrations of arthropods around them, or (2)
they could act as drift fences concentrating and
directing arthropods normally wandering across
the forest floor into traps near them. To determine
if large dead wood was a preferred habitat or
acted as a drift fence, we placed 3-m long logs of
longleaf pine (20-25 cm diameter) in the center of
each plot and installed pitfall traps along them
immediately after felling. Four pitfall traps con-
structed from 480-mL capacity plastic cups (Han-
ula & Franzreb 1998) were installed near the log
at plot center (2 on each side 0.5 m from each end)

Florida Entomologist 92(2)

as close to the log as possible. To decrease the
chances of an arthropod bypassing the pitfall by
moving between the log and the pitfall we in-
serted wedge-shaped pieces of aluminum sheet
metal into the space so that the metal sheeting
created a barrier from the point where the log
contacted the ground to the edge of the pitfall. A
second set of 4 pitfall traps was installed along a
3-m long aluminum sheet metal drift fence (15 cm
high). The drift fence was located 10 m from the
center log. Traps were constructed and placed in
the same manner as the log pitfalls except that
the edge of each trap was in direct contact with
the drift fence. Beginning in Jan 1995, pitfall
traps were opened for month-long periods 6 times
per year and covered with ceramic tiles during
months when not in use to prevent arthropods
from falling in. Arthropods collected in this way
were placed in 70% ethyl alcohol, sorted to mor-
phologically similar groups, and identified to ge-
nus or the lowest taxonomic level possible by us-
ing a reference collection and appropriate taxo-
nomic keys. Biomass estimates were obtained by
oven drying (40C for 72 h) and weighing at least
20-30 specimens of each taxon. The average
weight of these specimens was multiplied by the
number of individuals within a sample to esti-
mate sample biomass.

Statistical Analyses

Horn's simplification of Morisita's index of sim-
ilarity was used to compare arthropod communi-
ties captured in the 2 types of traps (Horn 1966).
We used richness and the Shannon diversity in-
dex to measure arthropod community diversity.
Evenness was calculated to give further informa-
tion about the arthropod communities among
burn treatments. Calculations were based on the
cumulative totals of each arthropod for each plot
for the entire study period.
Because 2 trap locations were designated
within each plot, the study was analyzed as a
split-plot design. Analyses of variance were con-
ducted with the SAS GLM procedure (SAS 1985)
to test for interactions of burn frequency and trap
location, and trap location effects. Model effects
were burn treatment, trap location, block, treat-
ment x block interaction, and burn treatment x
trap location interaction. Analyses were con-
ducted for taxa in which 30 or more individuals
were caught during the entire study. Digweed et
al. (1995) found evidence of depletion of certain
Carabidae when traps were 10 m apart, so we also
plotted annual captures of some common families
and species to determine if there was any deple-
tion effect from trapping 4 years at the same loca-
tion or from drift fence pitfalls being 10 m from
the pitfalls near logs.
In order to examine arthropod community re-
sponses, we used non-metric multidimensional

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June 2009

Hanula et al: Arthropods and Coarse Woody Debris


Trap Location

Variable Drift fence (Mean SE) Log (Mean SE) P > F

Number of arthropods 3950.1 192.9 2859.1 101.8 <0.0001
Biomass (g dry wt/plot) 38.9 3.5 27.4 2.6 <0.0001
Shannon Diversity (H') 3.61 0.06 3.54 0.06 0.086
Evenness (J) 0.67 0.01 0.66 0.01 0.21
Richness (genera/plot) 216.8 3.46 212.1 3.41 0.25

Analyses of variance were conducted by the SAS GLM procedure (SAS 1985).

scaling analysis (NMDS) of trends in arthropod
abundance in the study plots. We chose NMDS
because there appeared to be an "arch effect" in
a preliminary canonical correspondence analysis
and because this method is more robust to vari-
ability in underlying patterns in morphospecies
responses than are eigenvalue-based ordination
techniques (Clarke 1993; Gaiser et al. 1998).
Morphospecies that had less than 10 individuals
were excluded resulting in a total of 184 mor-
phospecies in the analysis. To examine responses
of the arthropod community to environmental
variables, we analyzed the relationship between
the NMDS scores and environmental variables
using vector analysis (Gaiser et al. 1998). Vector
analysis, a form of indirect gradient analysis, is
a regression of the environmental variables on
the NMDS scores with the results displayed as
vectors overlaid on a plot of NMDS scores. Be-
cause we were interested in coarse woody debris
effects on arthropod community structure, and
because fire frequency had such a strong effect
on several variables, we excluded those environ-
mental measures that were highly correlated to
fire frequency (Table 1). We included fire fre-
quency, gallberry biomass, small woody debris,
downed coarse woody debris, and standing
woody debris as the environmental variables in
the vector analysis.


We caught significantly more total arthropods
and a greater biomass of arthropods in pitfall
traps near drift fences than in those near logs (Ta-
ble 2). The similarity of what was caught in the 2
types of traps ranged from 64.4% (SE = 3.6) in the
annually burned plots to 69.2% (SE = 1.4%) in the
quadrennially burned plots, but there were no
significant differences in similarity among burn
treatments. Likewise, Shannon diversity, even-
ness, richness, and numbers of rare species were
not significantly different for traps regardless of
adjacent barrier.

We observed significant (P < 0.05) interactions
between fire and trap location in 31 taxa (Table 3).
Of those, 10 taxa had higher numbers captured in
traps nears logs in some burn treatments (Fig. 1).
However, there was no consistent pattern, i.e., in
some cases frequent burning resulted in more of a
given taxa near logs while infrequent or no burning
had the same result for other taxa. In only a few
cases did frequent burning result in concentrations
of arthropods near logs, despite much lower leaf lit-
ter, less live herbaceous vegetation biomass, and re-
duced plant structure on those plots compared to
unburned or quadrennially burned plots (Hanula &
Wade 2003). Total volumes of coarse woody debris
among burn treatments were not significantly dif-
ferent and ranged from a mean of 8.0 m3/ha (SE =
1.01) for unburned control plots to 9.1 m3/ha (SE =
1.73) on annually burned plots.
The Monte Carlo test results of the NMDS in-
dicated a two-dimensional solution was optimal
(final stress 11.73, P = 0.0323). The first 2 axes of
the NMDS ordination explained 91.4% of the orig-
inal variance, with the first axis explaining 69.6%
and the second axis explaining an additional
21.8% (Fig. 2). The vector analysis indicated that
fire frequency was correlated with both axes,
more so with Axis 1 (R2 = 0.678, P < 0.05) than
Axis 2 (R2 = 0.574, P < 0.05). DCWD was also sig-
nificantly correlated to Axis 1 (R2 = 0.197, P <
0.05) but not Axis 2. Other environmental vari-
ables were not correlated at a = 0.05.
We captured over 932 genera in 5 years of trap-
ping of which 297 were captured in sufficient num-
bers for analysis. Of those, 101 arthropod taxa were
captured in significantly higher numbers in one
trap type or the other (Table 3). When examining
this many individual taxa, one is certain to encoun-
ter some significant results simply by chance. How-
ever, the 101 taxa represent 34% of the total number
of arthropod taxa with 30 or more individuals. At a
= 0.05 one would only expect 5% to have been cap-
tured in statistically higher numbers by chance.
Seventy three different arthropods were captured in
significantly (P < 0.05) higher numbers in pitfalls

Florida Entomologist 92(2)

June 2009


Drift Fence Log

Order Family Genus or Lowest Taxa Mean SE Mean SE P> F












Herpyllus" *

F .r.. i... ...,- ....r
Scotinella" *





Conocephalinae "

Gryllinae" *

11.9 1.5 2.8 0.6 <0.0001
7.0 1.0 3.4 0.6 0.0003
1.8 0.6 0.4 0.2 -
9.6 1.9 4.9 1.3 0.004
10.3 2.5 7.2 1.7 0.04
0.9 0.2 5.7 1.2 -
1.5 0.4 0.4 0.2 0.04
1.7 0.6 0.3 0.1 0.02
68.3 7.6 36.7 3.4 <0.0001
1.8 0.4 0.3 0.4 -
17.4 4.5 6.0 1.3 -
21.9 2.6 3.8 0.6 <0.0001
1.3 0.3 0.3 0.1 -
11.3 3.4 7.5 2.9 -
0.5 0.2 2.3 1.2 -
3.3 1.1 0.8 0.3 -
48.0 9.1 24.1 4.0 0.0001
13.8 2.3 8.0 1.9 0.01
34.2 8.5 16.1 3.5 -
423.8 46.5 287.1 26.2 <0.0001
1.7 0.6 0.8 0.2 -
51.5 7.2 28.3 6.4 0.0006
112.2 23.2 32.0 7.9 -
16.2 4.1 5.0 0.8 0.02
60.2 7.1 86.6 7.3 0.0005
46.8 9.3 36.6 7.1 0.05
19.9 2.1 12.3 2.3 0.04
104.0 9.5 77.8 8.2 0.005
4.1 0.7 8.3 0.9 0.0002
1.2 0.4 0.2 0.1 0.02
1.0 0.3 0.2 0.1 0.003
2.8 0.7 0.8 0.2 -
21.7 1.9 6.6 1.5 <0.0001
32.1 4.6 26.9 3.4 -
7.3 1.8 5.0 1.3 -
2.5 0.4 0.6 0.2 <0.0001
18.4 1.5 12.9 1.2 0.001
1.8 0.3 0.4 0.2 0.002
4.5 0.70 2.5 0.49 0.02
0.9 0.22 0.3 0.95 -
11.2 3.3 3.9 1.1 0.005
304.4 23.7 126.2 11.6 <0.0001
6.9 1.1 3.9 1.5 -
4.4 0.6 0.9 0.2 <0.0001
2.1 1.1 14.7 1.7 -
21.7 1.7 13.6 1.4 <0.0001
24.7 3.4 9.3 1.4 -
1.9 0.5 3.1 0.5 -
2.8 0.5 1.5 0.4 0.02

Asterisks (*) denote groups that were captured in higher numbers near logs.
Taxa followed by an "" had a significant interaction of trap location with burn frequency.

Hanula et al: Arthropods and Coarse Woody Debris


Drift Fence Log

Order Family Genus or Lowest Taxa Mean SE Mean SE P> F


Repipta *


Delphacidae" *









218.7 18.7
10.3 1.4
3.4 0.6
0.1 0.06
25.0 2.5
4.7 0.7
0.9 0.3
5.3 1.0
2.0 0.5
2.9 0.6
0.2 0.1
6.0 0.9
88.2 8.3
2.4 0.6
2.7 0.5
38.0 4.6
2.3 1.1
5.9 1.8
0.08 0.06
0.2 0.1
11.8 2.6
1.8 0.7
1.2 0.3
5.6 1.7
24.9 2.3
0.8 0.8
0.08 0.08
3.5 1.1
0.13 0.07
190.8 38.1
0.0 0.0
4.2 0.7
66.6 17.4
104.9 21.6
3.6 1.2
1.63 0.3
0.3 0.1
3.7 0.9
0.04 0.01
0.3 0.3
0.1 0.06
26.1 2.8
5.9 2.2
1.8 0.6
0.6 0.2
0.54 0.2
2.0 0.4
20.2 2.0
10.5 1.2
0.04 0.04

77.6 9.1 <0.0001
5.6 1.1 0.01
1.9 0.4 0.05
0.4 0.1 0.04
32.1 2.5 0.04
2.8 0.6 0.01
2.3 0.5 -
12.9 1.4 <0.0001
3.6 0.8 -
9.2 1.5 0.0003
1.9 0.3 -
0.3 0.1 <0.0001
42.6 3.3 <0.0001
0.7 0.1 -
0.6 0.1 0.0009
14.0 1.9 <0.0001
0.3 0.1 -
3.1 1.1 0.002
1.7 0.4 0.0003
2.3 0.5 0.0002
5.8 1.3 0.006
0.4 0.2 0.02
0.3 0.1 0.01
3.7 1.5 0.0001
3.6 0.6 -
2.9 0.9 0.008
2.5 0.5 0.0002
0.6 0.2 0.007
4.0 0.9 0.0005
91.1 18.9 0.003
2.4 0.8 0.01
1.5 0.4 0.003
23.4 8.8 <0.0001
54.5 12.9 0.03
0.7 0.2 0.02
4.8 0.7 0.0001
1.0 0.2 0.01
0.2 0.2 -
1.4 0.4 -
2.2 0.6 0.003
1.7 0.5 0.003
36.7 3.6 0.04
0.4 0.1 0.01
20.0 2.9 <0.0001
2.1 0.4 0.003
1.2 0.2 0.02
0.7 0.2 0.006
10.2 1.1 <0.0001
4.8 0.7 0.0005
2.3 0.71 0.007

Asterisks (*) denote groups that were captured in higher numbers near logs.
Taxa followed by an "" had a significant interaction of trap location with burn frequency.

Oilarus" *

Catonia" *

Megacephala "
Piemus *
Pterostichini *








Florida Entomologist 92(2)

June 2009


Drift Fence Log

Order Family Genus or Lowest Taxa Mean SE Mean SE P> F









Orfelia "

Sciara *






Asterisks (*) denote groups that were captured in higher numbers near logs.
Taxa followed by an "" had a significant interaction of trap location with burn frequency.

near drift fences while 28 different arthropods were
captured in higher numbers near logs. We plotted
the numbers captured by year to see if there was ev-
idence of depletion from pitfall trapping for a long
period of time or competition between traps. Fig. 3
shows 6 examples of those plots. Four years of trap-
ping did not affect the abundance of arthropods in
the vicinity of our traps nor did we see evidence of
declining trap captures in one trap type as the other


Prescribed burning over a 40-year period sig-
nificantly changed the understory vegetation and

litter layer of the study plots (Hanula & Wade
2003; Glitzenstein et al. 2003). Despite reduced
litter and less shrub cover on annually burned
plots, we saw no increased use of logs as habitat
by ground dwelling arthropods. Andrew et al.
(2000) reported similar results for ants although
they suggested that a few rare species may bene-
fit from the presence of logs in areas where fre-
quent low intensity fires are commonly used. We
found community similarity, diversity, and rich-
ness were the same for the 2 trap locations re-
gardless of the frequency of winter burning. Like-
wise, we saw no increase in overall arthropod
abundance or biomass near logs but instead we
captured more in traps near metal drift fences.










Hanula et al: Arthropods and Coarse Woody Debris

Araneae: Gnaphosidae
Herpyllus spp.

Araneae: Corinnidae
Scotinella spp.

Orthoptera: Gryllidae
immature Gryllinae


I 0 0, o

Orthoptera: Gryllidae

Homoptera: Delphacidae

,. !


* S

Homoptera: Cixidae
Catonia spp.

o a

Homoptera: Cixidae
Oilarus spp.

Coleoptera: Endomychidae
Epipocus spp.


Diptera: Sciaridae
Sciara spp.

o o

Diptera: Mycetophildae

/ /y/

Bum Frequency

Fig. 1. Mean number (SE) of individual arthropod taxa collected in pitfall traps near logs (*) or drift fences (0)
between 1994 and 2000. Trapping occurred in study plots in a north Florida longleaf pine forest, undergoing differ-
ent controlled burn frequencies. Graphs are for taxa in which fire frequency significantly affected trap captures and
captures near logs were greater than those in traps near metal drift fences. Probabilities (P > F) are for the trap
location x fire frequency interaction term in analyses of variance (Proc GLM, SAS 1987).

Habitat heterogeneity and structural diversity community richness and diversity (Hutchinson
are considered important factors in determining 1959; Southwood et al. 1979; Tilman & Pacala

Florida Entomologist 92(2)

S.D 2



*1.S -1f J

Fig. 2. NMDS biplot of fire freqi
downed coarse woody debris (DCWI
in a north Florida longleaf pine fore
points represent individual species s
angles the plots scores and the labe
their fire frequency in years, with U
burned plots. The angle and length
cate the direction and strength of the
respective variable with the species

1993; Wright et al. 1993). Howe
added logs did not result in grea
dance or diversity in their vicinit
burning on some plots, which g
the structure of the habitat for a
ula & Wade 2003).
Arthropod community structu
fire frequency and reported by
(2003). The first axis of the NMD
explained almost 70% of the var
correlated to fire return interv
Downed coarse woody debris di
pact on overall insect communi
the effect was much weaker tha
(R2 = 0.197). Possibly, the small
volume of woody debris on the p
cient to affect the arthropod co
study time frame was too short (
sults of Gunnarsson et al. (2004)
former may be the case. They fou
and abundance of ground-active
lated to the size of slash piles le
ing. Thus, larger amounts of dea
location may result in greater div
dwelling arthropods in the vicii
et al. (2004) also point out that p
sure arthropod activity and not r
dance. In contrast, Jabin et al.
soil arthropods in litter samples
tant from them in forests that
the same volume of dead wood a
They found that CWD was an
tural component to a variety of sE
cies regardless of season of sar

other groups abundance near CWD was affected
1 by timing of collection. Studies are underway to
1 determine if large volume inputs of woody debris
over an extensive area affect ground-dwelling ar-
thropods and other organisms (McCay et al.
: '2002).
I .. The higher numbers of arthropods captured
Near drift fences, and the higher number of taxa
captured more frequently in traps along drift
2 fences, suggest that the drift fences were more ef-
ficient at directing arthropods into pitfall traps
than logs. Hansen & New (2005) found that pit-
falls with barriers were more efficient at captur-
ing beetles than those without. Our data suggest
S that traps near metal drift fences were more effi-
cient than those near logs despite our efforts to
uency (FREQ) and prevent that. However, Sanzone (1995) also
)) from study plots caught more arthropods overall and higher num-
st, 1994-2000. The bers of most arthropod orders in pitfalls away
cores. The open tri- from logs. Sanzone's order level identifications
is above represent make it difficult to compare her results to our
indicating the un- study, but greater trap efficiency was not a factor
f the vectors indi- in that study because pitfalls away from logs did
relationship of the
and plot scores not have drift fences. Our data, when considered
together with Sanzone (1995), suggest that many
common ground-dwelling arthropods negotiate
around logs in ways that allowed them to avoid
ver, in our study being trapped. Originally we hypothesized that
ter overall abun- many arthropods would prefer the moist, shaded
y, despite annual habitat near logs. Again, our data and Sanzone's
reatly simplified (1995) suggest this is not generally the case for
arthropods (Han- most ground-dwelling arthropods. Although we
captured 36 taxa more frequently near logs, most
re was driven by were known saproxylic species. For example, bark
Hanula & Wade beetles (Curculionidae: Scolytinae), woodborers
S ordination that (Cerambycidae), and click beetle larvae (Elat-
iance was highly eridae) were captured more frequently near logs
aal (R2 = 0.678). and all either directly feed in large dead wood or
d have some im- are commonly found in it. Likewise, Medetera spp.
ty structure but flies (Dolichopodidae) captured more frequently
in fire frequency near logs are known predators of bark beetles
differences in the (Goyer et al. 1980) and many crane flies (Tipul-
lots were insuffi- idae) and fungus gnats (Mycetophilidae) are
immunity or the saproxylic (McAlpine et al. 1981). Beetles in the
5 years). The re- Endomychidae, Melandryidae, and Euglenidae
suggest that the captured more frequently near logs are also
nd that diversity known to be associated with fungi and rotten logs
beetles were re- (Borror et al. 1989). Although the majority of
eft after harvest- ground beetles (Carabidae) were captured more
d wood in a given frequently along metal drift fences, members of
rersity of ground- the genus Piemus and unidentified members of
lity. Gunnarsson the tribe Pterostichini were captured more often
)itfall traps mea- near logs. Thiele (1977) points out that prefer-
lecessarily abun- ences for certain levels of humidity among the
(2004) examined Carabidae is one of the most important factors
near logs and dis- governing their choice of habitats. Despite
contained about Thiele's (1977) conclusion that most forest cara-
s our study area. bid species preferred darkness and moist condi-
important struc- tions, we found only a few were common near logs.
iprophagous spe- In contrast, most rove beetles (Staphylinidae)
npling, while for were captured in greater numbers in traps near

June 2009

Hanula et al: Arthropods and Coarse Woody Debris

140 -- Log ptfalls
0 Lycosidae onn pa





6 Araneae









1994 1995 1996

1997 1998 1999


25 Carabidae



120 Staphylinidae




100 Scarabaeidae




1994 1995 1996 1997 1996 1999 2000

Year sampled

Fig. 3. Mean number (SE) of individual arthropod taxa collected between 1994 and 2000 in pitfall traps in
study plots in a north Florida longleaf pine forest. Graphs are examples of the variation in trap captures over time
shown to demonstrate that pitfall trapping over a 5-year period did not deplete arthropod numbers.

logs. Although staphylinids have diverse habits,
they are frequently found near decaying material
or beneath logs, stones or other debris. These re-
sults support those of Jabin et al. (2004) who also
found staphylinids were more abundant close to
logs. Overall, our data are similar to those of
Evans et al. (2003) who found that some inverte-
brate groups increased in abundance in litter
samples close to logs while others decreased. They
speculated that logs might influence abundance

of various arthropods in several ways including
increased predation near logs, changes in the lit-
ter composition near logs as they decompose, and
alteration of the C:N ratio of the fermentation
layer near logs. Clearly arthropods are a diverse
group and, as Thiele's (1977) extensive studies of
carabids show, even within a single family they
have highly varied habitat requirements.
Throughout the world dead wood is a critical
habitat to a number of saproxylic species that are

Florida Entomologist 92(2)

important for maintaining overall forest diversity
(Elton 1966; McMinn & Crossley 1996; Braccia &
Batzer 2001; Grove 2002; Grove & Hanula 2006).
Experience in European forests demonstrates
that without it a number of saproxylic species
could go extinct or be reduced to critically low lev-
els (Berg et al. 1994). We found that relatively few
non-saproxylic macroarthropods were captured
more frequently near logs even on plots where for-
est structure was greatly simplified by annual
burning. Arthropod community structure might
have been affected by the overall volume of logs
on frequently burned plots but the effect was
weak in comparison to the impact of fire fre-
quency. Detailed studies are needed to clearly un-
derstand how deadwood contributes to the habi-
tat needs of those species or groups found associ-
ated with it in longleaf pine communities.


We thank Arthur Phalo for coordinating sample
collection, processing, and data entry. Numerous stu-
dent workers sorted samples day in and day out,
Frances Brookshire labeled sample vials and proofed
data, and Lee Reynolds identified many of the speci-
mens. We thank James Pitts for identifications of spi-
ders and Hymenoptera, and Cecil Smith for
taxonomic help on other groups. Keith Lawrence and
the staff on the Osceola National Forest provided in-
valuable assistance throughout this study. Stan Zar-
noch helped with statistical analyses, and Simon
Grove and Jeff Lemieux provided helpful comments
on previous drafts of this paper. Finally, we thank the
Nature Conservancy's Ecosystem Research Program
and the A. W. Mellon Foundation who provided fund-
ing for this work.


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Casagrande et al.: Biocontrol of Birch Leafminer


'Department of Plant Sciences, University of Rhode Island, Kingston, RI, 02881, USA

2PSIS/Division of Entomology, University of Massachusetts, Amherst, MA, 01003, USA

3New Jersey Department of Agriculture, Beneficial Insect Rearing Laboratory, Trenton, NJ, 08625-0330,USA

4USDA-ARS, Beneficial Insects Research Laboratory, Newark, DE, 19713-3814, USA

5Cornell Extension, Long Island Horticulture Research Laboratory, Riverhead, NY, 11901-1115

Parasitoid releases against the birch leafminer Fenusa pusilla (Lepeletier) (Hymenoptera:
Tenthredinidae) in eastern North America began in 1974, with releases in eastern Canada,
followed by others in the Middle Atlantic States and New England. Of 4 parasitoids re-
leased, only 1, the ichneumonid Lathrolestes nigricollis (Thompson), established and spread
widely. Studies of its preliminary impacts were made in several locations in the 1980s and
1990s, but full impact of the parasitoid on host density was not yet achieved in that period.
Here we report results of surveys in 7 states (MA, CT, RI, NY, PA, NJ, DE) in 2007 docu-
menting the current birch leaf miner levels (as % of leaves mined in spring) and parasitism.
Survey results show that the pest has declined dramatically to barely detectable levels in 5
states (MA, CT, RI, NY, PA) but that in southern NJ, the pest remains abundant (ca 50%
leaves mined) despite significant parasitism levels. Survey results, in context with previous
evaluations made when populations were still declining, show that the project has been com-
pletely successful in much of the northeastern USA, but that there is a southern limit to ef-
ficacy in mid-New Jersey. Possible reasons for lack of control in this area, in contrast to high
levels of control elsewhere, are discussed.

Key Words: Fenusa pusilla, Lathrolestes nigricollis, birch leafminer, Betula spp. biological
control, evaluation, efficacy


La liberaci6n de parasitoides usados contra el minador de abedul, Fenusa pusilla (Lepele-
tier) (Hymenoptera: Tenthredinidae), en el este de Norte America empez6 en 1974, con libe-
raciones en el este de Canada, seguidas por otras en los estados del Medio-Atlantico y Nueva
Inglaterra. De los 4 parasitoides liberados, solo 1, el ichneum6nidos Lathrolestes nigricollis
(Thompson), se establecio y se esparcio ampliamente. Se realizaron studios sobre su im-
pacto preeliminario en varias localidades en los aios de la d6cada de los 1980 y 1990, pero
el impact complete del parasitoide no fue realizado en este periodo.Aqui reportamos los re-
sultados de los sondeos en 7 estados (MA, CT, RI, NY, PA, NJ, DE) en 2007 documentando
los niveles actuales de el minador de abedul (como un % de las hojas minadas en la prima-
vera) y el parasitismo. Los resultados del sondeo muestran que esta plaga ha disminuido
dramaticamente a niveles apenas detectables en 5 estados (MA, CT, RI, NY, PA) pero en el
sur de NJ, la plaga sigue abundante (ca. 50% de las hojas minadas) a pesar de niveles de pa-
rasitismo significativos. Los resultados del sondeo, en context con evaluaciones previas he-
chas cuando las poblaciones todavia estuvieron disminuyendo, muestran que el proyecto ha
sido exitoso en much del noreste de los Estados Unidos, pero hay un limited sureno para su
eficacia en el medio de New Jersey. Se discuten las razones posibles para la falta de control
en esta area en contrast con los niveles altos de control en otros lugares.

Birch leafminer (Fenusa pusilla [Lepeletier]) makes ornamental birches unsightly, although
(= pumila Klug) (Hymenoptera: Tenthredinidae), damage does not normally cause tree death. In-
an invasive Palearctic leaf mining sawfly attack- vestigations on natural sources of mortality asso-
ing several species of Betula, has been present in ciated with the pest in Connecticut (Friend 1933)
North America for at least 85 years, and was first and Quebec (Cheng & LeRoux 1965, 1969, 1970)
recorded in Connecticut (Friend 1933).When high showed that parasitism of larvae was insignifi-
proportions of tree leaves are mined, browning cant (<5%) compared to the situation in the its na-

Florida Entomologist 92(2)

tive range in central Europe, where parasitism
was much higher (38-47%) (Eichhorn & Pschorn-
Walcher 1973). This difference prompted the ini-
tiation of classical biological control programs in
both Canada and the USA, leading to the release
of 4 parasitoids: the ichneumonids Lathrolestes
nigricollis (Thompson) and Grypocentrus albipes
Ruthe, the braconid Phanomeris sp., and the eu-
lophid CL', ......,; i nitetis (Walker). The ichneu-
monids were released in Canada (Quebec and
Newfoundland) from 1974-75 (Raske & Jones
1975; Guevremont & Quednau 1977), and all 4
species were released in NJ, DE, MD, and PA be-
tween 1976 and 1982 (Fuester et al. 1984). Later,
releases of just the 2 ichneumonids were made in
MA (1979-1980), in MA and RI (1989-1991) (Van
Driesche et al. 1997), and western Canada in the
1990s (Langor et al. 2000).
Here we present information on the efficacy of
these releases in the northeastern United States.
A preliminary evaluation (Van Driesche et al.
1997) of the status of the pest in MA and RI con-
ducted from 1990-1995, showed that the pest had
declined locally around the principal MA release
site in Amherst from 54% of leaves mined in
spring to just 3%. Furthermore, by 1995 a differ-
ence had developed between Amherst, MA (at
various non-release sites within 4 km of the 1980
release site) versus a similar town (Northampton)
located 15-20 km from the release point. In the
Northampton area, where the parasitoid would
have arrived later, 33% of spring birch leaves
were still being mined and only 14% of the larvae
were parasitized by L. nigricollis. In contrast, in
Amherst, closer to the original release site, only
6% of spring leaves were mined and 28% of the
larvae were parasitized (Van Driesche et al.
1997). This suggests that at that point in time,
there was a zone, centered on the original release
area, where the pest was controlled, while further
away control was still developing.
Preliminary data published previously, while
suggesting the existence by the 1990s of local areas
of pest suppression, do not reflect the full extent of
area-wide pest control caused by this classical bio-
logical control project. Here, we present results of a
set of state-level surveys carried out in spring of
2007 by the authors to determine the current
abundance of F pusilla mines on birch species in
the northeast USA. Surveys were run in RI and CT
(Casagrande), Massachusetts (Van Driesche), New
Jersey (Mayer), Pennsylvania and Delaware
(Fuester) and Long Island, New York (Gilrein).


Sampling in Rhode Island (1990-2008)

Park in North Kingstown. Samples consisted of
100 leaves, haphazardly selected from a stand of
small (ca. 2-4 m) grey birch, Betula populifolia
Marshall, on the edge of large block of deciduous
forest. Individual leaves were plucked (without
looking at them) from branches distributed
throughout the site from leaves accessible from
the ground and were then returned to the labora-
tory where larvae were removed and dissected.
Samples were timed to closely precede larval drop
from the leaves for pupation. We did not measure
leaf damage until 1996, as until then it appeared
that all leaves in the stand were damaged. Begin-
ning in 1996, we noted whether there was any
birch leafminer damage to the leaves collected for
parasitism measurements. The site was sampled
from 1990 to 2008 except for 1992-93 and 2001-
03. Leaf damage for 1990 was determined by ex-
amining photographs of the sample site taken in
late May of that year.

State Surveys in 2007

Surveys were done in 7 states (MA, RI, CT, NY
[Long Island only], PA, NJ, DE) (Fig. 1) in May
and Jun of 2007 (Table 1). The survey was con-
ducted along roadsides in urban, suburban, or ru-
ral residential areas where grey or white (Betula
papyrifera Marshall) birch trees were present in
landscapes or naturally occurring along road-
sides. Following haphazard routes in such areas,
birch trees were examined when encountered.
From each tree, 100 leaves were plucked without
looking at them and then examined to determine
the number of leaves bearing distinctive F pusilla
mines. Leaves with leafminer larvae (from these
samples and supplemental samples) were taken
to the laboratory and dissected to detect eggs or
larvae of L. nigricollis. State summaries were

Beginning in 1990, the second year of releases Fig. 1. Areas (ellipses) surveyed for birch leafminer
ofL. nigricollis in RI, Casagrande and colleagues (Fenusa pusilla) and its parasitoid Lathrolestes nigri-
sampled for parasitism at a release site in Ryan collis in 2007.

June 2009

Casagrande et al.: Biocontrol of Birch Leafminer

IN 2007.

No. sites Sample % leaves mined % parasitism
State examined dates (# mined leaves/total) (# parasitized/ no. dissected)

MA 40 May 25-28 0.2 (8/4200) NA (no larvae recovered')
RI 23 May 30-Jun 5 0.5 (7/2300) NA (1 unparasitized larva recovered)
CT 2 May 31 0.0 (0/200) NA (no mines)
NY 70 May 20-Jun 20 0.0 (0/7000) NA (no mines)
PA 5 May 17-Jun 9 0.2 (6/3700) 0% (0/12)
NJ-north2 14 May 25-29 24.9 (348/1400) 57.9% (114/197)
NJ-south2 10 May 25-30 84.8 (848/1000) 27.5% (25/91)
DE 5 May 12-21 10.4 (114/1100) 1.2% (5/408)
Total 183 6.4% (1331/20,900) NA

No larvae were recovered to dissect for parasitism because larvae had already emerged from those few mines found.
'Northern NJ defined as 40NL or above; southern NJ being below 40NL. The northern third of NJ was not surveyed.

computed as the percentage of leaves with F pu-
silla mines (summed over all sites) and the per-
centage of birch leafminer larvae parasitized
(summed over all larvae from all sites). One ex-
ception to this process was that data from NJ
were kept separate by location because strong dif-
ferences occurred between sites in northern ver-
sus southern NJ. For comparison, the state was
split into northern (40NL or above) and southern
(below 40NL) parts.


Birch leafminer populations have come under
complete biological control in all areas surveyed
above 40NL (mid-New Jersey). Mined leaves
were extremely difficult to find in MA, CT, RI, PA,
and NY (Long Island) (Table 1). In these areas,
birch leafminer larvae could not be recovered in
large enough numbers to estimate rates of para-
sitism. In New Jersey, density of the pest has been
suppressed at most sites north of 40NL, but
south of this latitude, pest densities remain high
(Fig. 2). Parasitism in sites above 40NL (57.9%,
114/197) was approximately double the 27.5%
(25/91) rate found in southern NJ. Infestations of
birch leafminer at Newark, DE, were highly vari-
able, with the incidence of mined leaves ranging
from 0 to 57% on different trees and averaging
10.4%, intermediate between New Jersey and the
other states. Parasitism was low (1.2%), but sam-
ples were taken when most of the mines were
small. Because Lathrolestes nigricollis females
prefer to attack semi-mature and mature larvae
(Eichhorn & Pschorn-Walcher 1993), sample tim-
ing may have caused parasitism to be underesti-
Some insight into the development of control
over several decades is available from the re-
peated sampling of one site in RI, where annual

120 -






s3o 3NJS
southem NJ

381 40 40.2 4A
northem NJ (latitude)

Fig. 2. Percentage of birch leaves mined in 2007 by
birch leafminer versus latitude in New Jersey, showing
lack of control in southern New Jersey (left) versus
much greater pest suppression in northern New Jersey

samples were taken in most years from 1990 to
2008 (Fig. 3), and at this location, it is clear that
the percentage of leaves mined fell from about
87% (avg. for 1990, 1996, 1997, 1998) to about
2.6% (average for 2004, 2005, 2006, 2007, and
2008). During this same period, the rate of larval
parasitism by L. nigricollis rose steadily from an
average of 5.5% (1990, 1991, 1992, 1995, and
1996) to a high of 38% in 2004. Thereafter, too few
mines were present to obtain larvae to examine
for parasitism.


Fenusa pusilla, which remained a high-density
pest in the northeastern United States through-
out the 20th century, has now come under com-
plete biological control in MA, CT, RI, NY (Long
Island), PA, and northern NJ due to the intro-


*. .'


Florida Entomologist 92(2)

* No Data


1990 1992 1994 1996 199 2000 2002 2004 2006 20
Years Sampled

0g n NoData
a o

1990 1992 1994 1996 1996 2000 2002 2004 2006 2000
Years Sampled

Fig. 3. Percentage of grey birch leaves mined by first
generation Fenusa pusilla and percentage of larvae par-
asitized by Lathrolestes nigricollis at Ryan Park in
North Kingston, RI, between 1990 and 2008. (Years
with no bars prior to 2004 are missing data. After 2005,
larvae, and hence parasitism, were undetectable).

duced parasitoid L. nigricollis. In southern NJ,
the pest remains abundant despite relatively
high levels of parasitism. In DE, populations of
the pest are variable but lower than in southern
NJ. However parasitism rates observed in our DE
samples were also low. Why the pest has not been
suppressed in southern NJ or DE is unknown. In
terms of time, these areas have been colonized for
a longer period than the more northerly regions
where the pest has declined, so it seems unlikely
to be merely a transitional phase moving toward
control. Poor control by the parasitoid in southern
NJ may relate to poor photoperiodic match be-
tween the region from which the parasitoid was
originally collected (Austria, at 47-48NL) and
our sample areas in southern NJ (<40NL), or
may be due to southern NJ's higher summer tem-
peratures. Mismatch of photoperiod can limit par-
asitoid impact if adult emergence is not well
timed to the availability of the correct host stage,
or if the percentages of the parasitoid population
that enter diapause in the first and second gener-
ations differ from the diapause rates of the pest in
the corresponding generations (Nealis 1985; Van
Driesche et al. 1997). In DE, scarcity of suitable
birch species also may be important. Only 2 spe-
cies of birch (Betula lenta L. and Betula nigra L.)
occur naturally in Delaware (Taber 1995) and of
these, B. nigra is not a preferred host for F. pu-

silla. Preferred species such as gray birch or Eu-
ropean species exist only as planted landscaping
trees, and these are often in stressed condition
due to climate. Further research is needed in NJ
and DE to clarify the importance of these factors.
The results of our surveys illustrate the impor-
tance of continued monitoring of a classical bio-
logical control project for a very extended period,
here 34 years since the first releases. Evaluations
on short time scales (3-6 years), typical of many
research projects, may be premature and under-
estimate ultimate impacts of new biocontrol


This research was supported by the Cooperative
State Research Extension, Education Service, U.S. De-
partment of Agriculture and the Agricultural Experi-
ment Stations of participating states with coordination
through the regional biological control project NE1032.


CHENG, H. H., AND LEROUX, E. J. 1965. Preliminary life
tables and notes on mortality factors of the birch leaf
miner, Fenusa pusilla (Lepeletier) (Hymenoptera:
Tenthredinidae), on blue birch, Betula caerulea
grandis Blanchard, in Quebec. Annal. Soc. Entomol.
du Qu6bec 11: 81-104.
CHENG, H. H., AND LEROUX, E. J. 1969. Parasites and
predators of the birch leafminer, Fenusa pusilla (Hy-
menoptera: Tentrhedinidae) in Quebec. Canadian
Entomol. 101: 839-846.
CHENG, H. H., AND LEROUX, E. J. 1970. Major factors in
survival of the immature stages of Fenusa pusilla in
southwestern Quebec. Canadian Entomol. 102: 995-
parasites of the birch leaf mining sawfly (Fenusa pu-
silla [Lep.]) (Hym.: Tenthredinidiae) in central Eu-
rope. Commonwealth Institute of Biological Control,
Tech. Bull. 16: 79-104.
FRIEND, R. B. 1933. The birch leaf mining sawfly, Fe-
nusa pumila Klug. Bull Conn. Agric. Station (New
Haven) 348: 291-364.
SON, R. M., AND BLUMENTHAL, E. M. 1984. Introduc-
tion of exotic parasitoids for biological control of the
birch leafminer (Hymenoptera: Tenthredinidae) in
the middle Atlantic states. J. Econ. Entomol. 77:
GURVREMONT, H. C., AND QUEDNAU, F. W. 1977. Intro-
ducion de parasites ichneumonides pour la lutte bi-
ologique contre Fenusa pusilla (Hymenoptera:
Tenthredinidae) au Qu6bec. Canadian Entomol. 109:
SPENCE, J. R., AND SAUNDERS, C. 2000. Establish-
ment and spread of two introduced parasitoids (Ich-
neumonidae) of the birch leafminer, Fenusa pusilla
(Tenthredinidae). BioControl 45: 415-423.
NEALIS, V. 1985. Diapause and the seasonal ecology of
the introduced parasite Cotesia (Apanteles) rubecula
(Hymenoptera: Braconidae). Canadian Ento-
mol.117: 33-342.

June 2009

Casagrande et al.: Biocontrol of Birch Leafminer

RASKE, A. G., AND JONES, J. M. 1975. Introduction of
parasitoids of the birch leafminer into Newfound-
land. Bi-Monthly Res. Notes 31(3): 20-21.
TABER. W. S. 1995. Delaware Trees, 3rd ed., Delaware
Department of Agriculture Forestry Service, Dover,
Delaware, 284 pp.
AND TEWKSBURY, L. 1997. Establishment, distribu-

tion, and impact in southern New England of Lath-
rolestes nigricollis (Thompson) (Hymenoptera: Ich-
neumonidae), an introduced parasitoid of the birch
leafminer, Fenusa pusilla (Lepeletier) (Hy-
menoptera: Tenthredinidae). Canadian Entomol.

Florida Entomologist 92(2)

June 2009


'Department of Entomology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061

2Invasive Insect Biocontrol and Behavior Laboratory, USDA, ARS, Plant Sciences Institute, Beltsville, MD 20705

3Department of Entomology, University of Florida, Tropical Research and Education Center, Homestead, FL 33031

4Alson H. Smith, Jr. Agricultural Research and Extension Center, Virginia Polytechnic Institute and State University,
Winchester, VA 22602


The pink hibiscus ni.. :,1 i..._ 1........ .......... hirsutus (Green), is a highly polyphagous pest
that invaded southern Florida in 2002 and is now widely established throughout most of the
state. Although Hibiscus rosa-sinensis L. is a preferred and economically important host of
M. hirsutus, the susceptibility and expression of feeding symptoms by different cultivars
have not been evaluated. Cultivars of H. rosa-sinensis were infested with M. hirsutus and
evaluated daily for 40 d for the onset and percentage of terminals expressing feeding symp-
toms. Under different initial densities of M. hirsutus, the cultivar'President' showed no dif-
ference in the latency to expression of feeding symptoms, which occurred between 7 and 15
d after infestation, but did show significant differences between initial density and percent-
age of terminals expressing feeding symptoms from 10 d onward. When infested with 20 fe-
males, 80% of 'President' terminals exhibited symptoms 30 d after infestation. Four other
cultivars initially infested with 10 female M. hirsutus showed significant differences in the
onset and severity of feeding symptoms. All plants of the cultivars 'Florida Sunset' and
'Joanne' expressed damage symptoms at 12 2 SE d and 10 + 1 d, respectively, following in-
festation. Only a single plant of the cultivars 'Double Red' and 'Snow Queen' showed such
symptoms at 19 and 30 d after infestation, respectively. Significant differences between cul-
tivar and the percentage of terminals expressing feeding symptoms were observed from 20
d onward. Terminals sampled from all plants after 40 d revealed that egg, nymph, and adult
female M. hirsutus were found on all plants, including those that did not exhibit feeding
symptoms. These data have shown that hibiscus cultivars differ in their expression of M.
hirsutus feeding symptoms, that M. hirsutus can reproduce on cultivars of hibiscus that do
not express feeding symptoms, and that feeding symptoms are not a reliable indicator of in-
festation by M. hirsutus, highlighting the need for further investigation of the mechanisms
underlying differences among cultivars.

Key Words: Maconellicoccus hirsutus, Hibiscus rosa-sinensis, host tolerance


La cochinilla rosada del hibisco, Maconellicoccus hirsutus (Green), es una plaga altamente
polifaga que invadi6 el sur de la Florida en 2002 y ahora esta extensivamente establecida en
el mayor parte del estado. Aunque Hibiscus rosa-sinensis L. es el hospedero preferido y eco-
n6micamente important de M. hirsutus, la susceptibilidad y la expresi6n de los sintomas
debido a su alimentaci6n en diferentes variedades no han sido evaluados. Variedades de H.
rosa-sinensis con M. hirsutus fueron infestadas y evaluadas diariamente por 40 dias para la
aparici6n y porcentaje de parties terminales que expresaron sintomas de la alimentaci6n.
Bajo diferentes densidades iniciales de M. hirsutus, la variedad 'President' no mostraron
una diferencia en la latencia de la expresi6n de los sintomas de la alimentaci6n, que ocurri6
entire los 7 y 15 dias despu6s de la ingestaci6n, pero mostraron diferencias significativas en-
tre la densidad initial y el porcentaje de los terminales que expresaron sintomas de alimen-
taci6n de los 10 dias en adelante. Cuando las plants fueron infestadas con 20 hembras, 80%
de los terminales de la variedad'President' mostraron sintomas 30 dias despu6s de la inges-
taci6n. Cuatro otras variedades que fueron infestados con 10 hembras de M. hirsutus mos-
traron diferencias significativas en la aparici6n y severidad de las sintomas de la
alimentaci6n. Todas las plants de las variedades 'Florida Sunset' y 'Joanne' expresaron sin-
tomas de dano a los 12 2 SE dias y 10 1 d, respectivamente, despu6s de la ingestaci6n.
Solamente una plant en cada una de las variedades 'Double Red' y 'Snow Queen' mostr6 es-

Vitullo et al.: Maconellicoccus hirsutus Feeding Symptoms

tos sintomas a los 19 y 30 dias despu6s de la ingestaci6n, respectivamente. Se observaron di-
ferencias significativas entire las variedades y el porcentaje de los terminales que expresaron
sintomas de la alimentaci6n de los 20 dias en adelante. El muestreo revelo que se encontra-
ron huevos, ninfas y hembras adults de M. hirsutus en los terminales en todas las plants
despu6s de 40 dias, incluyendo aquellas plants que no mostraron sintomas de dano por la
alimentaci6n. Estos datos han mostrado que las variedades de hibisco varian en su expre-
si6n de los sintomas provocados por la alimentaci6n de M. hirsutus, que M. hirsutus puede
reproducirse en variedades de hibisco que no expresan sintomas por la alimentaci6n, que los
sintomas de la alimentaci6n no son indicadores confiables para evaluar la ingestaci6n de M.
hirsutus, y destaca la necesidad de hacer mas investigaciones de los mecanismos que causan
las diferencias entire las variedades.

The pink hibiscus mealybug, Maconellicoccus
hirsutus (Green) (Hemiptera: Pseudococcidae), is
a highly polyphagous pest that invaded southern
Florida in 2002 (Amalin et al. 2003), Louisiana in
2006 (Anonymous 2006), Texas in 2007 (Borgan &
Ludwig 2007), and Georgia in 2008 (Horton
2008). It feeds in phloem tissue and injects saliva
that can result in malformed leaf and shoot
growth, stunting, and plant death (USDA 2001).
Many economically important agricultural and
horticultural crops and common native plants
that are potential hosts of M. hirsutus occur in
Florida (Hodges & Hodges 2005) and throughout
the southern USA. Reported hosts include more
than 300 plant species in 74 families (USDA 2001;
Borgan & Ludwig 2007) and damaging popula-
tions have been recorded on numerous fruits, veg-
etables, and ornamentals (Hall 1921; Hall 1926;
Ghose 1972; Persad 1995; Chang & Miller 1996).
Variations among H. rosa-sinensis cultivars in
their susceptibility to M. hirsutus or their expres-
sion of feeding symptoms may have important
ramifications for management programs. In re-
sponse to the arrival ofM. hirsutus in Florida, a
biological control program involving the release of
2 encyrtid parasitoids,Anagyrus kamali (Moursi)
and Gyranusoidea indica (Shafee, Alam and
Agarwal), and the coccinellid predator Cryptolae-
mus montrouzieri (Mulsant) was implemented by
the Florida Department of Agriculture and Con-
sumer Services, Division of Plant Industry. New
infestation sites for release of biocontrol agents
are based on visual scouting of preferred hosts in
the landscape (USDA 2001). While M. hirsutus
populations have been greatly reduced in Florida
through the establishment and actions of biologi-
cal control agents (Amalin et al. 2003), biological
control does not cause local extinction ofM. hirsu-
tus populations (Ranjan 2004) and the pest has
continued to spread in Florida. Sex pheromone
traps for pink hibiscus mealybug deployed contin-
uously between 18 May 2007 and 10 Oct 2008
have shown that males continue to be captured at
locations in southern Florida where parasitoids
were released in 2007 (Vitullo, unpublished data).
Given that management decisions for M. hirsutus
are often based on the results of surveys recording
the presence or absence of symptoms, under-
standing the variations in the expression or se-

verity of these symptoms among cultivars is im-
portant. Furthermore, differences in susceptibil-
ity among cultivars may be helpful toward sus-
taining hibiscus production in Florida.
Hibiscus rosa-sinensis L. (Malvaceae), is con-
sidered a preferred host of pink hibiscus mealy-
bug (Hall 1921; Mani 1989; Kairo 1998) and 15
Hibiscus species have been confirmed with dam-
aging populations (Stibick 1997). Based on the
historical importance of hibiscus to ornamental
production in south Florida, M. hirsutus has had
negative impacts on local nurseries. A zero toler-
ance policy has been in effect since 2004 (FDACS
2005) and numerous quarantine actions have oc-
curred (Ranjan 2004; Gaskalla 2006). Retail dis-
tributors of ornamental plants in regions of the
US not presently infested by pink hibiscus mealy-
bug have reduced hibiscus orders to mitigate the
risk of being sent infested stock (J. Cou, Garden
Depot, Miami, FL, personal communication), es-
pecially since 2004, when infested hibiscus plants
were shipped from a Homestead, FL nursery to 36
States (Hodges & Hodges 2005). Additionally,
landscape design firms in southern Florida have
reduced the planting of hibiscus since the inva-
sion by M. hirsutus, and nurseries in southern
Florida have significantly curtailed hibiscus pro-
duction (J. Cou, personal communication). As
with many other host plants, typical feeding
symptoms from M. hirsutus on hibiscus include
leaf curl and shortened internodes, leading to ro-
setteing or "bunchy top" (USDA 2001). However,
during the course of our research in southern
Florida, we noticed that certain cultivars of hibis-
cus growing in residential areas appeared to suf-
fer more damage from pink hibiscus mealybug
than others, which may be attributed to symp-
toms varying among hosts (Stibick 1997; Anony-
mous 2005), and/or M. hirsutus preferring certain
hibiscus varieties (Abdel-Moniem et al. 2005).
The following research addresses the response
of several commercially important Hibiscus rosa-
sinensis cultivars to feeding by pink hibiscus mea-
lybug. Specific objectives were to determine the
relationship between mealybug density and the
expression of feeding symptoms in a susceptible
cultivar and the expression of feeding symptoms
in 4 cultivars infested with a single density of

Florida Entomologist 92(2)


Insects and Test Plants

Experiments were conducted in a greenhouse
at 28C at the University of Florida, Tropical Re-
search and Education Center, Homestead, FL.
Greenhouse temperature varied by 1C with a rel-
ative humidity range of 40 to 98%. Hibiscus rosa-
sinensis plants, approximately 1 m high and 0.66
m wide, were grown in 12-L plastic pots and
maintained in accordance with standard horticul-
tural recommendations (Ingram & Rabinowitz
1991). Periodic releases of Aphidius colemani
Vier. (IPM Laboratories, Inc., Locke, NY) kept
aphid levels low in the greenhouse.
Pink hibiscus mealybugs were reared on pota-
toes in a controlled environmental chamber at
26C and a 12:12 L:D regime (Serrano et al. 2001).
Plants were infested by transferring adult female
mealybugs with attached egg sac to the apical bud
of terminals with a micro-pin tool.

Expression of Feeding Damage in Relation to Mealybug

To evaluate the expression of feeding damage
in a susceptible H. rosa-sinensis cultivar infested
with different numbers of female mealybugs, we
placed 16 plants of the cultivar 'President' in the
greenhouse on 2 July 2007. Plants were infested
with 1, 5, 10, or 20 female mealybugs per plant (n
= 4 plants per infestation level). Due to con-
straints on the availability of mealybugs, the 4
replicates were infested on successive weeks, be-
ginning 19 Jul.
The number of terminals on each plant was re-
corded (range = 14-22 terminals per plant) and
plants were evaluated daily for 40 d for the num-
ber of terminals exhibiting "bunchy top" feeding
symptoms. This experimental duration allowed
for the completion of the first infesting generation
of mealybugs and the dispersal of the second gen-
eration crawlers on each plant (Mani 1989).

Expression of Feeding Damage by Hibiscus Cultivars

To evaluate feeding damage expressed by dif-
ferent H. rosa-sinensis cultivars, 3 plants of'Dou-
ble Red', 'Snow Queen', 'Florida Sunset' and
'Joanne' were placed in the greenhouse on 18 Jan
2008. These cultivars were selected following con-
sultation with Mr. J. Cou (Garden Depot, Miami,
FL) and were based on horticultural differences
among them and their commercial popularity.
'Double Red' has cordate leaves densely packed
along the shoot and red full double blooms.'Snow
Queen' has variegated cordate leaves with a
sprawling habit and fringed single red blooms.
'Florida Sunset' has cordate leaves and yellow
regular single blooms with an orange eye zone.

'Joanne' has ovate leaves and orange cartwheel
overlapped single bloom with a dark orange eye
zone. Plants were pruned to have between 17 and
20 terminals, then infested with 10 female mealy-
bugs on 12 Mar and evaluated daily for 40 d for
the expression of feeding symptoms.
After 40 d, 6 terminals (15 cm long) were
pruned from each plant. Terminals could now be
grouped into strata by presence or absence of
feeding symptoms, and a stratified random sam-
ple was used (Ott & Longnecker 2001). When pos-
sible, 3 terminals exhibiting "bunchy top" were re-
moved from each plant, and the remaining termi-
nals that exhibited no feeding symptoms were se-
lected to complete the sample size of 6 terminals.
Terminals were examined under a dissecting ste-
reomicroscope at 250X magnification and the
number of adult females, nymphs, and egg sacs
per terminal was recorded.

Data Analysis and Statistics

The latency to the first expression of feeding
symptoms (days) for 'President' plants with differ-
ent initial infestation levels was analyzed by ran-
domized complete block design ANOVA with 4
replicated blocks (SAS Institute 2001). The la-
tency to the first expression of feeding symptoms
was compared by descriptive statistics among hi-
biscus cultivars initially infested with an equal
number of mealybugs because plants that did not
become infested could not be included in the anal-
Repeated measures ANOVA (PROC MIXED,
SAS Institute 2001) was carried out to determine
effects of initial infestation density, days after in-
festation and their interaction on severity of feed-
ing symptoms by application of compound sym-
metry (cs) and first-order autoregressive (ar(1))
models (Ott & Longnecker 2001). Severity of feed-
ing symptoms was then compared among 'Presi-
dent' plants with different initial infestation lev-
els at 10, 20, 30, and 40 d after infestation by a
randomized complete block design ANOVA (SAS
Institute 2001), and evaluated for block effect. As-
sessment of the severity of feeding symptoms was
based on the percentage of terminals exhibiting
feeding symptoms on each plant, analyzed as arc-
sine square-root transformed percentages (Zar
1999). Severity of feeding symptoms was com-
pared among cultivars infested with the same
number of females with the same tests as above.
The number of unhatched egg sacs, nymphs, and
adult female mealybugs recovered from terminals
sampled after 40 d, regardless of feeding symp-
tom, was compared among cultivars by ANOVA
(SAS Institute 2001). The number of mealybugs
at different developmental stages found on culti-
vars with varied feeding symptoms was compared
with descriptive statistics. Analyses of data from
all experiments were considered significant at

June 2009

Vitullo et al.: Maconellicoccus hirsutus Feeding Symptoms

<0.05. Prior to ANOVA, tests for normality with
PROC UNIVARIATE (SAS Institute 2001) were
performed. Means of significant effects in ANOVA
were separated by Tukey Honestly Significant
Difference (HSD) test.

Infestation of the cultivar'President' with pink
hibiscus mealybug resulted in the expression of
"bunchy top" feeding symptoms in all plants
within 15 d. Initial infestation levels (mean la-
tency in days SE) of 1 (11.0 1.8), 5 (8.5 1.2),
10 (7.3 0.8), or 20 (7.0 0.6) adult females and
their egg sac did not significantly affect the la-
tency to the first terminal exhibiting feeding
symptoms (F= 2.23; df = 3,12; P = 0.1369). Low
levels of variance are evidence of an insignificant
block effect.
There were negligible differences between
compound symmetry and first-order autoregres-
sive analysis for all repeated measures compari-
sons for severity of feeding symptoms and initial
infestation density of the'President' cultivar, days
after infestation and their interaction. Compound
symmetry analysis on severity of feeding symp-
toms demonstrated a significant difference for ini-
tial infestation density (F = 50.24; df = 3,48; P <
0.0001), day after infestation (F = 65.65; df= 3,48;
P < 0.0001) and their interaction (F = 2.17; df =
9,48; P = 0.0415). As such, these data provide a
good indication of how the severity of feeding
symptoms changes over time under differing ini-
tial infestation densities; cumulative plant injury
over time was density dependent (Fig. 1). On d 10,
plants infested with 1 and 20 females differed sig-



E 4v



nificantly (F = 4.98; df= 3,12; P = 0.0180). On day
20, plants infested with 10 and 20 females dif-
fered from those with 1 female (F = 9.37; df= 3,12;
P = 0.0018). Plants infested with 20 females dif-
fered from all other plants at day 30 (F = 12.65; df
= 3,12; P = 0.0005) and 40 (F = 11.79; df = 3,12; P
= 0.0007) (Fig. 1). When day was excluded, sever-
ity of feeding symptoms was different for infesta-
tion density (F = 9.98; df = 3,48; P = 0.0001), but
not for block (F = 1.14; df = 3,48; P = 0.3436), or
their interaction (F = 0.48; df = 9,48; P = 0.8811).
Infestation of 4 cultivars with 10 pink hibiscus
mealybug females and their egg sac did not result
in the expression of bunchy top feeding symptoms
on all plants. One of 3 'Double Red' plants exhib-
ited symptoms, first occurring 19 d after infesta-
tion and totaled 2 shoots during the trial. One of
3 'Snow Queen' plants exhibited feeding symp-
toms, with 1 terminal exhibiting bunchy top at 30
d after infestation. All 'Florida Sunset' and
'Joanne' plants exhibited bunchy top, with a mean
( SE) number of days to first expression of feed-
ing symptoms of 12 2 and 10 1, respectively.
Repeated measures comparisons by compound
symmetry analysis on severity of feeding symp-
toms demonstrated a difference for cultivar (F =
82.73; df = 3,32; P < 0.0001), day after infestation
(F = 19.29; df = 3,32; P < 0.0001), and their inter-
action (F = 3.72; df = 9,32; P = 0.0028). As such,
these data provide a good indication of the change
in severity of feeding symptoms over time for dif-
ferent cultivars (Fig. 2). The severity of feeding
symptoms and cultivar at 10 d intervals showed
differences from 20 d after infestation onward; 20
d (F = 28.55; df= 3,8; P < 0.0001), 30 d (F = 25.59;
df = 3,8; P = 0.0002) and 40 d (F = 15.51; df = 3,8;

0 10 20 30 40
Day after initial infestation
Fig. 1. Mean cumulative percent of terminals of 'President' hibiscus plants exhibiting feeding symptoms in re-
sponse to different initial infestation densities of M. hirsutus.

Florida Entomologist 92(2)


l 30

c 20


.E 10


10 20 30

Day after initial infestation
Fig. 2. Mean cumulative percent of terminals of 4 hibiscus cultivars exhibiting feeding symptoms in response to
an initial infestation density of 10 female M. hirsutus.

P = 0.0011).'Florida Sunset' and'Joanne' were not
significantly different, but were different from
'Snow Queen' and 'Double Red', which did not dif-
fer (Fig. 2).
When terminals were removed from cultivars
on day 40, M. hirsutus egg sacs, nymphs, and
adults were found on all cultivars with or without
feeding symptoms (Table 1). The number of M.
hirsutus found on cultivars regardless of feeding
symptom resulted in 'Florida Sunset' having sig-
nificantly more unhatched egg sacs present than
'Double Red' and 'Snow Queen', but not 'Joanne'
(F = 5.94; df= 3,68; P = 0.0012).'Snow Queen' had
fewer nymphs than all other cultivars (F = 10.50;
df = 3,68; P < 0.0001). 'Florida Sunset' had signif-
icantly more adults than 'Snow Queen', but not
'Double Red' or 'Joanne' (F = 5.67; df = 3,68; P =
0.0016). The mean (SE) number of pink hibiscus
mealybugs found on terminals without feeding
symptoms (n = 51) was 8 2 egg sacs, 379 + 56

nymphs, and 10 + 2 adults, and terminals with
symptoms (n = 21), had 44 6 egg sacs, 1279
140 nymphs, and 49 7 adults. The presence of
adults and egg sacs at d 40 indicate that M. hirsu-
tus was able to reproduce on the hosts, based on
lifecycle duration (Mani 1989).
During our research with pheromone traps for
assessing populations of M. hirsutus in residen-
tial areas in south Florida, we noticed that not all
hibiscus plants exhibited the typical feeding
symptoms associated with pink hibiscus mealy-
bug, despite capturing many males in traps sus-
pended from these hosts (Vitullo, unpublished). It
is important for growers and landscape managers
to know that the number of pink hibiscus mealy-
bugs it takes to elicit feeding symptoms and the
latency from first infestation to the expression of
symptoms is not the same for all cultivars. The la-
tency to the first terminal exhibiting feeding
symptoms for the'President' cultivar was not den-


Presence No.
Cultivar of feeding symptoms of terminals Nymph Egg sac Adult female

'Double Red' No 16 620 + 93 15 4 18 5
'Double Red' Yes 2 907 + 262 23 7 28 1
'Snow Queen' No 17 86 + 23 4 1 5 1
'Snow Queen' Yes 1 101 17 17
'Florida Sunset' No 9 690 169 8 4 11 6
'Florida Sunset' Yes 9 1421 160 63 + 10 68 10
'Joanne' No 9 194 39 2 1 2 1
'Joanne' Yes 9 1352 245 34 7 38 8

June 2009

Vitullo et al.: Maconellicoccus hirsutus Feeding Symptoms

sity dependent. A single female with her egg sack
produced "bunchy top" feeding symptoms in 15 d
or less.
Timing of management action can be greatly
aided with information on differences in the la-
tency to the expression of the first feeding symp-
toms among different Hibiscus cultivars. For
'President', 'Florida Sunset', and 'Joanne', symp-
toms occurred in 7 to 15 d following infestation,
while in'Double Red' and 'Snow Queen', symptom
expression occurred much later or did not occur at
all. There may be different characteristics that
confer degrees of susceptibility. Abdel-Moniem et
al. (2005) evaluated the vertical distribution ofM.
hirsutus on three Hibiscus sabdariffa L. varieties
in an Egyptian nursery and found that only 1 in 3
varieties tested was attacked. Although feeding
symptoms were not evaluated, they suggested
that morphological and physiological characteris-
tics of H. sabdariffa may increase its susceptibil-
ity to M. hirsutus. While much additional re-
search would be necessary to reveal the mecha-
nisms of tolerance or resistance of hibiscus culti-
vars to pink hibiscus mealybug, traditional
breeding for reduced susceptibility to feeding in-
jury may be possible.
Expression of feeding symptoms varies be-
tween cultivars, but may be dependent on the
growth of the plant. The feeding symptoms of M.
hirsutus are an inability of the ground meristem
ofleafprimordia to differentiate into palisade and
spongy mesophyll when leaves open (Babu et al.
2004), suggesting that feeding symptoms will
only occur during this developmental time. The
limiting factor for hibiscus growth is temperature
(Ingram & Rabinowitz 1991), suggesting that the
temperature in the region and time of year also
may have an effect on the appearance of feeding
symptoms during hibiscus growth periods.While
infestation density had no effect on the latency to
first expression of feeding symptoms, cumulative
plant injury over time was density dependent.
When the cultivar 'President' was infested with a
single female and her egg sack, 33% of the termi-
nals exhibited "bunchy top" after 40 d, whereas
feeding by twenty females deformed 90% of the
terminals. Of the cultivars tested, only 'Joanne'
infested with 10 females expressed similar levels
of damage as 'President' at d 20. There was little
difference in the severity of symptoms from d 20
to 40 for 'Double Red', 'Snow Queen', 'Florida Sun-
set', and 'Joanne'. Anecdotal observations suggest
that newly-hatched crawlers of M. hirsutus may
walk for considerable distances on a plant before
settling to feed (Misra 1920). Feeding symptoms
before d 20 were likely caused by the infesting
generation. Based on the duration of the M. hirsu-
tus lifecycle (Mani 1989), symptoms from d 20 to
40 were likely caused by the second generation.
The colonization and resulting feeding symptoms
from the second generation would likely depend

on the size and structure of the plant being eval-
Many plants have been evaluated and shown
to be suitable for the development and reproduc-
tion of pink hibiscus mealybug (Serrano &
Lapointe 2002), while others expressed damage
symptoms but were not suitable for reproduction
or development (Kairo et al. 2000). Maconellicoc-
cus. hirsutus can reproduce on cultivars of hibis-
cus without producing symptoms, including 'Dou-
ble Red' and 'Snow Queen'. Given that feeding by
an initial density of 1 pink hibiscus mealybug per
plant elicited feeding symptoms on 'President'
and that feeding by 10 females caused minimal or
no damage on 'Snow Queen', it appears that feed-
ing symptoms are not a reliable indicator of the
presence or level of infestation of all hibiscus cul-
tivars by M. hirsutus.
When making landscape design selections,
knowledge of how pink hibiscus mealybug infes-
tations affect different hosts will aid in creating
aesthetic injury thresholds. Selection of plants
that are tolerant to pink hibiscus mealybug feed-
ing symptoms may make it possible to create low
management input landscapes for M. hirsutus in-
fested areas, especially in conjunction with bio-
control programs.

We thank the USDA-CSREES Southern Region IPM
Program, Award 2007-34103-18140 for funding, Garden
Depot (Miami, FL) for donating plants, and H. Glen for
technical assistance. Thanks to R. Fell (Dept. of Ento-
mology, Virginia Tech) and D. Pfeiffer (Dept. of Entomol-
ogy, Virginia Tech) for advising expertise.

NGUYEN, R. 2003. Biological control of pink hibiscus
mealybug in south Florida: a one-year assessment.
USDA-APHIS manuscript.
ANONYMOUS. 2005. Maconellicoccus hirsutus. OEPP/
EPPO Bulletin 35: 413-415.
ANONYMOUS. 2006. LSU agcenter entomologist says
leave hibiscus mealybug management to experts, ht-
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APPA, B. K. 2004. Anatomical alterations associated
with tukra symptoms in the leaf and stem of mulber-
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BORGAN, C. E., AND LUDWIG, S. 2007. Pink Hibiscus
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CHANG, L. W. H., AND MILLER, C. E. 1996. Pathway risk
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GASKALLA, R. 2006. Pink hibiscus mealybug (PHM) advi-
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HALL, W. J. 1921. The hibiscus mealybug, Phenacoccus
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HALL, W. J. 1926. The hibiscus mealybug in Egypt in
1925 with notes on the introduction of Cryptolaemus
montrouzieri. Ministry of Agric., Egypt, Tech. & Sci.
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HODGES, G. S., AND HODGES, A. C. 2005. Pink hibiscus
mealybug training manual. Pink Hibiscus Mealybug
Workshop, Gainesville, Florida.
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work.org/pub/php/news/2008/PHMealybug/ Access-
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INGRAM, D. L., AND RABINOWITZ, L. 1991. Hibiscus in
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r.i.. i. 1i .. l ...... 7, .*.......... hirsutus (Green). Report
for the Pink Hibiscus Mealybug Technology Transfer
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Upper Saddle River, NJ.

June 2009

Tormos et al.: New Species, Preimaginal Phases and Venom Apparatus ofAntrusa 255


'Unidad de Zoologia, Facultad de Biologia, Universidad de Salamanca, 37071-Salamanca, Spain
E-mail: tormos@usal.es


Antrusa montecristiensis, a new species of Dacnusini from Montecristo island (Italy), is de-
scribed, illustrated, and compared with allied species. Specific keys of the genus Antrusa for
the West Palaearctic region are provided. The preimaginal phases and the venom apparatus
ofA. curtitempus Fischer, Tormos, Docavo & Pardo, are described, illustrated, and compared
with species of allied genera. The larva stages are similar to those of Dacnusa; the immature
larvae differ in the number and distribution of the setae of the abdominal and thoracic seg-
ments, and the mature larva in the type of the tegumental differentations of the thorax and
abdomen as well as in the number and size of the sensilla of the maxillary palpi. The venom
apparatus of this species is very similar to that ofAntrusa flavicoxa (Thomson), differing
from it in length of the reservoir and the number of gland filaments.

Key Words: Hymenoptera, Braconidae, new species, immature stages, venom apparatus,
Antrusa, Spain


Se describe, ilustra, y compare con las species mas pr6ximas, una nueva especie de Dac-
nusini de la isla de Montecristo (Italia):Antrusa montecristiensis. Se proporcionan claves di-
cot6micas para la separaci6n de las species del g6nero Antrusa presents en el oeste
paleartico. Adicionalmente, las fases preimaginales y el aparato del veneno de A. curtitem-
pus Fischer, Tormos, Docavo & Pardo, se described, ilustran, y comparan con las species de
los g6neros mas afines. Los estados larvarios de esta especie son similares a los de Dacnusa,
pudi6ndose separar las larvas inmaduras a partir del numero y distribuci6n de las setas de
los segments toracicos y abdominales, mientras las larvas maduras pueden caracterizarse
mediante el tipo de diferenciaciones tegumentarias presents en el t6rax y abdomen, asi
como a partir del numero y tamaio de los palpos maxilares. El aparato del veneno de An-
trusa montecristiensis difiere, del deAntrusa flavicoxa (Thomson), en el numero de filamen-
tos glandulares y longitud del reservorio.

Translation by the authors.

The taxonomic rehabilitation of the genusAn-
trusa Nixon was proposed by Fischer et al.
(2004).Antrusa can be characterized and delim-
ited from Exotela Fdrster, Dacnusa Haliday and
Chorebus Haliday by a combination of the follow-
ing character states: (a) mandibles three-den-
tate, (b) vein nr antefurcal, (c) T1 with medial
longitudinal carina, and (d) no sexual dimor-
phism of the pterostigma. The latter character is
significant for separation from Dacnusa, but
cannot be applied easily without having both
sexes available. The longitudinal carina of T1
may be helpful. The three-dentate mandibles
separate Antrusa from Chorebus, and the ante-
furcal nr separates it from Exotela in the re-
stricted sense. We discovered a new species of
this genus, Antrusa montecristiensis sp. n., de-
scribed below, in Italy (Tuscan Archipelago: isola
di Montecristo). Regarding the preimaginal
stages of the species of this genus, it should be

noted that the structures that allow character-
ization of the final larval instar have not been
described (although see Capek 1970). Detailed
studies addressing the variation in gland and
reservoir morphology of the venom apparatus in
species of Dacnusini have been conducted by
Quicke et al. (1997) and Tormos et al. (2003). In
this article, the preimaginal phases and venom
apparatus of Antrusa curtitempus Fischer, Tor-
mos, Docavo & Pardo, 2004 are described.


Adults ofAntrusa montecristiensis sp. n. were
obtained in Mar 2000 with use of malaise trap
placed in the Tuscan Archipelago, isola di Mon-
tecristo. To study the preimaginal phases of A.
curtitempus, at beginning of Aug 1992, we col-
lected larvae of C(..,r ... ., horticola
(Goureau) (Diptera, Agromyzidae) mining leaves

Florida Entomologist 92(2)

of C. arietinum Linnaeus and took them to the
laboratory. We had obtained adults of this para-
sitoid from parasitized host pupae at Ayora (Va-
lencia, Spain) 2 years previously. To collect the
larvae we picked leaves from plants infested
with the agromyzids and placed them in contain-
ers of suitable dimensions whose openings were
covered with gauze held in place with a rubber
band. These receptacles were kept under envi-
ronmental conditions of temperature, relative
humidity (RH), and photoperiod. The larval and
pupal hosts were dissected periodically. The dis-
sections allowed us to study the egg, 2 immature
instars, mature larva, and the pupa of this spe-
cies. All dissections were performed in 0.9% sa-
line. For microscopic preparation of the preimag-
inal phases, the methods of Tormos et al. (2003,
2004) were employed.
The venom apparatus was prepared and
drawn according to the method described by
Quicke et al. (1992, 1997) (clorazol black method)
for dry museum specimens. The venom apparatus
was treated with a sodium hydroxide solution, af-
ter which the soft tissue was removed. It was then
possible to observe the characteristics of the re-
maining chitinous gland intima, which are not ap-
parent from examination of an intact gland and
The material examined (adults, immature
stages, and venom apparatus) is deposited at the
"Torres Sala" Entomological Foundation (Valen-
cia, Spain).
The terms for body morphology and wing ve-
nation, together with the criteria for collecting bi-
ometric data of adults, follow Fischer (1973,
2002) with 2 modifications: (1) mesosoma vs. tho-
rax, and (2) setae vs. hairs. All the material ex-
amined is deposited at the Museo del Medio Am-
biente (Valencia, Spain). The following abbrevia-
tions have been used in the descriptions: a2 =
lower vein of B (brachius); B = brachial cell; cql =
first cubital cross-vein; cu2 = 2nd abscissa of cu (
= cubital vein); cu2' = second abscissa of cubital
vein of hind wing; culb = lower cubital-anal cross
vein (3rd discoideal segment); d = discoidal vein;
F, Fl, F2, etc.= flagellomere (s), flagellomere 1, 2,
etc.; Fm, Fp = middle flagellomere (s), penulti-
mate flagellomere; M = medial cell of hind wings;
np = parallel vein; r' = radiellus (radial vein of
hind wing); nr = recurrent vein; nr' = recurrent
vein of hind wing; nv = nervulus; R = radial cell;
r, rl, r2 = radial vein, first, second abscissa of ra-
dius; st = pterostigma; SM' = submedial cell of
hind wings; T, T1, T2, T3, T2 + 3 = tergite (s), first,
second, third tergite, first + second tergite. The
terminology used in the description of the differ-
ent structures of the immature stages is that
used by Tormos et al. (2003, 2004). The terminol-
ogy used for characteristics of the gland and res-
ervoir parts of the venom apparatus follows Tor-
mos et al. (2003).


Antrusa montecristiensis sp. n. (Figs. 2-3)

Female-Head (Fig. 1 and 2a): Head 1.6-times
as wide as long, 1.7 times as wide as face, 1.35
times as wide as mesoscutum; eyes as long as
temples, protruding, behind eyes as wide as at
eyes; toruli in normal position, occiput bayed in-
wards; upper side, occiput and in the ocellar area
smooth; distance between ocelli greater than
ocellus width, distance between an ocellus and
eye as long as width of ocellar area. Face 1.6
times as wide as high, slightly and evenly con-
vex, middle elevation almost missing, with fairly
evenly distributed, scattered setae, seta points
discernable, edges of eyes only slightly converg-
ing below, nearly parallel sided. Clypeus slightly
convex, 2.7 times as wide as high, with few out-
standing setae. Tentorial pits round, their diam-
eter as great as the distance from eyes. Labrum
triangular, protruding, with inconspicuous se-
tae. Mandible slightly longer than wide, lower
edge straight, upper edge slightly directed up-
wards, with 3 teeth almost equally developed,
tooth 1 rounded, tooth 2 pointed and slightly pro-
truding, tooth 3 rounded, an incision between
tooth 2 and 3, outer surface shiny-to-uneven and
a few scattered setae. Antennae 30 antenno-
meres, longer than body, the basal flagellomeres
about 3 times as long as wide, the following
slightly shorter, Fp about 2 times as long as
wide; the setae as long as the segment width; in
lateral view 3 placodes visible. Mesosoma: 1.3
times as long as high, upper side convex. Mesos-
cutum (Fig. 2c) about 1.4 times wider than long,
notauli developed almost to the centre and crenu-
late, lateral and central lobes and declivity setose,
dorsal slit well differentiated. Prescutellar furrow
rectangular, with 3 longitudinal ridges. Scutel-
lum triangular. Postaxillae and metascutum gla-
brous. Propodeum reticulate, shiny, with short
and scattered pubescence, with pentagonal area,
a longitudinal carina inside, with basal carina
and costulae. Furrows of sides of pronotum crenu-
late below. Prescutellar furrow broad, irregularly
striated, tapering anteriorly and reaching edge,
not reaching middle coxa, prepectal furrow nar-
row, passing into the crenulate anterior me-
sopleural furrow, posterior mesopleural furrow
simple, epicoxal area of middle coxa with a few
scattered setae only, sternaulus short (Fig. 2e),
fairly deep and crenulated, not extending to the
posterior margin. Metapleuron with long scat-
tered setae distributed regularly, delimited from
propodeum by an irregular lamella. Hind femur 5
times as long as wide, hind tarsus scarcely
shorter than hind tibia. Wings (Fig. 3): st paral-
lel-sided, reaching beyond middle of R, r arising
from base of st by a distance as long as rl, the
latter slightly longer than the width of st when

June 2009

Tormos et al.: New Species, Preimaginal Phases and Venom Apparatus ofAntrusa 257

c /- -




2 1mm



^S~?0-"---- '
3 0.5 mm
Figs. 1-3. Fig. 1.Antrusa curtitempus Fischer, Tormos, Docavo & Pardo (female): Head in dorsal view; Fig. 2.An-
trusa montecristiensis sp. n. (female); Fig. 2a. Head in dorsal view; Fig. 2b. T2+3; Fig. 2c. Mesoscutum; Fig. 2d. T1;
Fig. 2e. Sternaulus; Fig. 3. Anterior right wing

infolded, distal half of r2 nearly evenly bent, R
not reaching tip of wing, cu2 developed by a dis-
tance greater than cql long, nr clearly antefur-
cal, d slightly longer than nr, nv postfurcal, B
about twice as long as wide, closed by vein culb,
np arising from middle of B; r' and cu2' indicated
only as folds, nr' absent. Metasoma: T1 (Fig. 2d)
as long as wide, apically 1.5-times as wide as ba-
sally, longitudinal median keel rather weak, the
remainder practically smooth, with a few setae
on its sides. T2 smooth (Fig. 2b). Setae of T2 + 3
(Fig. 2b) not distributed over the entire surface;
a broad, bare area between T2 and T3 (Fig. 2b).
Ovipositor sheath as long as hind basitarsus,
reaching slightly beyond tip of metasoma. Color:

Black. Dark brown on metasoma, except T1. Yel-
low on anellus, mouth parts, wing venation, T1,
and legs. Body length: 1.4 mm. Male-Like fe-
male except antennae with 32 antennomeres.
Material Examined: Holotype: 2, ITALY: Tus-
can Archipelago: Montecristo island, 15-
30.III.2000 (obtained through malaise trap).
Paratype: ditto, 1 6, 1 .
Etymology: The specific name of this species
refers to Montecristo island, where it was cap-
Taxonomic Position: The new species can be
distinguished with the following keys which are
indicated continuously.

1. Head (Fig. 1) behind eyes strongly narrowed; temples about half as long as eyes. Body length: 1.5 mm. Spain
......................................... A. curtitempus Fischer, Tormos, Docavo & Pardo, 2004
- Head (Fig. 2a) behind eyes as wide as at eyes or wider; temples about as long as eyes ................... .2
2. Head behind eyes widened. T2 weakly sculptured; T2 + 3 setose all over. Antennae 29-32 segmented; scape and
pedicel yellow. Body length: 2.5 mm. England, Germany, Central Russia .......... A. vaenia Nixon, 1954
- Head behind eyes not or only slightly wider than at eyes (Fig. 2a). Setae ofT2 + 3 not distributed over the entire
surface; a broad, bare area between T2 and T3 (Fig. 2b). T1 sometimes longitudinally striated ......... 3

1 04 mm

Florida Entomologist 92(2)

3. r2 nearly evenly bent (Fig. 3).................................................................. 4

- r2 distally bisinuate .................. .................................................6

4. Mesoscutum setose only anteriorly, the rest predominantly bare. T1 longitudinally striated, weak points between the
striae, shiny. T2 bare, smooth. Hind femora 5.5-times as long as wide in apical third. T1 brownish. T2+3 dark
brown. Antennae 31-segmented. Body length: 2.2 mm. Central Russia ........ A. chrysotegula (Tobias, 1986)

- Mesoscutum nearly entirely covered with short, fine setae (Fig. 2c). Antennae 30-32 segmented ............ 5

5. Hind femora 4.5 times as long as wide. T1 rugose-punctate, matte. T2 weakly longitudinally rugose at base. T1 black,
T2 dark brown, rest of metasoma yellow. Body length: 2.3 mm. Moldavia ...... A. chrysogastra (Tobias, 1986)

- Hind femora 5 times as long as wide. T1 with the longitudinal median keel rather weak, laterally practically
smooth (Fig. 2d). T2 smooth (Fig. 2b). T1 yellow, rest of metasoma dark brown. Body length: 1.4 mm. Italy
................ .................... ............................ A. montecristiensis sp. n.

6. Head behind eyes widened. Antennae 23-36-segmented. T1 narrow, folds stronger. T2 smooth. Body length: 2.2-
2.3 mm. Western Europe; North-West and Central Russia; Azerbaijan ... A. melanocera (Thomson, 1895)

- Head behind eyes not widened. Antennae 28-34-segmented. T1 somewhat wider, the folds rather weak. T2 some-
times weakly sculptured. Body length: 2.0-2.4 mm. Western Europe; North-, Central and South-West Rus-
sia; Azerbaijan; Siberia (Irkutsk) ............... ................. A. flavicoxa (Thomson, 1895)


The egg and first instar were found in host lar-
vae at different developmental stages. The first
instar remained within the trophamnion until the
host formed its puparium. The trophamnion layer
consist of polygonal cells. Second and third in-
stars were only found in hosts that had pupated.
Second and third instars and pupa were found in
host puparia. A specimen of each stage was stud-
ied for description.
Egg. The egg (Fig. 4) is ovoid, slightly pointed
at one end, and hymenopteriform in shape.
Shortly after being laid, the egg varies between
130-160 pm (X (mean) SD = 141 12.02, n = 4)
in length and between 50-71 pm (X SD = 57.4 +
8, n = 4) in width. The developing eggs, during
segmentation, increase in size (430-517 pm, n =
2), and become more spherical, tending towards
an oblong shape at the end of their development.
Larva. First instar Body (Fig. 5): Length and
width (at the level of the mesothoracic segment):
0.8 x 0.25 mm, with head well defined and 13 body
segments, caudate, vermiform. Last abdominal
segment with a well-differentiated ventral lobe in
the form of a tail (1 = 90-85 pm), with 28 setae (1 =
60 pm) distributed in a fan around the anus. Seg-
ments 1-12 with a row of setae (1 = 45 pm) on their
posterodorsal part, the numbers corresponding to
10 (mesothorax), 16 (metathorax), and between
20 and 40 (abdominal segments). Cranium (Fig.
5a) (w = 140 pm) with pleurostomal processes and
the hypostoma heavily slerotized, and with an
area of small sclerotized spicules positioned an-
tero-ventrally. Mouthparts: Mandibles (Fig. 5a')
well defined, with an oblong molar lobe and one
blade (1 = 15 pm) sharp, curved, and well sclero-

Second instar. Body (Fig. 6): Length = 0.7 mm;
w (at the level of the mesothoracic segment) =
0.20 mm, cylindrical, long with respect to me-
sothoracic width, with a reduced tail. Integument
bare. Without prominent cephalic sclerites or
Third instar. Body (Fig. 7): Typical hymenop-
teriform (1 = 1.70, w = 0.60 mm), more grub-like,
wit the tail further reduced in length, yellowish.
Integument with small setae (1 = 3 pm) and mi-
crotrichiae covering the thoracic and abdominal
segments, except the intersegmental zones and
around the spiracles and anus. Nine pairs of spir-
acles (Fig. 7b) (diameter = 10 pm), with the
atrium and closing apparatus well differentiated,
1 pair on the prothorax and another on the ante-
rior edge of each of the first 8 abdominal seg-
ments. Cranium (Fig. 7a): Width (maximum) =
0.20 mm, height (taken from the base of the man-
dibles) = 0.10 mm, reduced, weakly sclerotized,
with setae (1 = 3 pm); orbital antennal circular (d
= 85 pm), weakly protuberant; pleurostoma, supe-
rior and inferior mandible processes, hypostoma
and stipital sclerite well differentiated and sclero-
tized; the latter joined to the labial sclerite, which
is weakly sclerotized. Mouthparts. Mandibles
(Fig. 7a'): Length of blade = 20 pm with broad
base and relatively long blade, curved, unarmed
(smooth) unidentate, sclerotized; maxillary and
labial palpi oval, slightly protuberant, with one
sensilla [d = 3 pm] in the case of the labial palpi,
and with 1 sensilla [d = 3 pm] and the other
minute [d = 2 pm] in the case of the maxillary
palpi; salivary orifice well defined (1 = 20 pm).
Pupa. Exarate. Without cocoon.
The preimaginal phases ofA. curtitempus are
similar to those described for Dacnusa. The differ-
ences mainly lie in the number and distribution of

June 2009

Tormos et al.: New Species, Preimaginal Phases and Venom Apparatus ofAntrusa

4 __

6 __
a.1 m



ia 1
/ 1

0.5 mm

0.05 mm

7a'6k- mm

0.2 5 m d'

7a _
0.05 mm

Figs. 4-7. Fig. 4. Egg; Fig. 5. First instar (lateral view) showing the distribution of the setae of the body and lobes
of the last abdominal segment; Fig. 5a. Cranium; Fig. 5a'. Mouthparts; Fig. 6. Second instar (lateral view); Fig. 7.
Third instar (lateral view) showing the distribution of the tegumental differentations of the body; Fig. 7a. Cranium;
Fig. 7a' Mandible; Fig. 7b. Second spiracle; Fig. 8. Venom apparatus with detail of reservoir, secondary venom duct,
venom gland and gland filaments (sacks).

the setae of the abdominal and thoracic segments
of the first instar larva and in the type of the teg-
umental differentiations of the thorax and abdo-
men, as well as in the number and size of the sen-
silla of the maxillary palpi of the mature larva.
Unlike what was reported by Capek (1970) for
Antrusa melanocera (Thomson 1895), the last in-
star of A. curtitempus has a well differentiated
and sclerotized stipital sclerite.


This species has a venom apparatus with the
character states specified by Quicke et al. (1997)
for Antrusa flavicoxa (Thomson 1895). Thus, A.
curtitempus has (a) an undivided reservoir, (b) a
reservoir neck region without narrowing, (c) a sec-
ondary venom duct absent, (d) an extensively
branched venom gland (in this case with 6 sacks,
inA. flavicoxa with 7), (e) a venom gland inserted
at the extreme posterior end of the reservoir, and
(f) a secondary venom duct that is not narrow. The
morphological differences with the venom appa-
ratus ofA. flavicoxa are (a) the reservoir inA. cur-
titempus is between 4-6 times longer than maxi-
mally wide and in A. flavicoxa is 8 times longer
than maximally wide, and (b) the venom gland in

A. curtitempus has 6 sacks and inA. flavicoxa the
venom gland has 7 sacks.

Financial support for this paper was provided from
the Junta de Castilla y Le6n, project SA012A05, and the
Fundaci6n Entomol6gica "Torres-Sala".

CAPEK, M. 1970. A new classification of the Braconidae
(Hymenoptera) based on the cephalic structures of
the final instar larva and biological evidence. Can.
Entomol. 102: 846-875.
FISCHER, M. 1973. Das Tierrich. Hymenoptera, Bra-
conidae, Opiinae (Palarktische Region). Lief. 91: I-
XII. Walter de Gryter, Berlin.
FISCHER, M. 2002. Ubersicht tiber die Gattungen der
Aspilota-Genusgruppe mit Neubeschreibung von
Grandilota no. gen. sowie Redeskription von Rege-
tus Papp (Hymenoptera, Braconidae, Alysiinae). Z.
Arb. Gem. Ost. Ent. 54: 99-108.
2004. A new species ofAntrusa and three new spe-
cies of Chorebus (Hymenoptera: Braconidae) from
the Iberian Peninsula. Florida Entomol. 87: 306-311.
MARSH, P. M. 1992. Venom gland apparatus in cy-

Florida Entomologist 92(2)

clostome braconid wasps with special reference to
the subfamily Doryctinae (Insecta, Hymenoptera,
Braconidae). Zool. Scr. 21: 403-416.
H. C. 1997. Comparative morphology of the venom
gland and reservoir in opiine and alysiine braconid
wasps (Insecta, Hymenoptera, Braconidae). Zool.
Scr. 26: 23-50.
GAYUBO, S. F. 2003. Descriptions of adults, imma-
ture stages and venom apparatus of two new species

of Dacnusini: Chorebus pseudoasphodeli sp. n., par-
asitic on Phytomyza chaerophili Kaltenbach and C.
pseudoasramenes sp. n., parasitic on Cerodontha
phragmitophila Hering (Hymenoptera: Braconidae:
Alysiinae; Diptera: Agromyzidae). European J. Ento-
mol. 100: 393-400.
A. 2004. Descriptions of the final instar of Eurytoma
nodularis and E. heriadi (Hymenoptera: Eurytomi-
dae). Florida Entomol. 87: 278-282.

June 2009

Chen & Tsai: New Genera of Delphacids from China


1Guizhou Key Laboratory for Plant Pest Management of Mountainous Region, Guizhou University, Guiyang,
Guizhou Province, P. R. China 550025

2Institute of Entomology, Guizhou University, Guiyang, Guizhou Province, 550025, P. R. China

3Fort Lauderdale Research and Education Center, IFAS, University of Florida 3205 College Avenue, Fort
Lauderdale, FL 33314, USA


Two new genera of Tropidocephalini (Hemiptera: Fulgoroidea: Delphacidae: Delphacinae)
are described from Hainan Province, China. They are Yuanchia Chen and Tsai gen. nov.
and Neocarinodelphax Chen and Tsai gen. nov. One new species, Yuanchia maculata Chen
and Tsai sp. nov. (China: Hainan: Diaoluoshan) and 1 new combination, Neocarinodelphax
hainanensis (Qin and Zhang 2005) comb. nov. (China: Hainan: Wuzhishan, Tongshi,
Yancheng) (transferred from Carinodelphax Ding and Yang) are described or redescribed
and illustrated.

Key Words: Hemiptera, Fulgoroidea, Delphacidae, Tropidocephalini, new genus, new spe-
cies, new combination, bamboo pests, China


Se described dos g6neros nuevos de Tropidocephalini (Hemiptera: Fulgoroidea: Delphaci-
dae: Delphacinae) de la Provincia de Hainan de China. Estos son Yuanchia Chen y Tsai gen.
nov. y Neocarinodelphax Chen y Tsai gen. nov. Se described y se ilustran una nueva espe-
cie, Yuanchia maculata Chen y Tsai sp. nov. (China: Hainan: Diaoluoshan) y una nueva
combinaci6n, Neocarinodelphax hainanensis (Qin y Zhang 2005) comb. nov. (China: Hai-
nan: Wuzhishan, Tongshi, Yancheng) (transferida de Carinodelphax Dingy Yang).

The delphacid tribe Tropidocephalini (Hemi-
ptera: Fulgoroidea: Delphacidae: Delphacinae)
was erected by Muir (1915) and it is the second
largest tribe of Delphacinae. Tribal characters in-
clude forewings with R, separated from Rs+M ba-
sally; hindwings with M stalked with Culat basal
half, anal areas with IA, IIA, and IIA2, all reach-
ing posterior margin; post-tibial spur large and
thick, concave on inner surface, without teeth
along the hind margin; spinal formula of hind leg
5-6-4; aedeagus often tightly connected with ven-
tral portion of anal segment; complicated asym-
metrical distortion in the basal part of the aedea-
gus with at least one elongated process arising
from this part (Asche 1985; Chen 2003; Ding
Muir (1915) included 6 genera in the Tropi-
docephalini when he proposed it as a tribe of the
Delphacinae, and he included species having a
solid post-tibial spur but with a concave inner
surface and no teeth along the edge. Asche (1985)
increased the number to 21 genera. The tribe
Tropidocephalini has now been expanded to in-
clude more than 31 genera. Most of species of the
tribe feed on bamboo (Bambusoideae) or other
grasses (Gramineae). They are mostly Oriental

with 26 genera restricted to that region and an-
other 2 shared with the Neotropical region. Two
genera are restricted to the Neotropical, 2 to the
Palaearctic and 1 to the Australian region. The
widespread genus Tropidocephala Stal is distrib-
uted through the Palaearctic, Afrotropical, Orien-
tal, Australian, and Pacific regions (Donaldson
1991; Chen & Li 2000b; Chen 2003; Ding 2006).
The fauna of Chinese tribe Tripidocephalini is
very abundant and 10 new genera were erected in
the last 3 decades, as follows: Bambusiphaga
Huang and Ding (Huang et al. 1979), Speciner-
uures Kuoh and Ding (Kuoh et al. 1980), Paranec-
topia Ding and Tian (Ding & Tian 1981),Neobelo-
cera Ding and Yang (Ding et al. 1986), Arcifrons
Ding and Yang (Ding et al. 1986), Malaxella Ding
and Hu (Ding et al. 1986), Carinodelphax Ding
and Yang (Ding & Yang 1987), Carinofrons Chen
and Li (Chen & Li 2000b),Mirocauda Chen (Chen
2003) and Gufacies Ding (Ding 2006). A key to
genera of the tribe Tropidocephalini from China
was provided by Chen (2003), who described a
new genus. Seventeen genera and 68 species, in-
cluding 1 new genus, 7 new species, and a revised
key were provided in a monograph on the Delpha-
cidae of China (Ding 2006). Recently, the revision

Florida Entomologist 92(2)

and description of new species of some genera in
the tribe Tropidocephalini were also reported
(Chen et al. 2006; Chen & Liang 2007; Chen et al.
2007a,b). Most species of Tropidocephalini from
China feed exclusively on Bambusoideae (Yang et
al. 1999; Chen 2003; Chen et al. 2007a,b; Ding
2006). To date, the Chinese tribe Tropidocephalini
has been increased to 18 genera, 3 subgenera and
82 species (Muir 1913, 1915; Fennah 1956; Huang
et al. 1979; Kuoh 1979, 1980, Kuoh et al. 1980;
Ding & Tian 1981; Ding 1982, 1987, 2006; Ding &
Hu 1982, 1991; Ding et al. 1986, 1999; Yang &
Yang 1986; Zhu 1988; Yang 1989, 1992; Qin &
Yuan 1998, 1999; Chen 1999, 2002, 2003; Chen &
Ding 2000; Chen & Li 2000a,b, 2002; Chen et al.
2000, 2006, 2007a,b; Qin & Zhang 2005; Chen &
Liang 2007).
In this paper 2 new genera, Yuanchia gen.
nov. and Neocarinodelphax gen. nov., 1 new spe-
cies, Yuanchia maculata sp. nov., and 1 new com-
bination, Neocarinodelphax hainanensis (Qin and
Zhang, 2005) comb. nov. (transferred from Car-
inodelphax Ding and Yang) from the Hainan
Province, China, are described or redescribed and


Morphological terminology used in this work
follows that of Yang & Yang (1986). The genital
segments of the examined specimens were macer-
ated in 10% KOH and drawn from preparations in
glycerin jelly with aid of a light microscope. Illus-
trations of the specimens were made with a Leica
MZ 12.5 stereomicroscope. Spinal formula means
the numbers of apical spines of the hind tibiae
and 1t and 2nd hind tarsomeres. The type speci-
mens and examined specimens are deposited in
the Insect Collection at the Institute of Entomol-
ogy, Guizhou University, Guiyang, Guizhou Prov-
ince, China (IEGU).


Yuanchia Chen and Tsai gen. nov. (Figs. 1-11, 21-25)

Type species. Yuanchia maculata Chen and
Tsai sp. nov., here designated.
Diagnosis. In profile, dorsum, including vertex,
pro- and mesonotum and posterior margin of
forewings, in almost the same plane (Fig. 21). Ver-
tex nearly trapeziform, lateral carinae distinctly
keeled and slightly converging distad (Figs. 1, 22).
Y-carina with stalk absent (present in Fig. 1).
Head truncate in lateral view, frons at right angle
to the longitudinal axis of body (Figs. 21, 23).
Frons rectangular, narrower at base, lateral mar-
gins parallel from middle to apex, lateral carinae
keeled, median carina forked at extreme base
(Figs. 2, 24). Antennae cylindrical, exceeding level
of frontoclypeal suture (Figs. 2, 24). Forewing

broadening apically, rounded at apex (Figs. 3, 21).
Anal segment of male with ventral margin sad-
dle-backed, overlaying base of phallobase and
aedeagus (Figs. 8, 9). Pygofer with opening longer
than broad (Fig. 6). Genital styles slender, diverg-
ing distad (Figs. 6, 10, 11).
Description. Head including eyes narrower
than pronotum, vertex trapeziform, with margins
well defined. Vertex at base wider than long in
middle line, distinctly narrower at apex than at
base; submedian carinae originating from near
apical 1/4 of lateral carinae, uniting at anterior
margin of vertex; lateral carinae distinctly keeled
and slightly converging distad (Figs. 1, 22); Y-car-
ina with stalk absent. Head truncate in lateral
view, frons at right angle to longitudinal axes of
body (Figs. 21, 23). Frons rectangular, narrower
at base, lateral margins parallel from middle to
apex, lateral carinae keeled, median carina
forked at extreme base (Figs. 2, 24), about 3 times
longer in middle line than widest part, widest
above level of ocelli to apex. Clypeus at base as
wide as frons at apex, tricarinate (Figs. 2, 24), in
profile with post-clypeus strongly curving caudad
(Fig. 23); rostrum almost reaching hind trochant-
ers. Antennae exceeding level of frontoclypeal su-
ture, cylindrical, basal segment slightly widening
distad, about 1.5 times as long as wide, second
segment about 2 times as long as first (Figs. 2,
24). Pronotum as long as vertex medially, poste-
rior margin concave medially, lateral carinae in-
curved, reaching to hind margin. Mesonotum
about 1.5 times as long as vertex and pronotum
together in middle line, median carina discontin-
uous medially (Figs. 1, 22). Forewings broadening
apically, rounded at apex, row of cross veins lo-
cated in apical half of wing (Figs. 3, 21). Spinal
formula of hind leg 5-6-4, post-tibial spur with
apical tooth (Fig. 25).
Male Genitalia. Pygofer with opening longer
than broad (Fig. 6), in profile much shorter dor-
sally than ventrally (Fig. 5). Two sides of dia-
phragm separated from each other (Fig. 7). Aede-
agus slender, tubular, phallobasal process arising
basally (Fig. 8). Genital styles slender, diverging
distad (Figs. 6, 10). Anal segment of male short
and ring-like, lateroapical angles separated, each
moderately produced ventrad in a rounded lobe
(Figs. 4, 8), ventral portion saddle-backed, over-
laying base of phallobase and aedeagus (Figs. 4, 8,
9). Anal style long (Figs. 8, 9).
Host Plant. Unknown.
Distribution. Oriental (Indo-Malayian) Region
(southern China).
Er I,.. .. .... The name is derived from translit-
eration of the Chinese "yuan-chi" meaning the
rounded apex of forewings, a feature seldom seen
in Tropidocephalini. This genus is feminine in
Remarks. The features of the post-tibial spur
and of the male genitalia place this genus in the

June 2009

Chen & Tsai: New Genera of Delphacids from China

,/ 2


Figs. 1-11. Yuanchia maculata Chen and Tsai sp. nov. 1. Head and thorax, dorsal view; 2. Frons and clypeus;
3. Forewing; 4. Anal segment and anal style, dorsal view; 5. Male genitalia, lateral view; 6. Male genitalia, posterior
view; 7. Pygofer, posterior view; 8. Anal style, anal segment, connective and genital style, lateral view; 9. Anal style,
anal segment and aedeagus, posterior and ventral view; 10. Genital styles, posterior view; 11. Left genital style, lat-
eral view. Scale bars = 0.5 mm (Figs. 1-2); 1 mm (Fig. 3); 0.2 mm (Figs. 4-11).

Tropidocephalini. It is closely related to the genus
Arcofacies Muir in the shape of vertex and frons,
but differs in: vertex with length in middle line
shorter than width of base about 0.79:1.00 (about
0.53-0.59:1.00 in the latter); frons longer in mid-
dle line than widest part about 3.25: 1.00 (about
1.75-2.17:1.00 in the latter); frons with median
carina forked at extreme base (not forked in the
latter); dorsum of body without median longitudi-
nal stripe from apex of vertex to end of scutellum
(in the latter, dorsum of body with a narrow stripe

along median longitudinal carina of vertex, pro-
and mesonotum, which are often white, bordered
with brown or fuscous stripe); forewings rounded
at apex acuatee at apex in the latter). This new ge-
nus is also related to Carinofrons Chen and Li,
but differs in: vertex distinctly narrower at apex
than at base, submedian carinae originating from
lateral carinae subapically, uniting at anterior
margin of vertex (in the latter, vertex at apex as
wide as at base, submedian carinae originating
from lateral carinae medially, not uniting at apex

Florida Entomologist 92(2)

of vertex); frons with lateral margins subparallel
from basal 1/3 to apex, median carina forked at
extreme base (in the latter, frons with lateral
margins convex in middle, median carina forked
above the level of ocelli); anal segment of male not
sunk into dorsal emargination of pygofer (in the
latter, anal segment of male deeply sunk into dor-
sal emargination of pygofer); sides of diaphragm
separated (dorsal margin of diaphragm with ar-
mature in the latter).

Yuanchia maculata Chen and Tsai sp. nov. (Figs. 1-11,

Description. Body length (from apex of vertex
to tip of forewing): male 3.50 mm, female 4.05
mm; forewing length: male 2.90 mm, female 3.45
General color pale yellowish brown (Figs. 21-
25). Vertex with median carina and adjacent lat-
eral areas pale yellowish white (Fig. 22). Frons
brown with several pale yellowish spots (Fig. 22),
lateral carinae brown, median carina pale yellow-
ish brown; apex fuscous. Clypeus yellowish
brown, brown at base. Genae fuscous, lateral
margins and 3 spots near lateral carinae of frons
pale yellowish white (Fig. 24), area above ocelli,
between eyes and lateral carinae of frons with 3
brown spots (Fig. 23). Eyes fuscous infused with
red, ocelli red. Antennae with segment I brown; II
with proximal portions and apex brown (Fig. 24).
Pro- and mesonotum with outer areas of lateral
carinae, brown, areas between lateral carinae yel-
lowish, median carina yellowish white, lateral
margin of scutellum with two fuscous spots (Figs.
1, 22). Forewing yellowish brown to brown, darker
apically, 11 fuscous spots distributing along mid-
dle of costal margin to nearly middle of posterior
margin, a larger fuscous spot on fork of Cu1 (Figs.
3, 21). Legs pale yellowish brown, darker distad
(Figs. 21, 25). Abdomen yellowish ventrally,
brown dorsally, lateral areas pale yellowish to yel-
lowish brown, genital segment brown to fuscous.
Structural features as in generic descriptions.
Vertex wider at base than long submedially
(about 1.27:1.00). Frons longer in middle line
than broad at widest part (about 3.25:1.00). An-
tennae with basal segment 1.60 times as long as
wide, second segment 2.13 times as long as first.
Mesonotum 1.62 times as long as vertex and
pronotum together at middle line. Forewing
longer in middle line than broad at widest part
Male Genitalia. Aedeagus moderately long, tu-
bular, thick at base, slender at apex, slightly curv-
ing ventrad, apex abruptly bent to left and form-
ing a scoop, tapering apically, orifice dorsally near
apex (Figs. 8, 9). Phallobasal process slender, sub-
equal length of aedeagus, arising from base of
aedeagus dorsally, directed ventrocaudad and
curving to right distad (Figs. 8, 9). Genital styles

slender, diverging distad, broader at base, taper-
ing apically, apical half sinuate, apex rounded
(Figs. 10, 11). Anal segment of male with lat-
eroapical angles slightly produced ventrocaudad
in nipple-like process, separated from each other
(Figs. 4, 6).
Host plant. Unknown.
Distribution. South China (Hainan Province).
Specimens Examined. Holotype male, CHINA:
Hainan, Diaoluoshan (1847'N, 109052'E), 16-VII-
2007, Q.-Z. Song; paratype 1 male, same data as
holotype; 1 female, Hainan, Jianfengling
(18042'N, 108047'E), 12-VII-2007, Q.-Z. Song
Er I .. ..,. The name is derived from the Latin
word "macula" (marking) referring to the spots on
the scutellum and margin of the forewing.

Neocarinodelphax Chen and Tsai gen. nov. (Figs. 12-
20, 26-29)

Type species. Carinodelphax hainanensis Qin
and Zhang, 2005 (here designated).
Diagnosis. Dorsum of body in profile not in the
same plane, posterior margin of forewings dis-
tinctly keeled (Fig. 26). Vertex trapeziform with a
well defined margin, lateral carinae strongly
keeled, Y-carina with stalk weak (Figs. 12, 27).
Frons in profile at an acute angle to longitudinal
axes of body (Figs. 26, 28), subrectangular, widest
at apex, lateral carinae distinctly keeled, median
carina simple (Figs. 13, 29). Antennae cylindrical,
extending to the level of frontoclypeal suture
(Figs. 13, 29). Forewings with acute apical angle,
outer margin approximately straight, oblique,
posterior margin sinuate, with irregular mark-
ings at apical half (Figs. 14, 26). Pygofer of male
with inner side of each lateral margin having a
stout process (Figs. 16, 17).
Description. Dorsum of body in profile not in
the same plane, posterior margin of forewing dis-
tinctly keeled (Fig. 26). Head including eyes nar-
rower than pronotum. Vertex trapeziform, at base
wider than long medially (about 1.6:1.0), lateral
carinae strongly keeled, diverging caudad, Y-car-
ina with stalk weak, submedian carinae originat-
ing from lateral margin subapically, uniting at
anterior margin of vertex (Figs. 12, 27). Frons in
profile at an acute angle to longitudinal axes of
body (Figs. 26, 28), subrectangular, widest at
apex, lateral carinae strongly keeled, median car-
ina simple (Figs. 13, 29), longer in middle line
than widest about 2.69: 1.00. Clypeus at base
slightly wider than frons at apex, tricarinate, in
profile with post-clypeus in same plane as frons,
ante-clypeus curving caudad (Fig. 28); rostrum
nearly reaching posterior trochanters. Antennae
cylindrical, extending to frontoclypeal suture,
basal segment slightly widening distad, about 1.8
times as long as wide, second segment about 1.5
times as long as first. Pronotum as long as vertex

June 2009

Chen & Tsai: New Genera of Delphacids from China

Figs. 12-20. Neocarinodelphax hainanensis (Qin and Zhang) comb. nov. 12. Head and thorax, dorsal view; 13.
Frons and clypeus; 14. Forewing; 15. Hindwing; 16. Male genitalia, lateral view; 17. Male genitalia, posterior view;
18. Lateral process of pygofer; 19. Anal style, anal segment, aedeagus, connective, and genital style, lateral view;
20. Right genital style, lateral view. Scale bars = 0.5 mm (Figs. 12-13); 1 mm (Fig. 14-15); 0.2 mm (Figs. 16-20).

medially, posterior margin concave medially, tri-
carinate, developed, lateral carinae recurved,
reaching to hind margin. Mesonotum tricarinate,
but not distinctly keeled, about 1.8 times as long
as vertex and pronotum together in middle line
(Figs. 12, 27). Forewings broadening apically,
cross veins located in apical half, with apical an-
gle acute, outer margin approximately straight,
oblique, posterior margin sinuate, with irregular
markings at apical half (Figs. 14, 26). Hindwing
subtriangle, M+Cula and Culb with a short com-

mon stalk, vein A with 3 branches (Fig. 15). Spi-
nal formula of hind leg 5-6-4, post-tibial spur with
apical tooth.
Male Genitalia. Pygofer in profile much
shorter dorsally than ventrally, laterodorsal an-
gles obtusely rounded, not produced caudad, lat-
eral margins excavate at middle, each very
strongly produced dorsad with medial pillar-like
projection (Fig. 16); posterior opening longer than
broad, medioventral process absent, ventral margin
concave (Fig. 17). Aedeagus slender, tubular; phal-

Florida Entomologist 92(2)

28 1 29 25

Figs. 21-29. Yuanchia maculata Chen and Tsai sp. nov. and Neocarinodelphax hainanensis (Qin and Zhang)
comb. nov. Yuanchia maculata (Figs. 21-25) Chen and Tsai sp. nov.: 21. Male adult, lateral view; 22. Male adult,
dorsal view; 23. Head and thorax, lateral view; 24. Frons and clypeus; 25. Hind leg; Figs. 26-29. Neocarinodelphax
hainanensis (Qin and Zhang) comb. nov.: 26. Female adult, lateral view; 27. Female adult, dorsal view; 28. Head
and thorax, lateral view; 29. Frons and clypeus.

lobasal process arising basally (Fig. 19). Genital
styles long (Fig. 17). Anal segment of male short
and ring-like, lateroapical angles not produced
(Figs. 16, 17). Anal style moderately long (Figs.
16, 19).

Host Plant. Unknown.
Distribution. Oriental Region (southern
Er I, .... .. The genus name, which is mascu-
line, is a combination of "neo" (new) and "carino-

June 2009


Chen & Tsai: New Genera of Delphacids from China

delphax" (name of the related genus), in recogni-
tion of its similarity to the genus Carinodelphax
Ding and Yang.
Remarks. This genus is related to Carinodel-
phax Ding and Yang, but differs in having frons
widest at apex (widest at level of ocelli in the lat-
ter); antennae relatively long, reaching the level
of the frontoclypeal suture (in the latter, antennae
rather short, not reaching frontoclypeal suture);
pro- and mesonotum with 3 carinae normal (in
the latter, 3 carinae of pro- and mesonotum, espe-
cially median carina of scutellum strongly
keeled); the color pattern of forewings is also dif-
ferent. This genus resembles Acrofacies Muir in
color pattern of dorsum and of frons, but differs in
having the frons longer in middle line than widest
part about 2.69:1.00, widest at apex (in the latter,
frons longer in middle line than widest part about
1.75-2.17:1.00, widest at a level of ocelli). In pro-
file, frons at an acute angle to longitudinal axes of
body, post-clypeus in the same plane as frons (in
the latter, frons at right angle to longitudinal axes
of body, post-clypeus strongly curving caudad);
forewings relatively long and narrow, longer than
broad about 2.69:1.00, costal margin relatively
straight (in the latter, forewings relatively short
and broad, longer than broad about 2.20-
2.26:1.00, costal margin curved); pygofer with an
elongated projection at inner side of lateral mar-
gin (absent in the latter); aedeagus with long spi-
nose phallobasal process (phallobasal absent or
degenerative in the latter).

Neocarinodelphax hainanensis (Qin and Zhang) comb.
nov. (Figs. 12-20, 26-29)

Carinodelphax hainanensis Qin and Zhang,
2005: 390.
Description. Body length (from apex of vertex
to the tip of forewing): male 3.25 mm, female 3.80-
3.85 mm; forewing length: male 2.75 mm, female
3.30 mm.
General color brown to fuscous (Figs. 26-29).
Median carina of frons and clypeus, lateral mar-
gins of genae, pale yellowish white; antennae yel-
lowish brown, middle and apex of basal segment,
base of second segment, with blackish bands,
apex of second segment brown (Fig. 29). A narrow
stripe from apex of vertex to the end of scutellum,
a narrow stripe along lateral carinae of pronotum,
pale yellowish white, narrowly edged fuscous
(Fig. 27); posterior half of lateral areas of prono-
tum, and propleura, basal half of humeral plate,
pale yellowish white (Figs. 12, 26-28). Forewings
with irregular hyaline markings (Figs. 14, 26).
Abdomen with dorsal and ventral surface, genital
segment, yellowish brown to brown; anal segment
yellowish brown.
Structural features as in generic descriptions.
Vertex wider at base than long submedially (1.60:
1.00). Frons longer in middle line than wide at

widest part (2.69: 1.00). Antennae with first seg-
ment 1.82 times as long as wide, second segment
1.5 times as long as first. Mesonotum 1.86 times
as long as vertex and pronotum together in mid-
dle line. Forewing longer in middle line than wide
at widest part (2.69:1.00).
Male Genitalia. Anal segment of male with lat-
eroapical angles not produced, conjoined basally
to the base of aedeagus (Fig. 19). Pygofer in poste-
rior view with opening longer than wide, ventral
margin concave broadly, medioventral process ab-
sent; inner side of lateral margin with a very long
projection dorsally, apex rounded and with many
minute teeth (Figs. 16-18); pygofer with posterior
margin medially concave in lateral view (Fig. 16).
Aedeagus tubular, slender, sinuate, thick at base,
narrowing apically, apex flattened, ventral side of
apical half concave, apex margin rounded, orifice
dorsally near apex (Fig. 19); phallobasal process
long, spinose, tapering distad, arising from base
of aedeagus dorsally, slightly decurved, longer
than half of aedeagus (Fig. 19). Genital styles
thick and long, reaching to ventral margin of anal
segment, in posterior view nearly parallel (Fig.
17), in profile broader at base, narrower at apex,
apical margin truncate, inner aspect with a hook-
like process, inner basal part and apex with
minute teeth (Fig. 20).
Host plant. Unknown.
Distribution. South China (Hainan Province).
Specimens examined. 1 male, CHINA: Hainan,
Wuzhishan (1853'N, 109041'E), 13-VII-2007,
lamping, Q.-Z. Song; 1 female, Hainnan, Wuzhis-
han, 15-VII-2007, Q.-Z. Song; 2 males, 1 female,
Hainan, Diaoluoshan (1847'N, 109052'E), 16-VII-
2007, Q.-Z. Song (IEGU).

We are grateful to Ms. Q. Z. Song (Institute of Ento-
mology, Guizhou University, Guiyang, Guizhou Prov-
ince, China) for providing valuable specimens. This
research was supported by the National Natural Sci-
ence Foundation of China (No. 30100015, 30560020), by
Program for New Century Excellent Talents in Univer-
sity, (NCET-07-0220) by China Postdoctoral Science
Foundation (No. 2005037111), by the Provincial Foun-
dation for Excellent Youth in Science and Technology
Field of Guizhou (No. 20050520), and by the Nomarch
Foundation for Excellent Talents in Science, Technology
and Education Field of Guizhou (No. 2005357).


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June 2009

Ovruski et al.: Larval-Pupal Parasitoids of Fruit Flies


PROIMI Biotecnologia-CONICET, Divisi6n Control Biol6gico de Plagas, Laboratorio Moscas de La Fruta. Avda.
Belgrano y Pje. Caseros s/n (T4001MVB) San Miguel de Tucuman, Tucuman, Argentina


Parasitoids ofAnastrepha fraterculus (Wiedemann) and Ceratitis capitata (Wiedemann) in-
festing wild guava (Psidium guajava L.) and peach (Prunus persica (L.) Batsch) fruits in the
southernmost extension of the Bolivian Yungas forest, in the Province of Tarija, were sur-
veyed in Dec (early summer), Feb (mid summer) and Mar (late summer) 1999 and 2000. The
abundance patterns and diversity of fruit fly larval-pupal parasitoid species were similar to
previously published data for the southern and northern Yungas forests of Argentina. A total
of 1,600 guavas and 800 peaches, weighing 57.713 kg and 24.544 kg, respectively, were col-
lected yielding 13,080 tephritid puparia, 78.4% of which were A. fraterculus and 21.6% Cer-
atitis capitata (Wiedemann). We reared 712 larval-pupal parasitoids of the following species:
Doryctobracon areolatus (Sz6pligeti), D. brasiliensis (Sz6pligeti), D. crawfordi (Viereck),
Opius bellus Gahan, Utetes anastrephae (Viereck) (all Braconidae, Opiinae),Aganaspis pel-
leranoi (Brethes) Odontosema anastrephae Borgmeier, and Lopheucoila anastrephae (Ro-
hwer) (all Figitidae, Eucoilinae). Utetes anastrephae, 0. anastrephae, and L. anastrephae are
new fruit fly parasitoid reports for Bolivia, as well as 0. anastrephae in the South American
Yungas forest. A thorough sampling of other C. capitata andAnastrepha host plants, mostly
native fruit species, will be necessary in the Yungas forest of Bolivia before further conclu-
sions on abundance and composition of all fruit fly parasitoids can be reached.

Key Words: fruit flies, parasitoids, Braconidae, Figitidae, Bolivia, Yungas


Se reportan los resultados de un relevamiento de parasitoides asociados conAnastrepha fra-
terculus (Wiedemann) y Ceratitis capitata (Wiedemann) en guayabas (Psidium guajava L.)
y duraznos (Prunus persica (L.) Batsch) silvestres. Estas frutas fueron sistematicamente co-
lectadas en un sector del extreme mas sureno de la selva de Yungas Boliviana, en la provin-
cia de Tarija. El studio se realize en los meses de diciembre, febrero y marzo de 1999 y 2000.
Los patrons de abundancia y la diversidad de las species de parasitoides larvo-pupales de
"moscas de la fruta" registrados en el present studio fueron similares a los datos previa-
mente publicados para los sectors sur y norte de la selva de Yungas en Argentina. Se colec-
taron en total 1.600 guayabas y 800 duraznos, cuyos pesos totales fueron 57,713 kg y 24,544
kg respectivamente. De estos frutos, se obtuvieron 13.080 puparios de tefritidos, de los cua-
les el 78,4% fueronA. fraterculus y el 21,6% restante fueron Ceratitis capitata. Se identifi-
caron 712 parasitoides larvo-pupales pertenecientes a las species Doryctobracon areolatus
(Sz6pligeti), D. brasiliensis (Sz6pligeti), D. crawfordi (Viereck), Opius bellus Gahan, Utetes
anastrephae (Viereck) (Braconidae, Opiinae), Aganaspis pelleranoi (Brethes) Odontosema
anastrephae Borgmeier, y Lopheucoila anastrephae (Rohwer) (Figitidae, Eucoilinae). Utetes
anastrephae, 0. anastrephae, y L. anastrephae son citadas por primera vez para Bolivia. Sin
embargo, es necesario efectuar en las Yungas de Bolivia muestreos mas meticulosos de otras
plants hospederas de C. capitata yAnastrepha spp., principalmente species de frutas na-
tivas, para llegar a conclusions mas precisas sobre la abundancia y composici6n de los pa-
rasitoides de "moscas de la fruta".

Translation by the authors.

The native South American fruit fly, Anas- capitata (Wiedemann), are some of the most im-
trepha fraterculus (Wiedemann), and the intro- portant pests affecting commercial fruit produc-
duced Mediterranean fruit fly (Medfly), Ceratitis tion in Bolivia (Zavaleta Castro 2003). Both te-

Florida Entomologist 92(2)

phritid species are commonly found throughout
Bolivia's southern region, including fruit-growing
areas of the Bolivian provinces of Tarija and
Santa Cruz, where fruit infestation levels in com-
mercial peaches (Prunus persica (L.) Batsch) vary
between 50 and 90% (Escalante 1995; Zavaleta
Castro 2003).
The original native vegetation in this Bolivian
region is a subtropical montane rainforest, locally
known as "Yungas" (Kessler & Beck 2001), and is
extended throughout Argentina's northwestern
region (Provinces of Jujuy, Salta, Tucuman, and
Catamarca) (Brown et al. 2001). This phytogeo-
graphical region is characterized by a high diver-
sity of native and exotic fruit fly host plants grow-
ing in the remaining stands of pristine forest and
in perturbed areas adjacent to commercial fruit
crops and orchards (Ovruski et al. 2005).
Even thoughA. fraterculus and C. capitata are
significant pests in the Bolivian fruit-growing re-
gions, little information on the fruit fly parasi-
toids in Bolivia is available. Preliminary surveys
in the Bolivian Yungas forest by Escalante (1995)
revealed that native hymenopterous parasitoids
commonly attack A. fraterculus larvae infesting
mainly exotic host plants, and five Anastrepha
parasitoid species native to the Neotropical re-
gion listed for Bolivia by Escalante (1995) and by
Rogg & Camacho (2000) were Doryctobracon bra-
siliensis (Szepligeti), D. areolatus (Szepligeti), D.
crawfordi (Viereck), Opius bellus Gahan, and
Aganaspis pelleranoi (Brethes). Nevertheless, lit-
tle is known about their abundance, parasitiza-
tion rates, host flies and host plant ranges, and
distribution patterns in Bolivia.
Between 1969 and 1971, classical biological
control was attempted through the introduction
of Psyttalia concolor (Szepligeti) (reported as Bio-
steres concolor), Diachasmimorpha longicaudata
(Ashmead) (reported as Biosteres or Opius longi-
caudatus) (Braconidae), Tetrastichus giffardianus
Silvestri, Aceratoneuromyia indica (Silvestri) (re-
ported as Syntomosphyrum indicum) (Eu-
lophidae), Pachycrepoideus vindemiae (Rondani)
(Pteromalidae), and Dirhinus giffardii Silvestri
(Chalcididae) into Bolivia (Pruett 1996; Ovruski
et al. 2000). These parasitoid species were re-
leased in limited numbers in fruit-growing areas
of the provinces of Cochabamba and La Paz
(northern portion of Bolivian Yungas forest) and
in the province of Santa Cruz (southern section of
Bolivian Yungas forest). All parasitoid species,
with the exception of D. giffardi, were recovered
immediately following releases in the three Boliv-
ian provinces (Pruett 1996). Later, between 1976
and 1978 new introductions of D. longicaudata
into Bolivia and releases of thousands of this fruit
fly parasitoid were made in coffee crops in the
Yungas of La Paz (Rogg & Camacho 2000). Unfor-
tunately, the failure of well planned follow-up
studies made it impossible to carry out a detailed

analysis of the real effect of parasitoid releases on
fruit fly populations (Rogg & Camacho 2000).
The present study reports the results of a fruit
fly parasitoid survey in which wild guava (Psid-
ium guajava L.) and feral peach fruits were sys-
tematically sampled in an area of the southern-
most extension of the Bolivian Yungas forest. The
objectives were to identify indigenous parasitoid
species, determine relative abundances and vari-
ations in parasitoid and fly numbers over time,
natural parasitization rates and fruit infestation
levels, and parasitoid distribution patterns.
We did not try to assess the impact of parasi-
toids as a mortality factor in the population dy-
namic of A. fraterculusIC. capitata but rather to
determine the natural degree of larval parasitiza-
tion. Comparative studies in neighboring areas of
the endangered Yungas forest in NW Argentina
(Ovruski & Schliserman 2003; Ovruski et al.
2004, 2005, 2006; Schliserman et al. 2004; Schlis-
erman 2005; Orofo & Ovruski 2007) recorded 2
exotic, 1 cosmopolitan, and 10 neotropical parasi-
toid species.


The collecting site was located between 415
and 612 meters above sea level at 22o43' to
2239'S latitude and 6420' to 6419'W longitude,
in the district of Bermejo, province of Tarija,
southern Bolivia. The selected area covered an
area of 25 km2 from the southern border with the
Argentinean province of Salta (district of Aguas
Blancas) to next to the district ofArrosales in the
northern Tarija, Bolivia. The study site is part of
to the environmental unit called Premontane for-
est (lower sector of the Yungas forest in elevation
zone), which ranges from 300 to 600 m in altitude
(Brown et al. 2001). The native vegetation of this
Premontane forest has been almost completely
eliminated and the area transformed into agricul-
tural use and orchards (Brown & Kappelle 2001).
Thus, patches of disturbed wild vegetation with
high abundances of exotic fruit plants, mostly an-
imal-dispersed fruit species, such as Citrus spp.
(Rutaceae), Morus spp., Ficus carica L. (fig) (Mo-
raceae), Prunus persica (peach), Pirus communis
L. (pear) (Rosaceae), and Psidium guajava
(guava) (Myrtaceae), commonly can be found in
proximity to orchards. The climate is temperate-
warm humid with a mean temperature of 22.5C,
and a mean annual rainfall of 1,323 mm
Approximately 92% of the annual rain falls dur-
ing the summer (from Oct to Apr) (Kessler & Beck
Ten guava and 10 feral peach trees were cho-
sen at random on each sample date in the study
area. Ten fruits were harvested from each one of
these guava or peach trees, and 10 fruits were
also collected from the ground below each tree

June 2009

Ovruski et al.: Larval-Pupal Parasitoids of Fruit Flies

canopy Consequently, each sample had 20 guava
or peach fruits originating from any 1 tree. Only
ripe peach or guava fruits were collected, and
fruit trees were only sampled during the peak
fruiting season. Thus, peach trees were sampled
every 2 weeks during Dec (early summer), 1999
and 2000, whereas guava trees were sampled ev-
ery 2 weeks from Feb (mid summer) to Mar (late
summer), 1999 and 2000. All peach and guava
trees sampled were located in patches of second-
ary forests that surrounded the orchards.
Each fruit sample was placed individually into
cloth bags, and fruits collected from the ground
were separated from those picked from the tree
canopy The bags containing fruit samples were
transported to CIRPON laboratory (Centro de In-
vestigaciones para la Regulaci6n de Poblaciones
de Organismos Nocivos) in San Miguel de Tu-
cuman (2650' S, 65"13' W, 426 m), province of Tu-
cuman, Northwestern Argentina. In the labora-
tory, fruit was processed as described by Ovruski
et al. (2005). Fruits of each sample were weighed
and rinsed with a 20% sodium benzoate solution,
and placed in closed styrofoam boxes (20 x 20 x 30
cm) with damp sand in the bottom as a pupation
substrate for fly larvae. Fruits were placed on a
10-mm mesh metal screen fitted about 10 cm from
the bottom. Samples were kept inside a room at 26
+ 2C, 65 + 10% relative humidity and a photope-
riod of 14:10 (L:D) h for 4 weeks, and the sand was
sifted weekly to collect pupae. Afterwards, fruits
were dissected to determine the presence of larvae
or pupae remaining in the pulp. Live larvae were
allowed to pupate and were then added to the
other pupae collected from the same sample. Pu-
pae were removed weekly and the A. fraterculus
and C. capitata pupae were separated based upon
external pupal characters (White & Elson-Harris
1992). After that, pupae were counted and placed
in plastic glasses (300 cm3) filled with sterilized
moist sand in the bottom and covered with or-
gandy cloth over the top. These glasses were in-
spected twice each week and every emerging adult
fly or parasitoid was removed and identified.
Fruit fly species were identified by S. Ovruski
by with Zucchi's (2000) taxonomic key. Parasitoid
specimens were identified to species by S. Ovruski
using Wharton and Marsh's (1978) and Canal and
Zucchi's (2000) keys for Braconidae, Opiinae, and
the taxonomic description by Wharton et al.
(1998) and the key by Guimaraes et al. (2000) for
Figitidae, Eucoilinae. The nomenclature for the
Opiinae follows Wharton (1997) and follows
Wharton et al. (1998) for the Eucoilinae. Voucher
specimens were placed in the entomological col-
lection of Fundaci6n Miguel Lillo (FML) (San
Miguel de Tucuman, Argentina).
The fruit infestation level reported was based
on the number of fruit fly larvae per fruit or on the
number of fruit pupae per kg of fruit. The parasit-
ization rates were calculated on the basis of the to-

tal number of parasitoids emerged from total fruit
fly pupae recovered. Means and Standard Devia-
tion (SD) were calculated as summary statistics
for the parasitism percentage and fruit infestation
level data. Differences in the number of parasi-
toids recovered from fallen fruits versus those col-
lected in the tree canopy were analyzed by the
non-parametric Mann-Whitney U-test (P < 0.05).


Altogether 1,600 guavas and 800 peaches,
weighing 57.713 kg and 24.544 kg respectively,
were processed during this study (Table 1). Out of
these, 800 (27.868 kg) guavas and 400 (11.828 kg)
peaches were fallen fruit collected from the
ground. Mean ( SD) individual weight of guava
and peach fruit per sample was 36.8 5.6 g and
32.2 4.7 g, respectively (n = 160 guava samples
and n = 80 peach samples).
Only 2 fruit fly species, A. fraterculus and C.
capitata, were recovered from the all fruits col-
lected during 1999 and 2000. Eight species of par-
asitoids, all native to the Neotropical region, were
recovered from A. fraterculus pupae obtained
mainly from guavas. These were Doryctobracon
areolatus, D. brasiliensis, D. crawfordi, Utetes
anastrephae (Viereck), Opius bellus (all Bra-
conidae, Opiinae), Aganaspis pelleranoi, Odon-
tosema anastrephae Borgmeier, and Lopheucoila
anastrephae (Rohwer) (all Figitidae, Eucoilinae).
The species D. areolatus, D. brasiliensis, U. anas-
trephae, and A. pelleranoi were also recovered
fromA. fraterculus pupae stemmed from peaches.
Only 1 parasitoid species, A. pelleranoi, was re-
covered from C. capitata pupae obtained from
peaches during both 1999 and 2000.
The abundance of each fruit fly and parasitoid
species recovered per month and year in associa-
tion with the fruit species is summarized in Table
1. Of all parasitoids recovered during the study,
opiine and eucoiline species represented 55.3%
and 44.7%, respectively.
The fruit infestation levels by A. fraterculus
and C. capitata, and parasitization rates on both
tephritid species per collecting month and year
are shown in Table 2. A significantly higher num-
ber of eucoiline parasitoids were obtained from
guava and peach samples collected from the
ground than from the tree canopy (U = 671.0, Z =
8.6, P < 0.0001, n = 80 for guava, and U = 370.0, Z
= 4.1, P < 0.0001, n = 40 for peach) (Fig. 1). On the
contrary, no significant differences were found in
opiine parasitoids (U = 2674.0, Z = 1.8, P = 0.07, n
= 80 for guava, and U = 683.0, Z = 1.1, P = 0.26, n
= 40 for peach).


The high levels of infestation in wild guava
and in feral peach recorded in this study reveal


Total No.

Parasitoid species

Sample Date Fruit Fruit Cc Cc Af All
(mo and yr) species (kg) pupae' adults pupae Af Adults2 Da Db Dc Ua Ob Ap Oa La parasitoids

Feb/1999 Guava 400 1 1 1,706 712 37 27 1 3 0 53 1 0 122
Mar/1999 Guava 400 2 2 2,462 955 62 33 0 19 0 72 0 0 186
Dec/1999 Peach 400 1,394 623 206 82 6 5 0 1 0 22 0 0 34
Total/1999 1,200 1,397 626 4,374 1,749 105 65 1 23 0 147 1 0 342

Feb/2000 Guava 400 10 2 2,397 902 35 21 0 10 1 63 0 0 130
Mar/2000 Guava 400 2 2 3,669 1,209 83 32 0 18 0 91 0 1 225
Dec/2000 Peach 400 1,067 451 164 65 0 0 0 0 0 15 0 0 15
Total/2000 1,200 1,079 455 6,230 2,176 118 53 0 28 1 169 0 1 370

Total (82,461) 2,476 1,081 10,604 3,925 223 118 1 51 1 316 1 1 712

Cc, Ceratitis capitata.
'Af, Anastrepha fraterculus.
'Da, Doryctobracon areaolatus; Db, D. brasiliensis; Dc, D. crawfordi; Ua, Utetes anastrephae; Ob, Opius bellus; Ap, Aganaspis pelleranoi; Oa, Odontosema anastrephae; La, Lopheucoila anas-

Ovruski et al.: Larval-Pupal Parasitoids of Fruit Flies

+1 +1 +1
00 00

+1 +1 +1
(0 C1 C1


C~l cO '-l
00 'l
+1 +1 +1
00 1
00 CI

0 C<1 't
in 10 Cl
+1 +1 +1

It .- C11
11 CO i'
+1 +1 +1
LO ,C C11

CO t- ,q
dl 0

+1 +1 +1
[- (0 Cl


CQ~ 000
~O~ 0
2 2'Cl

O Ground
* Tree canopy


Guava Peach



Fig. 1. Results of the Mann-Whitney U-test compar-
ing eucoiline parasitoid number obtained from guava
and peach fruits collected from the ground and from the
tree canopy. The small black bar represents Median
value, white and gray boxes represent Quartiles values
(25-75 percentiles), and whisker represents Maximum
value. Bars with different letters differ significantly (U-
test,P <0.05).

that these fruits are important reservoirs of A.
fraterculus and C. capitata, respectively, in per-
turbed Yungas forests growing along the moun-
tains of Tarija, southern Bolivia, and probably
contribute to their high populations in agricul-
tural environments, in accordance to that re-
ported by Ovruski et al. (2003, 2004, 2005) in the
Argentinean Yungas forests.
About 93% of all parasitoids were recovered
from P. guajava. Seemingly, wild guava fruits not
only would be a reservoir from whichA. fratercu-
lus would spreads to orchards, but also as a valu-
able native parasitoid source in the southern Yun-
gas forest of Tarija, Bolivia. Similar observations
were made by L6pez et al. (1999) and Aluja et al.
(1998) in a tropical rainforest in Veracruz, Mex-
ico, and by Ovruski et al. (2004, 2005) in the Yun-
gas rainforests of NW Argentina.
The parasitoid survey revealed 8 parasitoid
species (D. areolatus, D. brasiliensis, D. craw-
fordi, U. anastrephae, 0. bellus, A. pelleranoi, 0.
anastrephae, and L. anastrephae) attacking A.
fraterculus, and 1 parasitoid species (A. pellera-
noi) attacking C. capitata for Tarija, Bolivia. This
survey is also provides the first records of U.
anastrephae, 0. anastrephae, and L. anastrephae
for Bolivia, and the first evidence of 0. anas-
trephae in the South American Yungas forest. All
of these opiine and eucoiline species belong to the
fruit fly parasitoid guild number 2 described by
Ovruski et al. (2000), which is characterized by
koinobiont, solitary, larval-pupal endoparasitoids
ofAnastrepha spp.
Before this study, D. areolatus, D. brasiliensis,
D. crawfordi, 0. bellus, andA. pelleranoi were the









Florida Entomologist 92(2)

only native parasitoid species recorded from A.
fraterculus in the Yungas forest of Santa Cruz, a
province in southern Bolivia (Escalante 1995).
The exotic parasitoid D. longicaudata, which was
recovered 6 years after its last release in the
northern Yungas forest of La Paz, Bolivia, (Rogg
& Camacho 2000) was not found in this southern
All fruit fly parasitoid species recovered in this
survey, with the exception of 0. anastrephae, had
been previously recorded from A. fraterculus in
the Argentinean Yungas forests (Wharton et al.
1998; Ovruski et al. 2004, 2005). However, D.
crawfordi, which is a widespread neotropical opi-
ine species (Ovruski et al. 2000), is yet to be re-
corded in the southernmost portion of the Yungas
forest of Argentina. Probably, the natural distri-
bution in Yungas forest of this species includes
only the northernmost portion of the Argentinean
Yungas (Ovruski et al. 2005), which spreads as far
as southern Bolivia embracing Tarija and Santa
The temporal variations in parasitoid abun-
dance and parasitization rates recorded in this
study throughout the 3 summer months (Dec,
Feb, and Mar) in 1999 and 2000 were closely re-
lated to the changes that occurred over time in
the fruit infestation levels caused by the most im-
portant fly host, A. fraterculus, in wild guava
fruits. Thus, higher parasitism rates on A. frater-
culus and numbers of the 4 most commonly col-
lected parasitoid in this study occurred in late
summer (Mar). For example, A. pelleranoi, D. are-
olatus, D. brasiliensis, and U. anastrephae, in-
creased between 1.4- and 6.3-times in abundance
from Dec to Mar in both 1999 and 2000. This is
consistent with data reported by Ovruski et al.
(2005) for the northernmost sector of the Argen-
tinean Yungas forest. Probably the increasing
abundance of parasitoids from Dec to Mar in asso-
ciation with A. fraterculus population, has been
influenced by environmental factors, such as
heavier rainfall toward the end of the summer, a
climatic characteristic of the Yungas forest phyto-
geographical region (Brown et al. 2001; Kessler &
Beck 2001).
The eucoilidA. pelleranoi and the opiine D. ar-
eolatus were the most abundant parasitoid spe-
cies attackingA. fraterculus larvae in wild guavas
and feral peaches in the southern section of Yun-
gas forest in SW Bolivia. Apparently, these 2 na-
tive parasitoid species exhibit a less competitive
interaction in a host plant (Sivinski et al. 1997).
These authors found differences in the foraging
patterns between A. pelleranoi and D. areolatus.
Thus, D. areolatus, like others fruit fly opiine par-
asitoids, stays on the fruit surface searching for
host larvae, while A. pelleranoi mainly attacks
larvae in fallen fruit by entering through holes in
the fruit. The comparative analysis between eu-
coiline and opiine parasitoid numbers recovered

from fruit samples collected from the ground and
those from the tree canopy for this study, corrobo-
rated previous reports by Sivinski et al. (1997) for
Mexico, Ovruski et al. (2004) for Argentina, and
Guimaraes & Zucchi (2004) for Brazil, indicating
that eucoiline parasitoids predominantly parasit-
ize fruit fly larvae in fallen fruit.
The results of this study showed for first time
several parasitoid species associated with A.
fraterculus and only 1 with C. capitata in the
southernmost portion of the Bolivian Yungas for-
est. However, these considerations are prelimi-
nary because only 2 fruit fly host plants were
sampled in this extensive phytogeographical re-
gion of Bolivia. More exhaustive sampling of
other C. capitata and Anastrepha host plants,
mostly native fruit species, are therefore needed
in this distinctive type of Bolivian subtropical for-
est before more definitive conclusions on parasi-
toid abundance patterns and diversity of fruit fly
parasitoids species can be reached.


We express our gratitude to Eduardo Frias
(PROIMI-CONICET) for valuable technical support.
Special thanks to Patricio Fidalgo (CRILAR-CONICET)
for useful assistance during field work. We thank Mar-
tin Aluja (INECOL, A.C., Xalapa, Veracruz, M6xico) for
sharing with us his vast experience on ecology, biology,
and ethology of fruit fly parasitoids. We thank to 2 anon-
ymous reviewers for helping us produce a better article.
Financial support was provided by the Consejo Nacio-
nal de Investigaciones Cientificas y T6cnicas de la
Republica Argentina (CONICET) (Grant PIP Nos. 0702/
98, 02567/01, and 5129/05).

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Florida Entomologist 92(2)

June 2009


Department of Entomology and Nematology, University of Florida, Bldg. 970, Natural Area Drive,
Gainesville, FL, 32611-0629, USA

1Presently at Dipartimento di Protezione delle Piante, Sezione Entomologia Agraria, Universita di Sassari,
Via De Nicola 1, 07100 Sassari, Italy
E-mail: acocco@uniss.it


The red palm mite, Raoiella indica Hirst, an important pest of coconut, banana, and date
palms is a new invasive pest in the Western Hemisphere. The red palm mite (RPM) has been
observed attacking bananas and plantains in Dominica and in Florida (M. A. Hoy, A. Cocco,
personal observation). In order to develop an efficient method to rear the RPM in quarantine
for a classical biological control project, several banana and plantain varieties were tested
as hosts for the RPM. Bananas are more desirable than coconut (a favored host plant) be-
cause bananas are easier to rear in small cages and will produce new shoots quickly after
pruning. Red palm mite females did not establish on the banana and plantain varieties
(Dwarf Cavendish, Dwarf Nino, Gran Nain, Dwarf Zan Moreno, Dwarf Green, Truly Tiny,
Musa sumatrana x Gran Nain, Dwarf Puerto Rican, Rose, Nang Phaya, Misi Luki, Manzano,
Lady Finger, Glui Kai, and Ebun Musak) of leaf discs tested, but they established on coconut
leaf discs. The mites could not be reared on potted banana trees (Glui Kai, Dwarf Green, and
Nang Phaya varieties), but a multigenerational colony has been maintained on coconut trees
and leaf discs. No RPM females survived on native palms tested (saw palmetto, cabbage
palm, and dwarf palmetto), but RPM completed a generation on needle palm, with longer de-
velopment time, higher mortality, and lower fecundity than when reared on coconut discs.
Our results indicate that coconut leaf discs and trees are better hosts for rearing RPM in
quarantine than banana, plantain varieties, or native palms tested. Quarantine tests and
field observations suggest that the host range of RPM may not be as broad as some reports
indicate because plants from which RPM adults and/or eggs have been collected might not
be suitable for establishment of a multigenerational colony. More studies under natural con-
ditions need to be conducted to evaluate the ability of R. indica to establish and spread on
native and ornamental palms in natural landscapes in Florida.

Key Words: Raoiella indica, laboratory rearing, host plants, saw palmetto, dwarf palmetto,
cabbage palm, needle palm, coconut


El acaro rojo de la palmer, Raoiella indica Hirst, una plaga important del coco, banano, y
de la palmer datilera es una plaga nueva invasora del Hemisferio Occidental. El acaro rojo
de la palmer (ARP) ha sido observado atacando bananos y platanos en Dominica y en la Flo-
rida (M. A. Hoy, A. Cocco, observaci6n personal). Para desarrollar un m6todo eficiente para
criar el ARP en cuarentena para un proyecto de control biol6gico clasico, varias variedades
de banano y platano fueron probadas como hospederos para el ARP. Los bananos son mas de-
seables que los cocos (una plant hospedera preferida) por que los banano son mas faciles
para criar en jaulas pequenas y produce nuevos brotes rdpidamente despu6s de la poda.
Las hembras del acaro rojo de las palmeras no se establecieron sobre las variedades de ba-
nano y platano (Dwarf Cavendish, Dwarf Nino, Gran Nain, Dwarf Zan Moreno, Dwarf
Green, Truly Tiny, Musa sumatrana x Gran Nain, Dwarf Puerto Rican, Rose, Nang Phaya,
Misi Luki, Manzano, Lady Finger, Glui Kai y Ebun Musak) en los discos de hojas probados,
pero se establecieron en los discos de hojas del coco. Estos acaros no pudieron ser criados en
arboles de bananos en macetas (variedades Glui Kai, Dwarf Green y Nang Phaya), pero ha
mantenido una colonia multigeneracional sobre arboles de coco y discos de hojas. Ningun
hembra de ARP sobrevivi6 las palmas nativas probadas (saw palmetto, cabbage palm, and
dwarf palmetto), pero ARP complete una generaci6n sobre la "needle palm" palmera de
aguja), con un period de desarrollo mas largo, mortalidad mas alta y fecundidad mas baja
que cuando fue criado sobre discos de coco. Nuestros resultados indican que los discos de la
hoja de coco y los arboles de coco son mejores hospederos para criar ARP en cuarentena que
las variedades de platano, banano o las palmeras nativas probadas. Las pruebas en cuaren-

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite

tena y las observaciones indican que el rango de los hospederos de ARP posiblemente no sea
tan amplio como indican algunos informes porque las plants sobre que han recolectado
adults y/o huevos de ARP pueden ser no apropiadas para el establecimiento de una colonia
multigeneracional. Se necesitan realizar mas studios bajo condiciones naturales para eva-
luar la habilidad de R. indica para establecerse y esparcirse sobre palmeras nativas y orna-
mentales en areas naturales de la Florida.

The red palm mite, Raoiella indica Hirst (Ac-
ari: Tenuipalpidae) (RPM), is a serious pest of eco-
nomically important fruit-producing trees such as
coconut Cocos nucifera L. and banana Musa spp.
(Nagesha-Chandra & Channabasavanna 1984;
Welbourn 2006). In addition, significant infesta-
tions have been reported on the date palm Phoe-
nix dactylifera L., plantains Musa spp., and orna-
mental palms, including the Christmas palm
Adonidia (= Veitchia) merrillii (Becc.) H. E.
Moore, and the Mexican fan palm Washingtonia
robusta H. Wendl (Zaher et al. 1969; Etienne &
Fletchmann 2006). The RPM was found in the
Caribbean for the first time in Martinique and
Saint Lucia in 2004 (Fletchmann & Etienne
2004), and has now spread throughout the Carib-
bean islands and invaded Florida and Venezuela
(Kane et al. 2005; Etienne & Fletchmann 2006;
Gutierrez et al. 2007; Rodrigues et al. 2007). Rao-
iella indica was detected in southeastern Florida
in 2007 and is now established in 3 counties (Flor-
ida Department of Agriculture and Consumer
Services 2008). Plants reported by USDA-APHIS
to be hosts of R. indica in Florida include orna-
mental palms such as the Fiji fan palm Pritchar-
dia pacifica Seem. & H. Wendl., the Miraguama
palm Coccothrinax miraguama (Kunth) Leon,
and the endangered native Florida thatch palm
Thrinax radiata Lodd. ex J. A. & J. H. (K. M. Grif-
fiths, personal communication; Coile & Garland
The establishment ofR. indica in North Amer-
ica has caused concerns about the economic im-
pact of this new pest in palm nurseries, subtropi-
cal agriculture, and natural and urban land-
scapes. The biology ofR. indica was studied on co-
conut in India by Nagesha-Chandra &
Channabasavanna (1984) and on date palm in
Egypt by Zaher et al. (1969), while studies on or-
namental and landscape palms, bananas, or plan-
tains have not been reported. The potential sus-
ceptibility of native Florida palms to the RPM is
of interest in the spread of the pest, as well as for
its effects in parks and natural areas.
A classical biological control program for
Florida was initiated by identifying potentially
effective predatory mites from areas where the
RPM is endemic (Hoy et al. 2006). Predatory
mites (Acari: Phytoseiidae) were imported from
Mauritius and colonies were established in the
quarantine laboratory at the Department of En-
tomology and Nematology, University of Flor-
ida, in Oct 2007. Colonies of the RPM and pred-

atory mites are being reared in quarantine on
coconut leaf discs (H. Bowman & M.A. Hoy, un-
published). Bananas and plantains were ob-
served to be a suitable host for the RPM in Do-
minica and in Florida (M.A. Hoy, N. Commo-
dore, and A. Cocco, personal observation) and
were considered more appropriate for rearing in
small spaces than the coconut palm because
they can be grown in small pots, fit into small
cages, and produce new shoots quickly after
pruning so they can be reused several times in
Thus, we conducted rearing tests to determine
whether R. indica could be reared on banana and
plantain leaf discs or trees under quarantine con-
ditions outside the infestation zone. In addition,
the suitability of selected Florida native palms as
hosts for the RPM was investigated. Native palms
commonly occur in natural landscapes and the
fan-shaped palm leaves could provide wider leaf
discs than the split-leafed coconut palms for bio-
assays with the phytoseiids being evaluated in
quarantine. In addition, these trials could provide
new biological information about the host range
and the behavior of the RPM.


Source of Raoiella indica, Coconut, and Native Palms

Foliage containing the RPM was collected from
heavily infested banana and coconut trees of un-
known cultivars in Lake Worth, FL during 2008,
placed into ice chests with ice packs, and brought
into the quarantine facility at the Department of
Entomology and Nematology, University of Flor-
ida, Gainesville. The foliage samples were kept at
21.5-23.8C inside ice chests until RPM were used
in tests. Potted coconut trees were purchased
from nurseries in southern Florida, tested for pes-
ticide residues, and reared at the Department of
Entomology and Nematology. Leaf discs used in
the experiments were obtained from mature
leaves. The native palms tested as suitable hosts
were dwarf palmetto Sabal minor (Jacq.) Pers.,
saw palmetto Serenoa repens (Bartr.) Small, cab-
bage palm Sabal palmetto (Walter) Lodd. ex
Schult., and needle palm Rhapidophyllum hystrix
(Pursh). These native palm samples were col-
lected in Gainesville, FL on the campus of the
University of Florida and at Kanahapa Botanical
Gardens from mature leaves that had not been
treated with pesticides.

Florida Entomologist 92(2)

Raoiella indica Rearing on Banana Discs and Trees

To verify that the RPM can be reared on ba-
nana leaf discs (leaves cut into pieces of specific
size), 4 banana varieties (Dwarf Puerto Rican
[plantain], Dwarf Cavendish, Dwarf Nino, and
Gran Nain) obtained from Agri-Starts, Inc. (Apo-
pka, FL) were potted into 3.8-L pots and allowed
to produce 6-10 leaves before being evaluated,
with coconut leaf discs used as a positive control.
Discs 18 x 45 mm wide cut with a single-edge ra-
zor blade from mature leaves were cleaned with a
brush and inspected for undesirable predatory
mites or insects under a dissecting microscope.
Banana and coconut discs were placed on water-
soaked cotton in a plastic tray (13 x 13 x 2.5 cm),
with the abaxial surface of the leaves facing up.
The cotton was kept wet for the duration of the
bioassay by adding water periodically, and nar-
row paper strips (Kimwipe, Kimberly-Clark Cor-
poration, Roswell, GA) were placed along the edge
of each leaf arena to reduce the likelihood of mites
running off or under the discs. Five young RPM
females, field collected from coconut trees (un-
known cultivar), were placed on each disc; the
survivorship and the number of eggs laid were re-
corded under a dissecting microscope every 24 h
for 7 d. Body length and dimensions of dark
patches on the dorsum of the body were consid-
ered to estimate the age of the females (Hoy et al.
2006). Twelve replicates were conducted by plac-
ing 6 leaf discs for each treatment in 2 trays at
26.9-31.4C, 56-100% RH, under a 16L:8D photo-
Because the establishment of R. indica on ba-
nana leaf discs might be affected by the original
host of the mite, additional banana and plantain
varieties were screened with young females col-
lected from infested banana leaves (unknown va-
riety) in Lake Worth, FL. Additional small ba-
nana and plantain trees (ca. 20 cm tall) were ob-
tained from Agri-Starts, Inc. (Apopka, FL) and
Dwarf Zan Moreno, Dwarf Green, Truly Tiny,
Musa sumatrana x Gran Nain (hybrid), Dwarf
Puerto Rican (plantain), Rose, Nang Phaya, Misi
Luki, Manzano, Lady Finger, Glui Kai, and Ebun
Musak varieties were tested, with coconut as a
positive control. Discs from leaves about 2-3
weeks old were set up as described above, except
that each disc (ca. 70 x 70 mm wide) was placed in
a tray containing cotton saturated with water.
Ten young RPM females were placed on each disc
and left undisturbed for 11 d. Each female was
considered a replicate. The survivorship, the be-
havior (feeding, not feeding, drowned, or dead),
and the number of eggs laid in each disc were re-
corded under a dissecting microscope every 24 h,
at 27.8-33.6C, 44-100% RH, under a 16L:8D pho-
To investigate whether the female's behavior
was determined by some chemical or physical

modification of the newly prepared banana discs,
another bioassay was conducted with banana
discs from leaves about 2-3 weeks old and held on
the bioassay trays for 3 d before RPM were placed
on them. The experimental design was the same
as the above experiment, except that only the cul-
tivars Nang Phaya, Dwarf Green, and Glui Kai
were tested, with coconut used as a control and 25
females were added. Development of the progeny
was monitored until adulthood was reached. Leaf
discs were replaced after 3 weeks, when they be-
came yellow. Mites were moved from degraded to
new leaf discs with a sable-hair brush (size 0000).
These bioassays were carried out at 27.8-33.1C,
43-100% RH during the oviposition period and at
25.6-31.6C, 51-100% RH during the developmen-
tal period of progeny, with both under a 16L:8D
To determine whether live banana trees could
be used to rear the RPM under quarantine condi-
tions, potted banana trees (varieties Glui Kai,
Nang Phaya, and Dwarf Green) ca. 20-30 cm tall
were tested. Because of space limitations inside
the quarantine room, a single potted coconut tree
was used as a control. Banana trees were pruned
so that only a single leaf about 2-3 weeks old was
used as the test arena. The leaves were cleaned
with a brush for undesirable insects and preda-
tory mites, and 20 young RPM females from field-
collected coconut leaves were placed on the abax-
ial surface of the leaves and left undisturbed for 7
d. Trees were placed into PVC-frame cages cov-
ered with organdy cloth at 22.6-31.9C, 42-73%
RH, under a 16L:8D photoperiod. Live and dead
adults and the number of eggs and larvae on each
leaf were recorded under a dissecting microscope.
Each treatment was replicated 5 times on 2 dates.
If leaves were found that contained eggs or lar-
vae, they were cut and placed on water-soaked
cotton for further observations of developmental
Survival analysis was estimated with the
PROC LIFEREG procedure (SAS Institute 2002).
Pairwise comparisons were performed to evaluate
significant differences between survivorship pat-
terns. The proportion of mites feeding, not feed-
ing, or drowned/dead on each disc was compared
by logistic regression (PROC LOGISTIC, SAS In-
stitute 2002).
To evaluate the RPM oviposition rate, the
mean number of eggs laid on each of the banana
or plantain discs was compared to the oviposition
rate on coconut discs with the Mann-Whitney U
test (Proc NPAR1WAY, SAS Institute 2002). The
treatment means were not compared with each
other because we were interested in comparing
each to coconut only. Mortality rates of RPM eggs
and immatures were compared with the Fisher's
exact test (PROC FREQ, SAS Institute 2002). In
the first bioassay, replicate discs of each treat-
ment were combined and analyzed as 1 data set.

June 2009

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite

Two-Choice Test for Host Preference on Selected Palms

A two-choice test was conducted to determine
the host preference of RPM with coconut vs. coco-
nut, coconut vs. needle palm, coconut vs. saw pal-
metto, coconut vs. cabbage palm, and coconut vs.
dwarf palmetto. Discs 18 x 45 mm wide were cut
with a single-edge razor blade from mature leaves
and hand washed with tap water and allowed to
dry, which usually took 10-15 min. The 2 different
disc halves were sealed together by painting a 4-
5 mm wide stripe of melted paraffin wax with a
camel-hair brush on the abaxial (lower) surface
(Hoy & Smilanick 1981). The joined leaf discs
were placed on water-soaked cotton in plastic
trays as for the previous tests. A single young
RPM female was placed on the midline paraffin
stripe of each arena with a sable-hair brush (size
0000) and allowed to move freely. The location
(disc half type or paraffin wax stripe), the behav-
ior (feeding, not feeding, drowned, or dead), and
the number of eggs laid were recorded under a
dissecting microscope after 48 h. Mites were
checked under a dissecting microscope after 24 h
to confirm the survival of each female and to de-
termine whether females had moved from the
paraffin midline. RPM females were touched gen-
tly with a sable-hair brush and considered alive if
they moved or walked away, while dead females
were removed and replaced with live adults. The
experiment was replicated 35 times by placing 5
arenas for each type of two-choice leaf disc in 7
trays. Two bioassays were conducted with RPM
females collected either from coconut or banana
trees in the field in order to investigate whether
the original host plant affected host choice. The
experiments were conducted at 28.1-34.6C, 52-
100% RH when mites from coconut were used,
and at 28.5-33.4C, 42-100% RH when mites from
bananas were tested, both under a 16L:8D photo-
period. Temperature and RH were recorded with
a Traceable Digital Thermometer (Fisher Elec-
tronics, Pittsburgh, PA).
Exact logistic regression (PROC LOGISTIC,
SAS Institute 2002) was used to analyze R. indica
behavioral response (host choice) to host plants
because some observations had zero values or
separation of the data set occurred (Heinze &
Ploner 2003). The number of eggs laid by RPM fe-
males on discs of the same arena was compared
with the Mann-Whitney U test (Proc NPAR1WAY,
SAS Institute 2002).

Survival and Reproduction of Females in No-choice
Tests on Selected Ornamental and Native Palms

To determine whether RPM females were able
to establish and oviposit on selected native palms
of Florida, the behavior of single young females in
single leaf discs was observed on coconut, needle
palm, saw palmetto, cabbage palm, and dwarf

palmetto. The leaf discs were set up as for the
two-choice test, except that the discs were of a sin-
gle leaf type. The survivorship, behavior (feeding,
not feeding, drowned, or dead), and the number of
eggs laid by single young R. indica females on
each disc were recorded under a dissecting micro-
scope every 24 h for 8 d.
To determine whether immature stages were
able to establish on these native palm species
from the eggs deposited by the females above, egg
eclosion and survival of immatures were re-
corded. The entire experiment was conducted
with young RPM females collected from both in-
fested coconut and banana trees and was repli-
cated 50 times on 2 dates. The bioassays with
RPM from coconut were carried out at 28.1-
32.6C, 50-100% RH; those with RPM from ba-
nana at 28.8-33.4C, 52-100% RH, both under a
16L:8D photoperiod.
Survivorship patterns were compared by the
PROC LIFEREG procedure (SAS Institute 2002).
Pairwise comparisons were performed to evaluate
significant differences between treatments. The
proportion of mites feeding, not feeding, or
drowned/dead on each disc was compared by lo-
gistic regression (PROC LOGISTIC, SAS Insti-
tute 2002). The mean number of eggs laid during
8 d on each palm species was compared to that on
coconut discs Mann-Whitney U test (Proc
NPAR1WAY, SAS Institute 2002). The percent-
ages of egg, larval, and protonymphal mortality
were analyzed with exact logistic regression
(PROC LOGISTIC, SAS Institute 2002).

Survival of R. indica on Needle Palm Leaf Discs

To determine whether R. indica could estab-
lish on needle palm discs, the survival of RPM im-
matures on that host plant was investigated and
compared to survival on coconut discs. The leaf
discs were set up as described above, except that
they were 25 x 90 mm wide and were not washed
but cleaned with a brush to preserve the cuticle
characteristics. Fifteen young RPM females were
sampled from field-collected coconut leaves and
placed on the leaf discs for 6 d and then removed.
Dead or drowned females were replaced every 24
h to maximize the number of eggs laid on coconut
and needle palm leaf discs. The number of eggs
laid, the percentage of egg eclosion, and the sur-
vival rate of RPM immatures were recorded every
24 h until adulthood was reached. The mean egg
incubation time was estimated per each disc as
MIT = y egg eclosion date ; egg oviposition
date (1)
where MIT is the Mean Incubation Time, the
mean egg eclosion date is the mean between the
first and last day of egg eclosion, and the mean
egg oviposition date is the mean between the first
and the last day of egg oviposition. The mean de-

Florida Entomologist 92(2)

velopment time from larva to adult was estimated
per each disc as
MDT = j adult emergence date j egg eclo-
sion date (2)
where MDT is the Mean Development Time, the
mean adult emergence date is the mean between
the first and last adult emergence, and the mean
egg eclosion date is the mean between the first
and last day of egg eclosion. The emerging F, fe-
males were examined every 24 h to verify
whether they laid eggs or not. Leaf discs were re-
placed every 3-4 weeks, when discs appeared de-
graded. Mites were transferred from aged to new
leaf discs with a sable-hair brush. The bioassay
was replicated 8 times, at 27.8-32.9C, 48-72% RH
during the oviposition period and at 25.6-29.9C,
56-100% RH during the developmental period,
both under a 16L:8D photoperiod.
The mean egg incubation time, and the mean
development time from larva to adult were ana-
lyzed with the Mann-Whitney U test (Proc
NPAR1WAY, SAS Institute 2002) (Lee 1992).
Mortalities of eggs and immatures were evalu-
ated with one-way ANOVA in Proc GLIMMIX
(SAS Institute 2002).


Raoiella indica Rearing on Banana Discs and Trees

During the first experiment, RPM females did
not establish on the 4 varieties of banana or plan-
tain leaf discs, but they settled down on coconut
discs (Fig. 1). There were significant differences in
survival of RPM females on different leaf discs

(X2 = 166.28; df = 4; P < 0.0001). Survivorship of
RPM females on coconut leaf discs after 7 d was
52%, significantly different than the survival on
Dwarf Puerto Rican, Dwarf Cavendish, Dwarf
Nino, or Gran Nain discs (all pairwise compari-
sons: P < 0.0001). The survivorship pattern of
RPM females on Dwarf Nino and Gran Nain leaf
discs was not statistically different (X2 = 0.30; df=
1; P = 0.5831). Dwarf Cavendish (banana variety)
and Puerto Rican (plantain variety) leaf discs
were significantly less suitable than the other
hosts tested.
During the 7-day bioassay, 60 RPM females
laid a total of 261 eggs on coconut leaf discs, while
surviving females laid a total of 6, 3, 32, and 42
eggs on Dwarf Puerto Rican, Dwarf Cavendish,
Gran Nain, and Dwarf Nino discs, respectively
(data not shown). The pairwise comparisons be-
tween the mean oviposition rate of the RPM on
coconut discs (0.76 eggs/female/d) showed signif-
icant differences from these on Dwarf Puerto
Rican (0.17 eggs/female/d) (P = 0.0054), Dwarf
Cavendish (0.05 eggs/female/d) (P = 0.0033),
Gran Nain (0.25 eggs/female/d) (P = 0.0037), and
Dwarf Nino (0.27 eggs/female/d) (P = 0.0068) leaf
When RPM females that were field collected
from banana trees were tested on 12 different ba-
nana and plantain leaf discs, significant differ-
ences in survivorship were observed among the
treatments (X2 = 108.85; df = 4; P < 0.0001)
(Table 1). The banana variety Misi Luki appeared
to be the least suitable host, with all RPM females
dying or running off the discs by the second day of
the bioassay. After 4 d, 100% mortality was ob-
served on Truly Tiny, Dwarf Puerto Rican, Rose,

0 1 2 3 Days 4 5 6 7

Fig. 1. Survivorship of R. indica females collected from coconut on different banana and plantain leaf discs un-
der quarantine conditions (n = 60 for each plant type). Treatments followed by the same letter are not significantly
different (P < 0.05).

June 2009

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite


Female behavior (%)d
Comparisons of Mean no. of
survivorship over No. of Not Drowned eggs/9/d
Host plant 11 db observations" Feeding feeding or dead (SE)

Coconut a 95 94 a 2b 4c 0.97+ 0.18
Glui Kai b 61 61 b 23 a 16 b 0.15 + 0.05 *
Dwarf Green c 34 38 cd 32 a 30 ab 0.23 + 0.10 *
Nang Phaya c 31 49 bc 19 a 32 ab 0.26 0.22 *
M. sumatrana x Gran Nain cd 25 32 cd 28 a 40 ab 0.07 0.05 *
Manzano cd 25 20 d 40 a 40 ab 0.03 0.03 *
Rose cde 21 38 cd 14 a 48 a 0.08 0.08 *
ZanMoreno cde 17 29 cd 12 ab 59 a 0.22+ 0.16 *
Ebun Musak cde 21 28 cd 24 a 48 a 0.23 + 0.19 *
Truly Tiny de 17 23 cd 18 a 59 a 0.13 + 0.13 *
Puerto Rican e 13 15 d 8 ab 77 a 0 *
Lady Finger e 15 26 cd 7 ab 67 a 0.75 0.25 ns
Misi Luki e 15 26 cd 7 ab 67 a 0.30 0.30 ns

Laboratory conditions: 27.8-33.6C, 44-100% RH, under a 16L:8D photoperiod. Number of females tested for each plant type =
bSignificant differences among survivorship patterns compared with PROC LIFEREG, treatments followed by the same letter
within a column are not significantly different (P < 0.05).
'Discrepancies in number of observations (potentially 109 x 11 d = 110) are due to the different female survival rates.
Significant differences compared with PROC LOGISTIC, treatments followed by the same letter within a column are not sig-
nificantly different (P < 0.05).
'Significant differences between coconut and each host plant compared with Mann-Whitney U test (PROC NPAR1WAY), treat-
ment means with are significantly different compared to coconut (P < 0.05).

and Lady Finger leaf discs, while on Dwarf Green
and Ebun Musak the same mortality rate was ob-
served after 5 d. On Dwarf Zan Moreno, Nang
Phaya, and Manzano leaf discs, no live female
was observed after 6 d, while the longest survival
among banana varieties was observed on Glui Kai
leaf discs (11 d). By contrast, RPM females on co-
conut discs exhibited 40% mortality after 11 d.
During the leaf disc bioassay, the Glui Kai variety
appeared to be the most suitable banana host for
the RPM (Table 1). Survivorship patterns of RPM
females on Dwarf Green and Nang Phaya discs
were not different than on Musa sumatrana x
Gran Nain, Manzano, Rose, Dwarf Zan Moreno,
and Ebun Musak, while they were statistically
different from the survival rate on Truly Tiny,
Dwarf Puerto Rican, Misi Luki, and Lady Finger
leaf discs (Table 1).
Observations on feeding behavior showed that
RPM females fed significantly more frequently on
coconut than on banana or plantain leaf discs (P <
0.0001) (Table 1). Females of R. indica were ob-
served feeding on Glui Kai discs in 61% of the ob-
servations, significantly more than on Dwarf
Green, Musa sumatrana x Gran Nain, Manzano,
Dwarf Zan Moreno, Ebun Musak, Truly Tiny,
Dwarf Puerto Rican, Misi Luki, and Lady Finger
leaf discs (P < 0.05). The proportion of RPM fe-
males feeding on the host ranged from 15 to 38%
on Dwarf Green, Rose, Musa sumatrana x Gran

Nain, Manzano, Rose, Dwarf Zan Moreno, Ebun
Musak, Truly Tiny, Dwarf Puerto Rican, Misi
Luki, and Lady Finger, but the differences were
not significant (Table 1).
The oviposition rate of RPM females on coco-
nut was 0.97 eggs/female/d, which was signifi-
cantly higher than on Glui Kai, Dwarf Green,
Nang Phaya, M. sumatrana x Gran Nain, Puerto
Rican, Manzano, Rose, Zan Moreno, Ebun Musak,
Truly Tiny, and Puerto Rican leaf discs (all pair-
wise comparisons were between the oviposition
rate on each banana or plantain leaf disc and that
on coconut discs: P < 0.05) (Table 1). Raoiella in-
dica females laid 0.75 and 0.30 eggs/female/d on
Lady Finger and Misi Luki leaf discs, respectively,
which were not significantly different from that
on coconut discs. Only 15 behavior observations
were made on those discs because RPM females
survived only 4 and 2 d, respectively, suggesting
that females laid eggs immediately after being
placed on the discs and then ran off the discs.
When RPM females were tested on 3-d-old
Dwarf Green, Glui Kai, Nang Phaya, and coconut
leaf discs, the 4 treatments exhibited significantly
different survivorship patterns from one another (F
= 70.79;df= 3;P < 0.0001) (Table 2). No live RPM fe-
males were observed on Nang Phaya and Dwarf
Green after 5 and 7 d, respectively, while 100% mor-
tality was observed on Glui Kai discs after 11 d.
Consistent with the previous bioassay, female sur-

Florida Entomologist 92(2)


Mortality of R. indica (%)d Successful
Comparisons of Mean no. of development to
Host plant survivorship over 11db eggs/2/d (SE) Eggs- Immatures* adult (%)

Coconut a 0.93 0.12 9 57 40
Glui Kai b 0.26 0.06 5 69 30
Dwarf Green c 0.29 + 0.11 4 77 22
Nang Phaya d 0.13 + 0.06 0 100 0

"Laboratory conditions: 27.8-33.1C, 43-100% RH (oviposition period); 25.6-31.6C, 51-100% RH (developmental period), both un-
der a 16L:8D photoperiod. Number of females tested for each plant type = 25.
bSignificant differences among survivorship patterns compared with PROC LIFEREG, treatments followed by the same letter
within a column are not significantly different (P < 0.05).
'Significant differences between coconut and each host plant compared with Mann-Whitney U test (PROC NPAR1WAY), treat-
ment means with are significantly different compared to coconut (P < 0.005).
Treatments were compared with the Fisher's exact test, ns = no differences among treatments; *= significant differences among
treatments (P < 0.05).

vival on Glui Kai discs was significantly higher than
these on Dwarf Green or Nang Phaya. At the end of
the experiment, RPM females on coconut discs ex-
perienced 36% mortality. The mean fecundity of
RPM females on coconut leaf discs (0.93 eggs/fe-
male/d) differed significantly from these of females
on Glui Kai, Dwarf Green, or Nang Phaya discs
(0.26, 0.29, and 0.13 eggs/female/d, respectively)
(Table 2). All the pairwise comparisons between the
oviposition rate on coconut discs and on banana
discs indicated a significant difference with P <
0.05. The R. indica eggs experienced 0-9% mortality
during the bioassay, but differences were not signif-
icant (P = 0.8576) (Table 2). Mortality of RPM im-
matures was significantly different among treat-
ments (Fisher's exact test: P = 0.0381), ranging from
57% on coconut to 100% on Nang Phaya leaf discs.
The rate of successful development from egg to
adult on coconut, Glui Kai, Dwarf Green, and Nang
Phaya discs was 40, 30, 22, and 0%, respectively
(Table 2). The emerged RPM females did not deposit
eggs on Glui Kai, Dwarf Green, or Nang Phaya discs
and died within 7 d, while on coconut discs 88 RPM
females established and laid 97 eggs over 7 d (data
not shown).
RPM females also failed to establish on potted
Glui Kai, Nang Phaya, and Dwarf Green banana

trees. After 7 d, no females survived and a total of
only 17, 10, and 1 eggs were observed on Glui Kai,
Dwarf Green, and Nang Phaya leaves, respec-
tively, corresponding to mean oviposition rates of
0.17, 0.10, and 0.01 eggs/female/7 d, respectively
(Table 3). In the same climatic conditions, RPM
females established and laid eggs on coconut
leaves. A coconut leaf disc of ca. 25 cm2 sampled
randomly revealed 36 R. indica females alive and
154 eggs, with a mean fecundity of 4.3 eggs/fe-
male/7 d, and only 1 dead (3% mortality) (data not
shown). Mortality rates of RPM eggs on banana
leaves ranged from 0 (Nang Phaya) to 50% (Glui
Kai), while larvae experienced 75, 0, and 100%
mortality after 7 d on Glui Kai, Dwarf Green, and
Nang Phaya leaves, respectively (Table 3). Leaves
with eggs or larvae were cut and placed on water-
soaked cotton, and subsequent observations indi-
cated that all larvae died within 4 d and failed to
molt to the protonymphal stage (Table 3).

Two-Choice Test for Host Preference on Selected Palms

Raoiella indica females did not exhibit a pref-
erence between needle palm or coconut leaf discs
(Table 4, test Al: exact P = 1.0000; test Bl: exact
p = 0.8506). The proportion of females feeding on


Mortality of R. indica (%)
Total no. Mean no. % molting from
Host plant eggs/20 9 / 7 d of eggs/ /7 d (SE) Adults Eggs Larvae larvae to protonymphs

Glui Kai 17 0.17 + 0.09 100 6 75 0
Dwarf Green 10 0.10 + 0.05 100 50 0 0
Nang Phaya 1 0.01 + 0.01 100 0 100 0

"Laboratory conditions: 22.6-31.9C, 42-73% RH, under a 16L:8D photoperiod. Number of females in each treatment = 20. In the
same climatic conditions, RPM females on coconut leaves exhibited 3% mortality and a mean fecundity of 4.3 eggs/female/7 d.

June 2009

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite


Female behavior (%)
mean no.
Females observed Drowned of egg/ /48 h
Treatment' on each half (%)bc Feeding Not feeding or dead (SE)d

A Mites collected from coconut
1) Coconut vs 49 a 88 a 6 a 6 a 1.4 0.3 a
Needle palm 46 a 100 a 0 a 0 a 0.9 0.3 a
2) Coconut vs 66 a 91 a 0 b 9 a 1.8 0.3 a
Saw palmetto 9b 0 b 67 a 33 a 0 b
3) Coconut vs 40 a 93 a Ob 7 b 1.6 0.3 a
Cabbage palm 34 a Ob 42 a 58 a 0,
4) Coconut vs 60 a 95 a Ob 5 b 0.9 0.2 a
Dwarf palmetto 26 b 0 b 33 a 67 a 0 b
5) Coconut vs 46 a 94 a 0 a 6 a 1.5 0.3 a
Coconut 49 a 94 a Oa 6 a 2.1 0.3 a
B Mites collected from banana
1) Coconut vs 37 a 100 a 0 a 0 a 1.0 + 0.2 a
Needle palm 43 a 87 a 0 a 13 a 0.9 0.3 a
2) Coconut vs 66 a 91 a 4 a 4b 1.2 0.3 a
Saw palmetto 29 b Ob 10 a 90 a 0
3) Coconut vs 66 a 100 a 0 a Ob 1.0 + 0.2 a
Cabbage palm 14 b Ob 0 a 100 a 0
4) Coconut vs 60 a 100 a 0 a Ob 1.2 0.2 a
Dwarf palmetto 14 b 0b 0 a 100 a 0,
5) Coconut vs 46 a 88 a 0 a 12 a 1.4 0.3 a
Coconut 40 a 93 a 0 a 7 a 1.2 0.3 a

"Laboratory conditions: (A) 28.1-34.6C, RH 52-100%; (B) 28.5-33.4C, RH 42-100%, both under 16L:8D photoperiod. Number of
females observed after 48 h for each bioassay = 35.
bDiscrepancies in the percentage of females on tested halves are because some females were found on the midline.
'Significant differences compared with PROC LOGISTIC, treatments followed by the same letter within a column for each bio-
assay are not significantly different (exact P < 0.05).
Significant differences compared by the Mann-Whitney U test (PROC NPAR1WAY), means followed by the same letter within
a column for each bioassay are not significantly different (P < 0.05).

coconut or needle palm discs after 48 h during tests
Al and B1 ranged from 87 to 100%, but the differ-
ences were not significant. RPM females appeared
to prefer coconut over saw palmetto (test A2: exact P
< 0.0001; test B2: exact p = 0.0351) or dwarf pal-
metto (test A4: exact P = 0.0428; test B4: exact p =
0.0025). No mites were observed feeding on saw pal-
metto or dwarf palmetto leaf discs during the 48-h
experiments. Unexpectedly, RPM females did not
show a preference between coconut and cabbage
palm leaf discs in test A3 (exact p = 0.8450), while
there was a significant difference between these
host plants in test B3 (exactp = 0.009). However, no
mites were detected feeding on cabbage palm discs
during experiments A3 and B3, while all live fe-
males on the coconut halves of the test arenas ap-
peared established. During the two-choice tests, no
RPM female was observed feeding on saw palmetto,
cabbage palm, or dwarf palmetto leaf discs. Obser-

vations after 24 and 48 h revealed that mites did not
change their position after their original choice, per-
haps due to the width of the paraffin seal (4-5 mm).
RPM oviposition rates on coconut and needle
palm during the 48-h test were not significantly dif-
ferent, whether females were collected from coconut
or banana (Table 4, tests Al: x2 = 1.6770; df = 1;p =
0.1953; Bl: x2 = 0.4678;df = 1;p = 0.4940). However,
no eggs were laid by females on saw palmetto, cab-
bage palm, or dwarf palmetto disc halves (Tests A2,
A3, A4, B2, B3, B4), while RPM females on coconut
halves of the same test arenas produced 0.9 to 1.8
eggs/female /48 h.

Survival and Reproduction of Females in No-choice
Tests on Selected Ornamental and Native Palms

The native saw palmetto, cabbage palm, and
dwarf palmetto do not seem to be palatable hosts

Florida Entomologist 92(2)

for the RPM under quarantine conditions
(Table 5). Despite the fact that adult females from
the field-collected banana or coconut foliage were
assigned randomly to the test leaf discs, there
were significant differences in survivorship over 8
d (Table 5A: X2 = 218.2219; df = 4; P < 0.0001; Ta-
ble 5B: X2 = 241.0365; df = 4; P < 0.0001). Coconut
and needle palm discs appeared to be the most
suitable hosts for RPM females, while they did
not establish on saw palmetto, cabbage palm, or
dwarf palmetto discs. Survivorship ofR. indica fe-
males collected from coconut trees on needle palm
and coconut leaf discs after 8 d were 76 and 90%,
respectively, but the pairwise comparison was not
significantly different (X2 = 3.68; df = 1; P =
0.0550) (Table 5A). On saw palmetto discs, RPM
females survived significantly longer than on cab-
bage palm (x2 = 17.83; df= 1; P < 0.0001) or dwarf
palmetto discs (X2 = 15.19; df= 1; P < 0.0001) (Ta-
ble 5A). When RPM females collected from ba-
nana were used, the survivorship on coconut and
needle palm discs were 52 and 34%, respectively,
showing a marginally significant difference (X2 =
3.96; df = 1; P = 0.0467) (Table 5B). The survivor-
ship pattern of R. indica females collected from
infested banana trees on saw palmetto was signif-
icantly different than that on cabbage palm (X2 =
6.93; df= 1;P = 0.0085), while there was no differ-
ence in female survival on saw palmetto and
dwarf palmetto discs (X2 = 2.08; df= 1;P = 0.1491).
Survivorship patterns of R. indica on cabbage
palm and dwarf palmetto discs were not signifi-

cantly different in both experiments. Experi-
ments A and B (Table 5) were performed using
mites collected from infested coconut and banana
trees on two dates, so no statistical analysis were
performed to compare the two survivorship
curves. However, RPM females collected from co-
conut appeared to survive longer than females
collected from banana trees on coconut (90 and
55%, respectively) and needle palm (76 and 34%,
respectively) leaf discs (data not shown).
Behavior observations during the no-choice
test indicated that coconut was the better host for
the RPM females (Table 5). Significantly more R.
indica females were observed feeding on coconut
discs than on needle palm, saw palmetto, cabbage
palm, or dwarf palmetto discs in both experi-
ments (P < 0.0001 for both) (Table 5A, B). How-
ever, needle palm discs appeared to be palatable
to RPM females, which were observed feeding on
that host 72 and 59% of the observations (P <
0.0001, Table 5A, B). Red palm mite females were
observed feeding significantly less frequently on
needle palm than on coconut leaf discs after 8 d
(Table 5), while those differences in feeding be-
havior were not significant after 2 d (Table 4),
suggesting that longer observations are more re-
liable to determine the palatability of the host
Only 1 to 2% of the observations revealedR. in-
dica feeding on saw palmetto, cabbage palm, or
dwarf palmetto discs, whether using RPM fe-
males collected from coconut or banana trees, and


Female behavior (%)d

Comparisons of No. Drowned or
Host plant survivorship over 8 db observationsc Feeding Not feeding dead

A Mites collected from coconut
Coconut a 387 95 a 4c 1 b
Needle palm a 343 72 b 25 b 3b
Saw palmetto b 138 2 c 62 a 36 a
Cabbage palm c 90 1 c 43 a 56 a
Dwarf palmetto c 92 1 c 45 a 54 a
B Mites collected from banana
Coconut a 301 85 a 7c 8b
Needle palm b 259 59 b 28 b 13 b
Saw palmetto c 100 2 c 48 a 50 a
Cabbage palm d 77 1 c 34 ab 65 a
Dwarf palmetto cd 86 1 c 41 ab 58 a

aLaboratory conditions: (A) 28.1-32.6C, 50-100% RH; (B) 28.8-33.4C, 52-100% RH, both under a 16L:8D photoperiod. Number
of females observed on each host plant = 50 for 8 d.
bSignificant differences among survivorship patterns compared with PROC LIFEREG, treatments followed by the same letter
within a column in each experiment are not significantly different (P < 0.05).
'Discrepancies in number of observations (potentially 509 x 8 d = 400) are due to the different female survival rates.
Significant differences compared with PROC LOGISTIC, treatments followed by the same letter within a column are not sig-
nificantly different (P < 0.05).

June 2009

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite

there were no significant differences in RPM feed-
ing behavior on these discs (P > 0.05).
During the 8 d of the experiments, 50 R. indica
females collected from coconut trees laid a total of
360 eggs on coconut leaf discs, with an oviposition
rate of 0.92 eggs/female/d (Table 6A). RPM fe-
males from the same source laid a total 104 eggs
on needle palm leaf discs, corresponding to an ovi-
position rate of 0.28 eggs/female/d, while females
laid a total of only 1, 2, and 2 eggs (0.01 eggs/fe-
male/d) on dwarf palmetto, saw palmetto, and
cabbage palm, respectively, over 8 d. The fecun-
dity on coconut discs was significantly higher
than on needle palm (Mann-Whitney U test, P <
0.0001), saw palmetto (Mann-Whitney U test, P <
0.0001), cabbage palm (Mann-Whitney U test, P <
0.0001), or dwarf palmetto discs (all pairwise
comparisons between coconut vs. each native
palm with Mann-Whitney U test: P < 0.0001).
Likewise, when RPM females collected from ba-
nana trees were used, their oviposition rate on co-
conut discs (0.49 eggs/female/d) was significantly
higher than on other hosts (all pairwise compari-
sons: P < 0.0001). On needle palm discs females
laid a total of 36 eggs, at a rate of 0.11 egg/female/
d, while on saw palmetto discs only 2 eggs were
observed (0.01 egg/female/d) (Table 6B). No eggs
were deposited on cabbage palm and dwarf pal-
metto discs. RPM females field collected from co-
conut trees exhibited a higher fecundity than fe-
males sampled from banana trees (0.92 and 0.49
eggs/female/d, respectively) (Table 6A, B), sug-

gesting that coconut could be a more favorable
host than banana in the field. The fecundity of
RPM females on needle palm discs was signifi-
cantly lower than coconut discs after 8 d (Tables
6A, B) while the oviposition rate was not different
on coconut and needle palm discs after 2 d
(Table 4), suggesting that RPM females laid most
eggs on needle palm discs immediately after
transfer, while the oviposition rate on coconut
discs was more constant.
Mortality of eggs on coconut discs (11%) dif-
fered significantly from the egg mortality ob-
served on needle palm discs (24%) (Table 6A,
exact P = 0.0014). Mortality data of eggs from
saw palmetto, cabbage palm, and dwarf pal-
metto was excluded from the statistical analy-
sis because of the low number of eggs laid. Lar-
vae that hatched on coconut discs experienced
11% mortality, which was significantly lower
than the larval mortality on needle palm discs
(81%) (exact P < 0.0001). Mortality of proto-
nymphs on coconut discs was 21%, while all
protonymphs on needle palm discs died before
molting to the deutonymphal stage (exact P <
0.0001). Mortality rates of eggs laid by RPM fe-
males from banana trees on coconut and needle
palm discs were significantly different (8 and
36%, respectively) (Table 6B, exact P < 0.0001).
RPM larvae developed on coconut discs experi-
enced 63% mortality, while no larvae molted
successfully to the protonymphal stage on nee-
dle palm discs (exact P = 0.0007). RPM females


Mortality of R. indica (%)c
Mean no.
Host plant of eggs/$/d (SE)b Eggs Larvae Protonymphs

A Mites collected from coconut
Coconut 0.92 0.04 11b 33 b 21 b
Needle palm 0.28 0.03 24 a 81 a 100 a
Saw palmetto 0.01 0.01* -
Cabbage palm 0.01 0.01* -
Dwarf palmetto 0.01 0.01* -
B Mites collected from banana
Coconut 0.49 0.06 8 b 37 b 29
Needle palm 0.11 0.02 36 a 100 a -
Saw palmetto 0.01 0.01* -
Cabbage palm 0 -
Dwarf palmetto 0 -

aLaboratory conditions: (A) 28.1-32.6C, 50-100% RH; (B) 28.8-33.4C, 52-100% RH, both under a 16L:8D photoperiod. Number
of females observed on each host plant = 50 for 8 d.
bSignificant differences between coconut and each host plant compared with Mann-Whitney U test (PROC NPAR1WAY), treat-
ment means with are significantly different compared to coconut (P < 0.0001).
'Significant differences compared with PROC LOGISTIC, treatments followed by the same letter within a column are not signif-
icantly different (exact P < 0.05). Mortality of eggs and larvae from saw palmetto, cabbage palm, and dwarf palmetto were excluded
from the analysis because of the low number of eggs laid.

Florida Entomologist 92(2)

collected from coconut trees survived longer, ex-
hibited an higher oviposition rate, and were ob-
served feeding more often on coconut and nee-
dle palm discs than females collected from ba-
nana trees (Tables 5 and 6), perhaps due to the
different age of leaves tested or to the lower
suitability of banana for the RPM.

Survival of R. indica on Needle Palm Leaf Discs

During the 6-d assay, a total of 135 and 155 fe-
males were tested on coconut and needle palm
discs, respectively. Thirty five females were re-
placed on needle palm because they died or ran off
the discs, while 15 females were replaced on coco-
nut discs for the same reason. Despite the lower
number of females tested on coconut, a total of
648 eggs were laid on coconut discs and 365 on
needle palm. Mortality of eggs laid on coconut
discs was 3%, which was significantly lower than
egg mortality (7%) on needle palm discs (F =
30.67; df= 1, 14;P < 0.0001). During development
to adulthood, RPM immatures feeding on coconut
discs exhibited significantly lower mortality
(67%) than immatures growing on needle palm
(84%) (F = 33.87; df = 1, 14; P < 0.0001) (Table 7).
Although the determination of the exact develop-
ment time of R. indica was beyond goal of the ex-
periment, the Mean Incubation Time and the
Mean Development Time were assessed. The
mean Incubation Time ranged from 5.9 to 6.2 d on
needle palm and coconut, respectively, but the dif-
ference was not significant (X2 = 1.7340; df = 1; P
= 0.1879) (Table 7). The development time from
larva to adult on coconut discs averaged 12.1 d,
which was significantly lower than the Mean De-
velopment Time (25.9 d) on needle palm discs (X2
= 10.7299; df = 1; P = 0.0011). A total of 153 and
35 female progeny developed successfully on coco-
nut and needle palm discs, respectively. Discs
were examined every 24 h to verify the presence
of F2 eggs. On coconut discs, 21 d after the begin-
ning of the experiment, a total 961 eggs were
scored, with a 30% daily rate of increase which

made mite counts difficult and observations on co-
conut discs were stopped. However, on needle
palm discs, observations were stopped 64 d after
the beginning of the bioassay, and a total of only
49 eggs were scored.


The RPM established on coconut leaf discs and
potted trees, and small colonies have been main-
tained for many generations, while no stable col-
ony has been obtained on banana or plantain
discs or potted banana trees. Likewise, no RPM
females survived on the ornamental and native
palm discs tested, except on needle palm discs,
where the RPM completed a generation but expe-
rienced high mortality and a long development
It is unclear whether R. indica can actually
feed, reproduce, and develop within a normal
time period on all plants listed in Table 8 because
information about which RPM life stage was ob-
served on these plants was not always provided
(Fletchmann & Etienne 2004; Kane & Ochoa
2006; Mendonca et al. 2006; Welbourn 2006).
Pedigo (1996, p. 425) defines a host plant as "Suf-
ficiency of the plant as a host is finally determined
during feeding. If nutrients are adequate and no
toxicity occurs, the insect completes development
within a normal time period and becomes an
adult. Also sufficiency is indicated in normal
adult longevity and fecundity". It is possible that
the RPM was dispersed by wind currents to some
of these plants located beneath palm canopies,
and it is possible that gravid females could de-
posit a few eggs on these temporary hosts, but the
establishment of a multigenerational colony has
not always been documented.
Unknown varieties of bananas have been re-
ported to be suitable hosts for the RPM in Florida
(A. Cocco, personal observation), while in the
Eastern Caribbean significant multigenerational
infestations have been observed on the most
widely grown banana (Dwarf Cavendish, Giant


Mortality of R. indica (%)b Mean
Mean Development
Total no. of Y Total no. of Incubation Time larva to
Treatment" tested on 6 d eggs/6 d Eggs Immatures Time (d SE) adult (d SE)

Coconut 135 648 3 lb 67 6 b 6.2 0.1 a 12.1 0.2b
Needle palm 155 365 7 3 a 84 4 a 5.9 0.1 a 25.9 0.8 a

"Laboratory conditions: 27.8-32.9C, 48-72% RH (oviposition period); 25.6-29.9C, 56-100% RH (developmental period), both un-
der a 16L:8D photoperiod. Initial number of females for each of 8 replications = 15.
bTreatment means were compared with PROC GLIMMIX, means with the same letter within a column are not significantly dif-
'Significant differences compared by the Mann-Whitney U test (PROC NPAR1WAY); means followed by the same letter within
a column for each bioassay are not significantly different (P < 0.05).

June 2009


Family Plant species Referenceb


Acer sp.
Acoelorraphe wrightii (Grises. & H. Wendl.)
Adonidia merrilli (Becc.) Becc. (= Veitchia)
Aiphanes caryotifolia (Kunth) H. A. Wendl.
Aiphanes sp.
Archontophoenix alexandrae (F. Muell.) H. Wendl. & Drude
Areca sp.
Areca catechu L.
Bactris plumeriana Mart.
Beccariophoenix madagascariensis Jum. & H. Perrier
Bismarckia nobilis Hildebr. & Wendl.
Butia capitata (Mart.) Becc.
Caryota mitis Lour.
Chamaedorea spp.
Coccothrinax argentata (Jacq.) L. H. Bailey
Coccothrinax miraguama (Kunth) Becc.
Cocos nucifera L.
Corypha umbraculifera L.
Dictyosperma album (Bory) H. Wendl. & Drude ex Scheff.
Dypsis decaryi (Jum.) Beentje & J. Dransf.
Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. (= Chrysalidocarpus)
Elaeis guineensis Jacq.
Licuala grandis H. Wendl.
Livistona chinensis (Jacq.) R. Br.
Phoenix canariensis hort. ex Chabaud
Phoenix dactylifera L.
Phoenix reclinata Jacq.
Phoenix roebelenii O'Brien
Pritchardia pacifica B.C. Seem. & H. Wendl.
Pritchardia vuylstekeana H. Wendl. ?
Pseudophoenix sargentii H. Wendl. ex Sarg.
Pseudophoenix vinifera (Mart.) Becc.
Ptychosperma elegans (R.Br.) Blume
Ptychosperma macarthurii (H. Wendl. ex H .J.Veitch) H. Wendl. ex Hook. F.
Ptychosperma sp.

"Most host plants are reported with no information about which RPM stage was found. The establishment of a multigenerational colony has not always been documented.
bHost plants with are cited by Welbourn (2006) as by Pellegrano in press.

Mitrofanov & Strunkova (1979)
Welbourn (2006)*
Fletchmann & Etienne (2004)
K. Griffiths, personal communication
Kane et al. (2005)
K. Griffiths, personal communication
Pritchard & Baker (1958)
Nagesha-Chandra & Channabasavanna (1984)
Welbourn (2006)*
K. Griffiths, personal communication
Welbourn (2006)
K. Griffiths, personal communication
Etienne & Fletchmann (2006)
Welbourn (2006)
A. Cocco, personal observation
K. Griffiths, personal communication
Hirst (1924)
K. Griffiths, personal communication
Moutia (1958)
Welbourn (2006)
Kane et al. (2005)
Welbourn (2006)
Etienne & Fletchmann (2006)
K. Griffiths, personal communication
Etienne & Fletchmann (2006)
Sayed (1942)
Welbourn (2006)*
Welbourn (2006)
Etienne & Fletchmann (2006)
A. Cocco, personal observation
Welbourn (2006)
Welbourn (2006)*
K. Griffiths, personal communication
Etienne & Fletchmann (2006)
A. Cocco, personal observation


Family Plant species Referenceb


Rhapis excelsa (Thunb.) A. Henry ex Rehder
Roystonea borinquena O.F. Cook.
Schippia concolor Burret
Syagrus romanzoffianum (Cham.) Glassman
Syagrus schizophylla (Mart) Glassman
Thrinax radiata Lodd. ex J. A. & J. H. Schult.
Veitchia arecina Becc.
Veitchia sp.
Washingtonia filifera (L. Lind.) H. Wendl.
Washingtonia robusta H. Wendl.
Washingtonia sp.
Wodyetia bifurcata Irvine
Cassine transvaalensis Burtt-Davy
Phaseolus sp.
Heliconia sp.
Heliconia bihai (L.) L.
Heliconia caribaea Lam.
Heliconia psittacorum L.F.
Heliconia rostrata Ruiz & Pavon
Ocimum basilicum L.
Musa acuminata Colla
Musa balbisiana Colla
Musa uranoscopus Lour.
Musa x paradisiaca L. (= Musa sapientum L.)
Musa corniculata Rumph.
Musa spp.
Eucalyptus spp.
Eugenia sp.
Olea sp.
Pandanus utilis Bory
Pandanus sp.
Strelitzia reginae Banks ex Dryard
Ravenala madagascariensis Sonn.
Alpinia purpurata (Vieill.) K. Schum

aMost host plants are reported with no information about which RPM stage was found. The establishment of a multigenerational colony has not always been documented.
bHost plants with are cited by Welbourn (2006) as by Pellegrano in press.

Welbourn (2006)*
Welbourn (2006)*
K. Griffiths, personal communication
Kane et al. (2005)
Welbourn (2006)*
K. Griffiths, personal communication
A. Cocco, personal observation
K. Griffiths, personal communication
Welbourn (2006)
Etienne & Fletchmann (2006)
K. Griffiths, personal communication
Welbourn (2006)
Kane & Ochoa (2006)
Gupta (1984)
Pefia et al. (2006)
Welbourn (2006)*
Welbourn (2006)*
Welbourn (2006)
Etienne & Fletchmann (2006)
Chaudri et al. (1974)
Kane et al. (2005)
Kane et al. (2005)
Kane et al. (2005)
Kane et al. (2005)
Welbourn, (2006)
Etienne & Fletchmann (2006)
Kane & Ochoa (2006)
Kane & Ochoa (2006)
Kane & Ochoa (2006)
Welbourn (2006)
Kane & Ochoa (2006)
Etienne & Fletchmann (2006)
Welbourn (2006)
Etienne & Fletchmann (2006)

Cocco & Hoy: Laboratory Rearing of the Red Palm Mite

Cavendish, Robusta, and Williams) and plantain
(Apem, Cents Livre, Ordinary, Dwarf French, and
Horn) varieties (N. Commodore, personal commu-
nication). The banana and plantain varieties we
tested in our leaf disc and potted tree bioassays
Were different from those reported from the East-
ern Caribbean, except for the Dwarf Cavendish
banana variety. Despite these reports, RPM fe-
males did not establish and were often observed
not feeding on the banana and plantain varieties
Tested in our leaf disc and whole potted tree quar-
S8 antine bioassays. However, in the same experi-
= ments, RPM females established and fed continu-
5 -g 13 ously on coconut leaf discs. Among the banana
and plantain varieties tested, RPM females sur-
vived longer and were observed feeding more of-
ten on Glui Kai discs than on other varieties, sug-
Sgesting that Glui Kai is the most palatable ba-
o nana variety tested.
C The behavior of RPM females may be a better
Index of host suitability than the oviposition rate
because frequent observations of females not
) feeding, drowned or dead suggest that females
Share searching for a suitable host on which to es-
tablish and feed. The RPM progeny (F, females)
C reared on banana discs that reached adulthood
did not deposit eggs and no established colony of
the RPM was obtained on banana leaf discs or
potted banana trees.
The reasons) for the failure to establish RPM
females and immatures on banana trees and leaf
Sdiscs in quarantine are unclear. In our quaran-
S tine bioassays, the establishment of RPM fe-
males collected from coconut or banana trees
Swas evaluated on newly prepared and 3-d-old ba-
nana leaf discs, but neither the original host of
the RPM (coconut or banana) nor the age of the
0g leaf discs appear to promote the establishment of
54 g q .RPM colonies on banana leaf discs. Physical and/
S* or chemical modifications of banana and plan-
S t. tain leaf discs could have repelled RPM females,
4 B but the experiment with potted banana trees did
Snot result in the establishment of a stable colony
'S of the RPM. Characteristics of the cuticle or
H quantity of wax on the abaxial surface might
SS g make some banana or plantain varieties more
N suitable for the RPM than others. Consistent
With this hypothesis, while sampling the RPM
. ;c Jfrom 2 heavily infested banana trees of unknown
H ^ N K 1 variety(ies) in Lake Worth (Jun 2008), an unin-
t3 fested banana tree of unknown variety was ob-
Sserved less than 1 m away. A sprout from the
Base of the infested tree was collected, potted,
q and a new banana/plantain tree was grown in
t the quarantine laboratory in Gainesville. RPM
females were released on leaf discs and on young
Ss leaves of the growing shoot under quarantine
S8 g conditions, but RPM did not establish, perhaps
od because the shoot contained only young leaves
While the "mother" tree had RPM on mature
< .a leaves.

Florida Entomologist 92(2)

Field observations on both coconut and banana
trees revealed that mature leaves were more of-
ten infested by the RPM than young leaves (A.
Cocco and M. A. Hoy, personal observations).
Young leaves might be unsuitable for the RPM be-
cause of higher concentration of secondary plant
compounds than old leaves. For some trees, young
leaves are reported to have higher levels of sec-
ondary metabolites such as alkaloids, phenols,
flavonoids, and terpenoids than older leaves (Ber-
nays & Chapman 1994). The desert clicker Lig-
urotettix coquilletti McNeill (Orthoptera: Acrid-
idae) habitually feeds on older leaves of Larrea
tridentata because they contain a lower concen-
tration of the deterrent nordihydroguaiaretic acid
than young leaves (Chapman et al. 1988). Wood-
head (1983) observed that young leaves of some
varieties of Sorghum sp. are unsuitable for the
migratory locust Locusta migratoria L. (Ortho-
ptera: Acrididae) because they contain a specific
wax compound, while older leaves are accepted.
To clarify whether the leaf age affects the estab-
lishment of the RPM on banana or plantain trees,
young and old leaves of the same tree could be in-
fested under field conditions with known num-
bers of RPM females. In our experiments, because
RPM females established on coconut leaf discs
and trees under the same climatic conditions as
these of the banana discs and potted trees, we be-
lieve that abiotic factors such as temperature,
RH, and photoperiod did not affect the establish-
ment of the RPM.
Native and ornamental palms such as saw pal-
metto, cabbage palm, needle palm, dwarf pal-
metto, European and Chinese fan palms are com-
mon woody plants on the natural landscape of
Florida and are used for landscaping homes,
parks, and streets (Black 2003a, 2003b). In addi-
tion, saw palmetto and needle palm are economi-
cally important palms (Tanner et al. 2002; Coile &
Garland 2003). Extracts of saw palmetto fruits
are used to treat symptoms of benign prostatic hy-
perplasia (Gordon & Shaughnessy 2002). Our re-
sults indicate that saw palmetto, dwarf palmetto,
and cabbage palm leaf discs are not suitable hosts
for the RPM in quarantine. Preliminary labora-
tory tests indicated that the Chinese fan palm
Livistona chinensis (Jacq.) R. Br. and the Euro-
pean fan palm Chamaerops humilis L. also failed
to support establishment of RPM colonies (H.
Bowman & M. A. Hoy, unpublished).
Although the RPM completed a generation on
needle palm discs, it exhibited a doubled develop-
ment time and higher mortality of eggs and imma-
tures, and it is unclear if a multigenerational col-
ony on needle palm leaf discs can be established.
Anecdotal observations suggest that the RPM host
range needs additional studies. For example, ob-
servations in Broward County during Oct 2008 re-
vealed an uninfested needle palm in a botanical
garden ca. 50 m away from other infested palm

species (A. Cocco, personal observation), yet needle
palm might be a host based on our laboratory ob-
servations. At the same site, the cabbage palm and
the scrub palmetto Sabal etonia Swingle ex Nash
(closely related to the dwarf palmetto) were in-
spected, but no RPM was found, possibly confirm-
ing our finding that they are not hosts. Observa-
tions conducted in 13 counties in Florida until Aug
2008 report only the Florida thatch palm and the
Florida silver palm Coccothrinax argentata (Jacq.)
L. H. Bailey among native palms are a host of the
RPM; both palms are included on the Florida en-
dangered and threatened plant list (Coile & Gar-
land 2003) (A. Cocco, personal observation; K. M.
Griffiths, personal communication). The ability of
R. indica to establish and spread on native and or-
namental palms raises important questions about
the potential impact of the RPM on natural land-
scapes. Our quarantine experiments and field ob-
servations suggest that RPM adults can deposit
eggs and survive some days on unsuitable hosts, so
host range studies should report plants as suitable
hosts only when all stages of RPM are observed, in-
dicating that multigenerational colonies were es-
A multigenerational RPM colony has estab-
lished on coconut potted trees and stable colonies
can be maintained on coconut leaf discs by cutting
the discs into pieces every 3 weeks and placing
small portions of the infested old disc on new
discs, allowing the RPM to move from the old to
the new discs. Our results suggest that coconut
leaf discs and trees are the most suitable hosts for
RPM females and a better host on which to rear
the RPM in quarantine than the other hosts


The authors thank Hector Perez of the University of
Florida, Department of Environmental Horticulture,
for palm identifications and James Colee of the Univer-
sity of Florida, Department of Statistics, for statistical
consultation. Don Goodman of the Kanapaha Botanical
Gardens, Gainesville, FL, and Jerry Behan of the Deer-
field Beach Arboretum, Deerfield Beach, FL, kindly al-
lowed sampling of palms, Karolynne Griffiths of the
United States Department of Agriculture-APHIS-
PPQ suggested sites for sampling the RPM from coco-
nuts and bananas and provided information of hosts
from which RPM were collected. This research was sup-
ported in part by the Davies, Fischer and Eckes Endow-
ment in Biological Control to M.A. Hoy and a contract
(07-8312-0541-CA) from USDA-APHIS-PPQ.


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Florida Entomologist 92(2)

June 2009


'South American Biological Control Laboratory, USDA-ARS, Hurlingham, Argentina
E-mail: fmckay@speedy.com.ar


2Entomology & Nematology Department, Institute of Food & Agricultural Sciences, University of Florida,
P.O. Box 110620, Gainesville, Florida 32611-0620

'Invasive Plant Research Laboratory, USDA/ARS, University of Florida, 3225 College Avenue,
Ft. Lauderdale, Florida 33314


Brazilian peppertree (Schinus terebinthifolius Raddi, Anacardiaceae) is a perennial tree na-
tive to Argentina, Brazil, and Paraguay. The plant was introduced into the USA before 1900.
Originally grown as an ornamental, Brazilian peppertree is now considered an noxious
plant in Hawaii and Florida, where it is ranked among the most important threats to biodi-
versity in natural areas. Recent surveys conducted in northeastern Argentina recovered one
fungus associated with distorted leaves and 36 phytophagous insects collected on Brazilian
peppertree. A leaf-feeding notodontid moth, a new species of gracillariid leaf blotch miner,
and a stem-boring weevil have been selected for further studies to determine their potential
as biological control agents of Brazilian peppertree in the USA. The results of these surveys
are summarized herein and descriptions are included of the insects that are considered most
promising for biological control of this weed.

Key Words: foreign exploration; classical biological control; Schinus terebinthifolius; host
range; invasive weed; Florida Everglades


Schinus terebinthifolius Raddi, (Anacardiaceae) es un arbol perenne native de Argentina,
Brasil, y Paraguay. Introducido en los Estados Unidos antes de 1900, como especie ornamen-
tal, S. terebinthifolius es considerada una plant nociva en los estados de Florida y Hawaii,
donde amenaza la biodiversidad de areas naturales. Inspecciones de campo recientes en el
noreste de Argentina revelaron la presencia de un hongo asociado a hojas deformadas y 36
artr6podos fit6fagos recolectados de S. terebinthifolius. Un Notodontidae desfoliador, un
Gracillariidae minador de las hojas, y un Curculionidae minador del tallo han sido seleccio-
nados para realizar studios que determine su potential como agents de control biol6gico
de S. terebinthifolius en los Estados Unidos. El present trabajo resume el resultado de las
inspecciones de campo y las descripciones de los artr6podos considerados mas promisorios
para el control biol6gico de S. terebinthifolius en los Estados Unidos.

Translation by the authors.

Brazilian peppertree (Schinus terebinthifolius
Raddi, Anacardiaceae) is a Neotropical species
whose native range extends along the Atlantic
coast of Brazil, from Recife south to Rio Grande do
Sul, and west to north-eastern Argentina and ad-
jacent Paraguay (Barkley 1944, 1957, unpub-
lished data). This species has been introduced to
many countries around the world as an ornamen-
tal (Morton 1978; Panetta & McKee 1997). Cur-
rently, Brazilian peppertree is listed as a prohib-
ited plant and a noxious weed in Florida, and is

considered an invasive species in Florida, Califor-
nia, Texas, and Hawaii (Randall 2000; HSASC
2001; FLEPPC 2005; USDA, NRCS 2009). In its
exotic range, the tree decreases the biodiversity of
infested natural areas by aggressively invading a
variety of coastal and upland habitats (Mytinger
& Williamson 1987; Gann et al. 2001). In Florida,
infestations of Brazilian peppertree are estimated
to occupy over 283,400 ha in central and south
Florida (Wunderlin & Hansen 2003; Cuda et al.
2006). Brazilian peppertree constitutes not only a

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