Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00014
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 2006
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
 Record Information
Bibliographic ID: UF00098813
Volume ID: VID00014
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text

Deyrup: New Florida Dacetine Ant


Archbold Biological Station, P.O. Box 2057, Lake Placid, FL 33862 USA

The dacetine ant Pyramica boltoni is described from specimens collected in leaf litter in dry
and mesic forest in central and northern Florida. It appears to be closely related to P. dietri-
chi (M. R. Smith), with which it shares peculiar modifications of the clypeus and the clypeal
hairs. In total, 40 dacetine species (31 native and 9 exotic) are now known from southeastern
North America.

Key Words: dacetine ants, Hymenoptera, Formicidae


Se describe la hormiga Dacetini, Pyramica boltoni, de especimenes recolectados en la hoja-
rasca de un bosque m6sico seco en el area central y del norte de la Florida. Esta especie esta
aparentemente relacionada con P. dietrichi (M. R. Smith), con la cual compare unas modi-
ficaciones peculiares del clipeo y las cerdas del clipeo. En total, hay 40 species de hormigas
Dacetini (31 nativas y 9 exoticas) conocidas en el sureste de America del Norte.

The tribe Dacetini is composed of small ants
(usually under 3 mm long) that generally live in
leaf litter where they prey on small arthropods,
especially springtails (Collembola). The tribe has
been formally defined by Bolton (1999, 2000). Ne-
arctic dacetines may be recognized by a combina-
tion of features exemplified in Fig. 1: expanded,
lobed occipital area of the head, elongate, nar-
rowed projection of the head beyond the eyes, and
the elongate, narrow mandibles. Most species
have enlarged, spoon-shaped or otherwise modi-
fied hairs on the head, especially on the clypeus,
and whitish, spongy processes on the petiole and
post-petiole, as in Fig. 1. In spite of their striking
appearance, and a diversity of character states
that allow easy recognition of most species, the
dacetines remain poorly known. This can be at-
tributed to their small size and cryptic habits.
There are only two Nearctic genera of Dace-
tini: Strumigenys and Pyramica. Other genera
listed for this region, for example, in Bolton's
1995 catalog of ants, were synonymized by Bolton
(1999) in his reclassification of the genera of the
Dacetini. In addition, certain species that had
been assigned to Sr ...i.,.. i ... were referred to
Pyramica on the basis of a series of fundamental
character states. In practice, Nearctic Pyramica
may be recognized by their broad, well separated
mandibular bases, while S'r ....; -.. '.., have nar-
row mandibular bases that appear to be attached
near the midline of the head (Bolton 1999).
Pyramica species use their mandibles to seize and
hold prey until it can be stung, while S'r .... i-..-., ,
species are able to snap their mandibles shut with
such force that the prey may be killed outright
(Bolton 1999). Bolton (1999) presents a detailed

discussion of generic distinctions and the evolu-
tion of mandibular structure in the Dacetini.
Dacetine ants show their greatest diversity in
moist tropical regions. The revision of the tribe by
Bolton (2000) includes 872 species, only 43 of
which occur in North America north of Mexico.
Southeastern North America has the great major-
ity of Nearctic species, including, by my count, 31
native species and 9 introduced species. The na-
tive species appear to represent a Nearctic radia-
tion; only 1 native species has a range that extends
into the Neotropics. It has been suggested (Deyrup
1988) that the diverse southeastern fauna is com-
posed of species that persisted in mesic southeast-
ern refuges during the climatic shifts of the Pleis-
tocene, providing a partial glimpse of what was
once a much richer Arctotertiary woodland fauna.
With this background, it is not surprising that
additional species of dacetine ants are still being
discovered in the Southeast. Not only are these
ants small and cryptic, but some species may
have relictual geographic distribution in isolated
patches of habitat, or they may be dependent on a
specialized microhabitat that remains unknown.

Pyramica boltoni Deyrup, new species
Diagnosis of Worker (Fig. 1) and Queen
Distinguished from all other Pyramica by the
following combination of character states: clypeus
obtusely pointed, with four radiating, subapical,
and two decumbent, apical hairs; two large, curved
standing divergent hairs at apical third of clypeus;
mandibles with toothless area (= "diastemma")
basal to apical series of teeth barely visible in dor-
sal view. Otherwise, it is generally similar to P di-

Florida Entomologist 89(1)

Fig. 1. Pyramica boltoni, new species, worker: lateral habitus view and frontal view of head; length: 1.6 mm.

etrichi (M. R. Smith) (see Discussion below and
Fig. 2). On each mandible there are four enlarged
subapical teeth, of which the first (basal) in the se-
ries is widest at the base, the third is about half the
length of the two basal teeth, and the fourth is only
slightly shorter than the first two.

Description of Holotype Worker

Measurements in mm: total length 1.66 (=length
of head from clypeal apex to occipital margin +
length mesosoma + length petiole, postpetiole,
gaster); head length from clypeal apex to occipital
margin: 0.47; maximum head width: 0.33; length
of mesosoma: 0.45; length of petiole: 0.17; length
of postpetiole + gaster: 0.57. The features de-
scribed below are illustrated in Fig. 1: Clypeus in
frontal view obtusely pointed, in lateral view
slightly upturned apically; surface of clypeus
smooth, without small discal hairs; large clypeal
hairs as follows: two decumbent apical hairs ex-
tending laterally over mandibles; four radiating
subapical hairs; five pairs of sublateral recurved
hairs; one pair of stout, curved hairs, originating
at apical third of clypeus, directed upward and
outward. Frontal and occipital areas with sparse,
suberect, curved hairs: one pair at apical quarter
of antennal scrobe, one pair at sides of occipital
lobes, one pair lateral, about midway between the
other two. Mandibles with diastemma barely visi-
ble beyond clypeus in frontal view. Antennal

scapes each with five elongate, suberect hairs on
leading edge: a subbasal hair directed apically, a
hair at basal third directed slightly basally, three
hairs on apical half directed apically; scape other-
wise with shorter, subreclinate, apically directed
hairs. Pronotum shining, slightly rugose dorsally
along sides in front; a pair of elongate, irregularly
curved, fine hairs on dorsolateral carinae, another
pair on dorsal posterior angles. Mesosoma shining
laterally, obscurely reticulate-rugose dorsally;
propodeal teeth short, triangular, infradental lam-
inae weakly emarginate just below teeth. Petiole
and postpetiole with well developed subapical and
inferior spongiform lobes. Gaster shining, dorsal
surface with a few, elongate, fine, erect hairs.

Description of a Paratype Dealate Queen

Measurements in mm: total length (measured
as in worker):2.05; head length: 0.55; maximum
head width: 0.37; length of mesosoma: 0.55;
length of petiole: 0.25; length of postpetiole +
gaster: 0.70. Structural character states similar
to worker, except for presence of ocelli, large com-
pound eyes, and modifications of the mesosoma
associated with flight.

Type Material

Type localities and associated information, as
appear on specimen labels. All specimens, as far

March 2006

Deyrup: New Florida Dacetine Ant


Fig. 2. Pyramica dietrichi (M. R. Smith), worker: lateral habitus view and frontal view of head; length: 1.8 mm.

as I know, were extracted from leaf litter, with var-
ious types of Berlese funnels. I have not seen any
living specimens. Holotype worker: Florida: High-
lands Co., Archbold Biological Station, 8-X-2004,
M. Deyrup, at base of Pinus elliottii in pine and
oak copse near cottage one. Paratype dealate
queen used in description of queen: same site,
habitat, collector as paratype, 14-X-2004. Addi-
tional paratype material: all paratypes from Flor-
ida; collector's initials: L. D.: Lloyd R. Davis, Jr.;
M. D.: Mark Deyrup; C. J.: Clifford Johnson. Three
workers: same locality as types, 6-II-1984, M. D.,
Quercus geminata and Q. myrtifolia litter; 1
worker: same locality as types, 26-1-1984, M. D.,
Quercus laevis litter; 1 worker: same locality as
types, 18-1-1984, M. D., Carya floridana litter; 1
worker: same locality as types, 3-IX-1993, M. D.
Indian River Co.: Vero Beach, 7-II-1993, M. D.,
pine and oak hammock, 20 workers, 1 queen; Mar-
tin Co.: Jonathan Dickinson State Park, 2-X-1988,
M. D.,1 queen, 2 workers; Gilchrist Co.: Trenton,
1-X-1993, L. D., 1 worker; Dixie Co.: Old Town, 8
mi. north and 1.3 mi. east of Rt. 349, 11-V-1993, L.
D., 2 workers; Old Town, 11-X-1993, L. D., 1
queen; St. John's Co.: St. Augustine, 1 mi. south-
west on Rt. 207, 9-IV-1993, L. D.1 worker; Favor-
Dykes State Park, 3-V-1987, C.J., xeric upland
with Quercus laevis and Q. myrtifolia, 1 worker;
Favor-Dykes State Park, 21-III-1987, C. J., Quer-
cus laevis leaf litter, 1 worker; Favor-Dykes State

Park, II-11-1994, M. D., mesic forest near camp-
ground, 2 workers; Citrus Co.: Holden, 5 mi. west,
25-IX-1993, L. D., 1 worker; Wakulla Co.: Ochlock-
nee State Park, 7-III-1986, C. J., oak leaf litter
sample 637, 1 worker; Polk Co.: The Nature Con-
servancy Tiger Creek Preserve, 5-X-1989, M. D.,
leaf litter from Quercus laevis habitat, 2 workers;
Jackson Co.: Florida Caverns State Park, 30-5-
1988, Paul Skelley, 1 worker; Brevard Co.: Titus-
ville, State Rd. 405, 10-IV-2003, Zachary Prusak,
Enchanted Forest, leaf litter, 1 worker; Marion
Co.: Ocala, 2.5 mi. north on Rt. 441, 13-VI-1993, L.
D., 2 workers; Ocala, 9 mi. south southwest, 21-II-
1993, M. D., sand pine scrub habitat, Ocala Water-
way Scrub, 4 workers; Ocala National Forest, 23-
VII-1992, M. D., sand pine scrub, 3 mi. south Big
Scrub Campground on Rd. 588,1 worker; Ocala
National Forest, 2-IX-1985, C. J. 1 mi. west of Ju-
niper Springs on Rt. 40, sand pine scrub, 2 queens,
1 worker; Ocala, 1-II-1994, Zachary Prusak, State
Rd. 484, 1.3 mi. west of 1-75, sand pine scrub hab-
itat,1 queen; Volusia Co.: Spruce Creek Nature
Conservancy Preserve, 22-X-1994, M. D., 2 work-
ers; Putnam Co.: Rodman Reservoir, 3-IV-1988, C.
J., scrub just west of dam, sample 831, 4 workers;
Ordway Preserve, 20-XI-1993, L. D., 0.5 mi. from
main entrance, I worker; Ordway Preserve, 27-I-
1995, L. D., Berlese funnel OK-012795, 1 worker;
Florahome, 20-XII-1987, C. J., Quercus laevis san-
dhill 5 mi. north of Florahome, Rt. 100, 1 queen;


4h -,, I- .

Florida Entomologist 89(1)

Alachua Co.: Hawthorne, 8-VI-1986, C. J., sand
pine scrub 2.4 mi. east of town, 1 worker; Kanap-
aha Lake, 2-XI-1988, C. J., park near lake, oak lit-
ter, 1 worker; Cross Creek, 4-IV-1988, C. J., open
xeric pine forest 2-3 mi. north of Cross Creek, 1
worker; Cross Creek, 2 mi. southeast, 7-IX-1986,
C. J., hardwood litter sample 656B, 1 queen, 4
workers; Cross Creek, 4-VII-1985, C. J., 2 mi.
southeast of Cross Creek, oak-palmetto litter,
sample 370, 1 queen; Cross Creek, 4-VIII-1985, C.
J., 6 mi. north of town, sample 425B, 1 queen;
Gainesville, 31-XII-1988, C. J., flatwoods, county
fairgrounds, sample 790, 1 queen; Gainesville, 13-
VIII-1989, C. J., county fairgrounds, pine and pal-
metto litter, sample 1000, 1 worker.

Deposition of Type Material

Holotype, 13 workers, 2 queens: Museum of
Comparative Zoology, Harvard University, Cam-
bridge, MA; 8 workers, 2 queens: Florida State
Collection of Arthropods, Gainesville, FL; 12
workers, 1 queen: Los Angeles County Museum of
Natural History, Los Angeles, CA; remaining
paratypes: collection of the Archbold Biological
Station, Lake Placid, FL.


The species is named for Barry Bolton, whose
revisions of dacetine ants, culminating in his re-

vision of the tribe (2000) have brought organiza-
tion and logic to the group. He has enormously in-
creased the number of identified specimens in col-
lections, and has personally described several
hundred species. His work on dacetines is pre-
sented with easily used keys and numerous illus-
trations, so that the group is, for the first time, ac-
cessible to a wide range of entomologists.


Members of the genus Pyramica are usually
most easily identified by characters of the head,
especially the structures of the clypeus and man-
dibles and the modifications of the setae of the
clypeus and antennal scapes. Pyramica boltoni
shares clypeal character states with P dietrichi,
including the pointed clypeal shape with the tip
turned up away from the plane of the mandibles,
the decumbent apical setae, the set of radiating
subapical setae, and the pair of elongate, curved
setae arising near the apical third of the clypeus
(Fig. 2). A third species, P. ornata (Mayr), shares
these clypeal features, but the subapical setae are
short and strongly expanded apically (Fig. 3).
Pyramica boltoni is distinguished from P dietri-
chi by having four, rather than six, radiating sub-
apical setae, and by having the jaws protruding a
shorter distance beyond the clypeus, so that the
diastemma is barely visible in frontal view (Figs.
1 and 2). In addition, the sides of the pronotum of

Fig. 3. Pyramica ornata (Mayr), worker: lateral habitus view and frontal view of head; length: 1.8 mm.

March 2006


Deyrup: New Florida Dacetine Ant

P boltoni are primarily shiny, not reticulate as in
P dietrichi (Figs. 1 and 2). The pair of enlarged,
upturned discal setae at the apical third of the
clypeus are conspicuously larger than those of
most P. dietrichi, but the size of these setae in P.
dietrichi is somewhat variable, and it is possible
that there might be overlap with those of P. bolt-
oni in some populations. Pyramica boltoni keys to
P dietrichi in Bolton's (2000) key
Several lines of evidence strongly suggest that
P boltoni is not a variant of P. dietrichi. The diag-
nostic features listed above are consistent in all
the specimens examined from 38 separate collec-
tions spread over northern and central Florida.
Pyramica boltoni is sympatric with P dietrichi;
there is no intergradation, and there are five
known sites where both species occur. The diag-
nostic character states of the clypeal setae and the
length of the mandibles relative to that of the
clypeus are the kinds of character states that have
been used in distinguishing many species of
Pyramica, for example, in Bolton's key (2000). Un-
fortunately, these characters have not been associ-
ated with any natural history traits, but it is likely
that such traits exist, given the consistency of the
character states within each species of Pyramica.
Pyramica boltoni is known only from Florida,
ranging from Highlands and Martin Cos. in the
south-central Peninsula, north into St. John's Co.
in the northeast corner of the state, and west into
Jackson Co. in the central Panhandle. It might
well occur in southern Georgia near the Florida
border, but there are no known Georgia speci-
mens. In Florida this species does not seem to be
as widespread or abundant as P. dietrichi, which
occurs throughout the state, including the Keys,
north into Maryland and Illinois, and west into
eastern Texas. I have examined 307 specimens of
P dietrichi from Florida, Georgia, Alabama, Ar-
kansas, Oklahoma, Texas, and Illinois. Within the
area where both species are known to occur, there
is some evidence that P dietrichi may occupy a
wider range of habitats, specifically habitats that
are wet, such as low flatwoods and swamp forest.
Habitat information is available for 28 collections
of P boltoni and 49 collections of P dietrichi; all

specimens were extracted from leaf litter. Habi-
tats of P boltoni include xeric forest: 16 (57.1%);
mesic forest: 11 (39.2%); wet forest, wet flat-
woods: 1 (3.6%). Habitats of P dietrichi include
xeric forest: 23 (46.9%); mesic forest: 15 (30.6%);
wet forest, wet flatwoods: 11 (22.4%). Pyramica
dietrichi shows a significant difference in its
greater preference for, or tolerance of, wet habi-
tats (Chi square = 4.83, P value = 0.03).
Although the known distribution of P boltoni
is restricted relative to those of most native
southeastern Pyramica species and it is not par-
ticularly common within this range, it cannot be
considered a species that is rare or endangered. It
is known from a series sites where its habitat
might be expected to be protected including four
state parks, one county park, two Nature Conser-
vancy preserves, the Archbold Biological Station,
the Ordway Preserve (managed by the University
of Florida), and several sites in the Ocala Na-
tional Forest.


I gratefully acknowledge Lloyd R. Davis, Jr., Clifford
Johnson, Zachary Prusak, and Paul Skelley who gener-
ously donated specimens ofP. boltoni to the collection of
the Archbold Biological Station for this study. Leif Dey-
rup did the statistics on habitat differences. Two anon-
ymous reviewers provided useful suggestions and
corrections. I thank the governmental and private agen-
cies who protect and manage the natural areas in which
this interesting species occurs. This research was sup-
ported by the Archbold Biological Station.


BOLTON, B. 1995. A new general catalog of the ants of
the World. Harvard University Press, Cambridge,
Mass. 504 pp.
BOLTON, B. 1999. Ant genera of the tribe Dacetonini (Hy-
menoptera: Formicidae). J. Nat. Hist. 33: 1639-1689.
BOLTON, B. 2000. The ant tribe Dacetini. Mem. Ameri-
can Entomol. Institute 65: 1-1028.
DEYRUP, M. 1988. Smithistruma memorialis (Hy-
menoptera: Formicidae), a new species of ant from
the Kentucky Cumberland Plateau. Entomol. News
109: 81- 87.

Florida Entomologist 89(1)

March 2006


Departamento de Entomologia, Instituto de Ecologia A. C.
Km 2.5 carretera Antigua a Coatepec 351 Congregaci6n El Haya, 91070 Xalapa, Veracruz, M6xico
E-mail: delgadol@ecologia.edu.mx


A new Mexican and Guatemalan species, Onthophagus yucatanus, belonging to the Clypea-
tus group is described and illustrated. The distinctive characters of this species, its geo-
graphical distribution, and habits are described.

Key Words: Scarabaeidae, Onthophagus, New species, Mexico, Guatemala


Se describe e ilustra una nueva especie mexicana y guatemalteca, Onthophagus yucatanus,
perteneciente al grupo Clypeatus. Se comentan sobre sus caracteres distintivos, asi como su
distribuci6n geografica y habitos.

Translation provided by author.

The Clypeatus group of the worldwide genus
Onthophagus Latreille, represents a heteroge-
neous and taxonomically difficult group of Amer-
ican species. According to Zunino & Halffter
(1997) the Clypeatus group is formed by three
species complexes, named Clypeatus, Mirabilis,
and Nasicornis. However, delimitation of these
complexes, and of all the groups of this genus, is
vague and requires clarification based on phylo-
genetics approaches. At present we prefer, as did
Howden & Gill (1993) and Kohlmann & Solis
(2001), to exclude the species of theDicranius and
Mirabilis groups (sensu Howden & Gill 1993). In
addition, we also exclude the species of the Nasi-
cornis complex (sensu Zunino & Halffter 1997)
from the Clypeatus group, primarily on the basis
of the lack of tubercles and horns on the vertex, at
least in the males.
Thus, the Clypeatus group would include only
the following species: 0. clypeatus Blanchard
from Colombia, Ecuador, Peru, Bolivia, and
French Guyana; 0. rhinophyllus Harold from
Venezuela and Colombia; 0. rhinolophus Harold
from Mexico and Guatemala; 0. belorhinus Bates
from Mexico and Guatemala; 0. xanthomerus
Bates from Colombia, Ecuador, and Peru; 0. prae-
cellens Bates from Costa Rica, Panama, and Co-
lombia; 0. dicranoides Balthasar from Ecuador;
0. lojanus Balthasar from Ecuador; 0. maya
Zunino from Mexico and Belize; 0. propraecellens
Howden & Gill from Costa Rica and Panama; O.
andersoni Howden & Gill from Costa Rica; O. lu-
ismargaritorum Delgado from Mexico; 0. ver-
acruzensis Delgado from Mexico; 0. coriaceoum-
brosus Kohlmann & Solis from Costa Rica; 0. gra-

taehelenae Kohlmann & Solis from Costa Rica
and Panama; 0. limonensis Kohlmann & Solis
from Costa Rica; 0. nemorivagus Kohlmann &
Solis from Costa Rica; 0. singulariformis Kohl-
mann & Solis from Costa Rica; 0. viridivinosus
Kohlmann & Solis from Costa Rica and 0. no-
tiodes Solis & Kohlmann from Costa Rica (Zunino
& Halffter 1997; Delgado & Pensado 1998; Kohl-
mann & Solis 2001; Solis & Kohlmann 2003). Dis-
tribution of these species is almost restricted to
the tropical rain forests in areas generally below
1,000 m asl. In contrast to the extensive coproph-
agy of many species of Onthophagus, most species
of this group show a strong tendency towards
feeding on rotting fruit and carrion (Zunino &
Halffter 1997).
In two recent studies on the fauna of coleopter-
ous Scarabaeidae of the Peninsula de Yucatan,
Mexico (Peraza 2004, unpublished data), and of
the region of Peten, Guatemala (Cano 1998, un-
published data), several specimens were obtained
which represent an undescribed species of Ontho-
phagus. We describe it here in the Clypeatus

Onthophagus yucatanus, new species
(Fig. 1, 4)

Type Material

Holotype: "MEXICO: Yucatan, Tzucacab, Tigre
Grande, 17-X-2001. 0900-1730 horas, coprotrampa
humanno, L. N. Peraza-Flores col.". Allotype same
data as holotype, except: 20-VIII-2001. Both depos-
ited in the Entomological Collection (IEXA) of the

Delgado et al.: A New Onthophagus from Mexico and Guatemala

Institute de Ecologia, A. C. (Veracruz, Mexico).
Paratypes (116 S' 132 Y 2) same data as holo-
type, except: 19-VIII-2001 (1 6, 6 2 Y); 20-VIII-
2001 (14 6S, 6 Y Y); 17-18-IX-2001, 18:00-09:00
hrs (1 2); 18-19-IX-2001, 18:00- 09:00 hrs (4 S8,
10 Y ); 19-IX-2001, 09:00-18:00 hrs (14 S6 7 2 2);
19-20-IX-2001, 18:00-09:00 hrs (3 6S, 7 2 2); 17-
X-2001, 09:00-17:30 hrs (25 S 28 2 2); 17-18-X-
2001, 17:30-08:00 hrs (10 6S, 9 2 2); 16-XI-2001,
08:30-17:30 hrs (6 6S, 5 9 ); 16-17-XI-2001,
17:30-09:00 hrs (4 2 2); 17-XI-2001, 09:30-17:30
hrs (2 SS 6 2 2); 17-18-XI-2001, 17:30-07:00 hrs
(2 66); 15-16-XII-2001, 17:00-08:30 hrs (1 S, 2 2
2); 16-XII-2001, 08:30-17:20 hrs (2 S6 5 2 2); 13-
1-2002, 09.00-17:30 hrs (1 S, 3 2 2); 11-12-II-2002,
17:15- 08:30 hrs (1 S, 1 2); 12-II-2002, 08:30-17:30
hrs (3 S 4 2 2); 12-II-2002, 17:30-09:00 hrs (2
6S, 2 9 9); 13-II-2002, 09:00- 19:35 hrs (1 9); 13-
14-II-2002, 17:35-08:00 hrs (1 6); 11-12-III-2002,
17:30- 09:30 hrs (1 9); 12-III-2002, 09:30-18.00 hrs
(3 S 2 9 9); 13-III-2002, 09:00-18:00 hrs (1 S, 1
9); 13-14-III-2002, 18:00-09:00 hrs, (3 6S, 1 9);
19-IV-2002 (1 6); 14-V-2002, 10:00-18:30 hrs (3
SS 2 9 9); 14-15-V-2002, 18:30-10:00 hrs (1 S, 1
9); 10-12-VI-2002 (2 9 9); 10-12-VII-2002, 18:30-
08:30 hrs (3 6S, 5 9 9); 13-V-2002, excremento de
perro (1 9); 20-VIII-2001-17-IX-2001 NTP80 (1 9);
17-IX-2001-17-X-2001, NTP80 (1 9); 17-X-2001-
16-XI-2001, NTP80 (1 S); 16-XI-2001-16-XII-2001,
NTP 80 (1 9); 13-II-2002-11-III-2002, NTP80 (1 9);
15-V-2002-12-VI-2002, NTP80 (4 S6); 12-VI-2002-
12-VII-2002, NTP80 (1 9); 12-VII-2002-17-VIII-
2002, NTP80 (1 6). "GUATEMALA: Peten, Aldea
Carmelita, Campamento Chuntuqui, 24-25-II-
1996, bosque alto, 1732'N 90007'W, E. Cano col." (2
S S, 1 9); same data as anterior, except: Campa-
mento El Naranjo, 25-II-1996, heces de jaguar, E.
Cano col. (1 9); San Miguel la Palotada, 16-III-
1999, M. Jol6n col. (1 S, 1 9); San Miguel la Palo-
tada, 7-VIII-1999, M. Jol6n col. (1 9). Twenty
paratypes deposited in each one of the following
collections: Florida State Collection of Arthropods
(Gainesville, United States), Canadian Museum of
Nature (Ottawa, Canada) and Instituto de Bi-
ologia de la Universidad Nacional Aut6noma de
Mexico (Mexico City); 151 paratypes deposited in
the Instituto de Ecologia, A. C. (Veracruz, Mexico);
seven paratypes deposited in the Universidad del
Valle de Guatemala (Guatemala City); and ten
paratypes deposited in each one of the following
collections: Cuauhtemoc Deloya Collection (Ver-
acruz, Mexico), Lizandro N. Peraza Collection
(Yucatan, Mexico) and Leonardo Delgado Collec-
tion (Mexico City).


Holotype male (Fig. 1, 2). Length: 5.4 mm,
maximum width (at basal third of elytra): 3.3
mm. Small, ovate, dorsally glabrous; dorsal color
metallic dark green with very slight cupreous re-

flections on head, anterolateral regions of prono-
tum dull, venter with same color as dorsum but
with cupreous reflections more pronounced, fem-
ora and tibiae completely reddish green, tarsi red-
dish. Clypeus triangular, with apical third ob-
liquely reflexed, apex acuminated, sides slightly
and evenly sinuated to the genae; clypeus moder-
ately concave, front clypeal region feebly convex,
frons flattened, genae slightly convergent for-
ward; vertex with two slender, divergent horns,
bases of horns well separated and arising at level
of posterior portion of eyes, horns slightly below
the top of pronotum. Head with punctures small,
shallow and sparse, shallower and finer towards
apex and vertex.
Pronotum with anterior right angles and some-
what projected, lateral borders strongly curved in
front of middle, posterior angles almost evenly
rounded to base. Disc and posterolateral portions
of pronotum swollen; anterior third declivous with
two longitudinal, obtuse tubercles in front of disc,
tubercles slightly divergent and separated by a
shallow concavity widened forward; tubercles and
lateral portions of pronotum delimiting two very
feeble concavities. Pronotum with moderately
large, ringed punctures, denser and larger at ante-
rior angles; lateral concavities and anterior angles
with finely reticulate punctation; anterior concav-
ity almost smooth; pronotal base finely margined
with a row of small punctures. Elytra with evident
humeral and apical calli; elytral striae marked by
double line crenulated by medium-size punctures;
intervals with fine, shallow punctures and rugosi-
ties, denser to lateral intervals.
Metasternum convex with dense, ringed punc-
tures. Abdominal sternites shagreened, with a
row of small punctures adjacent to anterior mar-
gin, each puncture bearing a yellowish seta; sixth
abdominal sternite narrowed medially. Pygidium
strongly convex to apical third, surface with large
and deep punctures moderately dense, each punc-
ture bearing a small, yellowish seta. Protibiae
with inner border evenly curved, outer border
quadridentate with teeth situated at distal mid-
dle, apex with inner projection bearing a brush of
setae; apical spur elongate and curved outside.
Apex of meso and metatibiae with a row of small
spinules intermixed with long setae. Genitalia
with parameres strongly projected ventrally, dor-
sally flattened and with apices parallel.
Allotype female (Fig. 1, 4). Length: 5.3 mm,
maximum width (at basal third of elytra): 3.3
mm. Differs from holotype in the following re-
spects: anterolateral regions of pronotum scarcely
dull; clypeus semitrapezoidal, slightly emargin-
ated at middle, anterior margin very scarcely re-
flexed and with sides almost straight to the ge-
nae; frontoclypeal region carinated from side to
side; frons with two rounded, transversal tuber-
cles situated slightly behind of anterior border of
eyes; vertex without horns; clypeus with strongly

Florida Entomologist 89(1)

Fig. 1. Dorsal view of head and pronotum of Onthophagus spp. 1, male of 0. luismargaritorum Delgado. 2, male
of 0. yucatanus sp. nov. 3, female of 0. luismargaritorum Delgado. 4, female of 0. yucatanus sp. nov.

rugose punctation, genae with coarse punctures;
pronotum with anterior angles not projected, al-
most evenly convex, only with a small, central
concavity adjacent to anterior margin; pronotum
with finely reticulate punctation restricted to an-
terior angles; sixth abdominal sternite not nar-
rowed medially; protibiae slightly broader and

without inner projection and brush of setae; api-
cal spur longer.
Variation in the series of paratypes (113 6 6,
128 Y 2).-Length: 3.5-5.6 mm, maximum width
(at basal third of elytra): 2.3-3.6 mm. The dorsal
color varies from dark green to blue; size of punc-
tures on elytra varies moderately in both sexes; in

March 2006

Delgado et al.: A New Onthophagus from Mexico and Guatemala

the smaller males the clypeus is scarcely reflexed
and the cephalic horns are reduced to small,
transverse tubercles, the pronotum is nearly
evenly convex, with only a small concavitiy
flanked by two small rounded tubercles; in the
smaller females the pronotum is evenly convex,
lacking anterior concavity.

Type Locality
Mexico, Yucatan, Tzucacab, Tigre Grande
(19042'36"N 89002'28"W).

The specific epithet derives from Yucatan, the
name of the Mexican state where this species was

Taxonomic Remarks

Onthophagus yucatanus shares several charac-
ters with 0. luismargaritorum. Both species are
distinguished from the remaining species in the
Clypeatus group by the following combination of
characters: major males with clypeus obliquely re-
flexed and acuminated (not rectangular, rounded
or with a projection "T" shaped), the horns on the
vertex arising between the eyes (not arising behind
the eyes) and the protibiae short and wide (not
elongate and slender), and the females with the tu-
bercles on the vertex situated at level of the ante-
rior border of eyes (not at level of the posterior bor-
der of eyes), and the pronotum without tubercles.
The two species are separated by the following
characters: males and females of any size of
O. yucatanus by the dorsal color metallic dark
green or blue with very slight cupreous reflections
on head and larger and denser ringed punctures of
the pronotum, not with dorsal color strongly me-
tallic cupreous green and small and sparse ringed
punctures of pronotum as in 0. luismargaritorum;
major males of 0. yucatanus with the central tu-
bercles of pronotum obtuse and with the central
concavity widened to anterior margin (Fig. 1, 2),
not with tubercles rounded and the central con-
cavity with same wide from tubercles to anterior
margin as in 0. luismargaritorum (Fig. 1, 1). Fe-
males of any size of 0. yucatanus with the cephalic
tubercles transverse (Fig. 1, 4), not with cephalic
tubercles oblique as in females of 0. luismargari-
torum (Fig. 1, 3) and major females of 0. yucata-
nus with a small concavity adjacent to anterior
margin of pronotum, not with pronotum evenly
convex as in females of 0. luismargaritorum.

Onthophagus yucatanus is known from the
type locality, situated in the central region of the

Peninsula de Yucatan, Mexico, and from the re-
gion of Pet6n, Guatemala. Both areas present
tropical moist subdeciduous forest with low alti-
tude. Extensive sampling in other localities in the
northern peninsula with dry deciduous forests
produced no specimens of this species.


At the type locality, specimens of 0. yucatanus
were caught with traps baited with human excre-
ment and with traps baited with rotting squid.
Two hundred and thirty one specimens were ob-
tained in the coprotraps and only 11 in the ne-
crotraps. This feeding preference contrasts with
that of most species of the Clypeatus group, which
are captured primarily at rotting fruit and car-
rion. Only 0. luismargaritorum shares the same
preferences, with 83% of the specimens known of
this species collected at human and cow dung,
and only 17% with necrotraps (Delgado 1995).
Although many specimens were collected dur-
ing the day (124) in comparison to those caught
during the night (75), more data are needed to de-
fine the daily activity of this species.


The authors thank Enio Cano for loan of specimens
from Guatemala, Enrique Reyes for assistance in the
collecting trips to the localities of the Peninsula de
Yucatan, and Andr6s Trejo and Francisco Nava for par-
tial support during the collecting trips. This work is a
contribution to the projects "Ecology and systematics of
phytophagous and saprophagous insects" (902-08/044)
and "Interactions: Ecology and systematics" of the Insti-
tuto de Ecologia, A. C.


DELGADO, L. 1995. Onthophagus luismargaritorum,
nueva especie mexicana del grupo Clypeatus (Co-
leoptera: Scarabaeidae). Folia Entomol6gica Mexi-
cana 94: 57-61.
DELGADO, L., AND M. PENSADO. 1998. Una nueva espe-
cie mexicana de Onthophagus del grupo Clypeatus
(Coleoptera: Scarabaeidae). Folia Entomol6gica
Mexicana 103: 75-80.
HOWDEN, H. F., AND B. D. GILL. 1993. Mesoamerican
Onthophagus Latreille in the Dicranius and Mirabi-
lis groups (Coleoptera: Scarabaeidae). The Canadian
Entomol. 125: 1091-114.
KOHLMANN, B., AND A. SOLiS. 2001. El g6nero Onthoph-
agus (Coleoptera: Scarabaeidae) en Costa Rica. Gior-
nale Italiano de Entomologia 9: 159-261.
SOLiS, A., AND B. KOHLMANN. 2003. New species of dung
beetles (Coleoptera: Scarabaeidae: Scarabaeinae)
from Costa Rica and Panama. Zootaxa 139: 1-14.
ZUNINO, M., AND G. HALFFTER 1997. Sobre Onthopha-
gus Latreille, 1802 americanos (Coleoptera: Scara-
baeidae: Scarabaeinae). Elytron 11: 157-178.

Florida Entomologist 89(1)

March 2006


'University of Florida, Department of Entomology and Nematology, Tropical Research and Education Center
18905 SW 280th Street, Homestead, FL 33031

2U.S. Horticulture Research Laboratory, USDA-ARS, 2001 South Rock Road, Fort Pierce, FL 34945


Twelve pesticides used in citrus were tested for their contact toxicity to Aprostocetus va-
quitarum Wolcott (Hymenoptera: Eulophidae) a parasitoid of Diaprepes abbreviatus (L.)
(Coleoptera: Curculionidae). Sevin 80 WSP, Malathion 5 EC, and Imidan 70 WSB re-
sulted in the most rapid death ofA. vaquitarum adults. Admire@ 2F, Danitol 2.4EC, and
Surround WP were also very detrimental. Kocide 101 WP, Citrus Soluble Oil, Micromite@
80 WGS, Acramite@ 50 WS, Micromite 80 WGS + Citrus Soluble Oil, Aliette WDG, and
Agrimek 0.15 EC + Citrus Soluble Oil were slightly to non-toxic to A. vaquitarum. The rel-
ative toxicity of the pesticides was consistent up to four weeks after application. Signifi-
cantly fewer adult A. vaquitarum emerged from D. abbreviatus eggs laid on foliage treated
in the field with Sevin XLR and Imidan 70 WSB than emerged from the water treated
control. Field residues of Sevin XLR remained toxic for seven days while the effects of Im-
idan 70 WSB were no longer significant after one week. The number of A. vaquitarum
adults emerging from host eggs laid on treated foliage was not significantly different among
Micromite 80 WGS, Acramite 50 WS, and the control, but significantly fewer adults
emerged from foliage treated with either Micromite@ 80 WGS + Citrus Soluble Oil or Citrus
Soluble Oil alone. There were no significant differences between oviposition or new genera-
tion adults when A. vaquitarum was exposed to Micromite 80 WGS or a water control.

Key Words: insecticides, diflubenzuron, selectivity, toxicity, citrus IPM, biological control


Se estudi6 la toxicidad de various plaguicidas aplicados comunmente en citricos para Apros-
tocetus vaquitarum Wolcott (Hymenoptera: Eulophidae) un parasitoide de Diaprepes abbre-
viatus L. (Coleoptera: Curculionidae). Sevin 80 WSP, Malathion 5 EC, e Imidan 70 WSB
fueron los que mas rdpidamente causaron la muerte de A. vaquitarum. Admire@ 2F, Dani-
tol 2F, Danitol 2.4 EC y Surround WP fueron muy t6xicos para el parasitoide. Compa-
rados con el testigo absolute, Kocide 101 WP, aceite soluble de citricos, Micromite@ 80
WGS, Acramite 50 WS, Micromite@ 80 WGS + aceite soluble de citricos, Aliette WDG y
Agrimek 0,15EC + aceite soluble de citricos no resultaron significativamente t6xicos para
el parasitoide. La toxicidad relative de estos plaguicidas se mantuvo durante un period de
4 semanas. Emergieron significativamente menos adults de A. vaquitarum de huevos de
D. abbreviatus que habian sido depositados en hojas tratadas en el campo con Sevin XLR
e Imidan 70 WSB en comparaci6n con aqu6llos que emergieron de huevos depositados en
hojas tratadas con agua. Los efectos t6xicos de Sevin XLR continuaron por 1 semana, mien-
tras que los efectos de Imidan dejaron de ser significativos despu6s de 1 semana. No hubo di-
ferencias significativas entire el numero de adultosA. vaquitarum emergidos de huevos de D.
abbreviatus en follaje tratado con Micromite@ 80 WGS y Acramite 50 WS comparado con
el testigo absolute. Sin embargo, si las hubo con Micromite 80 WGS + aceite soluble de ci-
tricos o con el aceite soluble de citricos cuando este ultimo fue aplicado solo. No hubo dife-
rencias significativas ni en la puesta ni en la emergencia de una nueva generaci6n de
parasitoides cuando se expuso las hembras a Micromite@ 80WGS en comparaci6n con aqu6-
llas que se expuso al agua (testigo absolute).

Translation provided by the authors.

Pesticides are a critical component of insect species targeted. Of particular concern are bene-
pest management in citrus production. Insect ficial insects which play a vital role in suppress-
fauna affected by pesticide applications often en- ing pest insect populations. Non-selective pesti-
compasses a group extending beyond the pest cide application can disrupt beneficial insect pop-

Ulmer et al.: Pesticide Toxicity to A. vaquitarum

ulations and may lead to outbreaks of pest insects
(Barbosa & Schultz 1987). Much of Florida's cit-
rus production is intended for juicing, which re-
quires a less severe pesticide regime than the
fresh fruit market (Michaud & Grant 2003). How-
ever, populations of specific pests, such as the root
weevil, Diaprepes abbreviatus (L.) (Coleoptera:
Curculionidae), regularly necessitate the use of
broad spectrum insecticides (Timmer et al. 2005).
Diaprepes abbreviatus is native to the Carib-
bean and was presumably introduced from Pu-
erto Rico. It was first reported in Florida in 1964
and is established across the citrus-producing re-
gions of the state (Woodruff 1964). Diaprepes ab-
breviatus feeds on >270 species of plants from 59
families (Simpson et al. 1996). It is a significant
pest for ornamental growers and is economically
very important in the citrus industry where it is
estimated to cost producers over 70 million dol-
lars annually (Stanley 1996). Adult weevils feed
along the edges of leaves leaving characteristic
semi-circular notches but the most significant
damage is done by larvae feeding on the root sys-
tem which can weaken or kill the plant. Root feed-
ing also may leave the plant more susceptible to
root rot organisms such as Phytophthora spp.
(Timmer et al. 2005). Though there is evidence
that soil-applied pesticides may be effective
against larvae (McCoy et al. 1995), pesticide re-
gimes often target the adult stage with foliar ap-
plications, particularly during periods of new cit-
rus growth.
Biological control is a vital component in an
ongoing effort toward an integrated pest manage-
ment system for D. abbreviatus. In the late 1990s,
programs were initiated to introduce hymenop-
teran egg parasitoids from the Caribbean islands
into Florida. In 2000 the ecto-parasitoidAprosto-
cetus vaquitarum Wolcott (Hymenoptera: Eulo-
phidae) was introduced into Florida from the Do-
minican Republic (Jacas et al. 2005).Aprostocetus
vaquitarum has been mass reared and released in
several Florida counties since 2000 and is now
considered to be established in parts of southern
Florida (Jacas et al. 2005).Aprostocetus vaquitar-
tum is one of the principal parasitoids ofD. abbre-
viatus in its native range, and in areas where it
has become established in south Florida, egg mor-
tality rates of 70-90% have been observed (Peia
et al. 2005).
Very little is known about the toxicity of insec-
ticides used in citrus production to parasitoids of
D. abbreviatus. Two products were tested against
two other D. abbreviatus egg parasitoids (Amalin
et al. 2004) but no information exists on the rela-
tive toxicities of commonly used pesticides to
A. vaquitarum. Our study was initiated to deter-
mine the relative toxicities of several pesticides
used in Florida citrus production to A. vaquita-
rum. Four pesticides registered for control of
D. abbreviatus and three insecticides registered

for other citrus insect pests were examined, as
well as four mite control products and two fungi-
cides. The contact toxicity of the pesticides was
evaluated in the laboratory. Some products were
also tested further in the field. Information on the
relative potency of these crop protection products
could be of use in the development ofD. abbrevia-
tus management strategies aimed to minimize
adverse impacts on beneficial insects such as
A. vaquitarum.



Twelve pesticides labeled for use against
D. abbreviatus in Florida were tested for toxicity
to A. vaquitarum (Table 1). The application rate
for each pesticide was based on the 2005 Florida
Citrus Pest management Guide (Timmer et al.
2005). All commercially formulated pesticides
were diluted in water to an application rate of
935.4 liter/ha (100 gal/ac).

Pesticide Toxicity Trial

To test the toxicity of the 12 pesticides listed
above, female A. vaquitarum (<72 h old) were ex-
posed to the recommended field application rate
of each commercial formulation. Filter paper
(Fisherbrand Filter Paper, P4 Medium-Fine po-
rosity, slow flow rate, Cat. no.: 09-803-6G) was
dipped in a mixed pesticide or a water control for
3 seconds and air dried for 2.5 h. The treated filter
paper was then cut into 0.5 cm x 5.0 cm strips. Fe-
male A. vaquitarum were placed one individual
per tube into 10-ml test tubes. A smear of honey
was provided on the inner surface of each test
tube as a food source and the open end of the test
tube was covered with 2 ply of kimwipe (Kim-
berly-Clarke, Kimwipes EX-L) secured with
rubber tubing to allow ventilation. The kimwipe
was moistened with water daily. Pesticide-treated
and control filter paper strips were placed in the
test tubes and each parasitoid was examined for
signs of life 8, 16, 24, 48, 72, and 96 h after expo-
sure. Mortality was assessed under a 50x dissect-
ing microscope. An insect was recorded as dead if
it did not move or twitch in a 10-s period. The ex-
periment was repeated 4 times with 10 to 14 indi-
viduals in each replicate for each treatment. Two
of the four replicates also were monitored at 24-
hour intervals until 100% mortality was achieved
to assess longevity.
An experiment was carried out exactly as
above with pesticide strips tested at intervals of 7
d, 14 d, and 21 d after pesticide application for
treatments that resulted in higher mortality than
the control when tested 0 d after application. Pes-
ticide and control strips were weathered out of
doors and protected from precipitation and direct

Florida Entomologist 89(1)


Application rate
Trade name Class Active ingredient (product/1 litre water) Manufacturer

Sevin 80 WSP1 Carbamate Carbaryl (80%) 12.0 g Bayer CropScience
Sevin XLR1 Carbamate Carbaryl (44.1%) 10.0 ml Bayer CropScience
Malathion 5 EC1 Organophosphate Malathion (57%) 7.5 ml Micro Flo Company
Imidan 70 WSB1 Organophosphate Phosmet (70%) 2.4 g Gowan Company
Admire 2 F' Neonicotinoid Imidacloprid (22%) 2.5 ml Bayer CropScience
Danitol 2.4 EC1 Pyrethroid Fenpropathrin (30.9%) 1.6 ml Valent USA Corporation
Surround WP' Kaolin clay Kaolin clay (95%) 60.0 g Engelhard Corporation
Kocide 101 WP2 Copper fungicide Copper 1..I.-,l I... i ;; 14.4 g Griffin 1.1.c
Alliete WDG2 Phosphonate Aluminium tris (80%) 6.0 g Bayer CropScience
AgriMek 0.15 EC' Glycoside Abmectin (2%) 0.5 ml Syngenta
Citrus Soluble Oil3 Petroleum oil Petroleum oil (99.3%) 10.0 ml Platte Chemical Company
Micromite 80 WGS3 IGR Diflubenzuron (80%) 0.47 g Crompton Manufacturing
Company, Inc.
Acramite 50 W53 Unknown Bifenazate (50%) 1.2 g Crompton Manufacturing
Company, Inc.

Insecticide, Fungicide, Miticide.

sunlight in 1.5-liter wax paper dishes with perfo-
rated fitted plastic lids. All experiments were con-
ducted in a walk-in growth chamber with a 16:8
(L:D) photoperiod, 23C night:26C day tempera-
ture regime, and 58-60% RH. Percent mortality
was corrected with Abbott's Formula: 100 x (1 %
surviving on treatment/surviving on control) (Ab-
bott 1925). Longevity ofA. vaquitarum on the dif-
ferent treatments was compared by one-way
ANOVA. Mean separations were performed with
the Tukey HSD method (a = 0.05) (Statistix 8
Analytical Software, 2003).

Effect of Residual Pesticide on Parasitism

To test the residual effects of specific pesticides
on A. vaquitarum, products were applied out of
doors to 1-1.5 m tall green buttonwood (Conocar-
pus erectus L.) host plants. In the first of two ex-
periments Sevin@ XLR, Imidan 70 WSB, and a
water control were tested in a randomized com-
plete block design, and replicated 3 times with 3
green buttonwood plants per replicate. Treat-
ments were applied with a hand-gun sprayer op-
erating at 2413 kPa (350 psi) and delivering 935.4
liter/ha of finished spray (~3.79 liter/tree). Four
hours after treatment, 15-cm long branches from
each treatment (n = 30) were placed in 500-ml
plastic containers with water and offered to 150
field collected D. abbreviatus inside 30 x 30 x 30-
cm plexiglass cages. After being exposed toD. ab-
breviatus for 12 h, the branches with egg masses
were removed and placed inside a clean plexiglass
cage into which female A. vaquitarum (2 per egg
mass) were released. The A. vaquitarum females
were removed after 48 h. The branches remained
in water for a further 72 h after which the egg
masses were cut from the branches with scissors

and placed into individual 10-ml test tubes until
emergence. Host plant material from each treat-
ment was collected 7 and 14 d after being sprayed
and was exposed toD. abbreviatus andA. vaquita-
rum as above.
In a separate experiment Micromite 80 WGS,
Acramite 50 WS, Citrus Soluble Oil, Micro-
mite@ 80 WGS + Citrus Soluble Oil, and a water
control were tested with green buttonwood host
plants exactly as described in the preceding para-
graph. Data for both experiments were analyzed
with a Kruskal-Wallis ANOVA (Statistix 8 Ana-
lytical Software, 2003).

Micromite 80 WGS Fecundity Trial

To test the effects of the insect growth regula-
tor Micromite 80 WGS on fecundity, female
A. vaquitarum (<48 h old) were exposed to filter
paper strips (1 cm x 5 cm) treated with Micro-
mite@ 80 WGS or a water control and placed in 9-
cm diameter petri dishes. A smear of honey was
placed on the inner surface of the lid as a food
source, and a water soaked cotton disc (1 cm di-
ameter, 0.3 cm height) was placed in the dish. A 3-
cm diameter hole was cut in the lid and covered
with fine mesh to provide ventilation. Females
were exposed to each treatment for 70 h. After 70
h, one D. abbreviatus egg mass (<48 h old) was in-
troduced into the dish. The two leaves which con-
tained the egg mass were left intact and the peti-
oles were inserted into a water soaked block of flo-
rist's foam covered in aluminum foil. The egg
mass was exposed to a female for 24 h and then
removed. A total of three egg masses were offered
to each female at 24-h intervals. Fifty females
were used for each treatment; a sub-sample of egg
masses from 20 of the females on each treatment

March 2006

Ulmer et al.: Pesticide Toxicity to A. vaquitarum

was dissected 40 h after being exposed. The egg
masses from the other 30 females were reared to
emergence. Egg masses opened after 48 h were
examined under a dissecting microscope to deter-
mine the stage of A. vaquitarum development.
The number of adultA. vaquitarum and the num-
ber of D. abbreviatus eggs available in each egg
mass were calculated at the completion of the ex-
periment with aid of a dissecting microscope. The
experiment was conducted in a growth chamber,
with conditions as above. Data were subjected to
a Kruskal-Wallis ANOVA to analyze oviposition
and adult emergence as well as detect significant
differences between the means (a = 0.05) (Statis-
tix 8 Analytical Software, 2003).


Pesticide Toxicity Trial

When the pesticides were tested on the day of
application, Sevin@ 80 WSP, Malathion 5 EC, and
Imidan 70 WSB resulted in more rapid death of
A. vaquitarum than the other products or the con-
trol (F ,,,, = 27.5, P < 0.001) (Table 2). Females
survived only 15.3 1.7 h on Sevin 80 WSP. The
pesticides Kocide 101, Citrus Soluble Oil, Micro-
mite 80 WGS, Acramite 50 WS, Micromite 80
WGS + Citrus Soluble Oil, and Agrimek 0.15 EC
+ Citrus Soluble Oil were not significantly differ-
ent from the control on whichA. vaquitarum sur-
vived 122 10.1 h. Admire 2 F, Danitol 2.4 EC,
Surround WP, and Aliette were intermediate.
Seven days after application, Sevin 80 WSP,
Malathion 5 EC, and Imidan 70 WSB once again
resulted in the most rapid death ofA. vaquitarum
which survived only 18.4 1.4 h on Sevin 80

WSP (F,,,,0 = 40.5, P < 0.001) (Table 2). Aliette, Ko-
cide101, and Agrimek 0.15 EC + Citrus Solu-
ble Oil were not significantly different from the
control on which females survived a mean of
123.4 9.3 h. Admire 2 F, Danitol 2.4 EC and
Surround@ WP resulted in intermediate survival
times. Fourteen days after application Sevin 80
WSP resulted in the most rapid death (16.4 1.2
h) followed by Malathion 5 EC, Imidan 70 WSB,
Admire 2F, Kocide101, Danitol 2.4 EC, and
Surround WP. Agrimek 0.15 EC + Citrus Solu-
ble Oil was not different from the control on which
females survived for a mean of 124.8 17.3 h
(F8,11 = 16.7, P < 0.001) (Table 2). Twenty-one days
after application Sevin 80 WSP, Malathion 5
EC, and Imidan 70 WSB again resulted in the
most rapid death of A. vaquitarum, which sur-
vived 24.0 2.5 h on Sevin@ 80 WSP Admire@ 2
F, Danitol 2.4 EC, and Surround WP resulted
in intermediate survival time, while Kocide101
and Agrimek 0.15 EC + Citrus Soluble Oil were
not significantly different from the control on
which females survived for a mean of 173.2 21.2
h (F ,19 = 25.5, P < 0.001) (Table 2).
Sevin 80 WSP was the most toxic pesticide
whenA. vaquitarum was exposed to the pesticides
on the same day as pesticide application, result-
ing in 24% mortality after 8 h and 98% mortality
after 24 h (Table 3). Malathion 5 EC and Imidan
70 WSB were the only other products which re-
sulted in mortality after 8 h, and following Sevin
80 WSP were the most toxic of the products tested
resulting in 100 and 98% mortality, respectively,
after 48 h. All three products resulted in 100%
mortality of A. vaquitarum by 72 h. Admire 2 F
and Danitol 2.4 EC were the next most lethal
products resulting in 96 and 85% mortality by the


Time after application

Treatment Od 7d 14 d 21d

Sevin 80 WSP 15.3 (1.7) e 18.4 (1.4) c 16.4 (1.2) d 24.0 (2.5) d
Malathion 5 EC 26.0 (3.4) e 23.1 (2.4) c 33.2 (3.1) cd 25.4 (2.3) d
Imidan 70 WSB 23.0 (3.6) e 25.1 (2.4) c 45.6 (6.6) bcd 42.1 (4.6) d
Admire 2 F 62.0 (5.8) cd 58.3 (2.9) b 57.6 (3.2) bc 67.8 (5.4) bcd
Danitol 2.4 EC 95.0 (6.0) bc 59.1 (3.6) b 68.4 (6.1) bc 109.6 (7.8) b
Surround WP 91.0 (7.4) bc 66.9 (4.1) b 78.0 (8.7) b 93.9 (9.5) bc
Kocide 101 WP 108.0 (10.5) ab 103.7 (9.9) a 67.2 (4.1) bc 168.0 (20.2) a
Agrimek 0.15 EC & Oil 115.0 (9.4) ab 106.3 (8.0) a 126.0 (16.0) a 163.8 (15.1) a
Alliete WDG 93.0 (8.5) bc 132.9 (12.0) a
Citrus Soluble Oil 104.7 (6.1) ab
Micromite 80 WGS 115.0 (9.4) ab
Acramite 50 ws 139.0 (9.1) a
Micromite 80 WGS & Oil 135.0 (8.6) a
Control (water) 122.0 (10.1) ab 123.4 (9.3) a 124.8 (17.3) a 173.2 (21.2) a

Means within each time of application followed by the same letter are not significantly different (P = 0.05).

Florida Entomologist 89(1)

March 2006


Percent mortality Abbott's Corrected

Treatment 8h 16 h 24 h 48 h 72 h 96 h 72 h 96 h

0 d after application
Sevin80 WSP
Malathion 5 EC
Imidan 70 WSB
Admire 2 F
Danitol 2.4 EC
Surround WP
Kocide 101 WP
Agri-Mek 0.15 EC & Oil
Citrus Soluble Oil
Micromite 80 WGS & Oil
Micromite 80 WGS
Acramite 50 WS

7 d after application
Sevin80 WSP
Malathion 5 EC
Imidan70 WSB
Admire 2 F
Danitol 2.4 EC
Surround WP
Kocide 101 WP
Agri-Mek 0.15 EC & Oil
Alliete WDG

14 d after application
Sevin80 WSP
Malathion5 EC
Imidan 70 WSB
Admire 2F
Danitol 2.4 EC
Surround WP
Kocide 101 WP
Agri-Mek 0.15 EC & Oil

21 d after application
Sevin80 WSP
Malathion 5 EC
Imidan 70 WSB
Admire 2 F
Danitol 2.4 EC
Surround WP
Kocide 101 WP
Agri-Mek 0.15 EC & Oil

82.5 98.0
36.3 76.3
47.8 86.8
4.5 26.3
2.3 2.3
0 0
0 0
0 2.0
0 0
2.0 2.0
0 0
0 0
0 0
0 0

71.5 93.8
58.8 84.5
52.3 84.8
6.8 26.8
0 5.3
0 0
0 2.3
0 0
0 0
0 0

64.5 95.8
31.5 60.8
24.3 55.3
8.5 30.0
0 4.3
0 0
0 2.5
0 4.5
0 0

64.8 87.0
33.3 69.8
27.3 52.5
14.3 21.3
2.3 7.0
0 2.5
0 0
0 0
0 2.3

100 100 100
100 100 100
98.0 100 100
59.5 89.5 95.8
22.0 55.5 85.0
21.3 36.5 67.0
13.0 39.0 72.8
10.5 40.8 80.5
6.3 27.5 58.3
2.0 13.3 61.0
2.0 8.8 36.8
6.5 8.5 40.5
2.0 4.3 27.8
8.8 17.5 50.0

100 100 100
100 100 100
100 100 100
76.3 98.3 100
56.3 96.0 98.3
27.8 54.5 71.5
17.8 42.0 46.0
8.0 34.8 48.8
0 19.3 36.8
5.8 19.8 41.3

100 100 100
100 100 100
92.5 95.0 95.0
73.5 100 100
56.3 88.0 95.0
28.8 53.8 73.3
17.0 59.0 83.3
11.3 30.0 43.8
9.3 32.5 57.8

100 100 100
100 100 100
79.3 93.3 100
69.3 82.8 98.0
40.5 58.3 75.0
39.3 57.3 88.3
16.8 35.3 53.0
9.5 21.5 44.0
9.0 14.0 36.0

100 100
100 100
100 100
87.3 91.6
46.1 70.0
23.0 34.0
26.1 45.6
28.2 61.0
12.1 16.6
0 22.0
0 0
0 0
0 0
x x

100 100
100 100
100 100
97.9 100
95.0 97.1
43.3 51.5
27.7 8.0
18.7 12.8
0 0
x x

100 100
100 100
92.6 88.2
100 100
82.2 88.2
31.6 36.7
39.3 60.4
0 0
x x

100 100
100 100
92.2 100
80.0 68.8
51.5 60.9
50.4 81.7
24.8 26.6
8.7 12.5
x x

Treatments with survival equal to the control were not tested the following week.

completion of the 96-h experiment. Exposure to tality than that observed on the water control,
Surround@ WP, Kocide 101 WP, and Aliette re- while mortality on Agrimek 0.15 EC + Citrus
sulted in substantially higherA. vaquitarum mor- Soluble Oil was marginally higher than that on

Ulmer et al.: Pesticide Toxicity to A. vaquitarum

the control. Aprostocetus vaquitarum exposed to
Citrus Soluble Oil, Micromite 80 WGS, Micro-
mite 80 WGS + Citrus Soluble Oil, and Acra-
mite 50 WS had mortality rates marginally
lower than that observed on the control. These
four treatments did not appear to have an effect
onA. vaquitarum survival, thus these treatments
were not subjected to testing two to four weeks af-
ter application. The results were similar when the
pesticides were tested two weeks after pesticide
application. Sevin 80 WSP and Malathion 5 EC
resulted in 10% mortality after 8 h and allA. va-
quitarum were dead 48 h after exposure to Sevin
80 WSP, Malathion 5 EC, and Imidan 70 WSB.
Admire 2 F and Danitol 2.4 EC resulted in 100
and 98% mortality, respectively, after 96 h. Sub-
stantial mortality (72%) was also observed on
Surround@ WP after 96 h. Females exposed to Ko-
cide 101 WP and Agrimek 0.15 EC + Citrus
Soluble Oil showed slightly higher mortality than
those exposed to the water control while those ex-
posed to Aliette had a mortality rate slightly less
than the control. Testing three and four weeks af-
ter pesticide application yielded similar results to
weeks one and two. Sevin 80 WSP remained the
most toxic followed by Malathion 5 EC, Imidan
70 WSB, Admire 2 F, Danitol 2.4 EC, Sur-
round WP and Kocide 101 WP (Table 3). The
relative toxicity of the pesticides was strikingly
consistent over the four week period. Mortality
rates decreased slightly over the four weeks; how-
ever, the toxicity of each pesticide was generally
preserved. Though the pesticide treated substrate
was maintained out of doors, it was protected
from sunlight and precipitation.

Effect of Residual Pesticide on Parasitism

Survival of A. vaquitarum was affected by
Sevin XLR and Imidan 70 WSB applied to foli-
age in the field. Fewer adult A. vaquitarum

emerged from D. abbreviatus eggs laid on foliage
treated with Sevin XLR (0.0) and Imidan 70
WSB (2.3 1.3) than emerged from the water
treated control (17.9 1.9) when host eggs were
laid immediately following pesticide application
(F2,61 = 30.8, P < 0.001) (Table 4). The number of
adults emerging from eggs laid one week after ap-
plication was lower on the Sevin XLR treated fo-
liage than on foliage treated with either Imidan
70 WSB or water (F25, = 10.34, P < 0.001). The
number of adult A. vaquitarum emerging from
host eggs laid two weeks after pesticide applica-
tion was not different among the treatments (F2,76
= 0.54, P = 0.59). The number of D. abbreviatus
larvae emerging from eggs laid immediately after
pesticide application was higher on foliage
treated with Imidan 70 WSB than on either the
Sevin@ XLR or control treatment (F2,61 = 23.9, P <
0.001) (Table 4). There were no differences in the
number ofD. abbreviatus larvae emerging among
the treatments from eggs laid one week (F, 5 =
1.11, P = 0.33) or two weeks after pesticide appli-
cation (F216 = 0.94, P = 0.39).
Significantly more A. vaquitarum adults
emerged from host eggs laid immediately after
pesticide application on foliage treated with Mi-
cromite 80 WGS or Acramite 50 WS than from
the control. Fewer A. vaquitarum emerged from
foliage treated with Micromite 80 WGS + Citrus
Soluble Oil or Citrus Soluble Oil alone (F,1 =
29.3, P < 0.001) (Table 5). The number of A. va-
quitarum emerging from host eggs laid on foliage
treated one (F414 = 13.9, P < 0.001) and two weeks
(F4 14 = 11.7, P < 0.001) earlier was not signifi-
cantly different among Micromite 80 WGS, Ac-
ramite 50 WS, and the control, but significantly
fewer adults emerged from foliage treated with
either Micromite 80 WGS + Citrus Soluble Oil
or Citrus Soluble Oil alone. The number of neo-
nate D. abbreviatus larvae emerging was lower on
foliage treated with Citrus Soluble Oil than on


Mean number ( S.E.)

Treatment Day 0 Day 7 Day 14

AdultA. vaquitarum
Control (water) 17.9 (1.9) a 12.0 (1.9) a 15.6 (2.1) a
Sevin XLR 0.0 (0.0) b 2.1(1.4) b 12.6 (2.3) a
lmidan70WSB 2.3 (1.3) b 13.6 (1.6) a 13.7 (1.7) a
Neonate D. abbreviatus
Control (water) 0.6 (0.4) a 3.4 (2.3) a 0.6 (0.6) a
Sevin XLR 0.8 (0.5) a 4.3 (2.1) a 1.2 (0.6) a
lmidan70WSB 29.6 (5.4) b 0.7 (0.3) a 2.6 (1.5) a

Means followed by the same letter are not significantly different (P = 0.05).

Florida Entomologist 89(1)


Mean number ( S.E.)

Treatment Day 0 Day 7 Day 14

Adult A. vaquitarum
Micromite 80 WGS 22.6 (1.9) a 19.4 (2.9) a 12.9 (1.3) a
Acramite 50 WS 24.8 (3.3) a 20.8 (2.6) a 12.8 (1.8) a
Control (water) 9.3 (1.6) b 20.5 (2.7) a 14.2 (1.9) a
Micromite 80 WGS + Oil 0.8 (0.6) c 3.7 (1.0) b 3.3 (0.7) b
Citrus Soluble Oil 2.1 (1.1) c 1.4 (0.4) b 6.2 (1.8) b
Neonate D. abbreviatus
Micromite 80 WGS 0.0 (0.0) a 0.9 (0.9) a 0.3 (0.3) a
Acramite 50 WS 1.4 (0.7) a 0.3 (0.2) a 2.7 (1.4) a
Control (water) 2.6 (1.3)a 4.5 (2.2) a 2.7 (1.4) a
Micromite 80 WGS+ Oil 0.0 (0.0) a 0.5 (0.5) a 0.0 (0.0) a
Citrus Soluble Oil 23.6 (3.4) b 14.1 (2.5) b 12.8 (2.8) b

Means followed by the same letter are not significantly different (P = 0.05).

any of the other treatments when eggs were laid
immediately (F4,133 = 40.5, P < 0.001), one (F4,145
13.9, P < 0.001), or two weeks (F4145 = 11.7, P <
0.001) after pesticide application (Table 5).

Micromite@ 80 WGS Fecundity Trial

There were no differences between oviposition
by females exposed to Micromite 80 WGS (20.3 +
3.3) and those exposed to a water control (17.0 +
2.8) (F, ,= 0.21, P = 0.65) (Table 6). After 48 h,
45.8% of A. vaquitarum eggs laid by females ex-
posed to Micromite 80 WGS reached the first in-
star while 64.9% of eggs laid by females exposed
to a water control reached the first instar after 48
h. There was no difference in the number of new
generation A. vaquitarum adults emerging be-
tween Micromite@ 80 WGS treated females (13.4


Treatment Control Micromite 80 WGS

Eggs 6.0 (1.2) a 11.0 (1.8) a
Larvae 11.0 (1.9) a 9.3 (2.4) a
Total Oviposition 17.0 (2.8) a 20.3 (3.3) a
Host eggs 185.5 (15.6) a 168.5 (11.8) a
Adult Offspring 11.2 (1.0) a 13.4 (2.5) a
Host Eggs 167.9 (6.4) a 171.4 (7.8) a

Means followed by the same letter are not significantly dif-
ferent (P =0.05).

2.5) and those females exposed to a water con-
trol (11.2 1.0) (F,,56 = 0.34, P = 0.56). The number
of host D. abbreviatus eggs was not different be-
tween the treatments for either the dissected egg
masses (F1 = 0.98, P = 0.33) or those reared to
emergence (F ,56 = 0.29, P = 0.59).


The impact of the 12 pesticides tested against
A. vaquitarum ranged from harmless to highly
toxic. The most acutely toxic products tested were
Sevin@ 80 WSP carbamatee), Malathion, and Im-
idan 70 WSB (organophosphates). When tested
immediately after application, mortality was
more than twice as rapid as the next most toxic
pesticides. Admire@ 2 F (neonicotinoid) and Dan-
itol@ 2.4 EC (pyrethroid) were also toxic toA. va-
quitarum. These and many other neurotoxic in-
secticides used in citrus are known to be ex-
tremely detrimental to a range of hymenopteran
parasitoids and other beneficial insects (Easwar-
amoorthy et al. 1990; Villanueva-Jimenez & Hoy
1998; Jacas & Garcia-Marf 2001; Wakgari &
Giliomee 2003; Michaud & Grant 2003), including
Aprostocetus ceroplastae (Girault) (Wakgari &
Giliomee 2001). The regular use of these products
will impede the establishment and productivity of
A. vaquitarum and almost certainly have a nega-
tive impact on various other beneficial insects.
Surround@ WP (kaolin clay) is touted as non-
toxic and IPM-compatible. Though not as toxic as
the neurotoxic insecticides, Surround@ WP in-
creased mortality and reduced the longevity ofA.
vaquitarum compared to the control and the other
less harmful pesticides tested. The cause of death
was not clear in the present study, but cadavers

March 2006

Ulmer et al.: Pesticide Toxicity to A. vaquitarum

were often observed covered in kaolin particles.
Application of Surround WP to citrus has also
been observed to increase scale insect infestations,
most likely due to interference with parasitism
(S. L. Lapointe, unpublished). The effect of Sur-
round@ WP in the present experiment may have
been magnified by the exclusion of precipitation
from the weathering process. Under these condi-
tions, the Surround WP residue would be ex-
pected to remain intact and not decline in potency.
Aprostocetus vaquitarum females exposed to
Agrimek 0.15 EC (avermectin), Kocide 101 WP
(copper hydroxide) and Aliette WDG showed a
slight increase in mortality but longevity was not
significantly different than that observed on the
control. These products appear to be compatible
with this parasitoid. However, at recommended
rates Kocide 101 WP and Agrimek@ 0.15 EC
were shown to be detrimental toAgeniaspis citri-
cola (Logvinovskaya) (Hymenoptera: Encyrti-
dae), a parasitoid of the citrus leafminer (Phylloc-
nistis citrella (Stainton) (Lepidoptera: Gracillari-
idae) (Villanueva-Jimenez & Hoy 1998). Aver-
mectin can affect citrus predatory mites
negatively, including Euseius stipulatus (Athias-
Henriot) (Jacas & Garcia Marf 2001). Thus, fur-
ther study of the effects on other beneficial insects
in citrus is warranted.
Three of the products tested showed no contact
toxicity to adult females, including Acramite 50
WS, Micromite 80 WGS, Citrus Soluble Oil, and
Micromite 80 WGS + Citrus Soluble Oil. Acra-
mite 50 WS (bifenazate) also has been shown to
be compatible with the predatory mite Phytoseiu-
lus persimilis (Kim & Yoo 2002) and only moder-
ately harmful to ladybird beetles (James & Coyle
2001). Micromite 80 WGS (diflubenzuron) is
known to be toxic to several parasitoid species
(Zaki & Gesraha 1987; Zijp & Blommers 2001;
Amalin et al. 2004; Schneider et al. 2004), while it
appears relatively harmless to others (Willrich &
Boethel 2001; Amalin et al. 2004). Citrus Soluble
Oil used alone is known to be compatible with
other parasitoids and beneficial predators in cit-
rus production (Amalin et al. 2000; Villanueva-
Jimenez et al. 2000).
Results of the field studies were consistent
with the laboratory tests. Sevin XLR sprayed on
host plant material was extremely toxic, resulting
in no parasitoid reproduction when D. abbrevia-
tus eggs were laid immediately after spraying,
and continued to significantly reduce A. vaquita-
rum populations one week after application. It
was not until 14 d after treatment that Sevin
XLR no longer had a significant impact onA. ua-
quitarum. Imidan 70 WSB significantly reduced
A. vaquitarum reproduction but lost efficacy more
quickly than Sevin XLR and no longer reduced
A. vaquitarum reproduction after one week in the
field. Very low numbers of neonate D. abbreviatus
closed on the control, due to high levels of para-

sitism, or on foliage treated with Sevin XLR,
due to its toxicity to both the parasitoid and the
pest. However, treatment with Imidan 70 WSB
resulted in high mortality of the beneficial but not
D. abbreviatus. Though Imidan 70 WSB broke
down relatively quickly in the field, its negative
impact on A. vaquitarum and minimal effect on
D. abbreviatus make it a very poor candidate for
an IPM program.
Insect growth regulators such as Micromite
80 WGS (diflubenzuron) have generally been con-
sidered environmentally safer alternatives to
broad-spectrum insecticides. Diflubenzuron has
been shown to have a minimal impact on some hy-
menopteran parasitoids (Villanueva-Jimenez &
Hoy 1998; Willrich & Boethel 2001; Amalin et al.
2004). However, it is known to have devastating
effects on other parasitoid species (Zaki & Ges-
raha 1987; Zijp & Blommers 2001; Amalin et al.
2004; Schneider et al. 2004). Many pesticide tox-
icity tests are carried out on only one developmen-
tal stage of the parasitoid, usually the adults, but
Schneider et al. (2003a, 2003b) showed that di-
flubenzuron can have detrimental effects on de-
velopmental processes while appearing relatively
harmless to adult parasitoids. In the present
study, Micromite 80 WGS was not toxic to adult
A. vaquitarum but was further tested under field
conditions and in the laboratory to assess oviposi-
tion and development from egg to adult. Consis-
tent with the contact toxicity test with adult fe-
males, Micromite 80 WGS had no negative af-
fect onA. vaquitarum when host eggs were laid on
field treated host plants and tested 0 to 14 d after
application. Oviposition by females exposed to
Micromite 80 WGS in the laboratory was not
significantly different from the control, though
there was an indication that the time from ovipo-
sition to first-instar eclosure may be extended for
eggs laid by Micromite 80 WGS exposed fe-
males. If detrimental, diflubenzuron, a chitin syn-
thesis inhibitor, often inhibits egg hatch and dis-
rupts the molting process (Marx 1977; Weiland et
al. 2002). In the present study there were no ap-
parent differences in development from egg to
adult between the Micromite 80 WGS treatment
and the water control.
Micromite 80 WGS (diflubenzuron) is effec-
tive in reducing D. abbreviatus populations
(Schroeder 1996) and had no affect onA. vaquita-
rum in the present study. Micromite 80 WGS
use appears to be compatible with both A. va-
quitarum and Quadrastichus haitiensis (Gahan)
(Amalin et al. 2004), two of the primary parasi-
toids of D. abbreviatus now established in south
Florida. Diflubenzuron was also considered
harmless for citrus predatory mites (Jacas &
Garcia Marf 2001). However, diflubenzuron was
shown to adversely affect Ceratogramma eteinnei
(Delvare) (Amalin et al. 2004), an endoparasitoid
ofD. abbreviatus which was released and failed to

Florida Entomologist 89(1)

establish in south Florida, as well as several
other hymenopteran parasitoids (Zaki & Gesraha
1987; Zijp & Blommers 2001; Amalin et al. 2004;
Schneider et al. 2004). Its impact on the agroeco-
system merits further study.
Though Micromite@ 80 WGS alone had no neg-
ative impact on A. vaquitarum, Micromite 80
WGS + Citrus Soluble Oil and Citrus Soluble Oil
alone did significantly reduce A. vaquitarum re-
production. It appears that the D. abbreviatus egg
masses laid between leaves treated with Citrus
Soluble Oil do not remain closed and become ex-
posed to the elements. Aprostocetus vaquitarum
will not oviposit into an exposed egg mass and if
oviposition occurs before the egg mass opens, par-
asitoid eggs and larvae desiccate and die when the
egg mass opens (J. E. Pefia, unpublished). Though
Citrus Soluble Oil does not appear to be toxic to
A. vaquitarum, it does indirectly affect this para-
sitoid by reducing the efficiency of the adhesive
D. abbreviatus uses to secure and protect its eggs
between two leaves. Citrus Soluble Oil treatments
had a negative affect on C. eteinnei (Amalin et al.
2004). Aprostocetus vaquitarum, an ecto-parasi-
toid, is extremely vulnerable to the environment
and possibly even more severely impacted when
the host egg mass is opened and exposed.
The aim of this study was to evaluate the rela-
tive toxicity of citrus pesticides in an effort to pro-
mote the use of compounds with low toxicity lev-
els to A. vaquitarum and other beneficial insects.
At recommended rates, Sevin@ 80 WSP,
Malathion 5 EC, and Imidan 70 WSB, were ex-
tremely toxic to A. vaquitarum and will discour-
age the establishment of this insect. Admire@ 2F,
Danitol@ 2.4 EC, and Surround@ WP were also
moderately toxic to A. vaquitarum and regular
use would be detrimental to a control program
aimed at establishing and maintaining this para-
sitoid. Kocide 101 WP, Aliette WDG, and
Agrimek@ 0.15 EC + Citrus Soluble Oil were rel-
atively non-toxic. Micromite 80 WGS, Acra-
mite@ 50 WS, and Citrus Soluble Oil were non-
toxic to A. vaquitarum adults and these products
appear to be very suitable for an IPM program.
Micromite 80 WGS was also shown not to dis-
rupt development ofA. vaquitarum while Citrus
Soluble Oil, though not toxic, did reduce the suc-
cess ofA. vaquitarum by causing host egg masses
to become exposed. Given the restricted field eval-
uations conducted in the present study, further
research should focus on the impact of these pes-
ticides under field conditions.


We are grateful to J. Jacas (Universitat Jaume I) and
C. Mannion (University of Florida) for critical review of
the manuscript. Special thanks are extended to J.
Alegria, Z. Alegria, and D. Long (University of Florida)
for providing technical assistance throughout the inves-

tigation. The research was supported by University of
Florida-Institute of Food and Agricultural Sciences,
United States Department of Agriculture, and Coopera-
tive State Research Education and Extension Service.
Florida Agricultural Experiment Station Journal Series


ABBOTT, W. S. 1925. A method of computing the effective-
ness of an insecticide. J. Econ. Entomol. 18: 265-267.
2000. Selective toxicity of some pesticides to Hibana
velox (Araneae: Anyphanenidae), a predator of citrus
leafminer. Florida Entomol. 83: 254-262.
AMALIN, D. M., P. STANSLY, AND J. E. PENA. 2004. Effect
of Micromite@ on the egg parasitoids Ceratogramma
etiennei (Hymenoptera: Trichogrammatidae) and
Quadrasticus haitiensis (Hymenoptera: Eulophidae).
Florida Entomol. 87: 222-224.
BARBOSA, P., AND J. C. SCHULTZ. 1987. Insect Outbreaks.
Academic Press, Inc., San Diego, CA. pp. 291-292.
K. KURUP. 1990. Toxicity of certain insecticides to
Sturmiopsis inferens, a larval parasite of sugarcane
moth borers. Entomophaga. 35: 385-391.
JACAS, J. A. AND F. GARCIA-MARI. 2001. Side-effects of
pesticides on selected natural enemies occurring in
citrus in Spain. IOBC Bull. 24: 103-112.
JACAS, J. A., J. E. PENA, AND R. E. DUNCAN. 2005. Suc-
cessful oviposition and reproductive biology of
Aprostocetus uaquitarum (Hymenoptera: Eulophi-
dae): A predator of Diaprepes abbreviatus (Coleop-
tera: Curculionidae). Biol. Contr. (In press).
JAMES, D. G., AND J. L. COYLE. 2001. Which Pesticides
are safe to beneficial insects and mites? Agrichemi-
cal Environmental News. Feb. 2001. Issue No. 178.
KIM, S. S., AND S. S. Yoo. 2002. Comparative toxicity of
some acaricides to the predatory mite, Phytoseiulus
persimilis and the twospotted spider mite, Tetrany-
chus urticae. BioControl. 47: 563-573.
MARX, J. L. 1977. Chitin synthesis inhibitors; new class
of insecticides. Science. 197: 1170-1172.
1995. Effect of surface-applied and soil-incorporated
insecticides for the control of neonate larvae of Di-
aprepes abbreviatus in container-grown citrus. Proc.
Florida State Hort. Soc. 108: 130-136.
MICHAUD, J. P., AND A. K. GRANT. 2003. IPM-compati-
bility of foliar insecticides for citrus: Indices derived
from toxicity to beneficial insects from four orders. J.
Insect Sci. 3: 1-10.
CAN, AND A. HOYTE. 2005. Recovery of parasitoids
(Hymenoptera: Eulophidae and Trichogramma-
tidae) released for biological control of Diaprepes ab-
breviatus (Coleoptera: Curculionidae) in Florida.
Proc. Int. Citrus Congress (In press).
VINUELA. 2004. Action of insect growth regulator in-
secticides and spinosad on life history parameters
and absorption in third-instar larvae of the endopar-
asitoid Hyposoter didymator. Biol. Contr. 31: 189-198.
2003a. Toxicity and pharmacokinetics of insect

March 2006

Ulmer et al.: Pesticide Toxicity to A. vaquitarum

growth regulators and other novel insecticides on
pupae of Hyposoter didymator (Hymenoptera: Ich-
neumonidae), a parasitoid of early larval instars of
lepidopteran pests. J. Econ. Entomol. 96: 1054-1065.
Susceptibility of Hyposoter didymator (Hymenop-
tera: Ichneumondiae) adults to several IGR's pesti-
cides and spinosad by different exposure methods.
IOBC/wprs Bull. 26: 111-112.
SCHROEDER, W. J. 1996. Diflubenzuron residue: reduc-
tion of Diaprepes abbreviatus (Coleoptera: Curcu-
lionidae) neonates. Florida Entomol 79: 462-463.
ADAIR 1996. Diaprepes abbreviatus (Coleoptera:
Curculionidae): Host plant associations. Environ.
Entomol. 25: 333-349.
STANLEY, D. 1996. Suppressing a serious citrus pest.
Agric. Res. 44: 22.
8 User's Manual. Tallahassee Florida: Analytical
Software. ISBN 1-881789-06-3.
Florida citrus pest management Guide. University
of Florida Cooperative Extension Service, Institute
of Food and Agricultural Sciences. SP-43. Download
at: http://edis.ifas.ufl.edu
icity of pesticides to the citrus leafminer and its par-
asitoidAgeniaspis citricola evaluated to assess their
suitability for an IPM program in citrus nurseries.
BioControl. 43: 357-388.
DAVIES. 2000. Field evaluation of integrated pest man-
agement-compatible pesticides for the citrus leafminer
Phyllocnistis citrella (Lepidoptera: Gracillariidae) and
its parasitoid Ageniaspis citricola (Hymenoptera: En-
cyrtidae). J. Econ. Entomol. 93: 357-367.

WAKGARI, W., AND J. GILIOMEE. 2001. Effects of some
conventional insecticides and insect growth regula-
tors on different phenological stages of the white
wax scale, Ceroplastes destructor Newstead (Hemi-
ptera: Coccidae), and its primary parasitoid, Apro-
stocetus ceroplastae (Girault) (Hymenoptera:
Eulophidae). Int. J. Pest. Mang. 47: 179-184.
WAKGARI, W., AND J. GILIOMEE. 2003. Natural enemies
of three mealybug species (Hemiptera: Pseudococ-
cidae) found on citrus and effects of some insecti-
cides on the mealybug parasitoid Coccidoxenoides
peregrinus (Hymenoptera: Encyrtidae) in South Af-
rica. Bull. Entomol. Res. 93: 243-254.
CURT. 2002. A literature review and new observa-
tions on the use of diflubenzuron for control of
locusts and grasshoppers throughout the world. J.
Orthoptera Res. 11: 43-54.
WILLRICH, M. W., AND D. J. BOETHEL. 2001. Effects of
diflubenzuron on Pseudoplusia includes (Lepi-
doptera: Noctuidae) and its parasitoid Copidosoma
floridanum (Hymenoptera: Encyrtidae). Biol. Con-
trol. 30: 794-797.
WOODRUFF, R. E. 1964. A Puerto Rican weevil new to
the United States (Coleoptera: Curculionidae). Flor-
ida Dept. Agr., Div. Plant Ind., Entomol. Circ. 30: 1-2.
ZAKI, N., AND M. A. GESRAHA. 1987. Evaluation of zertel
and diflubenzuron on biological aspects of the egg
parasitoid, Trichogramma evanescens Westw. and
the aphid lion Chrysoperla carnea Steph. J. Appl.
Entomol. 104: 63-69.
ZIJP, J. P., AND L. H. M. BLOMMERS. 2001. The effect of
diflubenzuron on parasitism of Anthonomus po-
morum by Centistes delusorius. Entomol. exp. &
Appl. 98: 115-118.

Florida Entomologist 89(1)

March 2006


'University of Florida, Ft. Lauderdale Research & Education Center
3205 College Ave., Ft. Lauderdale, FL 33314

2Dow AgroSciences, 7257 NW 4th Blvd, #20, Gainesville, FL 32607


Four buildings (two high-rise condominiums, a single-family residential structure, and a
trailer) in Broward and Miami-Dade Counties, Florida, infested with Formosan subterra-
nean termites (FST, Coptotermes fomosanus Shiraki) were treated with baits containing
0.5% wt/wt noviflumuron. Each building represented a challenging treatment scenario for
liquid termiticides due to the location of the infestation within the structure and/or occupant
refusal to permit pesticide application in termite-infested living and activity areas. Marking
of FST by in-situ baiting with blank bait matrix treated with 0.5% wt/wt Neutral Red dye in-
dicated only one FST foraging population infested each building. Two FST infestations were
aerial in high-rise condominiums. Noviflumuron baits were applied to two buildings in
aboveground stations, one building with in-ground stations, and the remaining building
with both station types. All detected FST infestations were eliminated within 71-92 days af-
ter first application of noviflumuron baits. FST foraging populations with confirmed ground
contact consumed approximately 4-fold more bait than did aerial infestations; mean + SD,
242 74 g vs. 62 51 g, respectively. Termite feeding activity was monitored before, during,
and after bait application at two buildings with an acoustic emissions detector (AED) and in
one building with a microwave detector. Cessation of termite activity measured with these
devices corresponded with elimination of live FST previously observed in stations, infested
wood, and foraging tubes. No FST were observed in any monitoring station or building dur-
ing the 12-18 month inspection period following elimination of the detected FST infestation.

Key Words: SentriconTM System, RecruitTM termite bait, noviflumuron, chitin synthesis
inhibitor, Coptotermes formosanus


Cuatro edificios (dos condominios altos, una estructura residential para una sola familiar, y
una casa trailer en los Condados de Broward y de Miami-Dade, Florida, infestadas con la ter-
mita subterranea de Formosa (FST, Coptotermes fomosanus Shiraki) fueron tratadas con ce-
bos que contienen 0.5% peso/peso de noviflumuron. Cada edificio represent un desafio para
el tratamiento con los termiticidas liquidos debido al lugar de la infestaci6n dentro la estruc-
tura y/o el rechazo del occupante para permitir la aplicaci6n de un pesticide en areas infes-
tadas de sus hogares donde ellos tienen sus actividades. Al marcar los FST in-situ con una
matriz de cebo vacio tratado con 0.5% peso/peso de tinta Roja Neutral indico una sola pobla-
ci6n de FST forrajera infestando cada edificio. Dos infestaciones dereas de FST fueron pre-
sentes en los condominios. Se aplicaron cebos de Noviflumuron en puestos encima de la
superficie en dos edificios, en puestos subterraneos en un edificio, y en puestos de las dos cla-
ses en los otros edificios. Todas las infestaciones de FST detectadas fueron eliminadas dentro
de 71-92 dias despu6s de la primera aplicaci6n de los cebos de Noviflumuron. Las poblaciones
forrajeras de FST con contact confirmado con la tierra consumieron aproximadamente 4 ve-
ces mas cebo que las infestaciones dereas; con un promedio DS de 242 74 g vs. 62 51 g,
respectivemente. Se realize un monitoreo de la actividad de la alimentaci6n de las termitas
antes, durante y despu6s de la aplicaci6n de cebo en dos edificios con un detector de emissio-
nes acistico (AED) y en un edificio con un detector de microhonda. La paralizaci6n de la ac-
tividad de las termitas media con estos aparatos correspondia con la eliminaci6n de los FST
vivos observados anteriormente en los puestos de studio, madera infestada, y en los tubos
de forrajeo. Ningun FST fue observada en cualquier puesto o edificio durante el period de
inspecci6n de 12-18 meses despu6s de la eliminaci6n de la infestaci6n detectada de FST.

Hexaflumuron, a chitin synthesis inhibitor, in Reticulitermes flavipes (Kollar) and Coptoter-
was documented in the late 1980s to cause mes formosanus Shiraki (Su & Scheffrahn 1993).
ecdysis inhibition resulting in delayed mortality Since that discovery, many field studies have ver-

Cabrera & Thoms: Control of Formosan Subterranean Termites

ified that termite baits containing hexaflumuron
eliminated colonies or populations of 15 species of
subterranean termites in 15 states in the US, and
in Australia, Japan, Malaysia, Taiwan, France,
England, Italy, Cayman Islands, Puerto Rico, and
US Virgin Islands (Lee 2002; Sajap et al. 2002;
Su 2002a, b; Su et al. 2002; Su et al. 2003; Su &
Hsu 2003).
In 1995, Dow AgroSciences developed the
chitin synthesis inhibitor, noviflumuron, and be-
gan trials to assess its toxicity against common
household and structural pests, including subter-
ranean termites (Smith et al. 2001). In laboratory
trials in which R. flavipes were fed radiolabeled
noviflumuron or hexaflumuron, noviflumuron
demonstrated significantly faster speed of action,
greater potency, and nearly 4-fold slower clear-
ance from termites compared with that of hexaflu-
muron (Sheets et al. 2000; Karr et al. 2004). In
field trials conducted throughout the US from
1998-2000, 74 colonies of Reticulitermes spp.
baited with noviflumuron were eliminated in
about half the time as the 53 colonies baited with
hexaflumuron (mean = 107 days vs. 205 days, re-
spectively, Smith et al. 2001). Sajap et al. (2005)
eliminated five structural infestations of Coptoter-
mes gestroi (Wasmann) within 35-56 d with above-
ground (AG) baits containing 0.5% noviflumuron.
The purpose of the trials described in this
study was to determine if bait containing 0.5%
noviflumuron could eliminate structural infesta-
tions of FST. Although not a selection factor, each
building in this study represented a challenging
treatment scenario for liquid termiticides due to
the location of the infestation within the struc-
ture (dispersed in the upper story of high-rise con-
dominiums or in a difficult to access crawlspace)
and/or occupant refusal to permit pesticide appli-
cation in critical areas.


Termite Bait/Monitoring Stations

In-ground (IG) or AG stations used in the Sen-
tricon System (Dow AgroSciences, Indianapolis,
IN) were used to monitor termite activity and ap-
ply noviflumuron baits. IG stations, as described
by Su et al. (2002), contained monitoring devices
(two 1.4 x 2.8 x 17.5 cm wood slats, Fig. 1) when
first installed. To apply the bait, the wood moni-
toring devices were replaced by a bait tube con-
taining 0.5% wt/wt noviflumuron on a 35-g roll of
textured laminated cellulose.
An AG station consisted of a rigid, rectangular
plastic housing (14.8 cm long x 9 cm wide x 5 cm
deep) containing two 35-g laminated textured cel-
lulose rolls (Fig. 2). One side of the housing was
open to expose the matrix to the termite-infested
substrate; the other side was closed by a remov-
able lid. Blank bait containing no active ingredi-

Fig. 1. In-ground (IG) station for applying wood mon-
itoring devices (shown here) or bait tube containing 35-
g roll of laminated cellulose matrix; blank or impreg-
nated with 0.5% wt/wt noviflumuron.

ent was used to monitor termite activity in both
IG and AG stations.

Study Sites

Noviflumuron baits were evaluated at three
buildings in Broward County and one in Miami-
Dade County, FL that had active subterranean
termite infestations. Soldiers and/or alates were
collected and identified as FST with the key by
Scheffrahn (1994).
DAYCARE was a 195.2-m2, single-story, con-
crete block, slab-on-grade house in Hollywood
Hills, FL, converted to a daycare center for in-
fants and preschool children. In mid-May, 2001,
daycare employees reported that massive emer-
gence of FST alates forced them to evacuate the
daycare center and relocate the children. Subse-
quently, on 18 May, 2001, the first author found
live FST in a backyard mango tree (Fig. 3) and in
the garage in an aerial carton nest located be-
tween pieces of plywood propped over the expan-
sion joint abutting the house slab. The nest was
removed to expose many FST mud foraging tubes
emerging from the expansion joint. FST damage,

Florida Entomologist 89(1)

Fig. 2. Slicing bait of aboveground (AG) station con-
taining two 35-g rolls of laminated cellulose matrix;
blank or impregnated with 0.5% wt/wt noviflumuron.

including alate release slits, was found on the
garage rear entrance doorframe and the nearby
exposed header in the breezeway (Fig. 3). DAY-
CARE had not been treated with liquid termiti-
cides due in part to concerns of the staff about
pesticide exposure.






TRAILER was a small, wood frame, 40-m2
trailer with aluminum siding and a crawlspace. A
33.2-m2 addition consisting of three rooms (stor-
age, sewing, and entry area) and a porch on a
raised concrete slab were attached to the trailer
(Fig 4). According to detailed records maintained
by the owner, FST alate flights occurred in the
front rooms or bedroom between approximately
6:00 pm and 12:30 am on 48 different days from
May 27 through August 1, 2002. On June 5, 2002,
a pest control company applied PremiseTM ter-
miticide, containing 0.05% imidacloprid (Bayer
Environmental Science, Montvale, NJ), around
the trailer perimeter and in the crawlspace ac-
cording to the company service report and owner
observations. No evidence was found that the hol-
low block foundation supporting the raised con-
crete slab was drilled and treated with termiti-
cide. Access to this foundation through the crawl-
space was very restricted. During an inspection
on June 28, 2002, the first author found dead FST
alates in the bedroom and storage room, and FST
alate release slits in the wall paneling of the bed-
room and living room (Fig. 4). No evidence of live
FST was found in the crawlspace.


o o

I oooe ooo ...-- -
Ss O AY IG Inactive Monitor Station
0 IG Baited Station
SGAAGE I IG Active Monitor Station
EG AG Baited Station
Blank Station



SP ter box
o~f~o r^-

Fig. 3. Ground plan of DAYCARE showing placement of in-ground and aboveground stations and 0.5% noviflu-
muron bait. Gray shaded area indicates population foraging area based on presence of termites in the mango tree,
dyed termites in bait stations, and AED readings.

March 2006

n fR OD (-1

Cabrera & Thoms: Control of Formosan Subterranean Termites

AG Blank Active Station
AG Baited Station
~ Area with visible FST damagelalate emergence holes


Fig. 4. Floor plan of TRAILER showing placement of aboveground stations after inspection on October 25, 2002,
0.5% noviflumuron bait, and areas damaged by Formosan subterranean termites. Gray shaded area indicates pop-
ulation foraging area based on presence of dyed termites in bait stations.

CONDO was an eight-story, reinforced con-
crete, multi-unit residential building in Halland-
ale, FL. On April 19, 2002, the first author found
FST infesting extensive foraging tubes extending
up the west stairwell onto the 8th floor landing and
attic. There was no evidence of termite damage or
activity on the ground floor or around the building
perimeter. In mid-May, 2002, a husband and wife
reported FST infesting their 8th floor condomin-
ium adjacent to the west stairwell. Because the
wife was 6-mo pregnant and her infirm mother
was about to be moved into their condominium,
the occupants would not permit any pesticide
application inside their condominium. Subse-
quently, on May 30, 2002, in this condominium
the authors found termite damage and carton ma-
terial in the doorframes of the two bathrooms and
the second bedroom and in the master bedroom
closet wall next to the west stairwell, where ter-
mites remained active (Fig. 5).
PENTHOUSE was a 29-story, reinforced con-
crete, multi-unit residential building in Aven-
tura, FL. On November 1, 2002, the first author
found live FST infesting a palm tree stump and in
the soil around the base of another palm stump
inside a planter, and in foraging tubes and several
joists on the underside of the raised, wood patio
deck (Fig. 6). No termite damage was reported by
any other occupants or found indoors in the rooms
directly below the infested patio. FST have been

documented previously to establish aerial infes-
tations on flat rooftops of high-rise buildings sim-
ilar to CONDO and PENTHOUSE (Su et al. 1989;
Weissling & Thoms 1999).

Installation of Monitoring Stations

In-ground (IG) stations were installed follow-
ing the label directions for Recruit IIM termite
bait. Stations were installed around the perime-
ter of DAYCARE (Fig. 3) and TRAILER (Fig. 4)
more than 45 cm from the foundation to avoid
placement in soil previously treated with termiti-
cide. Spacing between stations did not exceed 6 m
where soil access was not restricted by driveways
and concrete patios (Fig. 3). Stations also were in-
stalled adjacent to areas with visible termite ac-
tivity, such as in the rooftop planter box of PENT-
HOUSE (Fig. 6). After initial IG station installa-
tion, one or two "auxiliary" IG stations were in-
stalled within 30 cm of each IG station with
termite activity if space permitted.
Aboveground (AG) stations containing blank
bait rolls were installed where live FST were vis-
ible in surface foraging tubes (DAYCARE expan-
sion joint, CONDO stairwell, and PENTHOUSE
deck joist) and damaged wood (TRAILER parti-
tion wall and PENTHOUSE palm stump, Figs. 3-
6). AG stations were installed adjacent to where
acoustic emissions detector (AED) counts indi-


Florida Entomologist 89(1)




Living Room

M AG Blank Active Station
AG Baited Station








IIjj-l IS

Fig. 5. Floor plan of CONDO showing placement of in-ground and aboveground stations, 0.5% noviflumuron
bait, and area monitored with an acoustic emissions detector. Gray shaded area indicates population foraging area
based on presence of dyed termites in bait stations and AED readings.

cated termite feeding but FST were not visible
(DAYCARE breezeway header and CONDO door
frames). In these locations, 2-mm diam holes were
drilled into the wood before attaching each AG
station to provide the termites access to the bait.
Each AG bait roll was sliced several times longi-
tudinally (Fig. 2) and then moistened with 30-60 ml
of water (DAYCARE), 5% sucrose water solution
drink (TRAILER; GatoradeTM, Gatorade Corp.,
Chicago, IL). The sliced bait rolls were placed di-
rectly in contact with the FST-infested surface. The
station housing was attached to the surface with
screws for wood surfaces and latex caulk (Poly-
seamsealTM, OSI, Mentor, OH). The removable lid
was sealed to the housing with screws and masking
tape. All IG and AG stations were numbered and
their locations were documented on site maps.

Use of Electronic Termite Detection Devices
At DAYCARE and CONDO, an Acoustic Emis-
sion Detector (AED, LocatorTM Insect Detection
Device, Dow AgroSciences, Indianapolis, IN) was
used to monitor FST feeding activity in exposed
wood. The AED has been documented to be a sim-

ple, non-disruptive method to quantify subterra-
nean termite feeding activity (Scheffrahn et al.
1993) before and after application of termite baits
(Su et al. 2000; Su et al. 2002; Su et al. 2003;
Weissling & Thoms 1999).
Thirteen locations on one doorframe and the
breezeway header at DAYCARE (Fig. 3) and 27 lo-
cations on three doorframes at CONDO (Fig. 5)
were monitored with an AED. The sensors were
attached by putty (HanditakTM, SuperGlue Corp.,
Hollis, NY) to monitored wood members. Each
AED location was marked and monitored once for
30 s in the noise reduction setting during each
visit before, during, and after bait application.
Based on user experience, readings of less than 5
counts usually do not indicate termite activity.
Readings were repeated if sensor movement or
detachment occurred. The last AED monitoring
was conducted at 16.5 mo (DAYCARE) and 1 mo
(CONDO) after elimination of all detectable FST
activity. Acoustic detection was not continued at
CONDO because damaged doorframes were re-
placed by the condominium owners after elimina-
tion of all detectable FST activity.
A microwave detector (TermaTracTM Archer-
field, Queensland, Australia) was used to detect

March 2006

Cabrera & Thoms: Control of Formosan Subterranean Termites

Fig. 6. Floor plan of PENTHOUSE showing placement of in-ground and aboveground stations and 0.5% noviflu-
muron bait. Gray shaded area indicates population foraging area based on presence of dyed termites in bait stations.

FST activity at TRAILER. Evans (2002) verified
that the TermaTrac had a 90% success rate for de-
tecting subterranean termites. The detector was
used to locate FST in and near areas of the wall
and floor of the trailer with visible termite dam-
age. For wall paneling, the sensor horn was held
flush against the surface while taking readings
for 30 to 60 s. On the floor, the horn was placed up-
right directly on the surface for the same amount
of time. The detector was used again on the same
areas of the floor one year after elimination.

Bait tube Installation in IG Stations

At least 20 ml of water was used to moisten the
matrix (blank, dyed or with noviflumuron) in a
bait tube before installation. Termites from in-
fested wood monitors or bait tube were gently
transferred to the chamber in the top of the bait
tube. The new bait tube was placed in the same
station from which the termites were extracted.

Replenishment of AG Stations

Dry AG bait was remoistened with 30-60 ml of
water on subsequent inspections after installa-

tion. If one third to one half of the matrix was con-
sumed, one additional AG station with moistened
matrix (blank, dyed or with noviflumuron) could
be attached on top of the existing station with
screws and masking tape.

Delineating Termite Populations with Dye Markers

When live termites were found in two or more
IG and/or AG stations at a test site, the bait dyed
with a 0.5% wt/wt Neutral Red in a bait tube or
AG station was installed as previously described.
On subsequent inspections, stations containing
dyed termites were documented, classified as be-
ing connected, and the termites were considered
to be from the same foraging population. Dyed
matrix was introduced into additional stations
containing dyed termites if the intensity of the
color in dyed termites or the number of dyed ter-
mites was very low.

Application of Noviflumuron Baits

Application of noviflumuron bait began when
two or more IG and/or AG stations with live ter-
mites were found to be connected based upon the

Florida Entomologist 89(1)

dye marker. Noviflumuron baits were not applied
in at least one connected station with live ter-
mites at each site to provide an independent mon-
itor for termite activity. Only blank baits were ap-
plied in FST-infested stations located within the
children's play yard and breezeway at DAYCARE
(Fig. 3) and in the condominium at CONDO (Fig.
5), due to occupants' requests not to have pesti-
cides applied in these areas. By three mo after
termite activity had ceased in all stations and in
previously detected infested locations within a
building, all noviflumuron baits were replaced
with wood monitors in IG stations and blank bait
in AG stations.

Station Inspections

Stations were inspected approximately every
1-4 weeks before and during application of novi-
flumuron. Wood monitors or baits that were
moldy, degraded, or more than 50% consumed
were replaced. The station type (IG or AG), type of
device in station (monitor, dye, active ingredient),
station device action (new, inspected, or replaced),
estimated consumption of device, presence or ab-
sence of termites, presence of dyed termites, and
estimated number of termites were recorded. The
amount of wood and bait consumed was calcu-
lated by multiplying the total estimated percent-
age of the device consumed by the known mean,
dry weight of whole devices: 62 g for wood moni-
tor, 35 g for IG bait roll and 70 g for AG bait roll
(Tables 1 and 2; DeMark & Thomas 2000;
Weissling & Thoms 1999).
Stations were inspected approximately every 3
mo for at least one year following removal of all
noviflumuron bait. During the final site inspec-
tion, the previously infested building or condo-
minium unit and landscape features, such as the
mango tree at DAYCARE, were visually inspected
for evidence of any new termite activity.


Dye marking indicated only one FST foraging
population infested each building. Dyed termites
were found in all IG and AG stations with termite
activity at each site. The cessation of termite ac-
tivity detected in foraging tubes, infested wood,
and by electronic monitoring following applica-
tion of noviflumuron in stations where dyed ter-
mites were found further indicate only one FST
foraging population infested each building. Huss-
eneder et al. (2003) demonstrated FST in New Or-
leans collected within a foraging territory delin-
eated by mark-recapture were genetically similar
and could be genetically differentiated from ter-
mites in adjacent territories.
Stations were rapidly infested by FST, so bait-
ing with noviflumuron began within 2-6 weeks af-
ter station installation, except at TRAILER. At
this site, 22 IG stations were installed on July 8,
2002, 17 IG stations were installed on July 26,
2002 along the sides of two neighboring trailers,
and three AG stations containing blank bait were
installed in the bedroom on August 2, 2002. These
stations were inspected through September 30,
2002, but no FST activity was detected in any sta-
tion. On October 2, 2002, the first author con-
ducted a thorough structural inspection, includ-
ing the crawlspace, with a flashlight, probing tool,
and the TermatracTM, but no FST were seen or de-
tected. Subsequently, all stations were removed
At the request of the property owner, the first
author visually re-inspected TRAILER on Octo-
ber 25, 2002. Live FST were found in a low-rise,
wood-paneled partition separating the kitchen
and entry areas (Fig. 4) along the juncture of the
trailer and raised slab addition. The termites
were likely entering through the hollow blocks of
the stem wall of the raised slab. AG stations con-
taining blank bait moistened with sucrose solu-


Date Total Total #
Date detected estimated Total noviflumur- Days post-
monitor Date 1" FST g dry wt estimated on bait elimination
stations bait infestation Days to bait matrix mgAI devices final
Site installed applied eliminated eliminate consumed consumed installed2 inspection'

DAYCARE 6/29/01 7/13/01 10/05/01 82 294 147 15IG, 6AG 499
TRAILER 7/08/02 2/21/03 5/02/03 71 189 95 6 AG 364
CONDO 5/30/02 6/28/02 9/30/02 92 98 49 6 AG 549
PENTHOUSE 11/27/03 1/13/03 3/26/03 73 26 13 4 IG 373

1Inspection of all monitoring stations and building
'Numbers of noviflumuron bait devices installed are greater than number of baited IG and AG stations indicated in Fig 3-6 be-
cause noviflumuron baits were replaced as required if consumed or degraded in IG and AG stations.

March 2006


Initial noviflumuron
application 1" Month 2nd Month 3rd Month
Type of
Site (station type2) matrix3 # FST (n)4 Total g (n)5 # FST (n) Total g (n) # FST (n) Total g (n) # FST (n) Total g (n)

DAYCARE (IG & AG) Blank 970 (10) 266 (8) 1570 (8) 85 (6) 100 (1) 231 (5) 0 0
0.5% AI 175 (4) 28 (3) 700 (4) 247 (7) 129 (3) 19 (1)
TRAILER (AG) Blank 183 (4) 172 (4) 1210 (5) 151(5) 400 (2) 105 (3) 2(1) 11(3)
0.5% AI 670 (3) 175 (3) 50 (1) 14 (2) 0 0
CONDO (AG) Blank 1595 (7) 221(7) 420 (3) 35 (2) 200 (1) 14 (1) 0 0
0.5% AI 500(4) 98(5) 7 (3) 0 0 0
PENTHOUSE (IG & AG) Blank 525 (5) 265 (5) 560 (3) 114 (5) 1280 (4) 168 (4) 0 14 (1)
0.5% AI 55(2) 26(2) 0 0 0 0

No termites or matrix consumption observed at any site after 3"d month.
IG = inground station, AG = aboveground station.
'Blank = IG wood monitoring device (62 g), blank IG bait (35 g), or blank AG bait (70 g). 0.5% AI (noviflumuron) = IG bait (35 g) orAG bait (70 g).
'n = total number of stations with live termites by inspection period.
5n = total number of stations with matrix consumption by inspection period.

Florida Entomologist 89(1)

tion were installed over live FST on this partition
wall on November 8, 2002. Termites did not con-
sume any of the bait through January 2003.
Blank bait was then moistened with a sports
drink on January 24, 2003 and two weeks later,
termites were found actively consuming the bait.
The previous termiticide application and ex-
cessive wood decay in the trailer may have inter-
fered with FST foraging and feeding on IG and
AG stations. The untreated soil under the slab in
the attached addition appeared to be the refugia
for the remaining FST infestation. FST workers
contacting the imidacloprid perimeter treatment
may have suffered lethal and sublethal effects,
preventing them from foraging in IG stations. The
trailer frame was damp and decayed as a conse-
quence of flooding several years before. Certain
species of wood-decay fungi have been shown to
make some wood species more palatable to C. for-
mosanus than non-decayed wood (Cornelius et al.
2003; Cornelius et al. 2004). The combination of
high wood moisture and fungal decay may have
made the wood more palatable to the termites
than the bait matrix until the sports drink was
added. Although this observation suggests this
product is a feeding stimulant, Cornelius (2005)
found no significant feeding preferences by C. for-
mosanus workers when offered filter paper disks
soaked with water or Gatorade.
Detected FST infestations were eliminated in
71 to 92 days (mean SD = 80 10 days) at all
buildings after first application of noviflumuron
bait (Table 1). The date of elimination was deter-
mined by cessation of all termite activity in IG
and AG stations, previously infested foraging
tubes and wood, and in electronically monitored
locations. There were some aberrant AED read-
ings at CONDO when monitored on September 6
and September 30. Four exceptionally high AE
counts (69, 171, 240, and 602) recorded from two
of the doorways on September 6, 2002 appear to
have been anomalies possibly due to a malfunc-
tion with the equipment and were excluded from
the mean counts for this date (Fig. 8). On that day,
75 dead soldiers were found in one AG station in-
side CONDO. On a previous inspection on August
29 of AG stations in the stairwell, only 7 live ter-
mites, all soldiers, were found. The absence of
workers and presence of only soldiers are two in-
dicators of a termite population in decline after
workers have fed on noviflumuron bait. A single
high reading (18 counts) recorded on September
30 inside the condo appeared to be aberrant based
on the absence of termites in stations. September
30, 2002 was determined to be the date for elimi-
nation of the detected FST infestation because no
consumption of AG bait occurred since the previ-
ous inspection and no live termites were ob-
served. The lack ofAE counts on October 25 2002,
the subsequent absence of termites in damaged
wood removed by the tenants, and lack of FST ac-

tivity in AG stations through the following 18 mo
until the final structural inspection further indi-
cate this FST infestation was eliminated.
Ground-based FST foraging populations at
DAYCARE and TRAILER ate approximately 4-
fold more bait matrix with noviflumuron com-
pared with that eaten by aerial populations at
CONDO and PENTHOUSE (mean + SD; 242 + 74
vs. 62 51 g, respectively). Weissling & Thoms
(1999) also found a ground-based FST colony con-
sumed, on average, 2.5-fold more termite bait
than two aerial FST colonies. This may be due to
differences in foraging population size. It has
been observed that the availability of food and
water can be limited for aerial termite colonies
(Su et al. 1997, 2001) which could restrict popula-
tion growth. Although we did not estimate the
population size, the furthest distances between
detected foraging locations were less for rooftop
infestations at PENTHOUSE (6 m) and CONDO
(9 m) than for DAYCARE (20 m).
No re-infestation by FST was found in any
building or monitoring station during the remain-
ing evaluation period following elimination of
FST activity. The final inspection of stations and
buildings occurred 12-18 mo after FST elimina-
tion (Table 1). During final inspections, no live
termites were found in any station, building, or
previously infested landscape feature, such as the
mango tree at DAYCARE. No consumption of ter-
mite monitors in stations or new termite damage
or signs of recent termite activity, such as forag-
ing tubes, were found.
Results indicate 0.5% noviflumuron is not a
feeding deterrent to FST. At DAYCARE during
the third and final month of bait application, live
termites and matrix consumption were observed
only in noviflumuron baits, not in blank baits or
wood monitors (Table 2). In addition, AED moni-
toring detected no termite feeding activity after
August 15, 2001 during the second month of bait
application (Fig. 7). Live termites and consump-
tion were observed in monitoring stations con-
taining blank monitors, but not noviflumuron,
during the second and third month after initial
application of noviflumuron at PENTHOUSE.
The reason for this is during the second month of
noviflumuron bait application, the owner reland-
scaped the planter bed removing the palm stumps
with AG stations and all but one noviflumuron-
baited IG station. The stacked AG stations at-
tached to the wood decking were not disturbed.
New IG stations containing wood monitors were
subsequently installed, but no further termite
activity was observed in any IG station. Despite
this disruption in the bait application process, the
detected FST infestation was eliminated at
This study demonstrates the versatility of cel-
lulose baits containing 0.5% noviflumuron for
eliminating structural infestations of FST. Novi-

March 2006

Cabrera & Thoms: Control of Formosan Subterranean Termites

7/13/2001 7/27/2001 8/15/2001 8/24/2001

9/7/2001 9/2112001 10/5/2001


Fig. 7. Total estimated number of termites (bars) and mean ( SE) acoustic emissions counts (line) per monitor-
ing location before, during, and after application of 0.5% noviflumuron bait, DAYCARE, Broward County, Florida,

E 500


E 200




20 0

15 C




5130/2002 6/7/2002 6/182002 612812002 7/12/2002 7/26/2002 819/2002 9/6/2002 9/30/2002 102512002

Fig. 8. Total estimated number of termites (bars) and mean ( SE) acoustic emissions counts (line) per monitor-
ing location before, during, and after application of 0.5% noviflumuron bait, CONDO, Broward County, Florida,



a 1600


o 1200

c 800

'* 400




140 C



80 u
60 |



-- -

flumuron baits can be applied IG and/or AG to
eliminate FST populations. Unlike liquid termiti-
cide treatments, all areas of soil access to the
building for a ground-based FST population and
all termite activity within the building for above-
ground infestations did not need to be treated
with noviflumuron baits to eliminate the struc-
tural infestation of FST. We also were able to
eliminate FST infestations in two situations
(CONDO and DAYCARE) where the occupants
would not permit application of pesticide to in-
door living or outdoor activity areas.


The authors sincerely appreciate the assistance of
K. Nitsch, D. O'Brien, K. Wheeler, C. Alles, and D. Reis-
inger for assisting with station and building inspec-
tions. The research would not have been possible
without the cooperation of the following building own-
ers, occupants, and managers; K. Kuehney and S. Davis
of DAYCARE, J. and D. Levy, B. Kassell, and D. Capps of
CONDO, D. Centauro of TRAILER, and R. Meyer-
Werner and B. Capaldo of PENTHOUSE. The authors
thank R. H. Scheffrahn, and W. H. Kern, Jr., University
of Florida-Institute of Food and Agricultural Sciences,
Fort Lauderdale Research & Education Center, and two
anonymous reviewers for reviewing and improving the
manuscript. This research was supported by the Florida
Agricultural Experiment Station and a grant from Dow
AgroSciences LLC.


CORNELIUS, M. L. 2005. Effect of bait supplements on
the feeding and tunneling behavior of the Formosan
subterranean termite (Isoptera: Rhinotermitidae) In
C.-Y. Lee and W. H. Robinson [eds.], Proc. 5th Int.
Conf. on Urban Pests, Singapore, 10-13, July 2005.
S. WILLIAMS, AND M. P. LOVISA. 2003. Responses of
the Formosan subterranean termite (Isoptera: Rhi-
notermitidae) to wood blocks inoculated with lignin-
degrading fungi. Sociobiology 41:513-525.
LAX. 2004. Effect of a lignin-degrading fungus on
feeding preferences of Formosan subterranean ter-
mite (Isoptera: Rhinotermitidae) for different com-
mercial lumber. J. Econ. Entomol. 97: 1025-1035.
DEMARK, J. J., AND J. D. THOMAS. 2000. Seasonal activ-
ity, wood consumption rates, and response to above-
ground delivery of hexaflumuron-treated bait to Reti-
culiterms flavipes (Isoptera: Rhinotermitidae) in Penn-
sylvania and Wisconsin. Sociobiology 36: 181-200.
VARGO, AND N.-Y. SU. 2003. Describing the spatial
and social organization of Formosan subterranen
termite colonies in Armstrong Park, New Orleans.
Sociobiology 41: 61-65.
EVANS, T. A. 2002. Assessing efficacy of TermaTrac-a
new microwave based technology of non-destructive
sampling for termites (Isoptera). Sociobiology 40(3):
2004. Laboratory performance and pharmacokinetics

March 2006

of the benzoylphenylurea noviflumuron in eastern
subterranean termites (Isoptera: Rhinotermitidae).
J. Econ. Entomol. 97: 593-600.
LEE, C.-Y. 2002. Control of foraging colonies of subter-
ranean termites, Coptotermes travians (Isoptera:
Rhinotermitidae) in Malaysia using hexaflumuron
baits. Sociobiology 39: 411-416.
Above-ground baiting for controlling the subterra-
nean termite, Coptotermes travians (Isoptera: Rhi-
notermitidae) in Selangor, Peninsular Malaysia.
Sociobiology 39: 345-352.
2005. Field evaluation of noviflumuron for control-
ling Asian subterranean termite, Coptotermes
gestroi (Isoptera: Rhinotermitidae) In C.-Y Lee and
W. H. Robinson [eds.], Proc. 5th Int. Conf. on Urban
Pests, Singapore, 10-13, July 2005.
SCHEFFRAHN, R. H. 1994. Keys to soldiers and winged
adult termites (Isoptera) of Florida. Florida Ento-
mol. 77: 460-474.
AND R. K. MUELLER 1993. Evaluation of a novel,
hand-held, acoustic emissions detector to monitor
termites (Isoptera: Kalotermitidae, Rhinotermiti-
dae) in wood. J. Econ. Entomol. 86: 1720-1729.
SHEETS, J. J., L. A. KARR, AND J. E. DRIPPS. 2000. Kinet-
ics of uptake, clearance, transfer, and metabolism of
hexaflumuron by eastern subterranean termites
(Isoptera: Rhinotermitidae). J. Econ. Entomol. 93:
SMITH, M. S., L. L. KARR, J. E. KING, W. N. KLINE, R. J.
flumuron activity in household and structural insect
pests, pp. 345-353 In S. C. Jones, J. Zhai, and W. H
Robinson [eds.], Proceedings of the 4th International
Conference on Urban Pests. Eds. Pocahontas Press,
Blacksburg, VA. pp 345-353.
Su, N.-Y. 2002a. Baits as a tool for population control of
Formosan subterranean termites. Sociobiology
40(2): 1-16.
Su, N.-Y. 2002b. Dimensionally stable sensors for a con-
tinuous monitoring program to detect subterranean
termite (Isoptera: Rhinotermitidae) activity. J.
Econ. Entomol. 95: 975-980.
SU, N.-Y., P. M. BAN, AND R. H. SCHEFFRAHN. 1997. Re-
medial baiting with hexaflumuron in above-ground
stations to control structure-infesting populations of
the Formosan subterranean termite (Isoptera: Rhi-
notermitidae). J. Econ. Entomol. 90: 809-817.
SU, N.-Y., P. M. BAN, AND R. H. SCHEFFRAHN. 2001.
Control of subterranean termites (Isoptera: Rhino-
termitidae) using commercial prototype above-
ground stations and hexaflumuron baits.
Sociobiology 37: 111-120.
SU, N.-Y., P. M. BAN, AND R. H. SCHEFFRAHN. 2002.
Control of subterranean termite populations at San
Cristobal and El Morro, San Juan National Historic
Site. J. Cultural Heritage 3: 217-225.
Control of the Formosan subterranean termite infes-
tations in historic Presbytere and the Creole House
of the Cabildo, French Quarter, New Orleans, using
baits containing an insect growth regulator, hexaflu-
muron. Studies in Conservation 45: 30-38.
FRAHN. 2003. Protecting historic properties from
subterranean termites: a case study with Fort Chris-

Florida Entomologist 89(1)

Cabrera & Thoms: Control of Formosan Subterranean Termites

tiansvaern, Christiansted National Historic Site,
United States Virgin Islands. American Entomol.
49(1): 20-32.
SU, N.-Y., AND E.-L. Hsu. 2003. Managing subterra-
nean termite populations for protection of the his-
toric Tzu-Su Temple of San-Shia, Taiwan.
Sociobiology 41: 529-545.
Su, N.-Y., AND R. H. SCHEFFRAHN. 1993. Laboratory
evaluation of two chitin synthesis inhibitors,
hexaflumuron and diflubenzuron, as bait toxicants
against Formosan and eastern subterranean ter-
mites (Isoptera: Rhinotermitidae). J. Econ. Entomol.
86: 1453-1457.

Su, N.-Y., R. H. SCHEFFRAHN, AND P. M. BAN. 1989.
Method to monitor initiation of aerial infestations by
alates of the Formosan subterranean termite
(Isoptera: Rhinotermitidae) near structures. J. Econ.
Entomol. 88: 1643-1645.
WEISSLING, T. J., AND E. M. THOMS. 1999. Use of an
acoustic emission detector for locating Formosan
subterranean termite (Isoptera: Rhinotermitidae)
feeding activity when installing and inspecting
aboveground termite bait stations containing
hexaflumuron. Florida Entomol. 82: 60-71.

Florida Entomologist 89(1)

March 2006


Entomology and Nematology Department, University of Florida, Gainesville, FL 32611-0620


The seasonal abundance of phytophagous scarabs in Gainesville and Fort Lauderdale, Flor-
ida, was documented with ultraviolet blacklight traps operated fromApril 2002 to November
2004. Over 44,000 adult scarabs were trapped and identified, including 30 species from 14
genera. Hybosorus illigeri Reiche was the most abundant species trapped (n = 12,306 or
27.9% of total trap catches). Phyllophaga was the most diverse genus with ten species col-
lected. Tomarus cuniculus (F.) and Dyscinetus morator (F.) adults were trapped every month
of the year.Anomala innuba (F.), Cyclocephala lurida (Bland), C. parallel Casey, H. illigeri,
and Phyllophaga bruneri Chapin exhibited bimodal flight patterns. Adults of these five spe-
cies combined represented 49.1, 56.5, and 64.6% of the collections in 2002, 2003, and 2004,
respectively. Species that occurred in both locations tended to be active earlier in Fort Lau-
derdale than in Gainesville. The flight activity and species composition of potential scarab
pests in Florida appears to be different from those in the midwestern and northern U.S., sug-
gesting that turfgrass and ornamental plant managers need to adjust their management
strategies accordingly.

Key Words: Scarabaeoidea, flight activity, blacklight trapping


La abundancia estacional de las species fitofagas de escarabajos (Familia Scarabaeidae) en
Gainesville y Fort Lauderdale, Florida, fue documentada usando trampas de luz negra ultra
violeta operadas de abril 2002 hasta noviembre del 2004. Mas de 44,000 adults fueron cap-
turados e identificados, incluyendo 30 species en 14 g6neros. Hybosorus illigeri Reiche fue
la especie mas abundante capturada (n = 12,306 o 27.9% del total de las species capturadas
en la trampa). Phyllophaga fue el g6nero mas divers con diez species recolectadas. Adultos
de Tomarus cuniculus (F.) y Dyscinetus morator (F.) fueron capturados en cada mes del aio.
Anomala innuba (F.), Cyclocephala lurida (Bland), C. parallel Casey, H. illigeri y Phylloph-
aga bruneri Chapin mostraron un patron bimodal de vuelo. Los adults de estas cinco espe-
cies juntas representaron 49.1, 56.5, y 64.6% de las colecciones en 2002, 2003 y 2004,
respectivemente. Las species presents en estos dos lugares tienden a ser tempranamente
mas activas en Fort Lauderdale que en Gainesville. La actividad de vuelo y la composici6n
de species de Scarabaeiidae que son plagas potenciales en Florida parece ser diferentes que
las en la region central y norte de los Estados Unidos, esto sugiere que los de negocios de c6s-
ped y de plants ornamentales deben ajustar sus estrategias del manejo segun el caso.

Through their root feeding, some immature
scarab beetles (Coleoptera: Scarabaeoidea) or
white grubs, are economic pests of turfgrasses,
corn, sorghum, grains, vegetables, conifers, and
ornamental plants (Forschler & Gardner 1990;
Vittum et al. 1999). Some adult scarabs do not
feed, while others partially or completely defoli-
ate hardwood trees and shrubs (Habeck &
Wolfenbarger 1968), make mounds while ovipos-
iting in the soil, or are nuisances by being abun-
dant and active on golf course greens and tees. In
addition, birds, raccoons, moles, armadillos, and
other animals can cause extensive damage when
digging to find white grub prey (Potter 1998).
White grubs were considered minor pests in
Florida while several chlorinated hydrocarbon and

organophosphate insecticides were available for use
on turfgrass (Ralph White, pers. comm.). Most of
these compounds are no longer available, and white
grub populations have increased and become locally
damaging along the Gulf and Atlantic Coasts. Sev-
eral lawn care firms have reported poor control of
several scarab species that are uncommon or do not
exist in other states (pers. obs.). Although lists of
scarabs collected in other states and their adult
phenologies have been published (see Forschler &
Gardner 1991), such data are incomplete in Florida.
Thus, the following blacklight study was conducted
to describe the seasonality of phytophagous scarabs
in Florida. This information will be used to develop
an integrated pest management program for white
grubs on urban turfgrasses.

Buss: Adult Scarab Seasonality


The blacklight traps (BioQuip, Rancho
Dominguez, CA) used in this study were AC-pow-
ered, had a 22-watt Circline bulb, and were oper-
ated day and night for the duration of the study.
Traps contained an insecticidal strip (dichlorvos,
Hot Shot No-Pest Strip@) to kill insects caught in
the bucket. The strip was replaced monthly or as
needed. A hole in the bottom and side of the
bucket allowed rain water to drain. All traps were
suspended ca. 2 m from the ground from a metal
One trap was located at the Gainesville Golf
and Country Club in Gainesville (Alachua Co.),
FL. The primary turfgrass on the golf course was
bermudagrass (Cynodon dactylon [L.] Pers.) vari-
eties 419, Ormond, and GN1. The soil was classi-
fied as Blichton-Urban land complex (loamy, sili-
ceous, hyperthermic Arenic Plinthic Paleaquults).
The trap was operated from 4 April 2002 until 22
November 2004. Scarabs were collected once or
twice a week and frozen.
The second trap was located at the Fort Lau-
derdale Research and Educational Center (Bro-
ward Co.), FL. It was placed near bermudagrass
and St. Augustinegrass (Stenotaphrum secunda-
tur [Walt.] Kuntze.) turfgrass research plots. The
soil was Margate fine sand (siliceous, hyperther-
mic Mollic Psamnaquent). Specimens were col-
lected once or twice a week from 30 April 2002 un-
til 15 November 2004, and frozen.
Scarab beetles were sorted, counted, and
stored in 70% ethyl alcohol (EtOH). Hybosorus il-
ligeri Reiche (Coleoptera: Scarabaeoidea: Hybo-
soridae) was also included because the adults are
nuisance pests on golf courses in Florida (pers.
obs.), but the larvae may not be phytophagous
(Woodruff 1973; Ocampo 2002). The genitalia of
Cyclocephala spp. and Phyllophaga spp. were ex-
tracted for species and gender determination.
Identifications were based on comparisons with
specimens at the Florida Collection of Arthropods
and keys (Gordon & Anderson 1981; Woodruff &
Beck 1989; Arnett et al. 2002). Identifications
were confirmed by M. Thomas and P Skelley, and
voucher specimens were deposited at the Florida
Collection of Arthropods at the Division of Plant
Industry, Gainesville, FL.


During this study, >44,000 adult scarabs were
trapped and identified, including 30 species from
14 genera. The total number of scarabs caught in
traps (Tables 1 and 2) varied somewhat each year
at both sites (11,334 beetles in 2002, 16,916 bee-
tles in 2003, and 15,791 beetles in 2004). The
number of species and genera also varied over
time (22 species and ten genera in 2002, 28 spe-
cies and 14 genera in 2003, and 26 species and 12

genera in 2004). The trapped scarabs were more
diverse at the Gainesville site (25 species) than at
the Fort Lauderdale site (14 species). However,
species that occurred in both locations tended to
be active earlier in Fort Lauderdale than in
Gainesville. The most abundant species captured
in the light traps in 2002 was Dyscinetus morator
(F.) (n = 4,550), but H. illigeri was the predomi-
nant species collected in 2003 (n = 4,942) and
2004 (n = 5,155). The species composition of phy-
tophagous scarabs in Florida had some slight
overlap with other southeastern states (Forschler
& Gardner 1991; Flanders et al. 2000; Harpoot-
lian 2001).
Three species of Anomala, A. innuba (F.), A.
marginata (F.), and A. undulata Melsheimer,
were collected, which represented 13.5% of the to-
tal catch during this study (n = 5,964). This was
the most abundant genus in 2003 and 2004.
Anomala innuba exhibited bimodal flight activity
at both locations (Fig. 1), with peak activity from
late April to mid-May and early August to mid-
September in Gainesville and from early April to
mid-May and late August to mid-October in Fort
Lauderdale. It had one flight peak in Alabama
and Kansas (Hayes 1925; Flanders et al. 2000).
The number of A. innuba collected in the Fort
Lauderdale trap appeared to be increasing each
year, which might indicate a growing pest prob-
lem there. Although not detected in this study,
Hall (1987) observed a bimodal flight pattern ofA.
marginata in a sugarcane field in south-central
Florida. This species has a unimodal flight pat-
tern in Kentucky and North Carolina (Brimley
1938; Ritcher 1966). Anomala spp. larvae attack
grass roots (Hayes 1925).
Cyclocephala spp. represented 13.0% of the to-
tal catch (n = 5,720). Several Cyclocephala spp.
have been considered among of the most damag-
ing white grub pests of sugarcane and turfgrass in
Florida (Reinert 1979; Hall 1987). The southern
masked chafer C. lurida (Bland.) had two flight
peaks each year in Gainesville (Fig. 2), which has
not been previously documented, and the species
was not collected in Fort Lauderdale. In other un-
published research, third instars were collected in
July, pupation occurred in late July and early Au-
gust, and the adults were identified (pers. obs.).
Its peak adult activity occurred from early May to
mid-June for the first generation and early Au-
gust to late September for the second generation.
Peak flight periods for C. lurida typically occur in
June and July in other states (Flanders et al.
2000). Cyclocephala parallel Casey had one
flight peak in Gainesville but had a bimodal flight
pattern in Fort Lauderdale (Fig. 3). Peak adult ac-
tivity for C. parallel was in early June in Gaines-
ville, and from late April to early June and mid-
August to late September in Fort Lauderdale.
However, in a study conducted in a south-central
Florida sugarcane field, peak C. parallel adult

Florida Entomologist 89(1)

March 2006


Species Year n Flight period Peak activity

Anomala innuba (F.)

Anomala marginata (F.)

Anomala undulata Melsheimer

Aphonus variolosus (LeConte)
Cyclocephala lurida (Bland.)

Cyclocephala parallel Casey

Cyclocephala seditiosa LeConte

Diplotaxis punctatorugosa Blanchard

Dyscinetus morator (F.)

Euetheola humilis rugiceps (LeConte)

Hybosorus illigeri Reiche

Pelidnota punctata (L.)

Phyllophaga crenulata (Froelich)

Phyllophaga debilis (LeConte)

Phyllophaga glaberrima (Blanchard)

















168 Apr. 12-Sep. 26 Apr. 25-May 9
Aug. 1-8
68 Apr. 21-Oct. 6 May 5-12
Aug. 21-Sep. 11
73 Apr. 29-Sep. 21 May 10-17
Aug. 13-30

33 May 2-July 18
58 Apr. 10-July 17
13 May 20-Aug. 19

38 Apr. 8-May 2
78 Feb. 25-June 9
22 Mar. 1-May 17

1 June 9

June 3-17
June 2-19
June 10-28

Apr. 12-May 2
Mar. 3-Apr. 8
Mar. 8-Apr. 12

1,579 Apr. 12-Dec. 5 May 2-June 17
Aug. 8-Sep. 26
955 Apr. 24-Nov. 17 May 5-June 16
Aug. 14-Sep. 29
1,075 Apr. 26-Nov. 1 May 20-June 21

39 May 20-July 15
20 May 22-June 26
16 June 10-July 12
1 June 5

8 Apr. 15-June 6
5 May 29-Aug. 14
1 Apr. 26


Apr. 8-Nov. 14
Feb. 10-Dec. 8
Jan. 5-Nov. 22
Apr. 8-Nov. 7
Mar. 13-Nov. 13
Mar. 8-Oct. 4

May 30-June 17
May 27-June 9
June 14-21

Apr. 12-May 20
Feb. 25-Apr. 8
Mar. 8-Apr. 12
Apr. 18-May 4
Mar. 17-Apr. 8
Mar. 29-Apr.19

587 Apr. 25-Oct. 7 May 2-June 17
Sep. 2-13
292 Apr. 8-Oct. 30 May 19-June 9
Aug. 14-Sep. 8
533 May 3-Oct. 22 May 24-June 28
Sep. 13-21

29 May 2-July 8
34 May 8-Aug. 14
72 May 10-Aug. 30

3 Apr. 8-May 9
5 Mar. 20-June 9
10 Mar. 15-May 6

1 May 30
1 May 24

39 May 9-Aug. 26
33 June 2-Aug. 25
36 May 24-Aug. 26

May 20-30
June 2-9
June 7-24

June 6-17
June 9-30
June 14-24

Buss: Adult Scarab Seasonality

FROM 2002 TO 2004.

Species Year n Flight period Peak activity

Phyllophaga latifrons (LeConte) 2002 109 May 2-July 11 May 30-June 17
2003 75 May 12-Aug. 14 June 2-12
2004 28 May 27-Aug. 2 June 14-July 6
Phyllophaga parvidens (LeConte) 2004 1 May 3 -
Phyllophaga prununculina (Burmeister) 2002 25 May 16-July 4 -
2003 32 June 2-July 14 -
2004 30 June 14-Aug. 2 -
Phyllophaga quercus (Knoch) 2002 121 May 30-Aug. 1 June 6-July 4
2003 115 May 27-Aug. 21 June 19-July 7
2004 204 June 7-Aug. 13 June 14-July 12
Phyllophaga tecta Cartwright 2003 22 Mar. 20-June 9
2004 9 Mar. 29-June 14
Phyllophaga uniforms (Blanchard) 2002 393 May 2-July 11 May 30-June 17
2003 568 May 8-Aug. 14 May 27-June 30
2004 852 May 10-July 29 June 7-28
Polyphylla occidendentalis (L.) 2002 3 Apr. 15-May 2 -
2003 5 Apr. 28-May 22 -
2004 2 May 10-June 17 -
Serica sericea (Illiger) 2003 3 Mar. 20-Apr. 10 -
Strategus antaeus (Drury) 2003 2 Aug. 14-Sep. 2 -
2004 1 June 28 -
Tomarus gibbosus DeGeer 2002 8 Apr. 25-May 20 -
2003 27 Apr. 21-Sep. 18 -
2004 21 Apr. 29-Sep. 28 -

activity only occurred from mid-April to mid-May
(Hall 1987). Trap catches contained primarily
male C. lurida (91.7%) and C. parallel (90.4%),
but only 43.5% ofC. miamiensis (Howden and En-
drodi) were male. Just one male C. seditiosa Le-
Conte was collected in this study.
The rice beetle, Dyscinetus morator (F.), was
abundantly collected (n = 9,493 or 21.6% of the to-
tal catch) nearly every month of the year, with
peak flight occurring between February and May
in Gainesville. Its peak flight period occurred in
May and June in Alabama and Georgia (Flanders
et al. 2000), but a second generation may occur in
August and September in Georgia (Forschler &
Gardner 1991). Woodruff (1970) also suggested
that D. morator was bivoltine in southern Florida,
based on the occurrence of greater adult activity
in the spring and fall and Smyth's (1915) data in-
dicating that other Dyscinetus spp. could go from
egg to adult in <144 days in Puerto Rico. However,
only one peak of adult activity was detected each
of the three years of this Florida study. It is
known to inhabit wet soils, marsh areas (Buck-
ingham & Bennet 1989), and compost (Ritcher
1966). Larvae feed on rice, pangola grass pas-

tures, crabgrass, water hyacinth, caladium bulbs,
and azaleas (Staines 1990).
Phyllophaga spp. represented 12.8% of the to-
tal catch during this study (n = 5,643). Fifty-four
species of Phyllophaga occur in Florida (Woodruff
& Beck 1989), but only nine species were collected
in Gainesville, compared to the three species col-
lected in Fort Lauderdale. Only P. glaberrima
(Blanchard) and P. latifrons (LeConte) occurred
at both sites in this study, and are known to have
a statewide distribution (Woodruff & Beck 1989).
The most abundant species was P. bruneri Chapin
(n = 2,240), which was only collected in Fort Lau-
derdale (Fig. 4) and is unique within this genus by
exhibiting a bimodal flight pattern (Fig. 4), repre-
senting two distinct generations, and is active ev-
ery month of the year in southern Florida
(Habeck & Wolfenbarger 1968). The earliest flight
activity in the year consistently began with
P crenulata (Froelich) and P tecta Cartwright in
March in Gainesville and P glaberrima and P lat-
ifrons in Fort Lauderdale. Species that were ac-
tive from May to August included P. glaberrima,
P latifrons, and P. quercus (Knoch), which have
similar flights in Alabama (Flanders et al. 2000)

Florida Entomologist 89(1)

March 2006

TO 2004.

Species Year n Flight period Peak activity

Anomala innuba (F.)

493 Apr. 30-Dec. 31

2003 1,952 Jan. 3-Dec. 26

2004 2,468 Jan. 6-Nov. 11

Apr. 30-May 14
Sep. 3-Oct. 8
Apr. 1-May 8
Aug. 26-Oct. 21
Apr. 9-May 4
Aug. 31-Oct. 16

Anomala marginata (F.)

Cyclocephala miamiensis (Howden & Endrodi)

Cyclocephala parallel Casey

69 Apr. 30-Dec. 31
80 Jan. 3-Dec. 26
71 Jan. 6-Nov. 15

4 Apr. 30-May 10
133 Apr. 22-May 16
35 Apr. 27-June 15

312 Apr. 30-Dec. 13

2003 1,004 Mar. 14-Nov. 21

547 Feb. 24-Oct. 25

May 10-28
Aug. 13-Sep. 28
Apr. 4-May 20
Aug. 19-Sep. 30
May 4-June 12
Aug. 31-Oct. 1

Dyscinetus morator (F.)

Euphoria sepulcralis (F.)

Euetheola humilis rugiceps (LeConte)
Hybosorus illigeri Reiche

2002 33 Aug. 6-Dec. 27
2003 59 Feb. 4-Dec. 30
2004 37 Jan. 9-Oct. 18
2003 10 Feb. 28-July 8
2004 22 Feb. 17-Aug. 10
2004 1 Jan.20

2002 1,622 Apr. 30-Dec. 27

2003 4,650 Feb. 11-Dec. 5

2004 4,622 Apr. 27-Nov. 4

May 6-June 6
Aug. 6-Sep. 17
May 6-June 3
July 22-Sep. 5
May 25-June 29
Aug. 10-Sep. 17

Pelidnota punctata (L.)

Phyllophaga bruneri Chapin

Phyllophaga glaberrima (Blanchard)

Phyllophaga latifrons (LeConte)

2 Apr. 30-May 10
38 Apr. 1-May 13
16 Mar. 23-July 9

760 Apr. 30-Dec. 24 Apr. 30-May 28
July 26-Aug. 20
617 Feb. 25-Dec. 26 Apr. 22-May 13
Aug. 19-Sep. 23
863 Jan. 2-Oct. 21 Apr. 23-June 1
Aug. 17-Sep. 17

4 May 10-Dec. 10
66 Mar. 25-Sep. 2
68 Mar. 19-Aug. 24

31 Apr. 30-July 18
222 Mar. 21-July 18
240 Mar. 23-Aug. 24

Apr. 29-May 13
May 25-June 12

Tomarus cuniculus (F.)



Tomarus subtropicus (Blatchley)

Apr. 30-Dec. 31
Jan. 3-Dec. 30
Jan. 28-Nov. 18

July 1

Buss: Adult Scarab Seasonality


5 -
0;: A._ IN___ L__A&

24 S A 't 7 t 7 1 '

B. Anomala innuba (Ft. Lauderdale)
8 700 -

f 500
s 300-

,0 ---, 0 1 .

Fig. 1. Flight activity ofA. innub at blacight traps located in Gainesville (A) and Fort Lauderdale, FL (B).

Fig. 1. Flight activity ofA. innuba at blacklight traps located in Gainesville (A) and Fort Lauderdale, FL (B).

and are probably univoltine (Luginbill & Painter
1953; Flanders et al. 2000). Genders for all adult
Phyllophaga spp. trapped from 2002 to 2004 were
identified. Species that had more males than fe-
males collected in traps, averaged over the three
years, included P crenulata (93.3% male), P glab-
errima (94.1%), P latifrons (64.4%), P parvidens
(LeConte) (100%), P prununculina (Burmeister)
(92.3%), P quercus (91.6%), P tecta (85.4%), and
P uniforms (Blanchard) (59.5%). Only P bruneri
(47.0% male) and P debilis (LeConte) (50%) had
more of a female bias. Nearly all of these species
can damage ornamental plant foliage as adults,

and many of the larvae infest nursery stock or
turfgrass (Woodruff & Beck 1989).
Tomarus spp. (formerly Bothynus and Ligyrus)
represented 10.2% of the total catch (n = 4,475).
The carrot beetle, T gibbosus (DeGeer), which is a
pest of various vegetable crops (Hayes 1917), was
collected only in Gainesville. Tomarus cuniculus
(F.), only collected in Fort Lauderdale, is an inva-
sive species that can cause significant turfgrass
damage along the Atlantic Coast of Florida.
Adults were abundantly collected in the Fort Lau-
derdale trap nearly every month of the year with-
out a distinct peak period of activity. Only one

Cyclocephala lurida (Gainesville)

rif -l

Fig. 2. Flight activity of the southern masked chafer, C. lurida, in Gainesville, FL.

. .xgt

Jfl. .tS



. 200

. 150


Z in-

171 L 6IS.& !-- ,n 6 r- Al,.& .N- .


Florida Entomologist 89(1)

3 12

Z 2

i 70
. 40
J 30

g6 I a C? 0
0. "7 =sx $~
is. S 6 d Si ,x S. S- E IP& i-S^ S
!2 Xo

Fig. 3. Flight activity of C. parallel at blacklight traps located in Gainesville (A) and Fort Lauderdale, FL (B).

T subtropicus (Blatchley) flew to a black light
trap. This species tends to be more abundant
along the Gulf Coast in St. Augustinegrass (pers.
obs.) and is considered a primary pest of sugar-
cane (Gordon & Anderson 1981). Adult T sub-
tropicus are active from April to July in sugarcane
in south-central Florida (Cherry 1985).
Hybosorus illigeri was collected at the light
traps from April to October in Gainesville and
nearly all year in Fort Lauderdale (n = 12,306
beetles from both sites, or 27.9% of the total
catch). Although Woodruff (1973) considered H. il-

ligeri to be univoltine, two peaks of activity (May
to June, August to September) were consistently
observed in this study at both locations (Fig. 5).
The smaller second flight peak suggests that not
all individuals fully complete a second genera-
tion. Little is known about its biology (Woodruff
1973; Ocampo 2002), but it has been collected at
light, in dung, in carrion, and has been observed
feeding on other scarabs (see Ocampo 2002). The
abundance of adults and the small mounds that
they make on golf course tees and greens are an-
noying to golfers and golf course superintendents,

160 Phyllophaga bruneri (Ft. Lauderdale)
I 60

i 6 i a o a e b
Z 2

Fig. 4. Flight activity of the Cuban May beetle, P. bruneri, in Fort Lauderdale, FL.

A. Cyclocephala parallel (Gainesville)

8h 8 i: 4 0a %% 4e
:sf 1 i. It

B. __Cqcl^o&ne parallela Ft.Laude-da Ale

March 2006

Buss: Adult Scarab Seasonality



170 B. Hybosorus illigeri (Ft. Lauderdale)
1250------- -
F 250
z 0 0 -10 Am aid

Fig. 5. Flight activity of H. illigeri at blacklight traps located in Gainesville (A) and Fort Lauderdale, FL (B).

but turf damage is not apparent even where den-
sities are high (pers. obs.).
The existence of a bimodal flight pattern for
several scarab species in Florida could be the re-
sult of several factors, which might be resolved by
collecting adults and rearing the subsequent gen-
eration(s) under controlled conditions. It is possi-
ble that two similar species or an undescribed in-
vasive species might coincide in an area, but
adults may not have been taxonomically sepa-
rated. Especially with individuals beginning their
flights early in the year, egg and/or larval develop-
ment might be at least initially slower due to
cooler and drier conditions (Gaylor & Frankie
1979; Potter 1981) than those individuals that fly
and lay eggs during the warmer and more humid
Florida summer. However, because some individ-
uals are simply active earlier, one generation may
have time to complete development and allow at
least a partial second generation to occur later in
the year. Turfgrass is an available and consistent
food source for grubs throughout the year in
southern Florida, but warm season grasses de-
cline in the fall and transition back in the spring
in northern Florida. Insect development time also
may be affected by changes in fertilization and
watering practices in winter months compared to
summer months. In addition, individuals or popu-
lations could diapause during adverse conditions.
A second flight peak of several scarab species may
not have been detected in studies specifically done

on sugarcane fields if flooding during the summer
or early fall was done to control grub populations
(Cherry 1984), if cane height reduced black light
visibility, or if the crop had been harvested.
More information is needed on the biology,
damage potential, and management of these key
scarabs. White grub populations are increasing in
importance in Florida turfgrass production and
maintenance industries. Older, broad-spectrum
insecticides, which may have kept white grub
numbers below damaging levels, have been re-
placed with products which lack efficacy against
root-feeding scarabs (e.g., pyrethroids, fipronil).
Management strategies based on application tim-
ing determined in more northern states have not
provided satisfactory results in Florida. According
to the data in this study, most scarab adults are
active from April to June, which may be the most
appropriate timing for preventive insecticide ap-
plications against young white grubs, if needed.

Field and/or laboratory assistance was provided by
B. Cabrera, L. Wood, P. Ruppert, S. Bledsoe, R. Davis, S.
Saha, A. Vincent, T. Duperron, and Y. Wang. M.C. Tho-
mas and P. Skelley, Florida Department of Consumer
Services, Division of Plant Industry, kindly provided
taxonomic assistance. An earlier draft of this manu-
script was thoughtfully reviewed by M. Branham, R.
Cave, D. Shetlar, and M. Thomas. Partial funding for
this project was provided by Bayer Environmental Sci-

A. Hybosorus illigeri (GanesviLe)

6 4 N ,

Ll I- ...

ence. This is the Florida Agricultural Experiment Sta-
tion Journal Series No. R-10985.


J. H. FRANK. 2002. American Beetles, Volume 2:
Polyphaga: Scarabaeoidea through Curculionoidea.
CRC Press. 880 pp.
BRIMLEY, C. S. 1938. The insects of North Carolina.
North Carolina Dept. of Agric. 560 pp.
BUCKINGHAM, G. R., AND C. A. BENNETT. 1989. Dyscine-
tus morator (Fab.) (Coleoptera: Scarabaeidae) adults
attack waterhyacinth, Eichhornia crassipes (Ponted-
eriaceae). Coleopterist Bull. 43: 27-33.
CHERRY, R. H. 1984. Flooding to control the grub Ligy-
rus subtropicus (Coleoptera: Scarabaeidae) in Flor-
ida sugarcane. J. Econ. Entomol. 77: 254-257.
CHERRY, R. H. 1985. Seasonal phenology of white grubs
(Coleoptera: Scarabaeidae) in Florida sugarcane
fields. J. Econ. Entomol. 78: 787-789.
Phyllophaga and related species (Coleoptera: Scara-
baeidae) collected in black-light traps in Alabama
pastures. J. Entomol. Sci. 35(3): 311-326.
FORSCHLER, B. T., AND W. A. GARDNER 1990. A review
of the scientific literature on the biology and distri-
bution of the genus Phyllophaga (Coleoptera: Scara-
baeidae) in the southeastern United States. J.
Entomol. Sci. 25: 628-651.
FORSCHLER, B. T., AND W. A. GARDNER 1991. Flight ac-
tivity and relative abundance of phytophagous Scar-
abaeidae attracted to blacklight traps in Georgia. J.
Agric. Entomol. 8(3): 179-187.
GAYLOR, M. J., AND G. W. FRANKIE. 1979. The relation-
ship of rainfall to adult flight activity; and of soil
moisture to oviposition behavior and egg and first in-
star survival in Phyllophaga crinita. Environ. Ento-
mol. 8: 591-594.
GORDON, R. D., AND D. M. ANDERSON. 1981. The species
of Scarabaeidae (Coleoptera) associated with sugar-
cane in south Florida. Florida Entomologist 64(1):
studies of the biology of the Cuban May beetle. Uni-
versity of Florida, Department of Entomology, Final
Report Grant 12-14-100-8030 (33). 93 pp.
HALL, D. G. 1987. Seasonal flight activity of adult sug-
arcane grubs in Florida. J. Amer. Soc. Sugar Cane
Technol. 7: 39-42.
HARPOOTLIAN, P. J. 2001. Scarab Beetles (Coleoptera:
Scarabaeidae) of South Carolina. Clemson Univer-
sity Public Service Publishing, Clemson, SC. 157 pp.

March 2006

HAYES, W. P. 1917. Studies on the life history of Ligyrus
gibbosus DeG. (Coleoptera). J. Econ. Entomol. 10:
HAYES, W. P. 1925. A comparative study of the history
of certain phytophagous scarabaeid beetles. Kansas
Agric. Exp. Sta. Tech. Bull. 15. 146 pp.
LUGINBILL, P., AND H. R. PAINTER 1953. May beetles of
the United States and Canada. United States Depart-
ment of Agriculture Technical Bulletin 1060: 1-102.
OCAMPO, F. C. 2002. Hybosorids of the United States
and expanding distribution of the introduced species
Hybosorus illigeri (Coleoptera: Scarabaeoidea: Hybo-
soridae). Ann. Entomol. Soc. Am. 95(3): 316-322.
POTTER, D. A. 1981. Seasonal emergence and flight of
northern and southern masked chafers in relation to
air and soil temperatures and rainfall patterns. En-
viron. Entomol. 10: 793-797.
POTTER, D. A. 1998. Destructive Turfgrass Insects: Biol-
ogy, Diagnosis, and Control. Ann Arbor Press, Inc.,
Chelsea, MI. 344 pp.
REINERT, J. A. 1979. Response of white grubs infesting
bermudagrass to insecticides. J. Econ. Entomol. 72:
RITCHER, P. 0. 1966. White Grubs and Their Allies: A
Study of North American Scarabaeoid Larvae. Ore-
gon State University Press, Corvallis. 219 pp.
SMYTH, E. G. 1915. Report of the south coast laboratory.
4th Rept. Board Comm. Agr. Puerto Rico (July 1914-
June 1915): 45-50.
STAINES, C. L., JR. 1990. Scientific note Dyscinetus mor-
ator (Coleoptera: Scarabaeidae) feeding on roots of
azaleas (Rhododendron spp.). Entomological News.
101(2): 98.
Turfgrass Insects of the United States and Canada.
Cornell University Press, Ithaca, NY. 422 pp.
WOODRUFF, R. E. 1970. The "rice beetle," Dyscinetus
morator (Fab.) (Coleoptera: Scarabaeidae). Entomol-
ogy Circular 103, Florida Dept. of Agriculture and
Consumer Services, Division of Plant Industry,
Gainesville, FL.
WOODRUFF, R. E. 1973. The Scarab Beetles of Florida.
Arthropods of Florida and Neighboring Land Areas
Vol. 8. Florida Dept. of Agriculture and Consumer
Services, Division of Plant Industry, Gainesville, FL.
200 pp.
WOODRUFF, R. E., AND B. M. BECK. 1989. The Scarab
Beetles of Florida (Coleoptera: Scarabaeidae) Part
II. The May or June Beetles (Genus: Phyllophaga).
Arthropods of Florida and Neighboring Land Areas,
Vol. 13. Florida Dept. of Agriculture and Consumer
Services, Division of Plant Industry. Gainesville, FL.
226 pp.

Florida Entomologist 89(1)

Dobbs & Boyd: Montandoniola moraguesi in the United States


'United States Department of Agriculture, Animal and Plant Health Inspection Service
Plant Protection and Quarantine, Miami Inspection Station, P.O. Box 59-2136, Miami, FL 33159

2United States Department of Agriculture, Agricultural Research Service, Southern Horticultural Laboratory
P.O. Box 287, Poplarville, MS 39470


The exotic anthocorid Montandoniola moraguesi (Puton) was intentionally introduced in
Hawaii and Bermuda for the control of thrips on outdoor plantings of ornamental Ficus.
These successful programs resulted in similar efforts to introduce this predator at several lo-
cations within the continental United States. Such attempts to establish the bug as a com-
ponent of biological control systems aimed at pest thrips apparently have been unsuccessful.
Our surveys and requests for museum records revealed detections of M. moraguesi in four
states: Alabama, Florida, Louisiana, and Mississippi. Circumstances surrounding detec-
tions in Alabama, Louisiana, and Mississippi suggest that viable populations may not cur-
rently exist in those states. M. moraguesi occurs widely throughout peninsular Florida,
wherever outdoor plantings of exotic, ornamental Ficus spp. are found. An updated distribu-
tion ofM. moraguesi is provided along with field observations and new thrips host records.

Key Words: biological control, predator, thrips, Thysanoptera, Florida


El antoc6rido ex6tico Montandoniola moraguesi (Puton) fue introducido intencionalmente
en Hawaii y Bermuda para el control de trips en las siembras de campo de plants ornamen-
tales del g6nero Ficus. Estos programs con exito resultaron en esfuerzos similares para in-
troducir este depredador en various lugares en el continent de los Estados Unidos. Los
intentos para establecer este chinche como un component de un sistema de control biol6gico
dirigido a las plagas de trips aparentemente no se han logrado. Nuestras busquedas y pedi-
dos de registros de museo revelaron que M. moraguesi fue detectado en cuatro estados: Ala-
bama, Florida, Louisiana, y Mississippi. Las circumstancias alredador de las detecciones en
Alabama, Louisiana, y Mississippi sugieren que poblaciones viables tal vez ya no existen en
estos estados. Montandoniola moraguesi esta ampliamente distribuida por la peninsula del
Florida, donde se encuentra siembras de campo de plants ornamentales ex6ticas de Ficus
spp. Se provee una distribuci6n mas actualizada de M. moraguesi adjunto con las observa-
ciones de campo y nuevos registros de los hospederos de trips.

Montandoniola moraguesi (Puton) (Hemiptera:
Anthocoridae) (Fig. 1) is an important predator of
several species of economically important thrips.
Although originally described in France, M. mora-
guesi now is thought to be native to Southeast
Asia (Herring 1967; Lattin 2000). Its reported dis-
tribution is essentially Old World. Populations are
known from Africa (Algeria, Egypt, Morocco,
Senegal, South Africa, Sudan, Western Sahara),
Asia (Japan, Israel, Philippines, Micronesia), Aus-
tralia, Europe (Canary Islands, France, Italy, Por-
tugal, Sicily, Spain) and Bermuda (Carayon & Ra-
made 1962; Funasaki 1966; Herring 1967; Lewis
1973; Muraleedharan 1977; Muraleedharan &
Ananthakrishnan 1978; Pericart & Halperin
1989; Postle et al. 2001). In the Western Hemi-
sphere, it has been reported only from South
America (Muraleedharan & Ananthakrishnan
1978), although it may exist throughout much of

the Caribbean and Latin America. Its prey in-
cludes more than 20 species of gall-forming thrips
(Table 1) from a wide variety of host plants (Mura-
leedharan & Ananthakrishnan 1978).
Because of its broad host range, M. moraguesi
sometimes is a useful biological control agent
against thrips. It has been successfully intro-
duced for the biological control of Gynaikothrips
ficorum (Marchal) (Thysanoptera: Phlaeothripi-
dae) in Bermuda (Leighton 1978) and Hawaii
(Funasaki 1966). In both areas, the bug became
established and provided good, long-term control,
but its establishment in Hawaii has caused biotic
interference (Reimer 1988). In the continental
United States, however, two attempted introduc-
tions in California (1965 and 1996) and at least
one in Texas (1992) apparently have not been suc-
cessful (Clausen 1978; Henry 1988; Paine 1992;
Hanlon & Paine 2003).

Florida Entomologist 89(1)


Fig. 1. Montandoniola moraguesi, dorsal view.

The primary pest target of M. moraguesi in the
United States has been the Cuban laurel thrips,
G. ficorum. Feeding by this thrips, a pest of Chi-
nese banyan Ficus microcarpa L. (Moraceae)
(Paine 1992), causes the leaves to fold upward
into galls where the thrips breeds and forms large
colonies. Recently, a second species G. uzeli Zim-
merman has become established in the United
States (Held et al. 2005). Gynaikothrips uzeli, a
pest of weeping fig, F benjamin (L.), was acci-
dentally introduced into Florida and is now being
spread throughout the southeastern United
States in shipments of ornamental weeping fig
originating from nurseries in South Florida (Held
et al. in press). The primary morphological differ-
ence between these thrips is the relative lengths
of the pronotal posterolateral pair of setae, but a
more practical way to distinguish G. ficorum from
G. uzeli is by host-plant association: G. ficorum

with F microcarpa and G. uzeli with F benjamin
(Mound et al. 1995). Ficus microcarpa survive in
plant zones 9-11, whereas F benjamin survive in
zones 10-11 (Turner & Wasson 1997).
The Cuban laurel thrips occurs in California,
Florida, and Texas (Denmark 1967) in the conti-
nental United States. Even though the thrips has
been known from Florida since at least 1887
(Denmark 1967), M. moraguesi was not detected
in that state until 1990. The discovery of this an-
thocorid in Florida was based on adults and
nymphs collected from curled and deformed Ficus
leaves in Palm Beach County (Bennett 1995). No
records of intentional introductions of M. mora-
guesi in Florida are available and its presence
there might be due to unintentional spread
through commerce or through natural means.
Although M. moraguesi has been detected in
Palm Beach County, Florida, exact locality data
have not been reported. Herein we confirm the es-
tablishment of M. moraguesi in South Florida 15
years after its initial detection, provide updated
information on distribution, report records from
museum searches in several states where the bug
potentially could become established outdoors,
summarize our field observations, and provide
new prey records.


In the continental United States, plant zones
9-11, i.e., those areas capable of supporting out-
door populations of ornamental Ficus spp., en-
compass peninsular Florida, coastal Louisiana,
southern areas of Texas and Arizona, and coastal
California. Based on the premise that the distri-
bution of M. moraguesi coincides with that of its
prey (Bennett 1995), we surveyed these areas
and/or requested specimen data from major ento-
mological museums.
We conducted surveys in Alabama, Arizona,
Florida, Louisiana, Mississippi, and Texas. Various
techniques were employed, including visual in-
spection and shaking of leaves and stems of orna-
mental Ficus spp. over a collecting net. Adults were
aspirated, preserved in alcohol, and transported to
the laboratory for curation and identification.
Museum records were solicited from Alabama
(Auburn University Entomology Museum, Au-
burn), Arizona (Arizona Department of Agricul-
ture, Phoenix; University of Arizona, Tucson),
California (California Academy of Sciences, San
Francisco; California Department of Agriculture,
Sacramento; San Diego Natural History Museum,
San Diego; University of California, Berkeley;
University of California, Davis; University of Cal-
ifornia, Riverside), Florida (Florida State Collec-
tion of Arthropods, Gainesville), Louisiana (Loui-
siana State University, Baton Rouge), Mississippi
(Mississippi State University, Starkville), Texas
(Texas A & M University, College Station), and

March 2006

Dobbs & Boyd: Montandoniola moraguesi in the United States


Thrips prey Host plant Reference

Alcothrips hadrocerus (Karny)
Androthrips flauipes Schmutz
Androthrips ramachandrai Karny
Aneurothrips punctipennis Karny
Arrhenothrips dhumrapaksha Ramak.
Arrhenothrips ramakrishnae Hood
Austrothrips cochinchinensis Karny
Brachythrips dantahasta Ramak.
Cercothrips nigrodentatus (Karny)
Crotonothrips gallarum Anan.
Frankliniella occidentalis (Pergande)
Gynaikothrips bengalensis Anan.
Gynaikothrips ficorum (Marchal)
Gynaikothrips flaviantennatus Moulton
Gynaikothrips malabaricus Ramak.
Gynaikothrips uzeli Zimm.
Holopothrips sp.
Liothrips ramakrishnae Anan. & Jag.
Liothrips africanus Vuil.
Liothrips citricornis Anan.
Liothrips ,.;......r. Bourn.
Liothrips indicus Anan.
Liothrips oleae Costa
Liothrips pallicrus (Karny)
Liothrips pallipes (Karny)
Liothrips urichi Karny
L ..- i, i... jambuvasi (Anan.)
Mesothrips extensivus Anan. & Jag.
Mesothripsjordani Zimm.
Nesothrips sp.
Phorinothrips loranthi Anan.
Psenothrips priesneri (Anan.)
Schedothrips orientalis Anan.
Tetradothrips foliiperda (Karny)
Teuchothrips longus Priesner
Thrips sp.
Thrips tabaci (Lindeman)
Trichothrips houardi Vuil.

Gymnosporia sp.
Ficus microcarpa
Cordia sp.
Ficus bengalensis
Mimusops elengi
Calycopterus floribundus
Memecylon sp.
Planchona valida
Memecylon sp.
Ficus benjamin
Ficus microcarpa
Cassearia tomentosa
Ficus bengalensis
Ficus benjamin
Tabebuia pallida
Schefflera racemosa
Guierra senegalensis
Maytenus sengalensis
. ..-.. .r virosa
Maytenus senegalensis
Olea europea
Vitis sp.
Peperomia sp.
Clidemia hirta
Terminalia sp.
Mallotus phillipinus
Ficus benjamin
Ficus aurea
Loranthus sp.
Walsura piscidea
Ventilago maderaspatana
Pothos scandans
Pavetta hispidula
Ficus craterostoma
Guierra senegalensis

Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
FSCA1 E2002-1796
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1971
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Sabelis & Van Rijn 1997
Muraleedharan & Ananthakrishnan 1978
Mound et al. 1995
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Mound et al. 1995
FSCA1 E2002-5207
Muraleedharan & Ananthakrishnan 1978
Carayon & Ramade 1962
Muraleedharan & Ananthakrishnan 1971
Carayon & Ramade 1962
Muraleedharan & Ananthakrishnan 1978
Carayon & Ramade 1962
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Reimer 1988
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
FSCA1 E2001-2090
Muraleedharan & Ananthakrishnan 1978
Muraleedharan & Ananthakrishnan 1978
Varadarasan and Ananthakrishnan 1981
Muraleedharan & Ananthakrishnan 1978
Varadarasan and Ananthakrishnan 1981
Carayon & Ramade 1962
Sabelis & Van Rijn 1997
Carayon & Ramade 1962

'Florida State Coll. of Arthropods, Florida Dept. Agr. and Cons. Serv, Div. of Plant Ind.

the District of Columbia (National Museum of
Natural History, Smithsonian Institution, Wash-
ington, D.C.). We also solicited data from the col-
lections of John D. Lattin (retired) (Oregon State
University, Corvallis, OR) and Tamera Lewis
(USDA, ARS, Wapato, WA), both of whom have
collected anthocorids from southern California.
Acronyms used are FSCA (Florida State Col-
lection ofArthropods, Gainesville, FL), LSU (Lou-
isiana State University Entomology Museum, Ba-
ton Rouge, LA), MIS (United States Department
of Agriculture, Animal and Plant Health Inspec-
tion Service, Plant Protection and Quarantine,
Miami Inspection Station, Miami, FL), NMNH
(United States National Museum of Natural His-

tory, Washington, D.C.), and SHL (United States
Department of Agriculture, Agricultural Re-
search Service, Southern Horticultural Labora-
tory, Poplarville, MS).


Based on field searches and museum records,
specimens of M. moraguesi are reported from Al-
abama, Florida, Louisiana, and Mississippi (Fig.
2). In Alabama and Mississippi, it was taken only
in retail garden centers, therefore, field popula-
tions might not occur in those states. Little is
known concerning the circumstances surround-
ing the single specimen housed at LSU. If popula-

Florida Entomologist 89(1)

M.Z) AL: Mobile Co.

LA: E. Baton Rouge Par. Nov 2004, first
Nov 1994, first collection collection in Alabama
in Louisiana MS: Pearl River Co.

Nov 2004, first
collection in Mississippi

,FL: Palm Beach Co.
Nov 1990, first
collection in Florida

Fig. 2. Current distribution of Montandoniola moraguesi in the southeastern United States based on field cap-
tures and museum records.

tions persisted in that state, one might expect the
LSU collection to contain additional specimens.
The following locality label data are provided
for M. moraguesi in the continental United States:
Alabama: Mobile Co., Mobile, Home Depot Garden
Center, 30.674N, 88.224W, 19, Ficus benjamin
infested with Gynaikothrips uzeli, 12-XI-2004, D.
Boyd (SHL). FLORIDA: ALACHUA CO., 36 12,
on Ficus sp., 21-V-2005, J. Brambila (MIS); Flor-
ida: Brevard Co., Indialantic, 1 Ficus retusa, 23-
XII-1991, K. Garret-Kraus (NMNH); Broward Co.,
Pompano Beach, 10 specimens, pred. on Gynaiko-
thrips ficorum, 30-VIII-1991, F. D. Bennett
(FSCA); Pompano Bch, 66, 69, Pr,-d G,..;.,...
thrips ficorum/Ficus, 19-XI-1991, F. D. Bennett
(FSCA); Pompano Beach, 6 specimens, pred. on
Gynaikothrips ficorum on Ficus sp., 19-XI-1991, F.
D. Bennett (FSCA); Collier Co., Everglades City, 2
specimens, ex Gynaikothrips ficorum on Ficus sp.,
3-V-1992, F. D. Bennett (FSCA); Hillsborough Co.,
Tampa, Busch Gardens, 36, 1 on Gynaikothrips
ficorum on Ficus, 6-XI-1992, F. D. Bennett
(USNM); Lee Co., Ft. Myers, 7 host Gynaiko-
thrips ficorum on Ficus, 2-V-1992, F. D. Bennett
(NMNH); Manatee Co., Bradenton, 26, 99, host
Gynaikothrips ficorum, on Ficus, 8-XI-1992, F. D.
Bennett (NMNH); Martin Co., Stuart, 76, 89,

host Gynaikothrips ficorum/Ficus microcarpa,
12-VIII-1992, F. D. Bennett (NMNH); Miami-
Dade Co., Miami, 3 specimens, 143 Ave., ex Ficus
aurea, 15-V-2001, Ed Putland FSCA# E2001-2090
(FSCA); Miami, SW 137 Ave. and 172 St., 6 speci-
mens, ex Tabebuia pallida, 14-X-2002, Holly
Glenn, FSCA# E2002-5207 (FSCA); Miami, 68 St.
and 102 Ave., 3 specimens, sweep net, 7-IV-2004,
J. Durand (FSCA); Miami, 68 St. and 102 Ave., 1
specimen, sweep net, 31-III-2004, J. Durand
(FSCA); Goulds, SW 232 Ave., 7 specimens ex
Ficus benjamin, 21-IV-2004, Eduardo Camero,
FSCA# E2004-2958 (FSCA); Homestead 232 St.
and 137 Ave., 1 specimen, ex Ficus microcarpa, 9-
V-2002, Mario Hernandez FSCA# E2002-1796
(FSCA); Miami, 143 Ave., on Ficus aurea Nutt.,
15-V-2001, Ed Putland, FSCA# E2001-2090
(FSCA); Miami, 68 St. at 102 Ave., 36, 22, ex Fi-
cus sp., 31-III-2004, J. Durand (MIS); Miami, 68
St. at 102 Ave., 36, 19, ex Ficus sp., 7-IV-2004,
J. Durand (MIS); Miami, 68 St. at 102 Ave., 76,
29, ex Ficus sp., 17-II-2005, T. Dobbs (MIS);
MONROE CO., Key Largo, 91421 U.S. 1, 16, 29,
ex Ficus sp. with Gynaikothrips sp., 5-IV-2005, T.
Dobbs (MIS); Key Largo, 103880 U.S. 1, 3 nymphs
ex Ficus sp. with Gynaikothrips sp., 5-IV-2005, T.
Dobbs (MIS); Palm Beach Co., West Palm Beach, 4

March 2006

Dobbs & Boyd: Montandoniola moraguesi in the United States

specimens, pred. on Gynaikothrips ficorum on
Ficus microcarpa, 23-III-1992, F. D. Bennett
(FSCA); West Palm Beach, 56, 5Y, on Gynaiko-
thrips ficorum Ficus microcarpa, 23-III-1992, F.
D. Bennett (NMNH); Pinellas Co., St. Petersburg,
9c, 119, host Gynaikothrips ficorum on Ficus, 6-
XI-1992, F. D. Bennett (NMNH). LOUISIANA: E.
Baton Rouge Par., Baton Rouge, 1 specimen, on Fi-
cus, 30-IX-1994, J. W. Tessmer (LSU). MISSIS-
SIPPI: PEARL RIVER CO., Poplarville, 1 speci-
men, pred. G. uzeli on F benjamin, 15-XI-2004,
D. W. Boyd (SHL); Poplarville, 1 specimen, pred on
G. uzeli on F benjamin, 14-XII-04, D. Held
(SHL); Poplarville, 1 specimen, pred. G. uzeli on F
benjamin, 05-1-2005, D. W. Boyd (SHL). We were
unable to locate the original specimens detected in
Palm Beach County in 1990.
Based on data from the Florida State Collec-
tion of Arthropods, Florida Department of Agri-
culture and Consumer Services, Division of Plant
Industry, M. moraguesi is associated for the first
time with Androthrips ramachandrai Karny on
F microcarpa, Holopothrips sp. on Tabebuia pal-
lida (Lindl.) Miers, and Nesothrips sp. on Ficus
aurea Nutt. (Table 1).
As stated by Bennett (1995), M. moraguesi is
widespread in Florida where outdoor plantings of
exotic ornamental Ficus spp. occur, and has now
been detected as far north as Gainesville. In Mi-
ami-Dade Co., the bugs were observed in direct
association with their thrips prey and were most
easily detected by searching for untrimmed Ficus
hedges with upcurled leaves. The anthocorids en-
ter and remain in the curled leaves while feeding
on all life stages of the thrips. Populations of Gy-
naikothrips and M. moraguesi were quite high in
some instances, yet the plants we observed nearly
always had significant new growth and showed no
outward signs of ill health aside from moderate
leaf distortion.
In Alabama, an adult M. moraguesi and associ-
ated nymphal exuviae were taken on a Ficus ben-
jamina plant in a retail garden center in Mobile
Co. No other specimens were located from that
state. The plants at the garden center were ob-
tained from a nursery in South Florida, and we as-
sume that the M. moraguesi may have hitchhiked
with plant material shipped from Florida to Ala-
bama. Two adults and a nymph of M. moraguesi
were collected in Pearl River Co., Mississippi, on F
benjamin plants infested with G uzeli. The
plants were traced to local retail nurseries that
had, similar to the case in Alabama, originally re-
ceived plant material from South Florida. An adult
was captured in East Baton Rouge Parish, Louisi-
ana, in 1994, two years after intentional releases
in neighboring Texas. This record predates by a full
decade similar finds in nearby Alabama and Mis-
sissippi. Whether the later records reflect a lack of
concentrated collecting in the interim is unknown.
We found no field populations of M. moraguesi in

any other states, nor did we find museum speci-
mens from other states. We found no specimens of
M. moraguesi from California or Texas, although
the bug has been introduced into those states to
control thrips on Ficus plantings (Bennett 1995;
Hanlon & Paine 2003). Even though outdoor Ficus
plantings with suitable thrips hosts are found in
other states, we suggest that in the continental
United States field populations of M. moraguesi
currently are restricted to peninsular Florida. Fur-
ther investigation will clarify this.
Collection of this anthocorid in Alabama and
Mississippi on plants shipped from Florida indi-
cates its potential for spread through commercial
trade. Its establishment along the Gulf Coast
could provide needed biological control of G. fi-
corum and G. uzeli. However, M. moraguesi has
been implicated in biotic interference in at least
two cases (Reimer 1988; Bennett 1995; Hanlon &
Paine 2003) and potentially can feed on thrips be-
ing used for the biological control of weed species.


The authors express gratitude to the following indi-
viduals for checking their respective institutional collec-
tions for specimens of M. moraguesi: Chris Baptista,
Arizona Department of Agriculture; Cheryl Barr, Uni-
versity of California, Berkeley; Victoria Bayless, Louisi-
ana State Arthropod Museum; Charles Bellamy,
California Department of Food and Agriculture; Paisley
Cato, San Diego Natural History Museum; Wayne
Clark, Auburn University; Susan Halbert, Florida State
Collection of Arthropods; Thomas Henry, National Mu-
seum of Natural History; Steven Heydon, University of
California, Davis; John Lattin, Oregon State University;
Tamera Lewis, USDA, ARS, Wapato, WA; Carl Olsen,
University of Arizona; Norman Penny, California Acad-
emy of Sciences; Edward Riley and Joseph Schaffner,
Texas A&M University. In addition, we thank Julieta
Brambila, Susan Halbert, Thomas Henry, John Lattin,
and A. G. Wheeler, Jr. for suggesting improvements to
the manuscript, Michael Ferro, Louisiana State Arthro-
pod Museum, for the photograph of M. moraguesi, Tho-
mas Henry for taxonomic support, and Susan Halbert
for providing M. moraguesi host records in Florida.

BENNETT, F. D. 1995. Montandoniola moraguesi (Hemi-
ptera: Anthocoridae), a new immigrant to Florida:
Friend or foe? Vedalia 2: 3-6.
CARAYON, J., AND F. RAMADE. 1962. Note sur la
presence en France et en Italie de Montandoniola
moraguesi (Puton) avec quelques observations sur
cet H6t6ropt6re Anthocorid6. Bulletin de la Soci6t6
Entomologique de France 67: 207-211.
CLAUSEN, C. P. 1978. Phlaeothripidae. Cuban Laurel
Thrips, pp. 18-19 In C. P. Clausen [ed.], Introduced
Parasites and Predators of Arthropod Pests and
Weeds. A World Review. U.S. Dept. Agric. Agric.
Res. Serv. Agric. Handb. 480.
DENMARK, H. A. 1967. Cuban Laurel Thrips, Gynaiko-
thrips ficorum, in Florida. Florida Dept. Agric. Ento-
mol. Circ. 59: 1-2.

FUNASAKI, G. Y. 1966. Studies on the life cycle and prop-
agation technique ofMontandoniola moraguesi (Pu-
ton) (Heteroptera: Anthocoridae). Proc. Hawaiian
Entomol. Soc. 19: 209-211.
HANLON, C. C., AND T. D. PAINE. 2003. Biological control
of Cuban laurel thrips (Thysanoptera: Phlaeothripi-
dae) in California. 1st International Symposium on Bi-
ological Control ofArthropods, FHTET-03-05: 474-478.
2005. Gynaikothrips uzeli (Thysanoptera: Phlaeo-
thripidae) in the Southeastern United States: Distri-
bution and Review of Biology. Florida Entomol. 88:
HENRY, T. J. 1988. Family Anthocoridae Fieber, 1837,
pp. 12-28 In T. J. Henry and R. C. Froeschner [eds.],
Catalog of the Heteroptera, or True Bugs, of Canada
and the Continental United States. St. Lucie Press.
Boca Raton, FL. 958 pp.
HERRING, J. L. 1967. Heteroptera: Anthocoridae. In-
sects of Micronesia 7: 391-414.
LATTIN, J. D. 2000. Minute pirate bugs (Anthocoridae),
pp 607-637 In C. W. Schaefer and A.R. Panizzi [eds.].
Heteroptera of Economic Importance. CRC Press.
Boca Raton, FL. 828 pp.
LEIGHTON, D. 1978. Thrips on Indian laurel. Bermuda
Dept. of Agric. and Fish. Mo. Bull. 48: 75-77.
LEWIS, T. 1973. Thrips, Their Biology, Ecology and Eco-
nomic Importance. Academic Press, London. 740 pp.
MOUND, L. A., C. WANG, AND S. OKAJIMA. 1995. Obser-
vations in Taiwan on the identity of the Cuban laurel
thrips (Thysanoptera: Phlaepothripidae). J. New
York Entomol. Soc. 103: 185-190.
MURALEEDHARAN, N. 1977. Some genera of Anthocori-
nae (Heteroptera: Anthocoridae) from south India.
Entomon 2: 231-235.

March 2006

1971. Bionomics of Montandoniola moraguesi (Pu-
ton) (Heteroptera: Anthocoridae), a predator on gall
thrips. Bull. Entomol. 12: 4-10.
1978. Bioecology of four species of Anthocoridae
(Hemiptera: Insecta) predaceous on thrips with key
to genera of anthocorids from India. Records of the
Zoological Survey of India 11: 1-32.
PAINE, T. D. 1992. Cuban laurel thrips (Thysanoptera:
Phlaeothripidae) biology in southern California: sea-
sonal abundance, temperature dependent develop-
ment, leaf suitability, and predation. Ann. Entomol.
Soc. Am. 85: 164-172.
PERICART, J., AND J. HALPERIN. 1989. The Anthocoridae
of Israel (Heteroptera). Phytoparasitica 17: 91-98.
Oriini (Hemiptera: Anthocoridae) new to Australia.
Aust. Jour. Entomol. 40: 231-244.
REIMER, N. J. 1988. Predation on Liothrips urichi
Karny (Thysanoptera: Phlaeothripidae): a case of bi-
otic interference. Environ. Entomol. 17: 132-134.
SABELIS, M. W., AND P. C. J. VAN RIN. 1997. Predation
by insects and mites, pp. 259-354 In T. Lewis [ed.],
Thrips as Crop Pests. CAB International, New York.
TURNER, R. G., JR., AND E. W. WASSON (eds.). 1997. Bo-
tanica: The Illustrated A-Z of Over 10,000 Garden
Plants and How to Cultivate Them. Random House
Australia, Publishers. 1020 pp.
Population dynamics and prey-predator/parasite re-
lationships of gall-forming thrips. Proc. Indian Nat.
Acad. B 47: 321-340.

Florida Entomologist 89(1)

Chu et al.: CC Traps for S. dorsalis


'USDA, ARS Western Cotton Research Laboratory, Phoenix, AZ 85040
E-mail: cchu@wcrl.ars.usda.gov.

2USDA, APHIS, PPQ, CPHST Pest Detection, Diagnostics and Management Laboratory, Edinburg, TX 78541

3National Pingtung University of Science and Technology, Neipu, Pingtung, 91201 Taiwan

4Ministry of Agriculture and Fisheries, St. Vincent and the Grenadines, Kingstown, St. Vincent, W.I.


Scirtothrips dorsalis (Hood) (Thysanoptera: Thripidae) is a recently identified invasive pest
to the Caribbean and poses a significant threat to agriculture and trade in the region. Meth-
ods are needed to detect the presence and to monitor populations of this pest so that it can
be effectively managed. Three different CC trap base colors (blue, yellow, and white) with or
without dichlorvos as a killing agent, and a newly developed and named the Blue-D trap
were studied in Taiwan and St. Vincent for attraction and capture of S. dorsalis. In lemons
in Taiwan, mean numbers of S. dorsalis caught in Blue-D traps were greater compared with
dichlorvos cube modified CC traps. In St. Vincent chili pepper plantings, the Blue-D traps
caught more Thrips palmi (Karny), Frankliniella sp., and Microcephalothrips abdominalis
(Crawford) than dichlorvos cube modified CC traps. More Frankliniella intonsa (Trybom),
Megalurothrips usitatus (Bagnall), T palmi, Frankliniella sp., and M. abdominalis were
caught in blue and white base CC traps than yellow base CC traps. Average captures per CC
trap per week were 0.07 and 0.02-0.09 S. dorsalis in Taiwan and St. Vincent, respectively.
There were no differences in S. dorsalis captures in white, blue, or yellow base CC traps. The
average weekly S. dorsalis catch for yellow sticky card traps was 19.8. CC traps can be used
for detection of S. dorsali and collecting intact S. dorsalis for taxonomic and genetic deter-
minations when a few of the species are found in a large commercial production area.Yellow
sticky traps can be used for monitoring S. dorsalis populations. A combination detecting sys-
tem of visual observation, yellow sticky traps, and CC traps may be an effective S. dorsalis
population detecting and monitoring system.

Key Words: Scirtothrips dorsalis, Frankliniella occidentalis, Thrips palmi, CC traps, Carib-
bean area


Scirtothrips dorsalis (Hood) (Thysanoptera: Thripidae) es una plaga invasora recien identi-
ficada en el Caribe y represent una amenaza significativa a la agriculture y comercio de la
region. Es necesario desarrollar m6todos para detectar la presencia de esta plaga y realizar
un monitoreo de su poblacion para lograr un manejo mas eficaz. Trampas de CC de tres co-
lores diferentes (azul, amarilla, y blanca) con o sin el pesticide dichlorvos como agent para
matar, y una trampa reci6n desarrollada y nombrada 'Blue-D' fueron estudiadas en Taiwan
y St. Vincent para ver su habilidad para atraer y capturar S. dorsalis. En limones en Taiwan,
el promedio del numero de S. dorsalis capturados en las trampas de Blue-D fue mas alto
comparado con las trampas de CC cubicas modificadas con dichlorvos. En siembras del chile
verde en St. Vincent, la trampa de Blue-D capture mas Thrips palmi (Karny), Frankliniella
sp., y Microcephalothrips abdominalis (Crawford) que la trampa de CC cubica modificada
con dichlorvos. Habian un mayor numero de Frankliniella intonsa (Trybom), Megalurothrips
usitatus (Bagnall), T palmi, Frankliniella sp. y M. abdominalis capturadas en trampas de
CC con la base de color azul o blanco que en las trampas con la base amarilla. El promedio
de los S. dorsalis capturados en las trampas de CC por semana fue 0.07 y 0.02-0.09 en
Taiwan y St. Vincent, respectivemente. No hubo ninguna diferencia en el numero de S. dor-
salis capturados en trampas de CC con la base de color blanco, azul o amarilla. El promedio
semanal de los S. dorsalis capturados con trampas de tarjetas pegajosas amarillas fue 19.8.
Se puede usar las trampas de CC para detectar la presencia de S. dorsalis y recolectar espe-
cimenes de S. dorsalis intactos para su identificaci6n taxon6mica y gen6tica cuando sola-
mente se encuentran pocos species en las areas grandes de producci6n commercial. Se puede
usar las trampas amarillas pegajosas para realizar un monitoreo de la poblaci6n de S. dor-

Florida Entomologist 89(1)

salis. Un sistema de detecci6n que combine la observaci6n visual, las trampas amarillas
pegajosas, y las trampas de CC puede ser efectivo para detectar y realizar monitoreos de
poblaciones de S. dorsalis.

Scirtothrips dorsalis (Hood) was described in
1916 as a new species collected from castor bean
and chili plants in Coimbatore, Southern India
(Hood 1919). S. dorsalis are polyphagous pests
that are widespread in habitat ranging from tem-
perate to tropical climate regions in Pakistan, Ja-
pan, and Australia (Mound & Palmer 1981). Pri-
mary hosts are onion, cashew nut, tea, chili, cot-
ton, tomato, mango, tobacco, and castor bean. The
insect has been reported by the Animal Plant
Health Inspection Service (APHIS) as one of the
thirteen most important pest species that could
become a serious threat to United States (US) ag-
ricultural crops if it becomes established in the
country (USDA-APHIS 2004). The Florida Nurs-
erymen and Growers Association (FNGA) also
consider S. dorsalis as an exotic pest with high po-
tential to damage their industry if it becomes es-
tablished in the state (FNGA 2003). Since 1984,
USDA-APHIS inspectors at various US ports of
entry have reported finding live S. dorsalis a total
of 89 times from imported plant materials of 48
plant taxa (USDA 2003). On July 6, 2003, a Plant
Protection and Quarantine (PPQ) officer in Mi-
ami, FL intercepted live S. dorsalis on chili pep-
pers shipped from St. Vincent, a Caribbean island
nation (Skarlinsky 2003). Subsequently, surveys
of St. Vincent and St. Lucia confirmed the pres-
ence of S. dorsalis on both islands (Ciomperlik &
Seal 2004). Thrips samples collected from both is-
lands were catalogued and submitted to the
USDA-Agricultural Research Service (ARS) Sys-
tematic Entomology Laboratory, Beltsville, MD,
and the Australian Commonwealth Scientific &
Industrial Research Organization (CSIRO) Ento-
mology Laboratory, Canberra, Australia.
A pest risk assessment by Venette & Davis
(2004) indicates that the potential geographic dis-
tribution ofS. dorsalis in the U.S. ranges from the
northeastern Atlantic area to Minnesota in the
northern latitudes and to Texas in the south.
Meissner et al. (2005) indicates that permanent
establishment would likely be limited to southern
and West Coast states. The species also seems ca-
pable of spreading throughout the entire Carib-
bean region. So far, it has become established on
the Caribbean islands of St. Lucia, St. Vincent
(Ciomperlik & Seal 2004), and Trinidad (USDA
Offshore Pest Information System 2004).
Current survey, detection, and monitoring
methods for S. dorsalis are laborious and require
significant manpower and technical training.
Most species in the genus Scirtothrips are pale in
color, minute, and must be cleared and slide
mounted for species identification. In addition,
the genus is confused taxonomically. The names

Heliothrips minutissimus (Bagnall), Anapho-
thrips andreae (Karny), Scirtothrips padmae (Ra-
makrishna), and S. fragaiae (Girault) appear in
the literature but are now considered to be syn-
onyms ofS. dorsalis (Mound & Palmer 1981). On-
going research and development methods that in-
corporate Polymerase Chain Reaction-Restriction
Fragment Length Polymorphism (RFLP-PCR) ge-
netic techniques (Toda & Komazaki 2002) are be-
ing conducted to explore the ribosomal ITS2 DNA
regions that can be used to rapidly distinguish be-
tween thrips species. These methods require that
individual insect samples be whole and undam-
aged, free of foreign contaminating substances,
and preferably without contaminant DNA. Based
on these requirements, sticky traps often used for
thrips detection and monitoring are unsuitable.
To obtain specimens for taxonomic and genetic
studies, terminal leaf samples can be collected in
ziplock bags and washed with ethanol to obtain
full intact specimens (Ciomperlik & Seal 2004).
The plastic cup trap (named CC trap) (Fig. 1) also
collects intact S. dorsalis specimens. It can be in-
stalled in a commercial production area where
the pest has been found or suspected and serviced
periodically for long periods of time. The CC trap
was designed and validated for monitoring sweet-
potato whitefly, Bemisia tabaci (Gennadius) B-
biotype, populations. Trap design was based on
the sweetpotato whitefly attraction to the color
yellow, flight patterns approaching plants, and
landing behavior (Chu et al. 1995; Chu & Henne-

1 External
2 Entrance
3 Internal
4 Plate
5 Cup inside

Fig. 1. CC trap with number trap surface identifica-

March 2006

Chu et al.: CC Traps for S. dorsalis

berry 1998). The white base CC traps caught more
S. dorsalis and Thrips palmi (Karny) in a peanut
field in India in 1996 than yellow base traps (Chu
et al. 2000). To extend the usefulness of the CC
traps for detection and monitoring S. dorsalis,
T palmi, and other thrip species under Carib-
bean island conditions, we are studying different
methods to increase trap efficacy.
White, yellow, and blue colors have been re-
ported attractive to S. dorsalis, T palmi, or F oc-
cidentalis (Beavers et al. 1971; Gillespie & Ver-
non 1990; Cho et al. 1995; Tsuchiya et al. 1995;
Chu et al. 2000; Hoddle et al. 2002; Chen et al.
2004). Chu et al. (2005) have modified CC white-
fly traps for detecting and monitoring western
flower thrips.
Objectives of the current study were (1) to
evaluate CC thrip trap modification with a killing
agent and a specimen preservative, (2) to evalu-
ate a modification of a commercially available
dichlorvos strip package for use as a thrips trap,
named the Blue-D trap in this report, and (3) to
evaluate sticky card traps.


Comparison of CC and Blue-D traps

The study was conducted in commercial farms
with randomized complete block designs with 15
and five replicates in Taiwan and St. Vincent,
respectively. Treatments were re-randomized
weekly in St. Vincent, but not in Taiwan. The four
CC trap treatments in Taiwan were trap base col-
ors (white, blue, and yellow with a 1 cm2 dichlor-
vos cube and 15 ml, 10% ethylene glycol) and the
Blue-D trap. In St.Vincent, the three trap base
colors with or without dichlorvos cube and with or
without ethylene glycol made a total of 12 treat-
ments. Blue-D trap was the 13th treatment. The
dichlorvos cubes in CC traps were not replaced
during the experimental periods. The ethylene
glycol treatment was used to preserve thrip spec-
imens. The CC traps were serviced weekly, disas-
sembled in the field, rinsed with 20 ml 85% etha-
nol to dislodge insects that were retained in the
trap bases, and stored in labeled glass vials for
identification in the laboratory.
The Blue-D traps were Hot Shot No-Pest
Strip dispensers (United Industry Corp., St.
Louis, Mo.) fitted with plastic bags attached to the
dispenser bottoms for collecting dead insects. Ver-
tically oriented blue plastic strips (two 2.5-cm
wide strips spaced at 2.5 cm apart) were attached
inside the front and back surfaces of the dichlor-
vos dispensers packages (Fig. 2). The Blue-D
traps were replaced weekly.
Experiment 1. Taiwan (dry season). The exper-
iment was conducted in a 0.6-ha lemon (Citrus li-
mon L., cv. Eureka) orchard for eight weeks from
26 November 2004 to 30 January 2005 in Neipu,

Fig. 2. Dichlorvos dispenser modification with blue
stripe and sample collecting bag placed in the top can-
opy of a lemon tree in Neipu, Pingtung County, Taiwan.

Taiwan. The traps were suspended in trees 1.9 m
above the ground.
Experiment 2. St. Vincent (wet season). The
experiment was conducted about one mile inland
from Caribbean sea in Georgetown, St. Vincent
Island in two (~0.2 ha each) geographically sepa-
rated chili pepper (Capsicum chinense L.) fields.
Plants were set 1 m within and between row spac-
ings in both fields. Plants in one field were Scotch
Bonnet variety and in the other field the West In-
dies Red variety. Traps were placed 1 m apart on
wooden stakes placed within the plant rows. The
trap bases were located about 22 cm below the
tallest plant terminals. The experiment was con-
ducted for six weeks from 14 October to 29 No-
vember, 2004.
Experiment 3. St. Vincent (dry season). The ex-
periment was conducted in two Scotch Bonnet va-
riety (~0.2 ha each) fields, as described in Experi-
ment 2. The experiment was conducted for six
weeks from 23 March to 4 May, 2005.
All thrips in the samples were identified and
counted. Scirtothrips dorsalis was readily sepa-
rated from the other species by the small size (0.7-
1.0 mm), pale yellow color, and the presence of mi-
crotrichia extending along the median area of the
abdominal sternites (Hoddle & Mound 2003). The
remaining species were separated from each other

Florida Entomologist 89(1)

by characters found in Nakahara's key to Thripi-
dae (unpublished). Representative individuals of
each thrips species were slide mounted to confirm
the species identity. Voucher specimens from St.
Vincent were deposited in the USDA-ARS Sys-
tematic Entomology Laboratory and in the St.
Vincent Ministry of Agriculture and Fisheries.

Seasonal Weather during the Wet and Dry Seasons in
St. Vincent, 2004-2005

Daily rainfall data and air temperatures dur-
ing both seasons were obtained from weather sta-
tion records housed at Rabacca Field Station in
Georgetown, St. Vincent. The weather station
was located about 0.4 km from the experimental
chili pepper fields. CC trap captures were exam-
ined in relation to average rainfall and tempera-
ture in the wet and dry seasons.

Comparison of Sticky Card Trap Colors (Dry Season)

Experiment 4. The experiment was conducted
during the dry season in 2005 in the same com-
mercial pepper fields in St. Vincent described for
Experiment 3. The thrips trap captures compared
sticky card traps with the CC and Blue-D trap
captures. The sticky card traps were 10.0 by
10.5 cm in size. White sticky card traps were
constructed by coating both sides with brush-on
Tanglefoot formula (Tanglefoot Co., Grand Rap-
ids, MI). Blue sticky card traps of the same di-
mensions were Takitrap obtained commercially
(Oecos Ltd., Kimpton, Hertfordshire, England).
Yellow sticky card traps also of the same dimen-
sions were custom made commercially (Olson
Products, Medina, OH). The sticky card traps
were placed in chili pepper rows 1 m apart. Traps
were oriented vertically with wire loops attached

to 25 cm long wooden stakes. Traps were placed 5-
10 cm above the plant terminals.

Statistical Analysis

Numbers of thrips caught were averaged over
sampling periods for each experiment. Data were
analyzed by t-tests, ANOVA orthogonal compari-
sons, or three factor factorial analysis (Anonymous
1989). Means were separated by Tukey's HSD.


Experiment 1. Comparison of CC and Blue-D
traps in Taiwan (dry season). Blue-D traps caught
more S. dorsalis than the CC traps (F = 4.8, df =
1, 58, P = 0.034). Mean captures for F intonsa or
T hawaiiensis (Morgan) for the two trap types
were not statistically different (Table 1). Catches
of S. dorsalis by the three different color CC trap
bases were not significantly different. White base
CC traps caught more T hawaiiensis (F = 14.5, df
= 2, 28, P = <0.001), and blue base CC traps
caught more M. usitatus (F = 49.0, df = 2, 28, P =
<0.001) compared with other trap base colors.
White and blue base CC traps caught more F in-
tonsa (F = 7.3, df = 2, 28, P = 0.028) compared
with yellow base CC traps. The four thrips species
are considered economic pests in Taiwan (Chang
Experiment 2. Comparison of CC and Blue-D
traps in St. Vincent (wet season). Blue-D traps
caught more T palmi, Frankliniella sp., and
M. abdominalis than CC traps (F = 42.4 99.2, df
= 1, 58,P = <0.001). Mean captures ofS. dorsalis for
the two trap types were not significantly different.
Captures of S. dorsalis in CC traps for different
trap base colors were not significantly different.
Blue and white base CC traps caught more T palmi


Mean numbers/trap/week

S. dorsalis F. intonsa T hawaiiensis M. usitatus

Trap type
Blue-D 0.34 0.20 a' 0.50 0.33 a 1.78 1.39 a 0.44 0.17 a
CC traps 0.07 0.02 b 0.11 0.02 a 0.36 0.05 a 0.76 0.17 a
F, P 4.8, 0.034 3.9, 0.053 3.0, 0.089 3.0, 0.089
CC trap (base color)
Whiteb 0.10 0.03 a 0.10 0.02 ab 0.63 0.09 a 0.18 0.04 b
Blue 0.02 0.01 a 0.20 0.05 a 0.28 0.06 b 2.07 0.28 a
Yellow 0.09 0.04 a 0.03 0.01 b 0.18 0.04 b 0.03 0.03 b
F, P 3.7, 0.047 7.3, 0.028 14.5, <0.001 49.0, <0.001

"Dichlorvos dispenser plus blue stripes (Blue-D).
bCC-trap base color with dichlorvos killing agent and ethylene glycol preservative.
'Means in a column not followed by the same letter are significantly different by orthogonal comparison for Blue-D vs. CC traps,
df = 1, 58 and by Tukey's HSD for CC traps, df = 2, 28.

March 2006

Chu et al.: CC Traps for S. dorsalis

and Frankliniella sp. (F = 4.1 and 5.3, df = 2, 44, P
= 0.024 and 0.009, respectively) than yellow base
CC traps. Blue base CC traps caught more M. ab-
dominalis than white and yellow base CC traps (F
= 99.2, df= 2,44, P = 0.003). Dichlorvos cubes in CC
traps increased captures of all thrips species (F =
6.0 20.6, df= 1, 44, P = 0.018 <0.001) except S.
dorsalis. Ethylene glycol in the CC trap bases had
no effect on trap catch numbers. However, thrips
captured in ethylene glycol equipped traps were
well preserved, with less damage to antennae and
less desiccation than traps without ethylene glycol.
There were significant treatment interactions
for CC trap base colors and dichlorvos treatments
for T palmi trap catches (F = 6.6, df = 2, 44, P =
0.003), but not for the three other thrips species (Ta-
ble 2). Thrips captures for the ethylene glycol treat-
ment, or the interactions with CC trap base color
and dichlorvos were not significantly different.
Frankliniella sp. captures were identified by
Systematic Entomology Laboratory as Franklin-
iella cephalica (Crawford) and Frankliniella insu-
laris (Franklin).

Experiment 3. Comparison of CC and Blue-D
traps in St. Vincent (dry season). Mean numbers
of T palmi, Frankliniella sp., and M. abdominalis
caught in Blue-D traps were greater compared
with CC traps (F = 8.2 10.4, df= 1, 58, P = 0.002
- 0.006). Mean captures ofS. dorsali were not sig-
nificantly different for the two trap types (Table
3). dichlorvos cubes in CC traps increased cap-
tures of all four thrips species (F = 8.6 72.2, df=
1, 44, P = 0.005 <0.001). On average, blue and
white base CC traps caught more T palmi, Frank-
liniella sp., and M. abdominalis than yellow base
CC traps (F = 9.5 16.7, df = 2, 44, P = 0.002 -
<0.001). The addition of ethylene glycol in traps
resulted in greater captures of S. dorsalis, Frank-
liniella sp., and M. abdominalis (F = 7.4 10.5, df
= 1, 44, P = 0.009 0.002), than the mean cap-
tures of T palmi. Blue and white base CC traps
with ethylene glycol caught more Frankliniella
sp. (F = 4.0, df = 2, 44, P = 0.024), but not the
other three thrips species. There were no signifi-
cant differences in thrips captures for other treat-
ment interactions.


Mean numbers/trap/week

S. dorsalis T palmi Frankliniella sp. abdominalis

Trap type
Blue-D 0.00 0.00 ab 0.47 0.16 a 0.71 0.17 a 0.82 0.10 a
CC traps 0.02 0.01 a 0.08 0.01 b 0.11 0.03 b 0.15 0.02 b
F, P 1.3, 0.259 42.4, < 0.001 48.9, < 0.001, 99.2, < 0.001
CC trap (base color)
White 0.02 0.01 a 0.09 0.03 a 0.10 0.03 ab 0.11 0.04 b
Blue 0.03 0.01 a 0.10 0.03 a 0.20 0.06 a 0.24 0.04 a
Yellow 0.02 0.01 a 0.03 0.01 b 0.03 0.02 b 0.09 0.03 b
F,P <0.1, None 4.1,0.024 5.3,0.009 6.9,0.003
Dichlorvos in CC traps
Yes 0.02 0.01 a 0.12 0.02 a 0.17 0.05 a 0.22 0.03 a
No 0.03 0.01 a 0.03 0.01 b 0.06 0.02 b 0.07 0.02 b
F, P 0.2, None 17.5, < 0.001 6.0,0.018 20.6, < 0.001
White-yes 0.02 0.01 a 0.17 0.05 a 0.13 0.05 a 0.15 0.06 a
White-no 0.02 0.02 a 0.02 0.01 b 0.08 0.04 a 0.07 0.03 a
Blue-yes 0.02 0.01 a 0.18 0.03 a 0.32 0.11 a 0.36 0.05 a
Blue-no 0.03 0.01 a 0.03 0.02 b 0.09 0.04 a 0.11 0.04 a
Yellow-yes 0.02 0.01 a 0.02 0.02 b 0.06 0.04 a 0.16 0.04 a
Yellow-no 0.03 0.01 a 0.04 0.01 a 0.00 0.00 a 0.03 0.01 a
F, P <0.1, None 6.6,0.003 1.6,0.214 2.3, 0.108

Dichlorvos dispenser plus blue stripes (Blue-D).
bMeans in a column of the same variable not followed by the same letter are significantly different by orthogonal comparison for
Blue-D vs. CC traps, and df = 1, 58, and by Tukey's HSD for CC traps, and df = 1 or 2, 44. Means of three way interactions were
not significantly different. Ethylene glycol treatments and other interactions were not statistically different.

Florida Entomologist 89(1)

March 2006


Mean numbers/trap/week

S. dorsalis T palmi Frankliniella sp. abdominalis

Trap type
CC trap
CC trap (base color)

Dichlorvos in CC traps

Ethylene glycol in CC traps
Base-ethylene glycol

0.05 0.03 ab
0.09 0.02 a
0.7, None

0.06 0.02 a
0.13 0.04 a
0.08 0.02 a

0.13 0.03 a
0.05 0.01 b

0.08 0.04 b
0.04 0.02 b
0.23 0.06 a
0.03 0.01 b
0.08 0.02 b
0.08 0.03 b

0.13 0.02 a
0.05 0.02 b

0.10 0.04 a
0.03 0.02 a
0.15 0.05 a
0.11 0.05 a
0.13 0.02 a
0.03 0.01 a
0.4, None

1.06 0.33 a
0.49 0.08 b

0.53 0.13 a
0.78 0.16 a
0.17 0.04 b
16.7, <0.001

0.82 0.12 a
0.17 0.03 b
55.3, <0.001

0.89 0.18 b
0.18 0.07 c
1.35 0.19 a
0.22 0.06 c
0.22 0.06 c
0.12 0.04 c
11.7, <0.001

0.51 0.09 a
0.48 0.12 a
0.2, None

0.57 0.15 a
0.50 0.21 a
0.74 0.21 a
0.83 0.26 a
0.23 0.06 a
0.11 0.04 a
0.5, None

0.21 0.09 a
0.08 0.02 b

0.08 0.02 ab
0.14 0.04 a
0.02 0.01 b
9.5, <0.001

0.14 0.03 a
0.02 0.01 b
33.9, <0.001

0.13 0.03 b
0.03 0.02 bc
0.26 0.06 a
0.02 0.01 c
0.04 0.02 bc
0.01 0.01 c

0.11 0.03 a
0.05 0.02 b

0.10 0.03 ab
0.06 0.03 b
0.21 0.07 a
0.07 0.03 b
0.02 0.01 b
0.03 0.02 b

0.33 0.06 a
0.17 0.03 b
8.2, 0.006

0.16 0.04 ab
0.25 0.06 a
0.10 0.03 b
7.2, 0.002

0.31 0.04 a
0.04 0.01 b
72.2, <0.001

0.28 0.06 b
0.03 0.03 c
0.46 0.07 a
0.04 0.03 c
0.18 0.05 bc
0.03 0.01 c
6.4, 0.004

0.22 0.04 a
0.12 0.04 b
10.5, 0.002

0.24 0.07 a
0.08 0.03 a
0.27 0.08 a
0.23 0.09 a
0.16 0.05 a
0.05 0.02 a

aDichlorvos dispenser plus blue stripes (Blue-D).
bMeans in a column of the same variable not followed by the same letter are significantly different by orthogonal comparison for
Blue-D vs. CC traps, and df = 1, 58, and by Tukey's HSD for CC traps, and df = 1 or 2, 44. Means of three way interactions were
not significantly different. Ethylene glycol treatments and other interactions were not statistically different.

Seasonal Weather Effects on CC Trap Catches during
the Wet and Dry Seasons in St. Vincent, 2004-2005

CC trap captures were 4.5 and 6.1 fold greater
for S. dorsalis and T palmi, respectively, during
the dry season compared with the wet season.
Rainfall averaged 1.1 mm per day during the dry
season and 18.0 mm per day during the wet sea-
son (Table 4). Air temperatures in the dry season
was 1.3C higher compared with the wet season.

Experiment 4. Comparison of Sticky Card Trap
Colors (Dry Season). Significantly more S. dor-
salis were caught on yellow sticky card traps
compared with blue sticky card traps. Yellow
and blue sticky card traps caught more S. dor-
salis than white sticky card traps (Table 5).
More T palmi, Frankliniella sp. and M. abdom-
inalis were caught on blue sticky card traps
compared with white and yellow sticky card

Chu et al.: CC Traps for S. dorsalis


Mean numbers/trap/week"

Rainfall Rainy Air Micro-
Mean total days temperature Frankliniella cephalothrips
Season mm/day (mm) total C S. dorsalis T palmi sp. abdominalis

Wet (W) 18.2+ 4.8 855.1 34/47 28.4 0.03 0.02 0.00 b 0.08 0.01 b 0.11 0.03 a 0.15 0.02 a
Dry (D) 1.1 0.4 48.1 10/43 29.7 0.01 0.09 0.01 a 0.49 0.09 a 0.08 0.01 a 0.17 0.02 a
t,P 57.8, <0.001 22.3,0.002 0.8, None 0.6, None
D/W ratio 0.58 0.47 0.23 1.05 4.5 6.1 1.4 1.1

"Means in a column followed by the same letter are not significantly different by t-test, df= 1.


Blue-D traps caught more S. dorsalis than the
CC traps in Taiwan, but approximately equal
numbers in St. Vincent. Similarly, Blue-D traps
consistently caught more of the other three thrips
species in the study compared with the CC trap.
Overall, the addition of dichlorvos cubes in-
creased CC trap captures ofS. dorsalis in St. Vin-
cent. The blue base CC trap with dichlorvos cubes
caught more S. dorsalis than the other treatment
combinations only during the dry season in St.
Vincent. Unfortunately, previous trap studies
conducted in India on S. dorsalis did not include
blue base CC trap comparisons with white and
yellow base traps (Chu et al. 2000). The addition
of ethylene glycol to CC traps increased trap
catches of S. dorsalis and M. abdominalis during
the dry season in St. Vincent. The addition of eth-
ylene glycol resulted in better preserved speci-
mens for taxonomic and genetic studies.
Numbers of S. dorsalis captured in CC traps
with dichlorvos were low in both Taiwan and St.
Vincent. We reported earlier that blue sticky card
traps caught more F occidentalis in a broccoli
field than yellow sticky card traps (Chen et al.
2004). Blue sticky card traps also captured
greater numbers of T palmi, Frankliniella sp.,
and M. abdominalis than yellow or white traps in
the current studies. Our results from St. Vincent

indicate that yellow sticky card traps were more
attractive to S. dorsalis than white or blue sticky
card traps. Similarly, Hoddle et al. (2003) re-
ported that Scirtothrips perseae (Nakahara) was
more attracted to yellow than white or blue sticky
card traps.
The Blue-D trap did not consistently capture
greater numbers of S. dorsalis than CC traps. Its
potential toxicity in the environment is of con-
cern. Although the CC trap captures fewer S. dor-
salis, the quality of the captured specimens is
high. They are easily recovered from the trap and
stored in ethanol for later taxonomic and genetic
analysis. Yellow sticky traps capture more thrips
than the CC traps but they also capture a large
number of non-target insects. In addition, thrips
that are captured on the sticky trap are not easily
removed and stored for later studies. Sticky traps
seem to be less labor intensive, require less com-
ponent assembly and therefore less expertise in
trap placement than the CC traps.
Seal et al. (2005) have determined from direct
plant sampling that economic damage to chili
peppers by S. dorsalis occurs at densities of 0.5
to 2 individuals (larvae or adults) per terminal
leaf. This sampling method requires nine sam-
ples per 24-48 m2 area in order to achieve the
90% precision level. This method may be too la-
bor intensive to use in large scale survey and de-
tection efforts. Alternatively, visual observation

MENT 4).

Mean numbers/trap/week"

Sticky trap color S. dorsalis T palmi Frankliniella sp. Microcephalothrips sp.

White 1.41 + 0.11 b 8.04 1.45 b 2.08 0.34 b 4.39 0.57 b
Blue 3.72 0.37 b 27.11 2.20 a 8.75 0.85 a 21.40 1.79 a
Yellow 14.10 + 1.06 a 7.38 0.62 b 1.73 0.18 b 4.30 0.45 b
F, P 111.4, <0.001 57.411.7, <0.001 50.0, <0.001 77.6, <0.001

"Means in a column not followed by the same letter are significantly different by Tukey's HSD, and df = 2 or 18.

Florida Entomologist 89(1)

for plant damage symptoms like curled, de-
formed, or yellow leaves coupled with placing
sticky card traps can be utilized as a preliminary
detection tool. Positive detections would then be
followed by direct plant sampling to capture
large number of individual specimens for taxo-
nomic verification. APHIS guidelines for survey
are based on the principle of finding one S. dor-
salis in a suspected area. The guidelines suggest
2,280 CC traps for initial survey that will be
placed in one square mile areas for detecting
S. dorsalis (USDA-aphis, 2004). If one or more
S. dorsalis is found the second phase of survey
the survey area will be expanded to the eight
surrounding square miles. The presence of a sin-
gle S. dorsalis in the second survey will lead to
an expansion of the survey area to 80 surround-
ing square miles for the third phase of survey.
Results of our studies estimate that CC traps
would catch 46 and 205 for wet and dry seasons,
respectively. These would translate to the cap-
tures of 46 and 205 S. dorsalis in CC traps for
the initial survey, 148 and 666 for the second
survey, and 1069 and 4813 for third phase of sur-
vey during wet and dry seasons, respectively, in
St. Vincent.
Current methods employed for detection of S.
dorsalis are inefficient as demonstrated in the
present report. Studies ofS. dorsalis behavior, in-
cluding the development of attractants and pher-
omones as potential lures, are being conducted to
develop more efficient trap systems for detection
and monitoring of this insect pest.
During preparation of this manuscript, S. dor-
salis was detected on roses in Palm Beach, FL and
in multiple retail garden centers on hot pepper
seedlings by the Florida Department of Agricul-
ture and Consumer Services (FDACS) (Wayne
Dixon, pers. comm.). Further surveys of retail
garden centers in the Lower Rio Grande Valley of
Texas likewise revealed the presence of this new
invasive species on hot pepper seedlings (M. Ci-
omperlik, unpublished data). Specimens were
confirmed as Scirtothrips dorsalis by the USDA
ARS Systematic Entomology Laboratory. The cur-
rent distribution of S. dorsalis in the Caribbean is
limited to a few islands. It has recently invaded
the US, and is expected to spread over time
through agricultural trade and tourism (Venette
& Davis 2004; Meissner et al. 2005). These obser-
vations also indicate an alarming potential for
rapid spread of this pest thrip species through in-
terstate movement of ornamentals and plant
seedlings in the US. The potential impact of this
thrips on agriculture in the United States alone
has been estimated at approximately $3.6 to $6.0
billion a year (Lynn Garrett, USDA APHIS PPQ
CPHST, pers. comm.). Effective survey and detec-
tion methods are needed to monitor the spread,
and manage populations, ofS. dorsalis both in the
Caribbean and the US.


The authors thank Tsung Tao Hsieh of National
Pingtung University of Science and Technology in Tai-
wan for technical assistance during the studies. In addi-
tion, we thank Jason Carlson and Juan Rodriguez
(USDA, APHIS) for technical support. We also thank
Laurence Mound (CSIRO) and Steve Nakahara (USDA,
ARS, SEL) for identification of thrips specimens.


ANONYMOUS. 1989. MSTATC. A microcomputer program
for the design, management, and analysis of agro-
nomic research experiments. Michigan State Univ.
Color and height preference of the citrus thrips in a
navel orange grove. J. Econ. Entomol. 64: 1112-1113.
CHANG, N. T. 1995. Major pest thrips in Taiwan, pp. 105-
108 In B. L. Parker, M. Skinner and T. Lewis [eds.],
Thrips Biology and Management, Proceedings of the
1993 International Conference on Thysanoptera. Ple-
num Press, New York and London. NATO ASI Series.
NEBERRY. 2004. Trap Evaluations for thrips (Thysan-
optera: Thripidae) and hoverflies (Diptera: Syrphidae).
J. Environ. Entomol.: 1446-1420.
KENNEDY. 1995. Comparison of colored sticky traps
for monitoring thrips populations (Thysanoptera:
Thripidae) in staked tomato fields. J. Entomol. Sci.
30: 176-190.
Bemisia argentifolii (Homoptera: Aleyrodidae): host
preference and factors affecting oviposition and feed-
ing site preference. J. Environ. Entomol. 24:254-360.
CHU, C. C., AND T. J. HENNEBERRY. 1998. Development
of a new whitefly trap. J. Cotton Sci. 2: 104-109.
M. SHREPATIS. 2000. Use of CC traps with different
trap base colors for silverleaf whiteflies (Homoptera:
Aleyrodidae), thrips (Thysanoptera: Thripidae), and
Leafhoppers (Homoptera: Cicadellidae). J. Econ. En-
tomol. 93: 1329-1337.
EXANDER, AND T. J. HENNEBERRY. 2005. Variations
in CC trap catches ofthrips associated with different
colors with or without dichlorvos cubes, pp. 1173-
1175 In P. Dugger and D. Richter [eds.], Proc. Belt.
Cotton Conf., New Orleans, LA.
CIOMPERLIK, M. A., AND D. SEAL. 2004. Surveys of St.
Lucia and St. Vincent for Scirtothrips dorsalis
(Hood), Jan. 14-23, 2004. USDA APHIS PPQ, Tech-
nical Report. 19 pp.
FNGA. 2003. Report of the Florida Nurserymen &
Growers Association Pest and Disease Task Force.
May 15, 2003. 4 pp.
GILLESPIE, D. R., AND R. S. VERNON. 1990. Trap catches
of western flower thrips (Thysanoptera: Thripidae)
as affected by color and height of sticky traps in ma-
ture greenhouse cucumber crops. J. Econ. Entomol.
83: 971-975.
HOOD, J. D. 1919. On some new Thysanoptera from
southern India. Insecutor Inscit. Menstir. 7., 90-103.
traction of thrips (Thysanoptera: Thripidae and Ae-

March 2006

Chu et al.: CC Traps for S. dorsalis

olothripidae) to colored sticky cards in a California
avocado orchard. Crop Protection. 21: 383-388.
HODDLE, M. S., AND L. A. MOUND. 2003.The genus Scir-
tothrips in Australia (Insecta, Thysanoptera, Thripi-
dae). Zootaxa 268: 1-40.
BRODEL, AND T. DOBBS. 2005. Evaluation of Possible
Pathways of Introduction for Scirtothrips dorsalis
Hood (Thysanoptera: Thripidae) from the Caribbean
into the Continental United States. Plant Protection
and Quarantine. Center for Plant Health Science
and Technology. 124 pp.
MOUND, L. A., AND J. M. PALMER 1981. Identification,
distribution and host plants of the pest species of
Scirtothrips (Thysanoptera: Thripidae). Bull. Ento-
mol. Res. 71: 467-479.
NAKAHARA, S. (Unpublished). Key to species and genera
of Thripinae (Thripidae) intercepted at agricultural
quarantine. Unpublished USDA, APHIS, PPQ docu-
ment. 11 pp.
KLASSEN. Distribution of the chili thrips, Scirto-
thrips dorsalis Hood (Thysanoptera: Thripidae),
within pepper plants and within pepper fields on St.
Vincent. Florida Entomol. (in press).

SKARLINSKY, T. L. 2003. Survey of St. Vincent pepper
fields for Scirtothrips dorsalis Hood. USDA, APHIS,
PPQ. 5 pp.
TODA, S., AND S. KOMAZAKI. 2002. Identification ofthrips
species (Thysanoptera: Thripidae) on Japanese fruit
trees by polymerase chain reaction and restriction
fragment length polymorphism of the ribosomal ITS2
region. Bull. Entomol. Res. 92: 359-363.
attraction of yellow tea thrips (Scirtothrips dorsalis
Hood). Japanese J. App. Entomol. Zool. 39: 299-303.
USDA. 2003. Port Information Network (PIN-309):
quarantine status database. U.S. Department of Ag-
riculture, Animal and Plant Health Inspection Ser-
vice, Plant Protection and Quarantine, Riverdale,
MD. Restricted access database.
USDA-APHIS. 2004. New Pest Response Guidelines:
Chili Thrips Scirtothrips dorsalis. June 15, 2004.
Scirtothrips dorsalis Confirmed as Present in Trin-
idad and Tobago, reported December 23, 2004,
USDA Restricted access database.
VENETTE, R. C., AND E. E. DAVIS. 2004. Mini Pest Risk
Assessment Chili thrips/yellow thrips, Scirtothrips
dorsalis Hood (Thysanoptera: Thripidae). Univ. of
Minnesota, St. Paul. 32 pp.

Florida Entomologist 89(1)

March 2006


Laboratory of Forest Zoology, Division of Agriculture and Agricultural Life Science
University of Tokyo, Tokyo, Japan

Present address: Research Center for Biodiversity, Academia Sinica, Nankang, Taiwan 115, R.O.C.


Egg hatching of the maple aphid, Periphyllus californiensis Shinji, was observed on saplings
ofAcer amoenum Carriere in two microhabitats, i.e., the understory of a maple stand (a
shaded site) and an open area in a nursery (a sunny site), over a 2-year period. Buds of
A. amoenum opened earlier at the shaded site than at the sunny site and eggs ofP. californ-
iensis also hatched a little earlier at the shaded site. To test whether oviposition timing or
microhabitat characteristics affected the timing of egg hatching, eggs were collected during
four periods in December to observe egg hatching in the laboratory. Hatching occurred ear-
lier at the shaded site than at the sunny site only for eggs laid in early December. The dura-
tion of egg hatching was shorter for eggs laid earlier compared with those laid later. The
duration of the egg stage (estimated as the median oviposition date to the median egg hatch-
ing date) was negatively correlated with the time when the eggs were laid. These results sug-
gest that differences in timing of egg hatching between habitats may be affected by the
microhabitat and date of oviposition.

Key Words: oviposition, egg duration, host plant phenology, synchrony


La eclosi6n de huevos del afido arce, Periphyllus californiensis Shinji, fue observads en re-
nuevos deAcer amoenum Carriere en dos micro habitares, o sea, la parte abajo de los arboles
de arce (un sitio con sombra) y una area abierta de un vivero (un sitio con sol), durante un
period de 2 aios. Los brotes deA. amoenum abrieron mas temprano en el sitio con sombra
que en el sitio bajo el sol y los huevos de P. californiensis tambi6n esclosionaron un poco mas
temprano en el sitio de sombra. Para probar si el tiempo de la oviposici6n o las caracteristi-
cas del micro habitares afectaron el tiempo de la eclosi6n de los huevos, se recolectaron hue-
vos durante cuatro periods en el mes de diciembre para observer la eclosi6n de huevos en
el laboratorio. La eclosi6n fue mas temprana en el sitio de sombe que en el sitio bajo el sol
solamente para los huevos colocados durante el inicio del mes de diciembre. La duraci6n de
la eclosi6n de huevos fue mas corta para los huevos puestos tempranamente que en compa-
raci6n con los huevos puestos mas tarde. La duraci6n del estadio de huevo (calculado de la
fecha median de oviposici6n hasta la fecha median de la eclosi6n de huevos) fue negativa-
mente correlacionada con el tiempo cuando los huevos fueron puestos. Estos resultados su-
gieren que las diferencias en el tiempo de la eclosi6n de huevos entire los habitares puede ser
afectados por el micro habitat y la fecha de la oviposici6n.

Insect performance is strongly influenced by
the environments in which the insects and their
host plants grow. In forests, the environment
around a plant may vary both temporally and
spatially (Bazzaz 1979). The phenology of dor-
mancy, leaf emergence, and leaf senescence of
trees of the upper layer may result in seasonal
changes in the quality and quantity of sunlight
reaching different regions of the forest floor, e.g.,
forest edges, gaps, understory, and open areas
next to forests (Denslow et al. 1990; Uemura
1994; Gill et al. 1998; Seiwa 1998; Kato & Komi-
yama 2002). These differences in quality and
quantity of sunlight within the forest may result

in variable temperature, humidity, food quality,
and predation by natural enemies, and may con-
sequently influence the development, growth,
survival, and abundance of insects (Rausher
1979; Lowman 1992; Shure & Wilson 1993; Dudt
& Shure 1994; Louda & Rodman 1996; Bergman
1999; McDonald et al. 1999).
Host plant phenology, including the timing of
budburst and leaf senescence, may be affected by
environmental conditions; for example, by exposure
to sun or shade (Furuta 1990; Lowman 1992; Seiwa
1999). This is particularly important in the early
spring when synchrony between the time of egg
hatching and the budburst plays an important role

Wang: Egg Hatching of Maple Aphid

in the performance and population growth of insects
(Dixon 1976; Holliday 1977; Wint 1983; Watt & Mc-
Farlane 1991; Hunter 1992; Akimoto & Yamaguchi
1994; Quiring 1994; Lawrence et al. 1997; Van Don-
gen et al. 1997; Martel & Kause 2002).
The maple aphid, Periphyllus californiensis
Shinji, dwells on maple trees year-round. In the
early spring, stem mothers (fundatrix), the first
parthenogenetic generation appearing from fertil-
ized eggs (Miyazaki 1987), hatch from overwinter-
ing eggs and give rise to one or more winged or
wingless spring generations by parthenogenesis.
This aphid feeds on growing buds, leaves, shoots,
and the inflorescence in spring. As leaves expand,
the soluble nitrogen concentration in the phloem
declines, and aestivating dimorphs are produced. In
the summer, aestivating dimorphs remain as first
instars, mostly on leaves, until autumn when they
resume growth and become wingless adults as the
food quality improves once again. In spring and au-
tumn, winged females disperse among maple trees
(Furuta 1987). In spring and autumn, the perfor-
mance of the maple aphid is closely attuned to the
budburst and leaf senescence phenology of its host
trees. In spring, the numbers of stem mothers and
their survival rates are higher on early-budding
trees than on late-budding trees (Furuta 1987).
Therefore, reproduction of stem mothers is mostly
observed on early-budding trees, and their winged
progeny disperse to late-budding trees where they
reproduce in turn (Furuta et al. 1984). In autumn,
the population increases first on early-senescing
trees, and the winged female progeny of the aesti-
vating dimorphs then disperse to late-senescing
trees on which they then reproduce. As a result, ovi-
parae are produced earlier on early-senescing trees
than on late-senescing trees (Furuta 1986). The
budburst and leaf senescence phenology of the ma-
ple tree, Acer palmatum, is influenced by the light
conditions experienced by the trees (Furuta 1990).
Environmental differences in exposure to sun and
shade may thus affect development of the egg stage
of the maple aphid, and the host tree phenology
may also affect the reproductive schedules of the
autumnal population and subsequent egg hatching.
In this study, the egg hatching ofP. californien-
sis was studied on Acer amoenum Carriere sap-
lings in two microhabitats with different light
conditions and microclimates, i.e., the understory
of a maple stand (a shaded site) and an open area
in a nursery (a sunny site) over a 2-year period.
Two questions were examined. First, do microcli-
matic differences between sites cause differences
in the timing of egg hatching between microhabi-
tats? Second, does the timing of oviposition have
an effect on the timing of egg hatching?


The study was conducted from spring 1997 un-
til spring 2000 in the Forest Experimental Sta-

tion at Tanashi (35N; 139E; 60 m elev.), situated
in Nishitokyo-shi, Tokyo, Japan. Two study sites,
a maple stand understory and an open area in a
nursery, were selected in order to observe the phe-
nology of the egg hatching of P californiensis and
the budburst of A. amoenum. The maple stand
was shaded by the trunks and branches of over-
story trees in winter and early spring (hereafter
called the 'shaded site'). The nursery was in an
open field with no shading (hereafter called the
'sunny site'). The two study sites were separated
by about 100 m.

Egg Hatching and Budburst in the Field

Potted saplings,15-40 cm high, ofA. amoenum
from the same provenance randomly placed in ei-
ther the sunny or shaded site from the spring of
1997 were used for observing the budburst phe-
nology. At the sunny site, 28 and 19 saplings were
observed, while 25 and 12 saplings were observed
at the shaded site in 1999 and 2000, respectively.
Budburst (defined as the time when leaves first
become visible from opening buds) was monitored
every 2-3 days from the beginning of February in
both years. The median date of budburst was de-
termined from counts of all buds on all saplings.
The cumulative percentage ofbudbursts was esti-
mated by averaging the accumulated percentage
of budbursts across all saplings.
In autumn 1998, only saplings at the shaded
site had established natural maple aphid colo-
nies. In order to permit observation of egg hatch-
ing at both study sites, fourth instar-adult ovi-
parae and males collected from Acer spp. in the
field were artificially placed onto saplings at both
sites on December 11, 1998.
Eggs laid by oviparae in autumn 1999 were ob-
served in 2000. Adult oviparae were observed on
saplings at the sunny and shaded sites at the be-
ginning and the end of December 1999, respec-
tively. In spring 2000, all hatched stem mothers
were removed after each observation. Observa-
tions were made every other day beginning in
February and ending in April in both years.
Unlike other studies that indicated high over-
wintering mortality of aphid eggs (Leather et al.
1995; Wade & Leather 2002), no obvious mortal-
ity of eggs was observed in this study. Since larval
syrphids are the primary predators of the maple
aphid in spring at the study sites, the eggs of syr-
phids were regularly removed from the study sap-
lings whenever they were found in order to pre-
vent predation of newly-hatched aphids.

Effect of Microhabitat and Oviposition Period

Oviparae were collected from maple trees in
the field during four periods in 1999 on December
1-2, 7-10, 15-18, and 23-27. These oviparae were
maintained in the lab on four to eight 15-30-cm-

Florida Entomologist 89(1)

high saplings of A. amoenum growing in the
shade. Eggs were collected from these oviparae
over short periods in the laboratory to minimize
the effects of host plants or environments on the
oviparae. On the last day of each period, all ovi-
parae were removed, and the saplings with eggs
were transferred to the field where they were
placed in either the sunny or shaded site. Egg
hatching was then recorded every 2 days during
spring 2000, and all hatching stem mothers were
removed during each observation.

Data Transformation and Statistical Analysis

Most statistical analyses in this study were
performed with SYSTAT (version 8, SPSS, Chi-
cago, IL, USA). The means and variation of the
timing of egg hatching were compared between
study sites by t-test and F-test (Elliott 1971), re-
spectively. Patterns of egg hatching were com-
pared by plotting regression lines of logit-trans-
formed proportion of eggs hatched against degree-
days within each study site. Differences between
slopes and elevations of the regression lines were
compared (Zar 1999). Slopes were compared first,
and elevations were only analyzed when slopes
showed significant differences. Degree-days were
calculated by the Sine method (Frazer & Gilbert
1976; Pruess 1983; Raworth 1994) and were accu-
mulated above 4.58C from 1 February 2000
(Wang & Furuta 2002). Daily minimum and max-
imum temperatures were obtained from the For-
est Experimental Station at Tanashi. Weekly
maximum and minimum temperatures at the
sunny and shaded sites were recorded by hanging
a maximum/minimum thermometer about 40 cm
above the ground on the north side of a wooden
box from February to the beginning of April 2000.
The relationships between these weekly maxi-
mum or minimum temperature data within each
study site (y) and weekly maximum or minimum
temperature data obtained from the Tanashi Ex-
perimental Station (x) were calculated by linear
regression. Then, the daily temperature data from
the Tanashi Experimental Station were applied to
the equations to obtain estimated daily maximum
and minimum temperatures for each study site.


Egg Hatching and Budburst in the Field

Egg hatching at the shaded site occurred ear-
lier than that at the sunny site in both years (Fig.
1). Egg hatching occurred about 10 and 6 days
earlier at the shaded site in 1999 and 2000, re-
spectively (t = 19.339, df = 783, P < 0.001; t =
9.408, df= 823, P < 0.001), but variation in time of
egg hatch between sites was the same in both
years (in 1999: F434349 = 1.107, P > 0.05; in 2000:
F,,,3 = 1.474, P > 0.05).

(a) 1999


0.5 L.

20 30 40 so 60 70 80 90
No. of days from February I

10 20 30 40 so 60
No. of days from March 1
Figure 1. Proportion of eggs hatched (circles) and
buds burst (triangles) in the sun (open) and shade
(solid) against time in (a) 1999 and (b) 2000.

The median dates of budburst were about 14
and 13 days earlier at the shaded site than at the
sunny site in 1999 and 2000, respectively. The de-
gree of synchrony between egg hatching and bud-
burst varied between sites and years. At the
shaded site, budburst began when 56% and 97%
of the eggs had hatched in 1999 and 2000, respec-
tively. At the sunny site, budburst began when
97% and 100% of the eggs had hatched in 1999
and 2000, respectively. The interval between the
date when the first egg hatched and the date of
the first budburst at the shaded site was 13 and
16 days in 1999 and 2000, respectively. At the
sunny site, the interval was 28 days in both years.
The interval between the median dates of egg
hatching and budburst at the shaded site was 26
and 16 days in 1999 and 2000, respectively, and
24 and 21 days at the sunny site.

Effect of Microhabitat and Oviposition Period

The timing of egg hatching varied between
sites and among oviposition times (by two-way
ANOVA: Site, F1721 = 5.312, P < 0.05; Time, F3721 =
7.218, P < 0.001). Between sites within each ovipo-
sition period, eggs laid on December 7-10 began to
hatch 6 days earlier at the shaded site than at the
sunny site. There were no differences between
sites for eggs laid in other periods. The median

March 2006

Wang: Egg Hatching of Maple Aphid

date of egg hatching was 4 days earlier at the
shaded site than at the sunny site for eggs laid on
December 1-2 and 15-18 (Table 1). The variation
in the timing of egg hatching between sites dif-
fered for eggs laid on December 7-10 (F105110 =
1.620, P < 0.05). Patterns of egg hatching between
sites had significantly different slopes for eggs laid
on December 1-2 and 7-10, and similar slopes but
significantly different elevations for eggs depos-
ited during the other two periods (Fig. 2, Table 1).
When oviposition periods were compared
within sites, eggs laid on December 1-2 began to
hatch about 8-14 days later than those laid on De-
cember 7-27. The duration of egg hatching (calcu-
lated as the number of days from when the first to
the last eggs hatched) was shortest for eggs laid
on December 1-2 (23-25 days), and longest for
eggs laid on December 23-27 (39-41 days). The
median duration of the egg stage (calculated as
the number of days from the median oviposition
date to the median egg hatching date) was nega-
tively correlated with the date of oviposition (cor-
relation coefficient r = -0.982, n = 8, P < 0.001).
The longest egg stage duration was 109-113 days
for those laid on December 1-2, and the shortest
was 86 days for those laid on December 23-27.
When patterns of egg hatching were compared
within each study site, only the eggs laid on De-
cember 7-10 and 23-27 hatched at the shaded site
revealed the same regression lines, as did eggs
laid on December 15-18 and 23-27 hatched at the
sunny site (Table 1).


Eggs at the understory site hatched a little
earlier and tended to require fewer thermal units
for egg development than those at the open site.
Differences in patterns of egg hatching between
sites tended to be larger for eggs laid earlier in
December than those laid later. Egg diapause ter-
mination can be affected by two thermal features,
the length of the lower temperature exposure and
the actual temperatures eggs experience (Leather
et al. 1995). Therefore, it is possible that greater
extremes of temperature in the open site com-
pared with the understory site may have influ-
enced the thermal conditions for egg diapause
termination (Tauber & Tauber 1976; Day 1984;
Tauber et al. 1986; Fisher et al. 1994; Wang & Fu-
ruta 2002), and the speed of egg development dur-
ing the post-diapause stage (Augspurger & Bar-
tlett 2003), which generated differences in the
timing of egg hatching between sites. Further-
more, differences in microclimates between sites
might be larger when the deciduous canopy is still
closed than they are after overstory trees lose
their leaves (Gill et al. 1998; Kato & Komiyama
2002). The last tree shorter than 2 m high to shed
its leaves at the shaded site did so by December
21, 1999 (Wang 2002). Therefore, differences be-
tween sites may have decreased over time in De-
cember and have resulted in larger environmen-
tal differences between sites for eggs laid early in
December than for those laid later.


Egg hatching Between
period N Period Median date Regression equation r2 Sites* Periods*

Dec 1-2
Shaded 35 Mar 7-29 Mar 19 y = -10.2 + 0.0662 x 0.954
Sunny 50 Mar 7-31 Mar 23 y = -9.91 + 0.0501 x 0.976
Dec 7-10
Shaded 105 Feb 22-Mar 29 Mar 21 y = -6.03 + 0.0384 x 0.960 dE
Sunny 110 Feb 28-Mar 31 Mar 19 y = -10.0 + 0.0564 x 0.991
Dec 15-18
Shaded 118 Feb 22-Mar 31 Mar 15 y = -5.72 + 0.0410 x 0.974 a dF
Sunny 153 Feb 22-Mar 31 Mar 19 y = -6.96 + 0.0401 x 0.996 a G
Dec 23-27
Shaded 94 Feb 22-Mar 31 Mar 19 y = -6.33 + 0.0419 x 0.992 b EF
Sunny 64 Feb 22-Apr 2 Mar 19 y = -6.03 + 0.0376 x 0.975 b G

*Test results from all regression lines by Tukey's test. Only those between sites within periods and among periods within sites
are shown. Slopes and elevations of values with the same capital letters do not differ. Slopes of values with the same small letters
do not differ, but the elevations differ. Slopes of values without the same letters differ.

Florida Entomologist 89(1)





a Weather Association 1997). Because higher tem-
peratures and changing day length have been im-
(a) plicated in the production of eggs with more-in-
Stensive diapause, i.e., entering a longer diapause,
S~ for some insects (Tauber et al. 1986; Masaki
1996), it is possible that oviparae might be stimu-
6 lated by the higher temperatures and longer day
Lengths in early December thus producing eggs
with more-intensive diapause than those pro-
0 50 10 150 200 250 300 duced later in the winter. In addition to the direct
T effects of microhabitats and oviposition times on
eggs examined in this study, determining other
potential factors which might have affected egg
(b) conditions through oviparae, e.g., genetic varia-
I tion (Komatsu & Akimoto 1995), maternal effects
(Mousseau 1991; Bradford & Roff 1993; Cherrill
S2000; Roff & Bradford 2000; Denlinger 2002), and
Shot plant quality (Hunter & McNeil 1997), may
require further detailed investigations.
SThe duration of egg hatching was shorter and
0 50 TOO 150 200 250 300 .
0 10 20 its onset was about 1-2 weeks later for eggs laid
on December 1-2 than for those laid later. Fewer
S- eggs were used in the experiment on December 1-
(c) 2 as oviparae could only be found on early-senesc-
ing trees in small numbers. Therefore, eggs laid
on December 1-2 may reflect the hatching pattern
for those laid on early-senescing trees. After De-
cember 1-2, oviparae became increasingly abun-
dant on both early- and late-senescing trees and
were collected on both kinds of trees. Eggs laid
0 50 100 150 200 250 300 during the period December 7-27 may represent a
combination of both late- and early-hatching
eggs, and this would be consistent with a longer
(d) period of egg hatching for this cohort.
Acer palmatum growing in the shade tends to
break buds earlier and enter senescence later
Than those in full sun (Furuta 1990). Saplings of
Acer saccharum are known to break buds earlier
and enter senescence later in the understory than
in gaps (Augspurger et al. 2003), and the results
0 50 100 150 200 250 300 of the present study are consistent in this regard.
Changes in a plant's phenology in different light
Degree-days environments may result from understory trees
avoiding canopy shade in order to maximize net
e 2. Proportion of eggs hatched in the sun carbon gain (Uemura 1994). Because aphids are
d shade (solid) for eggs laid during (a) Decem- sap-sucking insects, the soluble nitrogen in the
b) 7-10, (c) 15-18, and (d) 23-27 against degree- sap is critical for their growth (Dixon 1998). The
emulated above 4.58C from February 1. maple aphid can only feed and grow on develop-
ing buds until leaf expansion is complete. They
produce normal winged or wingless offspring
duration of the egg stage was longer for when food quality is high, and aestivating dimor-
d earlier in December than for those laid phs when food quality declines (Hashimoto & Fu-
i a laboratory study (Wang & Furuta ruta 1988). In spring, most stem mothers are
ggs ofP. californiensis deposited earlier in found on early-budding trees, and their progeny,
er also exhibited delayed hatching com- which develop into winged adults, disperse to
o those deposited later. The period from late-budding trees and reproduce there (Furuta
er 7-18, 1999, in this study seemed to be 1987). When food quality becomes poor, only di-
cal oviposition period for the timing of egg morphs are produced. These aestivating first in-
g. In the Tokyo region, the temperature stars will aestivate on leaves for several months
ly decreases in December, and day length until leaf senescence begins in autumn. Autum-
est between December 16 and 26 (Japan nal populations build up earlier on early-senesc-



(open) an
ber 1-2, (
days acci

eggs lai
later. Ii
2002), e
pared t(
the criti
is short

March 2006

Wang: Egg Hatching of Maple Aphid

ing trees than on late-senescing trees, and winged
individuals maturing on early-senescing trees
can also colonize and reproduce on late-senescing
trees. This results in the earlier appearance of
oviparae on early-senescing trees than on late-se-
nescing trees. Thus the entire life cycle of the ma-
ple aphid is driven by the host plant phenology,
including the production of oviparae and eggs.
Egg hatching of the maple aphid occurs earlier
than the budburst. This phenomenon has also
been observed in the gall-forming aphid Horma-
phis hamamelidis which hatches in advance of
the budburst (Rehill & Schultz 2002). In the early
spring, stem mothers of the maple aphid can stay
on the bud scales before bud growth begins, but
they will not molt to second instars until the buds
start to swell (Furuta 1990). Because stem moth-
ers can survive starvation conditions for a time
(Wang 2002), hatching earlier than bud swelling
may permit immediate initiation of growth when
suitable food becomes available, although it also
incurs the cost of a longer period of exposure to
natural enemies (Price et al. 1980). In addition,
early hatching increases the chance that multiple
generations can be completed on both early-bud-
ding and other late-budding trees. Thus the high-
est potential fitness will be obtained by stem
mothers hatching early on early-budding trees.


I thank Dr. K. Furuta and Dr. S. Lawson for helpful
comments on an early version of manuscript and the offic-
ers of the Tanashi Experimental Station of the Tokyo Uni-
versity Forests for their help with the field investigations.


AKIMOTO, S., AND Y. YAMAGUCHI. 1994. Phenotypic se-
lection on the process of gall formation of a Tetraneura
aphid (Pemphigidae). J. Anim. Ecol. 63: 727-738.
AUGSPURGER, C. K., AND E. A. BARTLETT. 2003. Differ-
ences in leaf phenology between juvenile and adult
trees in a temperate deciduous forest. Tree Physiol.
23: 517-525.
BAZZAZ, F. A. 1979. The physiological ecology of plant
succession. Annu. Rev. Ecol. Syst. 10: 351-371.
BERGMAN, K. 1999. Habitat utilization by Lopinga
machine (Nymphalidae: Satyrinae) larvae and ovipos-
iting females: implications for conservation. Biol.
Conserv. 88: 69-74.
BRADFORD, M. J., AND D. A. ROFF. 1993. Bet hedging
and the diapause strategies of the cricket Allonemo-
bius fasciatus. Ecology 74: 1129-1135.
CHERRILL, A. 2002. Relationships between oviposition
date, hatch date, and offspring size in the grasshopper
Chorthippus brunneus. Ecol. Entomol. 27: 521-528.
DAY, K. 1984. Phenology, polymorphism and insect-
plant relationships of the larch bud moth, Zeira-
phera diniana (Guen6e) (Lepidoptera: Tortricidae),
on alternative conifer hosts in Britain. Bull. Ento-
mol. Res. 74: 47-64.
DENLINGER, D. L. 2002. Regulation of diapause. Annu.
Rev. Entomol. 47: 93-122.

R. STRAIN. 1990. Growth responses of tropical shrubs
to tree fall gap environments. Ecology 71: 165-179.
DIXON, A. F. G. 1976. Timing of egg hatch and viability
of the sycamore aphid, Drepanosiphum platanoidis
(SCHR.), at bud burst of sycamore, Acer pesudopla-
tanus L. J. Anim. Ecol. 45: 593-603.
DIXON, A. F. G. 1998. Aphid Ecology. Chapman & Hall,
London, UK. 300 pp.
DUDT, J. F., AND D. J. SHURE. 1994. The influence of
light and nutrients on foliar phenolics and insect
herbivory. Ecology 75: 86-98.
ELLIOTT, J. M. 1971. Some Methods for the Statistical
Analysis of Samples of Benthic Invertebrates. Fresh-
water Biological Association, Scientific Publication
No. 25, Windermere, UK. 144 pp.
FISHER, J. R., J. J. JACKSON, AND A. C. LEW. 1994. Tem-
perature and diapause development in the egg of
Diabrotica barberi (Coleoptera: Chrysomelidae). En-
viron. Entomol. 23: 464-471.
FRAZER, B. D., AND N. GILBERT. 1976. Coccinellids and
aphids: a quantitative study of the impact of adult
ladybirds (Coleoptera: Coccinellidae) preying on
field populations of pea aphids (Homoptera: Aphid-
idae). J. Entomol. Soc. Br. Columbia 73: 33-56.
FURUTA, K. 1986. Host preference and population dy-
namics in an autumnal population of the maple
aphid, Periphyllus californiensis Shinji (Homoptera,
Aphididae). J. Appl. Entomol. 102: 93-100.
FURUTA, K. 1987. Amounts of favourable feeding mate-
rials in spring for the maple aphid, Periphyllus cali-
forniensis Shinji, estimated from the phenological
relations between the aphid and host trees. J. Appl.
Entomol. 104: 144-157.
FURUTA, K. 1990. Early budding ofAcerpalmatum caused
by the shade; intra-specific heterogeneity of the host
for the maple aphid. Bull. Tokyo Univ. For. 82: 137-145.
The effect of budding and flowering of maple trees on
the development of the maple aphid, Periphyllus cal-
iforniensis Shinji (Homoptera, Aphididae) popula-
tion. Z. Angew. Entomol. 98: 437-443.
Leaf phenology, photosynthesis, and the persistence
of saplings and shrubs in a mature northern hard-
wood forest. Tree Physiol. 18: 281-289.
HASHIMOTO, H., AND K. FURUTA. 1988. Reproduction of
maple aphid (Periphyllus californiensis) in spring in
relation to phenology of host trees. Japanese J. Appl.
Entomol. Zool. 32: 169-175.
HOLLIDAY, N. J. 1977. Population ecology of the winter
moth (Lepidoptera: Geometridae) on apple in rela-
tion to larval dispersal and time of budburst. J. Appl.
Ecol. 14: 803-814.
HUNTER, M. D. 1990. Differential susceptibility to vari-
able plant phenology and its role in competition be-
tween two insect herbivores on oak. Ecol. Entomol.
15: 401-408.
HUNTER, M. D. 1992. A variable insect-plant interac-
tion: the relationship between tree budburst phenol-
ogy and population levels of insect herbivores among
trees. Ecol. Entomol. 17: 91-95.
HUNTER, M. D., AND J. N. MCNEIL. 1997. Host-plant
quality influences diapause and voltinism in a
polyphagous insect herbivore. Ecology 78(4): 977-986.
data base "Himawari" CD-ROM 98. Japan Weather
Association, Tokyo, Japan.

KATO, S., AND A. KOMIYAMA. 2002. Spatial and seasonal
heterogeneity in understory light conditions caused
by differential leaf flushing of deciduous overstory
trees. Ecol. Res. 17: 687-693.
KOMATSU, T., AND S. AKIMOTO. 1995. Genetic differenti-
ation as a result of adaptation to the phenologies of
individual host trees in the galling aphid Kalten-
bachiella japonica. Ecol. Entomol. 20: 33-42.
1997. White spruce and the spruce budworm: defin-
ing the phenological window of susceptibility. Cana-
dian Entomol. 129: 291-318.
The Egg of Insect Overwintering. Cambridge Uni-
versity Press, Cambridge, UK. 255 pp.
LOUDA, S. M., AND J. E. RODMAN. 1996. Insect herbivory
as a major factor in the shade distribution of a native
crucifer (Cardamine cordifolia A. Gray, bittercress).
J. Ecol. 84: 229-237.
LOWMAN, M. D. 1992. Leaf growth dynamics and her-
bivory in five species of Australian rain-forest can-
opy trees. J. Ecol. 80: 433-447.
MARTEL, J., AND A. KAUSE. 2002. The phenological win-
dow of opportunity for early-season birch sawflies.
Ecol. Entomol. 27: 302-307.
MASAKI, S. 1996. Geographical variation of life cycle in
crickets (Ensifera: Grylloidea). European J. Ento-
mol. 93: 281-302.
1999. CO2 and light effects on deciduous trees:
growth, foliar chemistry, and insect performance.
Oecologia 119: 389-399.
MIYAZAKI, M. 1987. Forms and morphs of aphids, pp. 27-
50 In A. K. Minks and P. Harrewijn [eds.], Aphids:
Their Biology, Natural Enemies and Control, Vol. A.
Elsevier Science Publisher, Netherlands.
MOUSSEAU, T. A. 1991. Geographic variation in mater-
nal-age effects on diapause in a cricket. Evolution
45: 1053-1059.
PRUESS, K. P. 1983. Day-degree methods for pest man-
agement. Environ. Entomol. 12: 613-619.
QUIRING, D. T. 1994. Influence of inter-tree variation in
time of budburst of white spruce on herbivory and
the behaviour and survivorship of Zeiraphera ca-
nadensis. Ecol. Entomol. 19: 17-25.
RAUSHER, M. D. 1979. Larval habitat suitability and
oviposition preference in three related butterflies.
Ecology 60: 503-511.
RAWORTH, D. A. 1994. Estimation of degree-days using
temperature data recorded at regular intervals. En-
viron. Entomol. 23: 893-899.

March 2006

REHILL, B., AND J. SCHULTZ. 2002. Opposing survivor-
ship and fecundity effects of host phenology on the
gall-forming aphid Hormaphis hamamelidis. Ecol.
Entomol. 27: 475-483.
ROFF, D. A., AND M. J. BRADFORD. 2000. A quantitative
genetic analysis of phenotypic plasticity of diapause
induction in the cricket Allonemobius socius. Hered-
ity 84: 193-200.
SEIWA, K. 1998. Advantages of early germination for
growth and survival of seedlings ofAcer mono under
different overstorey phenologies in deciduous broad-
leaved forests. J. Ecol. 86: 219-228.
SEIWA, K. 1999. Changes in leaf phenology are depen-
dent on tree height inAcer mono, a deciduous broad-
leaved tree. Ann. Bot. 83: 355-361.
SHURE, D. J., AND L. A. WILSON. 1993. Patch-size effects
on plant phenolics in successional openings of the
Southern Appalachians. Ecology 74: 55-67.
TAUBER, M. J., AND C. A. TAUBER 1976. Insect seasonal-
ity: diapause maintenance, termination, and postdia-
pause development. Annu. Rev. Entomol. 21: 81-107.
TAUBER, M. J., C. A. TAUBER, AND S. MASAKI. 1986. Sea-
sonal Adaptations of Insects. Oxford University
Press, New York. 411 pp.
UEMURA, S. 1994. Patterns of leaf phenology in forest
understory. Canadian J. Bot. 72: 409-414.
A. A. DHONDT. 1997. Synchronization of hatching
date with budburst of individual host trees (Quercus
robur) in the winter moth (Operophtera brumata) and
its fitness consequences. J. Anim. Ecol. 66: 113-121.
WADE, F. A., AND S. R. LEATHER 2002. Overwintering of
the sycamore aphid, Drepanosiphum platanoidis.
Entomol. Exp. Appl. 104: 241-253.
WANG, C. 2002. Effects of the host tree (Acer amoenum)
phenology on the ecology of the maple aphid (Periph-
yllus californiensis). Ph.D. dissertation, University
of Tokyo.
WANG, C., AND K. FURUTA. 2002. Diapause termination,
developmental threshold and thermal requirements
of eggs of the maple aphid, Periphyllus californiensis
SHINJI. J. For. Res. 7: 1-6.
WATT, A. D., AND A. M. MCFARLANE. 1991. Winter moth
on Sitka spruce: synchrony of egg hatch and bud-
burst, and its effect on larval survival. Ecol. Ento-
mol. 16: 387-390.
WINT, W. 1983. The role of alternative host-plant spe-
cies in the life of a polyphagous moth, Operophtera
brumata (Lepidoptera: Geometridae). J. Anim. Ecol.
52: 439-450.
ZAR, J. H. 1999. Biostatistical Analysis. 4th ed. Prentice
Hall, NJ. 663 pp.

Florida Entomologist 89(1)

Lewter et al.: Fall Armyworm Genetics


'Department of Entomology, University of Arkansas, Fayetteville, AR 72701

2USDA-ARS Center for Medical Agricultural and Veterinary Entomology, Gainesville, FL 32608


Limited information exists on molecular genetic variation and distribution of the corn and
rice strains of the fall armyworm, Spodoptera frugiperda (J.E. Smith). This study was con-
ducted to investigate the genetic structure of S. frugiperda across a part of its range in the
United States. A 608-base-pair portion of the mitochondrial cytochrome oxidase I and II
genes was sequenced from 71 individuals resulting in three corn and four rice strain haplo-
types. Genetic divergence between the two strains ranged from 0.66 to 0.99%. A 562-base-
pair region of the nuclear ITS-1 gene was also amplified and sequenced from 17 individuals
representing both corn and rice strains. No variation was detected in any of the samples for
the ITS-1 region. Analysis of molecular variance was conducted on the resulting mtDNA
haplotypes from the Arkansas and Florida populations and as a hierarchical analysis be-
tween populations in the two states. Results indicate a significant overall (O, for all popula-
tions with the hierarchical analysis revealing that this significant (0 is due to structuring
of the populations between states. The observed genetic structure is possibly due to the dis-
tribution of fall armyworm strains.

Key Words: COI, COII, ITS-1, DNA sequence, genetic variation, population genetics,
Spodoptera frugiperda


Existe informaci6n limitada sobre la variaci6n gen6tica molecular y distribuci6n de las razas
del cogollero, Spodoptera frugiperda (J.E. Smith) de maiz y de arroz. Este studio fue reali-
zado para investigar la estructura gen6tica de S. frugiperda a trav6s de una parte de su
rango de distribuci6n en los Estados Unidos. Una porci6n de los 608 pares de bases de los ge-
nes I y II del citocromo-c-oxidasa mitocondrial fueron secuenciados de 71 individuos resul-
tando en tres haplotipos de la raza de cogollero en el maiz y cuatro haplotipos de la raza de
cogollero en el arroz. La divergencia gen6tica entire las dos razas de cogollero fue de 0.66 a
0.99%. Una region de 562 pares de bases del gene nuclear ITS-1 tambi6n fue amplificada y
secuenciada de 17 individuos representando ambas razas de maiz y de arroz. Ningun varia-
ci6n fue detectada en las muestras para la region ITS-1. Un andlisis de variancia fue reali-
zado usando los haplotipos resultantes de ADNmt de las poblaciones de Arkansas y de
Florida, al igual que un andlisis dejerarquia entire las poblaciones de los dos estados. Los re-
sultados indican una 0~ total significativa para todas las poblaciones con el andlis de jera-
quia revelando que esta (O significativa es debido a la estructura de las poblaciones entire los
dos estados. La estructura gen6tica observada posiblemente es debido a la distribuci6n de
las razas de cogollero.

The fall armyworm, Spodoptera frugiperda
(J.E. Smith), is a major pest on corn, sorghum,
and bermudagrass in the southeastern United
States (Knipling 1980; Pashley 1986; Sparks
1979). The preferred host plants of the fall army-
worm came under new scrutiny in 1986 when
Pashley proposed that the fall armyworm consists
of two morphologically undistinguishable strains,
a corn strain that prefers corn, cotton, and sor-
ghum, and a rice strain that prefers rice and ber-
mudagrass (Pashley 1986, 1988a). The range of
S. frugiperda is known to cover most of the West-
ern hemisphere, and the range of each strain,
however, has been examined from Louisiana down

through Central America and in the Caribbean to
Brazil (Pashley et al. 1985; Pashley 1986, 1988b).
Despite the possible benefits that population
genetic analysis of the fall armyworm may pro-
vide towards understanding dispersal, monitor-
ing the spread of insecticide resistance, and the
implementation of area-wide control programs,
relatively little research in this area has been
conducted. A survey of 22 allozyme loci by Pash-
ley et al. (1985) indicated significant heterogene-
ity between populations at five of 11 polymorphic
loci, due in large part to the distinctness of a sin-
gle Puerto Rican population collected from rice.
The phylogenetic relationships between the two

Florida Entomologist 89(1)

strains were further examined with three of these
polymorphic allozymes (Hbdh, PepF, and Est3;
Pashley 1988b). The majority of the genetic stud-
ies have focused on differentiating the rice and
corn strains with polymerase chain reaction re-
striction fragment length polymorphism (PCR-
RFLP), strain specific PCR, RFLP, amplified frag-
ment length polymorphism (AFLP) and allozyme
markers (Meagher & Gallo-Meagher 2003; Levy
et al. 2002; Nagoshi & Meagher 2003; McMichael
& Prowell 1999; Pashley et al. 1985; Lu et al.
1992; Adamczyk 1993; Lu & Adang 1996; Pashley
1989). A genetic variation study by Lu et al.
(1992) involving RFLP of a random genomic li-
brary from six populations (five of which were lab
colonies) from Louisiana, Mississippi, and Geor-
gia revealed high levels of genetic variation
within and among populations. However, no pop-
ulation genetic analysis was conducted in that
study, which focused on finding diagnostic mark-
ers for the corn and rice strains.
Mitochondrial-DNA (mtDNA) analysis is gen-
erally assumed to be more powerful than allo-
zyme analysis for revealing population structure,
and has been used for numerous population ge-
netic studies (Avise 1994). The cytochrome oxi-
dase I (COI) and cytochrome oxidase II (COII) re-
gions of the mtDNA genome have proved useful
for measuring genetic variation in numerous in-
sect taxa (Szalanski & Owens 2003; Austin et al.
2002; Taylor et al. 1997; Brower & Jeansonne
2004). Comparison of mtDNA variation with a
nuclear genetic variation can provide insight into
current versus historical gene flow in a species.
For example, high levels of mtDNA variation com-
bined with a lack of nuclear DNA variation may
indicate unidirectional mating between strains.
We investigated the extent of genetic variation
within and between races of fall armyworm using
DNA sequences of a portion of the mitochondrial

COI and COII genes, and the nuclear rRNA first
internal transcribed spacer (ITS-1) region.


Larval fall armyworm samples were collected
from sorghum and cotton in Raymond, MS and
Colfax, LA, respectively (Table 1). Additional lar-
val samples were obtained from southern Florida
and Altheimer, Arkansas, and larval and pupal
samples from lab colonies maintained at the Uni-
versity of Mississippi and the University of Flor-
ida also were obtained. Larval species identifica-
tion was confirmed with morphological keys of
Peterson (1962), and samples were designated as
corn or rice strain based on the host from which
they were collected (Table 1). Fall armyworm
adults were collected with pheromone traps
through summer and fall of 2001 to 2003 from
three locations in Arkansas: Tillar, Foreman, and
Fayetteville (Table 1). The traps at Tillar were lo-
cated on the border of experimental research
plots of different field crops (cotton, corn, soy-
bean, and sorghum). The adjacent landscape was
predominantly cotton with limited acreages of
soybean, rice, and corn. A large commercial field
of coastal bermudagrass was located within 14
mile of the traps. The location at Foreman was on
a grain farm and the predominant crops were
corn, soybean, peanuts, and sorghum. Some lim-
ited areas of commercial pasture were near the
sample areas. The location at Fayetteville was on
an agricultural research farm located in an ur-
ban/suburban area. Diverse crops and grasslands
were located nearby. Adult fall armyworm identi-
fication was confirmed by comparing DNA se-
quences to larval fall armyworm and other noc-
tuid DNA sequences (unpublished data).
DNA was extracted from individual moths, lar-
vae, and pupae with the Puregene DNA isolation


Location Strain* C1 C2 C3 R1 R2 R3 R4 n

Fayetteville, Washington Co., AR 11 2 3 16
Tillar, Drew Co., AR -6 1 3 10
Foreman, Little River Co., AR -7 3 1 11
Altheimer, Jefferson Co., AR R 4 4
Starkville, Oktibbeha Co., MS C 2 2
Raymond, Hinds Co., MS C 2 2
Colfax, Grant Parish, LA C 2 2
Gainesville, Alachua Co., FL C 2 2
Ona, Hardee Co., FL R 4 3 7
Miami-Dade Co., FL R 6 1 7
Collier Co., FL R 5 5
Broward Co., FL R 2 1 3
n 32 2 1 4 29 1 1 71

*Strain designation based on host from which larvae were collected.

March 2006

Lewter et al.: Fall Armyworm Genetics

kit D-5000A (Gentra, Minneapolis, MN). Voucher
specimens are maintained at the Arthropod Mu-
seum, Department of Entomology, University of
Arkansas, Fayetteville, AR. DNA vouchers, pre-
served on filter paper according to Owens & Sza-
lanski (2005), are maintained at the Insect Genet-
ics Laboratory, Department of Entomology, Uni-
versity of Arkansas, Fayetteville, AR.
PCR reactions were conducted with 1 pl of the
extracted DNA with New England Biolabs (Ips-
wich, MA) Taq DNA polymerase with thermopol
buffer. Approximately 608 bp of a mtDNA region
containing the COI, tRNA leucine, and COII
genes was amplified with the primers C1-J-2797
et al. 1994) and C2-N-3400 (5'-TCAATATCAT-
TGATGACCAAT-3') (Taylor et al. 1997). The
mtDNA marker was amplified with a thermal cy-
cler profile consisting of 35 cycles of 94C for 45 s,
46C for 45 s and 72C for 45 s according to Sza-
lanski et al. (2000). A 562-bp section of the nu-
clear 3' portion of 18S rDNA, all of ITS-1, and the
5' portion of 5.8S were amplified with the primers
(Vrain et al. 1992) and rDNA 1.58S (5'-GCCAC-
CTAGTGAGCCGAGCA-3') (Cherry et al. 1997)
with a thermal cycler profile consisting of 40 cy-
cles of 94C for 45 s, 53C for 1 min and 72C for
1 min as described by Szalanski & Owens (2003).
Amplified DNA from individual moths was puri-
fied and concentrated with minicolumns accord-
ing to the manufacturer's instructions (Wizard
PCRpreps, Promega). Samples were sent to The
University of Arkansas Medical School DNA Se-
quencing Facility (Little Rock, AR) for direct se-
quencing in both directions.
Consensus sequences were derived from both
of DNA sequences from an individual with Bioedit
5.09 (Hall 1999) to verify nucleotide polymor-
phisms, and sequences were aligned by
CLUSTAL W (Thompson et al. 1994) for both
mtDNA and nDNA sequences. Mitochondrial
DNA haplotypes were aligned by MacClade v4
(Sinauer Associates, Sunderland, MA). GenBank
accession numbers were AY714298 to AY714304
for the different fall armyworm haplotypes. Gene-
alogical relationships among mtDNA haplotypes
were constructed with TCS (Clement et al. 2000)
and the method described by Templeton et al.
(1992). The distance matrix option of PAUP*
4.0b10 (Swofford 2001) was used to calculate ge-
netic distances according to the Kimura 2-param-
eter model of sequence evolution (Kimura 1980).
Tests for population differentiation were con-
ducted by AMOVA as implemented in Arlequin v.
2.0 (Schneider et al. 2000). An analog of FST, sD,
was calculated from the haplotypes frequencies
and Tajima and Nei (1984) genetic distances (Ex-
coffier et al. 1992). Initially, AMOVA was used to
test mtDNA genetic differentiation among all Ar-
kansas and Florida populations sampled (4)T).

Subsequently, a hierarchical AMOVA was con-
ducted in which populations were grouped into
states to determine differentiation between states
(ocT) and among populations within states (gD T).
Pairwise comparisons, calculated independently
for all Arkansas and Florida population pairs of
O~, also were calculated. Permutations of the data
set were used to determine statistical significance
of the pairwise comparisons (P < 0.05).


Mitochondrial DNA sequencing of 71 fall ar-
myworm samples revealed an amplicon size of
608 bp. Nucleotide positions 1 to 222 were COI,
223 to 289 tRNA-leu, and 290 to 608 COII. The
average base frequencies were A = 0.36, C = 0.13,
G = 0.09, and T = 0.42. Corn haplotype C1 was the
most common haplotype for the corn strain and
occurred in all of the sampled locations where the
corn strain occurred (Table 1). The other two corn
strain haplotypes were found only in Arkansas.
Rice strain haplotype R2 was the most common
haplotype and occurred in every location where
the rice strain was found. Rice strains R1 and R4
were found only in Florida, while strain R3 was
found in both Arkansas and Florida.
Nine nucleotide sites were variable among the
observed three corn and four rice strain haplo-
types (Table 2). Three variable nucleotide sites
were located in the COI gene and the remainder
were located in the COII gene. Tajima-Nei dis-
tances (Tajima & Nei 1984) among the fall army-
worm haplotypes ranged from 0.164 to 0.329% for
the corn strain, 0.164 to 0.329% for the rice
strain, and 0.658 to 0.987% between strains. Fig.
1 shows the 95% parsimony network for the seven
haplotypes (Posada & Crandall 2001). Missing
haplotypes probably represent sampling gaps.
DNA sequencing of the nuclear ITS-1 region
from 17 FAW samples (Table 3) revealed an ampl-
icon size of 562 base pairs. No sequence variation
was detected in any of the 17 individuals and the
base frequencies were A = 0.23, C = 0.24, G = 0.26,
and T = 0.27.
AMOVA detected a significant overall 4,ST
(0.493, P < 0.001) when comparing mtDNA ge-
netic variation among populations (Table 4). The
amount of variation was almost equal within ver-
sus among populations (within 50.74%, among
49.26%). Hierarchical AMOVA conducted between
Arkansas and Florida populations detected a sig-
nificant ,CT (0.387, P < 0.005) between the two
states (Table 4). The comparison among groups ac-
counted for 38.70% of the observed variation.


This genetic investigation of the fall army-
worm mtDNA revealed significant levels of ge-
netic differentiation among populations both

Florida Entomologist 89(1)


Haplotype 55 76 79 361 367 403 421 511 529

Corn1 A C A T T A C T C
Corn 2 T
Corn 3 C
Rice 1 T C C G A
Rice 2 T C C A
Rice 3 T C C T A
Rice 4 T C C C A

within and between the two fall armyworm
strains. This research also represents the first at-
tempt to determine the geographical distribution
of fall armyworm haplotypes from mtDNA se-
quence data as well as determining the extent of
genetic variation within each strain. A haplotype
or allele is defined by one unique form of the gene
and differs from any other gene by at least one
nucleotide. Haplotype diversity or gene diversity
quantifies the number of haplotypes in relation to
their relative frequency to each other, and haplo-
type diversity is described as the probability that
two sequences randomly selected from a popula-
tion are different (Nei 1987).
Four haplotypes were observed for the rice
strain and three haplotypes were found for the
corn strain, although it is likely more haplotypes
may be discovered for each strain. Observed ge-
netic variation between strains was approxi-

Fig 1. Genealogical relationships among 7 haplo-
types of fall armyworm estimated by TCS (Clement et
al. 2000). The size of the ovals corresponds to haplotype
frequency, and a unit branch represents one mutation.
Small ovals indicate haplotypes that were not observed.

mately 0.66%. Estimated time of divergence be-
tween corn and rice strain is approximately
287,000 years based on a molecular clock rate of
2.3% divergence per million years (Brower 1994).
Populations of nearly all species, social or other-
wise, exhibit at least some degree of genetic dif-
ferentiation among geographic locales (Ehrlich &
Raven 1969). This observation becomes more dif-
ficult to accurately discern when dealing with a
migratory species such as the fall armyworm;
however, more studies such as this one could help
determine the migratory paths of the insect.
One of the purposes of the research presented
herein was to estimate the baseline genetic vari-
ation which occurs both within and between fall
armyworm strains. As with other animal popula-
tions, additional genetic structure normally is to
be expected over increasing spatial scales, where
populations can show additional differentiation
due to spatial habitat structure, isolation by dis-
tance, or other factors (Avise 1994). There may be
temporal differences in the occurrence of the rice
and corn. Temporal data, obtained by sampling
the same area throughout a season and over a pe-
riod of years, also may provide insight into the
specific migratory patterns of the fall armyworm.
Comparing mtDNA sequences with nuclear
markers can provide evidence of inter-strain mat-
ing within a species. The lack of variation in the
nuclear rDNA ITS-1 region combined with previ-
ously conducted laboratory-based mating studies
(Pashley & Martin 1987;Whitford et al. 1988; Na-
goshi & Meagher 2003) suggests that inter-strain
mating does occur in the field. However, the lack
of genetic variation in the rDNA ITS-1 region


County State mtDNA haplotype (n)

Washington AR C1(3), C2(1), R2(1)
Little River AR C1(3)
Drew AR C1(5)
Jefferson AR R2(2)
Collier FL R2(1)
Broward FL R3(1)

March 2006

Lewter et al.: Fall Armyworm Genetics


Hierarchy Categories %Variation ( estimate

One Group Among Populations 49.26 (s, = 0.493*
Within Populations 50.74

Two Groups Among Groups 38.70 (0, = 0.387*
Among Populations 19.45 (O, = 0.582*
Within Groups
Within Populations 41.85

must be approached with caution, because this
marker has no power to detect gene flow and this
invariant region may be ancestral to strain subdi-
vision. Prowell et al. (2004) also reported a lack of
variation in the ITS-1 region of the fall army-
worm, but it was cited as unpublished data.
Based on this study, there appears to be suffi-
cient genetic variation both within and between
populations to substantiate a more comprehen-
sive population genetics study on this species,
and we would recommend also that temporal data
be taken into consideration.


We thank Ralph Bagwell, Don Parker, and Clint
Allen for providing samples, and John W. Jones for tech-
nical assistance. Research was supported by a Univer-
sity of Arkansas, Arkansas Agricultural Experiment
Station research initiation grant.


ADAMCZYK, J. 1993. The UBS DNA Isolation Kit is a re-
liable method for extracting total nucleic acid from
insect tissue. USBiochemical Comments 20: 25-26.
AND A. KENCE. 2002. A comparative genetic analysis
of the subterranean termite genus Reticulitermes
(Isoptera: Rhinotermitidae). Ann. Entomol. Soc.
Amer. 95: 753-760.
AVISE, J. C. 1994. Molecular Markers, Natural History
and Evolution. Chapman & Hall, New York, NY. 511
BROWER A. V. Z. 1994. Rapid morphological radiation
and convergence among races of the butterfly Heli-
conuius erato inferred from patterns of mitochon-
drial DNA evolution. Proc. Natl. Acad. Sci. USA 91:
BROWER, A. V. Z., AND M. M. JEANSONNE. 2004. Geo-
graphical populations and "subspecies" of new world
monarch butterflies (Nymphalidae) share a recent
origin and are not phylogenetically distinct. Ann.
Entomol. Soc. Amer. 97: 519-527.
POWERS. 1997. The internal transcribed spacer re-
gion of Belonolaimus (Nemata: Belonolaimidae). J.
Nematol. 29: 21-29.

TCS: a computer program to estimate 6 genealogies.
Mol. Ecol. 9: 1657-1659.
EHRLICH, P. R., AND P. H. RAVEN. 1969. Differentiation
of populations. Science 165: 1228-1232.
Analysis of molecular variance inferred from metric
distances among DNA haplotypes: application to hu-
man mitochondrial DNA restriction data. Genetics
131: 479-491.
HALL, T. A. 1999. BioEdit: a user-friendly biological se-
quence alignment editor and analysis program for
Windows 95/98/NT. Nucl. Acids. Symp. Ser. 41:95-98.
KIMURA, M. 1980. A simple method for estimating evo-
lutionary rate of base substitutions through compar-
ative study of nucleotide sequences. J. Molec. Evol.
16: 111-120.
KNIPLING, E. F. 1980. Regional management of the fall
armyworm-a realistic approach? Florida Entomol.
63: 468-480.
NIAK. 2002. Strain identification of Spodoptera fru-
giperda (Lepidoptera: Noctuidae) insects and cell
line: PCR-RFLP of cytochrome oxidase subunit I
gene. Florida Entomol. 85: 186-190.
LU, Y., AND M. J. ADANG. 1996. Distinguishing fall ar-
myworm (Lepidoptera: Noctuidae) strains using a
diagnostic mitochondrial DNA marker. Florida En-
tomol. 79: 49-55.
KOCHERT. 1992. RFLP analysis of genetic variation
in North American populations of the fall armyworm
moth Spodoptera frugiperda (Lepidoptera: Noctu-
idae). Mol. Ecology 1: 199-208.
MCMICHAEL, M., AND D. P. PROWELL. 1999. Differences
in amplified fragment-length polymorphisms in fall
armyworm (Lepidoptera: Noctuidae) host strains.
Ann. Entomol. Soc. Amer. 92: 175-181.
Identifying host strains of fall armyworm (Lepi-
doptera: Noctuidae) in Florida using mitochondrial
markers. Florida Entomol. 86: 450-455.
NAGOSHI, R. N., AND R. L. MEAGHER. 2003. FR tandem-
repeat sequence in fall armyworm (Lepidoptera:
Noctuidae) host strains. Ann. Entomol. Soc Amer.
96: 329-335.
NEI, M. 1987. Molecular evolutionary genetics. Colum-
bia University Press, New York.
OWENS, C., AND A. L. SZALANSKI. 2005. Filter paper for
preservation, storage and distribution of insect and
pathogen DNA samples. J. Med. Entomol. (in press)

PASHLEY, D. P. 1986. Host associated genetic differenti-
ation in fall armyworm (Lepidoptera: Noctuidae): a
sibling species complex? Ann. Entomol. Soc. Amer.
79: 898-904.
PASHLEY, D. P. 1988a. Quantitative genetics, develop-
ment and physiological adaptation in sympatric host
strains of fall armyworm. Evolution 42: 93-102.
PASHLEY, D. P. 1988b. Current status of fall armyworm
host strains. Florida Entomol. 71: 227-234.
PASHLEY, D. P. 1989. Host-associated differentiation in
armyworms (Lepidoptera: Noctuidae): An allozymic
and mitochondrial DNA perspective, pp. 103-114 In
H. D. Loxdale and J. den Hollander [eds.], Systemat-
ics Association Special Volume No. 39. Claredon
Press, Oxford.
PASHLEY, D. P., AND J. A. MARTIN. 1987. Reproductive
incompatibility between host strains of the fall ar-
myworm (Lepidoptera: Noctuidae). Ann. Entomol.
Soc. Amer. 80: 731-733.
Genetic population structure of migratory moths:
the fall armyworm (Lepidoptera: Noctuidae). Ann.
Entomol. Soc. Amer. 78: 756-762.
PETERSON, A. 1962. Larvae of Insects. Edward Bros.,
Inc., Ann Arbor, MI.
POSADA, D., AND K. A. CRANDALL. 2001. Performance of
methods for detecting recombination from DNA se-
quences: computer simulations. Proc. Natl. Acad.
Sci. USA 98: 13757-13762.
2004. Multilocus genetic analysis of host use, intro-
gression, and speciation in host strains of fall army-
worm (Lepidoptera: Noctuidae. Ann. Entomol. Soc.
Amer. 97: 1034-1044.
Arlequin: A software for population genetic data. Ge-
netics and Biometry Laboratory, University of
Geneva, Switzerland.
AND P. FLOOK. 1994. Evolution, weighting, and phy-
logenetic utility of mitochondrial gene sequences and
a compilation of conserved polymerase chain reaction
primers. Ann. Entomol. Soc. Amer. 87: 651-701.

March 2006

SPARKS, A. N. 1979. A review of the biology of the fall ar-
myworm. Florida Entomol. 62: 82-87.
SWOFFORD, D. L. 2001. PAUP*: Phylogenetic analysis
using parsimony (*and other methods), ver. 4.0b10.
Sinauer, Sunderland, MA.
SZALANSKI, A. L., AND C. B. OWENS. 2003. Genetic vari-
ation of the southern corn rootworm, Diabrotica un-
decimpunctata howardi (Coleoptera: Chrysomelidae).
Florida Entomol. 86: 329-333.
FRITZ. 2000. Population genetics and phylogenetics
of the endangered American burying beetle, Nicro-
phorus americanus (Coleoptera: Silphidae). Ann.
Entomol. Soc. Amer. 93: 589-594.
TAJIMA, F., AND M. NEI. 1984. Estimation of evolution-
ary distance between nucleotide sequences. Mol.
Biol. Evol. 1: 269-285.
J. J. PETERSEN. 1997. Mitochondrial DNA variation
among Muscidifurax spp. (Hymenoptera: Pteromal-
idae), pupal parasitoids of filth flies. Ann. Entomol.
Soc. Am. 90: 814-824.
1992. A cladistic analysis of phenotypic associations
with haplotypes inferred from restriction endonu-
clease mapping and DNA sequence data. III. Cla-
dogram estimation. Genetics 132: 619-633.
1994. CLUSTAL W: improving the sensitivity of pro-
gressive multiples sequence alignments through se-
quence weighting, position-specific gap penalties
and weight matrix choice. Nucleic Acids Res. 22:
R. I. HAMILTON. 1992. Intraspecific rDNA restriction
fragment length polymorphism in the Xiphinema
americanum group. Fundam. Appl. Nematol. 15:
W. LEE. 1988. Oviposition preference, mating com-
patibility and development of two fall armyworm
strains. Florida Entomol. 71: 234-243.

Florida Entomologist 89(1)

Saarinen & Daniels: Miami Blue Butterfly Larvae and Ants


'Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611-0620, USA

2McGuire Center for Lepidoptera and Biodiversity Research, Florida Museum of Natural History
University of Florida, Gainesville, FL 32611-2710, USA


Historical, anecdotal records of the state-endangered Miami blue butterfly, Cyclargus tho-
masi bethunebakeri (Comstock & Huntington) (Lepidoptera), have mentioned larval associ-
ations with the Florida carpenter ant, Camponotus sp. Recent population studies confirm
that C. t. bethunebakeri larvae associate with Camponotus floridanus (Buckley) as well as
another member of the genus, Camponotus planatus (Roger). Additionally, caterpillars have
been observed tended by Crematogaster ashmeadi (Emery), Forelius pruinosus (Roger), and
Tapinoma melanocephalum (Fab.). Field surveys of remaining Miami blue habitat and re-
cent butterfly reintroduction sites reveal other potential ant associates, Paratrechina longi-
cornis (Latreille) and Paratrechina bourbonica (Forel), and a host of possible predaceous ant
species. The corresponding conservation implications are discussed. Detailed information is
also presented about larval ant-associated organs and their mediation of this facultative

Key Words: Ant organs, butterfly-ant relationship, facultative symbiosis


Registros hist6ricos y anecd6ticos de la mariposa en peligro de extinci6n Miami blue, Cyclar-
gus thomasi bethunebakeri (Comstock y Huntington) (Lepid6ptera) mencionan la asociaci6n
de sus larvas con la hormiga carpintera, Camponotus sp. Estudios recientes poblacionales
confirman que las larvas de C. t. bethunebakeri estan asociadas con Camponotus floridanus
(Buckley), como tambi6n con otro miembro del g6nero, Camponotus planatus (Roger). Adi-
cionalmente, se observaron orugas atendidas por Crematogaster ashmeadi (Emery), Forelius
pruinosus (Roger), y Tapinoma melanocephalum (Fabricius). Analisis de campo del habitat
remanente de la mariposa Miami blue y de localidades de reintroducciones recientes, reve-
laron asociaciones potenciales con otras species de hormigas, Paratrechina longicornis (La-
treille) y Paratrechina bourbonica (Forel), y un hospedero de posibles species de hormigas
depredadoras. Las implicaciones de conservaci6n son discutidas en este articulo. Asi mismo,
se present informaci6n detallada sobre los 6rganos involucrados en la asociaci6n larva-hor-
miga y su intervenci6n en esta simbiosis facultativa.

Translation provided by the authors.


The Miami blue, Cyclargus thomasi bethune-
bakeri (Comstock & Huntington) (Lycaenidae:
Polyommatinae), represents one of Florida's rarest
endemic butterflies and is currently listed as state-
endangered. Once commonly found in tropical
coastal hammocks, beachside scrub, and tropical
pine rocklands from the southern Florida main-
land south through the Florida Keys to Key West
and the Dry Tortugas, the species' overall distribu-
tion and numerical abundance has been reduced to
a single remaining metapopulation within the
boundaries of Bahia Honda State Park in the

Lower Keys (Klots 1964; Kimball 1965; Lencze-
wski 1980; Minno & Emmel 1993; Ruffin & Glass-
berg 2000; Calhoun et al. 2002). Developing larvae
of C. t. bethunebakeri have been shown to be tended
by ants in the genus Camponotus but the extent of
the relationship remains poorly understood (Minno
& Emmel 1993). Recent population studies of the
butterfly at Bahia Honda State Park and addi-
tional reintroduction sites within Everglades
National Park confirm a continued association.
Over 75 percent of lycaenid larvae with known
life histories associate with ants (Pierce et al.
2002). Such myrmecophilous relationships may
be mutualistic to varying degrees or even para-

Florida Entomologist 89(1)

sitic whereby larvae are predatory in ant nests
(Pierce & Mead 1981; Fiedler & Maschwitz 1988;
New 1993). The resulting communication be-
tween larvae and ants is mediated by a complex
array of tactile, chemical, and often audible sig-
nals (DeVries 1990). Specifically, larvae possess
highly specialized organs that can extrude alarm,
reward, or appeasement chemicals. In response,
tending ants often protect the surrounding larvae
from a variety of natural predators and parasi-
toids, and thus can potentially provide a benefit
for survival (Thomas 1980; Webster & Nielson
1984; Pierce & Easteal 1986; Savignano 1994).
Cushman & Murphy (1993) suggest that ant as-
sociations also may play an important role in the
persistence of lycaenid populations. They addi-
tionally propose that species with a dependence
on ants, whether facultative or obligatory, display
an increased sensitivity to environmental change,
and thus are more susceptible to endangerment
than species that lack ant associations. Here, we
identify additional ant associates and potential
predatory ant species and discuss the correspond-
ing implications for the conservation and recov-
ery of the Miami Blue butterfly, a critically imper-
iled butterfly.


Field surveys of ant species were conducted at
Bahia Honda State Park and the Flamingo
Campground, Rowdy Bend Trail, and Bear Lake
Road sites in southern portions of Everglades Na-
tional Park during daylight hours on 24-27 May,
2004 and 31 July-2 August, 2004. These areas
contain low numbers of Cyclargus thomasi be-
thunebakeri, either as part of a remaining natural
metapopulation or as reintroduced individuals.
Hand-collecting and baiting were used to survey
ants on and around patches of the butterfly's lar-
val host, Caesalpinia bonduc (L.) Roxb. (Fa-
baceae). Sugar baits consisting of index cards
with approximately 10 g of crushed pecan cookie
were placed along transects at the base of C. bon-
duc plants. Baits were left in the field for one
hour, at which time all cards were collected in Zi-
ploc-style plastic bags. Additionally, when C. t. be-
thunebakeri larvae were found in association
with ants, 1-2 ant specimens were collected from
the tended larvae.
Finally, to provide additional detail on the
structure of the larval ant organs, three C. t. be-
thunebakeri larvae from a captive colony main-
tained at the University of Florida were pre-
served and used for SEM and Auto-Montage pho-
tographic analysis. Larvae were placed in near
boiling water for 60 seconds, transferred to 25%
ethanol for two hours, 50% ethanol for another
two hours, and stored in 75% ethanol before being
photographed. No additional preparation or gold
coating was done to prepare specimens.


Eighteen ant species were collected in Ever-
glades National Park and Bahia Honda State
Park (Table 1). Of these, Camponotus floridanus,
Camponotus planatus, Crematogaster ashmeadi,
Forelius pruinosus, and Tapinoma melanoceph-
alum were confirmed to tend larvae of Cyclargus
thomasi bethunebakeri. Late instars were always
found in association with ants but early instars,
prepupae, and pupae were frequently found with-
out ants present. Camponotus floridanus tended
larvae for the majority of the observations and all
other ants were encountered 1-2 times, with no
two species tending larvae simultaneously. Two
ants typically tended a larva at a time, with the
exception of Crematogaster ashmeadi which often
tended in higher numbers (Fig. 1).
We name two additional species, Paratrechina
longicornis and Paratrechina bourbonica, as po-
tential ant associates. The former species was
found in proximity to C. t. bethunebakeri larvae
and appeared to tend them although encounters
were brief. The latter species was observed tend-
ing larvae of another lycaenid, S'r ....i martialis
(Herrich-Schaffer), on Caesalpinia bonduc at Ba-
hia Honda State Park. No predation by these ants
was observed.
Details of the ant organs of C. t. bethunebakeri
are shown in Fig. 2. Second through fifth instars
possess a dorsal nectary organ (=honey gland)
with associated perforated cupola organs on ab-
dominal segment A7 and a pair of eversible tentac-
ular organs on abdominal segment A8. Abdominal
segments A7 and A8 are fused dorsally. Tentacular
organs were observed to evert independently in
the field when stimulated by attendant ants, and
liquid droplets from the dorsal nectary organ were
actively imbibed by all species of ants. Campono-
tus floridanus became excited and agitated, evi-
denced by increased body and antennal move-
ments, when the tentacular organs were everted.


This study documents Camponotus floridanus
to be the primary ant species attending Miami
blue larvae. Camponotus floridanus is a native
ant species primarily active at night throughout
Florida; they are commonly found foraging on
C. bonduc and tending C. t. bethunebakeri larvae
in both the Everglades and Bahia Honda loca-
tions. Camponotus planatus is a diurnal species,
but is not commonly encountered in association
with larvae, having never been found tending lar-
vae in the Everglades and only once in Bahia
Honda. It is possible that in higher densities
C. planatus may more regularly tend larvae and
could potentially be important at protecting lar-
vae during the day. Buckley & Gullan (1991) have
shown that more aggressive ants provide better

March 2006

Saarinen & Daniels: Miami Blue Butterfly Larvae and Ants


Park Location

Everglades Bahia Honda Ant Status

Subfamily Pseudomyrmicinae
Pseudomyrmex elongatus (Mayr) 1 2 P
Pseudomyrmex gracilis (Fab.) 1 P
Pseudomyrmex simplex (Smith) 1 P
Subfamily Myrmicinae
Crematogaster ashmeadi (Emery) 1, 2 S
Monomorium floricola (Jerdon) 1 2 u
Pheidole dentata Mayr 1 2 u
Pheidole floridana Emery 1 1 u
Solenopsis invicta Buren 1, 2 P
Solenopsis geminata (Fab.) 1 1, 2 u
Tetramorium simillimum (F. Smith) 1 u
Wasmannia auropunctata (Roger) 1 1, 2 P
Subfamily Dolichoderinae
Forelius pruinosus (Roger) 1 1, 2 S
Tapinoma melanocephalum (Fab.) 1 S
Subfamily Formicinae
Brachymyrmex obscurior Forel 1 1, 2 u
Camponotus floridanus (Buckley) 1, 3 1, 2 S
Camponotus planatus (Roger) 1 1 S
Paratrechina bourbonica (Forel) 1 pS
Paratrechina longicornis (Latreille) 1 1, 2 pS

Collectors/Authors: 1 = present study; 2 = Deyrup et al. (1988), 3 = Ferster and Prusak (1994).
S = confirmed symbiont ofC. thomasi bethunebakeri (present study); pS = potential symbiont; u
ator (noted as a predaceous ant in included literature).

protection for soft scales and mealybugs, and as a
corollary the large and potentially aggressive
Camponotus species may prove effective in deter-
ring predators and parasitoids (Axen 2000).
Crematogaster ashmeadi were observed tend-
ing individual larvae in Bahia Honda but have
not yet been observed with larvae in the Ever-
glades. While not commonly found tending larvae,
interactions involving C. ashmeadi were charac-
terized by a minimum of five individuals. This be-
havior demonstrates the quality of their trailing
and recruitment signals. Other Crematogaster
species have been found worldwide to tend lycae-
nid larvae and this genus seems predisposed to
lycaenid symbioses (Atsatt 1981; Fiedler 1991;
Pierce et al. 2002; Saarinen 2005). These ants are
equipped with a flexible abdomen and attached
sting; despite their small size they are potentially
capable of defending larvae from other ants or
harmful invertebrates.
Both Forelius pruinosus and Tapinoma mel-
anocephalum may be opportunistically imbibing
food rewards from C. t. bethunebakeri larvae.
Field observations suggest that their behavior of-
fers little or no protection for the larvae they tend;

unknown; P= potential pred

Fig. 1. Crematogaster ashmeadi ants tending a late
instar Cyclargus thomasi bethunebakeri larva. Several
other C. ashmeadi ants were present but not visible in
this photo. Photo by Jaret Daniels.

Florida Entomologist 89(1)

March 2006



Fig. 2. Details of Cyclargus thomasi bethunebakeri fifth instar. a, dorso-posterior abdominal segments (A7-A8)
showing ant organs; b, detail of cuticular setae; c, tentacular nectary organ inverted; d, tentacular nectary organ
everted (cuticular setae omitted); e, dorsal nectary organ bordered by perforated cupolas. Figures by Emily

d I'

';ALt N

*" "' L it 13' 'i

^ ^..75~^ ,t-,:..;;' ^
( 1I ''/ ' '

F '-^ ^ rfA^',' i / t ^
'. 1
+ !) ^ ^ .1 *i , / ,
I i/I '
,i .~

-r 6-
I -

1 0* m mi

1.0 mm

Saarinen & Daniels: Miami Blue Butterfly Larvae and Ants

however merely their presence may deter preda-
tors. Both Paratrechina longicornis and P. bour-
bonica, along with T melanocephalum, have been
referred to as "tramp ants" (Passera 1994). While
such species may not provide demonstrative pro-
tection for larvae, at the very least they tolerate
nearby larvae and coincidentally tend instars
feeding on C. bonduc flowers and buds adjacent to
where the ants are also gathering nectar.
The facultative ant associations of C. t. be-
thunebakeri encompass four genera (five includ-
ing Paratrechina) and three subfamilies; Formici-
nae, Myrmicinae, and Dolichoderinae. These ly-
caenid larvae may secrete "non-specific" ant semi-
ochemicals as attractants to various ant species,
as proposed by Henning (1983). These chemicals,
primarily from the tentacular organs and poten-
tially from the perforated cupola organs, may
serve to alarm, excite, or appease ants. Further
study into the chemical secretions of all ant or-
gans may clarify the "intentions" of the larvae in
their emissions. Further comparisons of each ant
species' alarm and attractant pheromones with
those isolated from lycaenid volatiles may further
elucidate ant-larval relationships, including if
certain ants are chemically targeted and if others
are simply opportunistic tenders.
No interactions between other identified ant
species and C. t. bethunebakeri larvae were ob-
served. Several of these ants, however, may be
predating larvae at other times. All three
Pseudomyrmex species may be predators, possi-
bly excepting P simplex (Smith) due to its small
size. Miami blue larvae are always found in prox-
imity to abundant colonies of Camponotus flori-
danus and further field observations, especially
at night when C. floridanus are most active, need
to be carried out to assess interactions within the
ant mosaic of symbionts and predators.
This study also shows the persistence of Was-
mannia auropunctata (Roger) on Bahia Honda
State Park (first recognized there by Deyrup et al.
1988). This invasive tramp ant is native to the
New World tropics and its presence in the Florida
Keys may be a cause for concern. W auropunc-
tata, also known as the little fire ant is an oppor-
tunistic feeder that forages day and night and
bears a painful sting. Both W auropunctata and
the red imported fire ant, Solenopsis invicta Bu-
ren, have been implicated in the displacement of
endemic species, resulting in a loss of biodiversity
(Meier 1994; Wojcik 1994). Neither ant has been
found near C. t. bethunebakeri in the Everglades,
nor have they been observed harvesting imma-
tures or predating adults in Bahia Honda. Sole-
nopsis invicta mound density is perhaps not high
enough to impact the C. t. bethunebakeri metapo-
pulation on Bahia Honda because they do not ap-
pear as large or extensive in area as those found
in more disturbed habitats (personal observa-
tion). Further field work will need to examine pre-

dation rates by ants and the specific impact that
these invasive ants may have on endemic butter-
fly species, especially species of special concern.
Ant attendance may be critical to the long-term
survival of lycaenid taxa by impacting larval de-
velopment time, larval weight gain, and other de-
velopmental responses (Robbins 1991; Wagner
1993). The presence of an ant guard has led to
larger, more fecund adults in the related butterfly
Hemiargus isola (Reakirt) (Wagner 1993). How-
ever in the Australian species Jalmenus evagoras
(Donovan), ant-tended larvae pupate at a smaller
size, pupate for a shorter duration, and develop
into smaller adults (Pierce et al. 1987). In an as-
sessment of potential ant partners, it was shown
that Tapinoma sessile (Say) is a "neutral partner"
for the widely distributed North American lycae-
nid Gl. !...... i.. .lygdamus (Doubleday), provid-
ing no significant cost or benefit (Fraser et al.
2001). Researchers of the critically imperiled
European lycaenid butterfly Maculinea rebeli
(Hirschke) have repeatedly emphasized "the im-
portance of identifying local host ant species prior
to further management conservation strategies in
order to avoid failure of management programs or
even damage to populations on the edge of extinc-
tion" (Steiner et al. 2003). Ant attendance, obligate
or facultative, is not trivial; it can have profound
effects on the length of time individuals spend in
vulnerable immature stages as well as the result-
ing fecundity of adults. Both symbiotic ant part-
nerships and the negative impacts of predaceous
ants should be addressed in management plans
for the conservation of endangered lycaenid taxa.


This research was conducted under permits EVER-
2004-SCI-0038 and 5-04-58 and we thank Sue Perry of
Everglades National Park and Bob Rundle of Bahia
Honda State Park for assistance with permitting. We
thank Mark Deyrup and Lyle Buss for assistance with
the ant identifications and ant life histories, Paul Skel-
ley for assistance with the SEM at the Division of Plant
Industry in Gainesville, Florida, and Tom Emmel for
useful comments on the manuscript.

ATSATT, P. R. 1981. Lycaenid butterflies and ants: selec-
tion for enemy-free space. Am. Nat. 118: 638-654.
AXEN, A. H. 2000. Variation in behavior of lycaenid lar-
vae when attended by different ant species. Evol.
Ecol. 14: 611-625.
BUCKLEY, R., AND P. GULLAN. 1991. More aggressive
ant species (Hymenoptera: Formicidae) provide bet-
ter protection for soft scales and mealybugs (Ho-
moptera: Coccidae, Pseudococcidae). Biotropica
23(3): 282-286.
2002. The rise and fall of the tropical blues in Florida:
Cyclargus ammon and Cyclargus thomasi bethunebak-
eri (Lepidoptera: Lycaenidae). Hol. Lep. 7(1): 13-20.

CUSHMAN, J. H., AND D. D. MURPHY. 1993. Conservation
of North American lycaenids-an overview, pp. 37-44
In T. R. New [ed.], Conservation Biology of Lycae-
nidae (Butterflies). IUCN, The World Conservation
Union, Gland, Switzerland.
DEVRIES, P. J. 1990. Enhancement of symbiosis be-
tween butterfly caterpillars and ants by vibrational
communication. Science 248: 1104-1106.
1988. A review of the ants of the Florida Keys. Flor-
ida Entomol. 71(2): 163-176.
FERSTER, B., AND Z. PRUSAK. 1994. A preliminary check-
list of the ants (Hymenoptera: Formicidae) of Ever-
glades National Park. Florida Entomol. 77(4): 508-512.
FIEDLER K. 1991. Systematic, evolutionary, and ecologi-
cal implications of myrmecophily within the Lycae-
nidae (Insecta: Lepidoptera: Papilionoidea). Bonner
Zool. Monogr. 31: 1-210.
FIEDLER, K., AND U. MASCHWITZ. 1988. Functional anal-
ysis of the myrmecophilous relationships between
ants (Hymenoptera: Formicidae) and lycaenids (Lep-
idoptera: Lycaenidae). II. Lycaenid larvae as tropho-
biotic partners of ants-a quantitative approach.
Oecologia 75: 204-206.
FRASER, A. M., A. H. AXEN, AND N.E. PIERCE. 2001. As-
sessing the quality of different ant species as partners
of a myrmecophilous butterfly. Oecologia 129:452-460.
HENNING, S. F. 1983. Chemical communication between
lycaenid larvae (Lepidoptera: Lycaenidae) and ants
(Hymenoptera: Formicidae). J. Entomol. Soc. S. Afr.
46: 341-366.
KIMBALL, B. 1965. The Lepidoptera of Florida. An anno-
tated checklist. Arthropods of Florida and Neighbor-
ing Land Areas 1:1-363, Florida Department of
Agriculture, Division of Plant Industry, Gainesville.
KLOTS, A. B. 1964. A Field Guide to the Butterflies of
North America, East of the Great Plains. Houghton
Mifflin, Boston.
LENCZEWSKI, B. 1980. Butterflies of Everglades National
Park. National Park Service, South Florida Research
Center, Homestead, Florida. Report T-588. 110 pp.
MEIER, R. E. 1994. Coexisting patterns and foraging be-
havior of introduced and native ants (Hymenoptera
Formicidae) in the Galapagos Islands (Ecuador), pp.
44-62 In D. F. Williams [ed.], Exotic Ants: Biology,
Impact, and Control of Introduced Species. Westview
Studies in Insect Biology, Boulder, CO.
MINNO, M. C., AND T. C. EMMEL. 1993. Butterflies of the
Florida Keys. Scientific Publishers, Gainesville.
NEW, T. R. 1993. Conservation Biology of the Lycae-
nidae (Butterflies). IUCN Publ, Rep. 8, Gland, Swit-

March 2006

PASSERA, L. 1994. Characteristics of tramp ant species,
pp. 23-43 In D. F. Williams [ed.], Exotic Ants: Biology,
Impact, and Control of Introduced Species. Westview
Studies in Insect Biology, Boulder, CO.
The ecology and evolution of ant association in the
Lycaenidae (Lepidoptera). Annu. Rev. Entomol. 47:
PIERCE, N. E., AND S. EASTEAL. 1986. The selective ad-
vantage of attendant ants for the larvae of a lycaenid
butterfly, Glaucopsyche lygdamus. J. Anim. Ecol. 55:
TAYLOR, AND K. F. BENBOW. 1987. The costs and ben-
efits of cooperation between the Australian lycaenid
butterfly, Jalmenus evagoras, and its attendant ants.
Behav. Ecol. Sociobiol. 21: 237-248.
PIERCE, N. E., AND P. S. MEAD. 1981. Parasitoids as se-
lective agents in the symbiosis between lycaenid
butterfly larvae and ants. Science 211: 1185-1187.
ROBBINS, R. K. 1991. Cost and evolution of a facultative
mutualism between ants and lycaenid larvae. Oikos
62: 363-369.
RUFFIN, J., AND J. GLASSBERG. 2000. Miami blues still
fly. American Butterflies 8: 28-29.
SAARINEN, E. V. 2005. Life history and myrmecophily of
Neomyrina nivea periculosa (Lycaenidae: Thecli-
nae). J. Lep. Soc. 59(2): 112-115.
SAVIGNANO, D. A. 1994. Benefits to Karner Blue butter-
fly larvae from association with ants, pp. 37-46 In
D.A. Andrew, R. J. Baker, and C. P. Lane [eds.],
Karner Blue Butterfly: A Symbol of a Vanishing
Landscape. Minn. Agric. Exp. Sta., St. Paul.
2003. Host specificity revisited: new data on Myr-
mica host ants of the lycaenid butterfly Maculinea
rebeli. J. of Insect Cons. 7: 1-6.
THOMAS, J. A. 1980. Why did the large blue become ex-
tinct in Britain? Oryx 15: 243-247.
WAGNER, D. 1993. Species-specific effects of tending
ants on the development of lycaenid butterfly larvae.
Oecologia 96:276-281.
WEBSTER, R. P., AND M. C. NIELSON. 1984. Myrmeco-
phily in the Edward's hairstreak butterfly Satyrium
edwardsii (Lycaenidae). J. Lepid. Soc. 38: 124-133.
WOJCIK, D. P. 1994. Impact of the red imported fire ant
on native ant species in Florida, pp. 269-281 In D. F.
Williams [ed.], Exotic Ants: Biology, Impact, and
Control of Introduced Species. Westview Studies in
Insect Biology, Boulder, CO.

Florida Entomologist 89(1)

Sanborn: Central American Cicadas


Barry University, School of Natural and Health Sciences
11300 NE Second Avenue, Miami Shores, FL 33161-6695, USA


Analysis of museum specimens has added to the cicada fauna of Belize, El Salvador, Guate-
mala, and Honduras. Information on the cicada fauna reported in the literature as well as the
first records of cicada species to the fauna are reported here to provide a more accurate un-
derstanding of cicada diversity in each country and the region. The new records represent an
increase of 75, 14, 110, and 320%, respectively, to the cicada faunal diversity of each country.

Key Words: cicadas, biodiversity, Central America


Un studio de los especimenes de museos han incrementado la fauna de las chicharras (He-
miptera: Cicadidae) de Belize, El Salvador, Guatemala y Honduras. Informaci6n sobre la
fauna de las chicharras reportadas en la literature, los nuevos registros de las species men-
cionadas en este articulo estan reportados para proveer un entendimiento mas precise de la
diversidad de las chicharras en cada pais de la region. Estos nuevos registros representan un
aumento de 75, 14, 110, y 320%, respectivamente, en la diversidad de la fauna de chicharras
en estos paises.

The Central American cicada fauna has re-
ceived little study since Distant's Biologia Cen-
trali-Americana (Distant 1881, 1883, 1900, 1905).
Davis (1919, 1928, 1936, 1941, 1944) described
new cicada genera and species, primarily from
specimens he received from Mexico. Since that
time, most work on Central American cicadas has
focused on the ecology of Costa Rican (Young
1972, 1976, 1980, 1981) and Panamanian (Wolda
1984, 1993; Wolda & Ramos 1992) cicadas with
limited work being done on the Mexican fauna
(Moore 1962, 1996; Sueur 2000, 2002; Sanborn
2006). The lack of knowledge was illustrated in
the paper by Sanborn (2001), who identified the
first cicadas to be reported from El Salvador. The
taxonomic position of some of the Central Ameri-
can species has been altered (Boulard & Marti-
nelli 1996; Moulds 2003) and the process of de-
scribing new species (Sueur 2000; Sanborn et al.
2005) has begun but there are still many species
to be described (Sanborn unpublished).
I have come across multiple species in various
museum collections that have not been described
as being part of the cicada fauna in several Cen-
tral American countries as published in the Cica-
doidea bibliographies (Metcalf 1963a, b, c; Duffels
& van der Laan 1985) or more recent literature. I
have now identified specimens from several col-
lections and individuals that represent additions
to the cicada fauna of Belize, El Salvador, Guate-
mala, and Honduras. These new additions to the
cicada fauna of the region are identified along
with a listing of previously identified species from

the various countries to provide a current view of
the cicada fauna for the region.


Specimens for this study were found among the
undetermined material in the collections of the
Florida State Collection of Arthropods (FSCA), the
Smithsonian Institution, United States National
Museum (USNM), San Diego Natural History
Museum (SDMC), Bohart Museum of Entomology
at the University of California at Davis (UCDC),
Carnegie Museum of Natural History (CMNH),
University of Mississippi Insect Collection
(UMIC), William R. Enns Entomological Museum,
University of Missouri (UMRM), University of
Connecticut (UCMS), University of Georgia
(UGCA) and three individuals who donated their
specimens to the author. Original specimens are
housed in the collections above with vouchers of
most species and the specimens donated to the au-
thor in the author's collection. The number of spe-
cies previously attributed to each country was de-
termined from the cicada bibliographies (Metcalf
1963a, b, c; Duffels & van der Laan 1985) and the
more recent literature. Original references can be
located in these materials.


The regional cicada fauna for Belize, El Salva-
dor, Guatemala, and Honduras is summarized
here. Species identified as new to a country in-

Florida Entomologist 89(1)

elude available collection information. Biblio-
graphic information is provided for species that
have been described previously from a country.
There are currently four species that have
been collected in Belize, one of which is a recently
described new species (Sanborn et al. 2005).
Three species are added to the cicada fauna with
this report. The cicada fauna of El Salvador was
unknown until I reported on representatives of
seven species collected in the country (Sanborn
2001). One additional species was found in the
collection of the USNM. There are currently ten
species attributed to Guatemala. Eleven addi-
tional species are added to the fauna in this re-
port. There are currently five species reported to
inhabit Honduras. One of these is a recently de-
scribed species (Sanborn et al. 2005). Sixteen new
species records are added in this report.

Family Cicadidae

Subfamily Tibiceninae Atkinson, 1886

Tribe Zammarini Distant, 1905

Odopoea signoreti Stal, 1864. Specimens in the
UMIC collected at Honduras, Olancho, La Union,
Parque Nacional La Muralia, 15.07N 86.45W,
17-V-1996. The species is described from Mexico
(Metcalf 1963a).
Miranha imbellis (Walker, 1858). Specimens in
the FSCA collected at Honduras, Cortes, Parque
Nacional Cusuco, 1529'47'N 8812'43"W, 1600 m,
1-VII-2000. It has been reported previously from
Guatemala and Central America (Metcalf 1963a).
Zammara smaragdina Walker, 1850. Speci-
mens in the FSCA were collected in Guatemala,
Peten at the Tikal Ruins. Specimens in the SDMC
were collected at Honduras, Atlantida, El Pino,
Morraias Arriba, 12-VIII-1979 and specimens in
the UGCA were collected at Honduras, Olancho,
Dulce Mombre de Culmi, Montaia de Malacate,
26-VII-2001 and 11-VI-2003. The species is de-
scribed from Central America (Metcalf 1963a).
Zammara smaragdula Walker, 1850. Speci-
mens from Guatemala, Peten, Morajan, 4.8 km
East of Poptun, IV-V-1993 were given to the au-
thor by Br. Leon Cook. The species is described
from Central and South America (Metcalf 1963a).
Zammara tympanum (Fabricius, 1803). A
specimen collected in Belize, Cayo District, Maya
Mountain Lodge, 19-VII-1993 was given to the
author by Vince Golia. The FSCA has specimens
from Guatemala, Izabal, Puerto Barrios Cerro,
San Gil, 1,000 m, 13-IV-1992. The UGCA contains
specimens from Honduras, Cortez Merend6n,
1500 m, adjacent to Parque Nacional De Cusuco,
N15030'12", W88011'54", collected 19-V-2002 and
24-VII-2001. The species is described from South
and Central America (Metcalf 1963a; Duffels &
van der Laan 1985).

Tribe Tibicenini Distant, 1889

Diceroprocta belizensis (Distant, 1910). Speci-
mens in the UMRM are from Guatemala, Es-
cuintla, Nueva Concepcion, 30-VII-1985. Speci-
mens have been reported previously from Belize,
Honduras (Metcalf 1963a) and El Salvador (San-
born 2001).
Diceroprocta bicosta (Walker, 1850). Speci-
mens have been reported from Honduras (Metcalf
1963a) and El Salvador (Sanborn 2001).
Diceroprocta bulgara (Distant, 1906). A female
in the UCMS was collected in Guatemala, Sacate-
pequez, Cerro Alux, 2,000 m, X-2002. The species
is described from Mexico (Metcalf 1963a).
Diceroprocta pusilla Davis, 1942. The FSCA
has specimens collected at Guatemala, Guate-
mala, 11-V-1991; Honduras, Atlantida, RVS Cu-
ero y Salado, Salado Barra, 15046'N 89059'W, 2 m,
1-VIII-2000; Olancho, 1.1 km North of El Cerro,
750 m, 15008'48"N 85033'19"W, 19-IV-1999; and
Honduras, Cortes, 9.3 km NNW Cofradia, 800 m,
15029'14"N 88011'22"W, 16-VI-1999. The species
is described from Mexico (Metcalf 1963a).
Diceroprocta ruatana (Walker, 1850). Speci-
mens have been reported from Honduras (Metcalf
Cacama maura (Distant, 1881). The FSCA has
specimens from Honduras, Olancho, Culuco,
Aguan Valley, 29-III-1978. The species is de-
scribed from Mexico (Metcalf 1963a).

Tribe Fidicinini Distant, 1905

Proarna insignis Distant, 1881. The FSCA has
specimens from Guatemala, Izabal, La Graciosa,
15-IV-1995. The FSCA also has specimens col-
lected at Honduras, El Paraiso, 7 km North of
Oropoli, 30-IV-1993 and Atlantida, RVS Cuero y
Salado, Salado Barra, 15046'N 89059'W, 2-5 m, 22-
IV to 1-VIII-2000. There is a specimen from Hon-
duras, El Paraiso, Yuscaran, 1-VI-2003 in the
UGCA. The species has been reported in Central
America (Metcalf 1963a).
Proarna olivieri Metcalf, 1963. The UCDC con-
tains specimens collected at Guatemala, Retalhu-
leu, Retalhuleu, 18-23-VI-1986 and Retalhuleu,
Retalhuleu, El Asintal, 6-V-1989. The UCMS con-
tains a male from Guatemala, Chimaltenango,
Pochula Fca El Rosario, 15-IV-2003. The species
has been reported in Central America (Metcalf
Proarna sallaei Stal, 1864. The UGCA has a
specimen collected at Honduras, Atlantida, ~20
km SW La Ceiba, base of Pico Bonito, 16-VII-
2001. The species is described from Mexico (Met-
calf 1963a).
Pacarina champion (Distant, 1881). The
FSCA has specimens from the Belize, Toledo Dis-
trict, Punta Gorda, 5-8-VI-1990. Specimens in the
CMNH were collected at Honduras, Rio Grande.

March 2006

Sanborn: Central American Cicadas

The species has been reported from Guatemala
and Central America (Metcalf 1963a).
Pacarina puella Davis, 1923. The species has
been reported from Guatemala and Central
America (Metcalf 1963a).
Pacarina schumanni Distant, 1905. Specimens
collected in Belize, Cayo District, Maya Mountain
Lodge, 19 and 20-VII-1993 were given to the author
by Vince Golia. Additional specimens collected by
Charles Bartlett were collected in the Belize, Cayo
District, Teakettle Bank, Pooks Hill, 1709.257'N
8851.094'W, 294 ft., 8-VII-2003 and given to the
author. Specimens in the SDMC were collected at
Belize, Chaa Creek, 18-21 August 1987. Specimens
in the FSCA were collected at Honduras, La Paz,
San Martin, 1-V-1988 and at Honduras, Colon,
Trujillo, 22-VII-1968. Specimens have been re-
ported from El Salvador (Sanborn 2001).

Sub-tribe Fidicinina Boulard & Martinelli, 1996

Fidicina cachla Distant, 1899. Specimens from
Honduras, El Paraiso, 8.3 km SE Capire, 675 m,
13058'54"N 85049'25"W, 16-IV-1999 are in the
FSCA. The species is described from Costa Rica
(Metcalf 1963a).
Fidicinoides determinata (Walker, 1858). The
FSCA has specimens from Guatemala, Guate-
mala, 1-V-1994. There are also specimens in the
FSCA from Honduras, Yoro, Parque Nacional Pico
Bonito, El Portillo, 640 m, 15026'27"N 87008'09"W,
11-III-2000; and Honduras, Atlantida, Parque Na-
cional Pico Bonito, El Manchon, 350 m, 15029'18"N
87007'39"W, 18-III-2001. The species has been re-
ported from El Salvador (Sanborn 2001).
Fidicinoides pronoe (Walker, 1850). The FSCA
has specimens collected in Honduras, Olancho, 14
km east of La Colonia, 610 m, 28-IV-1993; and
Honduras, Lempira, Montana de Puca, 14042'00"N
88034'07"W, 1150 m, 28-VI-2000. Specimens have
been reported previously from El Salvador (San-
born 2001) and Guatemala (Metcalf 1963a).

Sub-tribe Guyalnina Boulard & Martinelli, 1996

Dorisiana amoena (Distant, 1899). Specimens
from Guatemala, Peten, Morajan, 4.8 km East of
Poptun, IV-V-1993 were given to the author by Br.
Leon Cook. The species is described from Costa
Rica (Metcalf 1963a).
Majeorona truncata Goding, 1925. A specimen
from Honduras, Atlantida, PN Pico Bonito, Esta-
cion CURLA, 17-VII-2001 is in the UGCA. The
species is described from Ecuador (Metcalf 1963a).

Tribe Hyantinii Distant, 1905

Quesada gigas (Olivier, 1790). It has been re-
ported from Central America, Belize, Guatemala,
Honduras (Metcalf 1963a) and El Salvador (San-
born 2001).

Tribe Cicadini Oshanin, 1907

Neocicada centramericana Sanborn, 2005. The
species is reported from Belize, Guatemala and
Honduras (Sanborn et al. 2005).
Cicada pennata (Distant, 1881). The species is
described from Guatemala (Metcalf 1963b). The
taxonomic position of the species remains unclear
based on the description of a single female speci-

Subfamily Tibicininae Distant, 1906
Tribe Dazini Distant, 1905

Daza montezuma (Walker, 1850). Specimens in
the FSCA from Guatemala, Peten, Cam. Yaxha-
Nakum, 180-300 m, 30-VI-1992. The species is de-
scribed from Mexico (Metcalf 1963a).

Tribe Carinetini Distant, 1905

Carineta trivittata Walker, 1858. The FCSA
has specimens from Honduras, Cortes, Cofradia,
Cusuco, 26-VIII-1994. The species has been re-
ported from Guatemala and Central America
(Metcalf 1963c).
Herrera ancilla (Stal, 1864). The FSCA has
specimens collected in Honduras, El Paraiso, 5.3
km N Cifuentes, N 1508'04" W 8535'36", 13-14-
VI-1999, Honduras, EAP, 30 km ESE of Tegus, 23-
V-1983 and EAP 35 km Este Teg., 20-VII-1983.
The species has been reported from Central
America, Belize, Guatemala (Metcalf 1963c) and
El Salvador (Sanborn 2001).

Tribe Taphurini Distant, 1905

C', ". .' i. guatemalena (Distant, 1883). The
species is described from Guatemala (Metcalf
Dorachosa explicata Distant, 1892. A single
specimen collected at El Salvador, San Salvador,
San Salvador, 3-6-VI-1958 is in the USNM. The
species is described from Panama (Metcalf


This work has added significant numbers of
representatives to the cicada fauna of the north-
ern Central American countries (Fig. 1). However,
there are probably many additional cicada spe-
cies present in each country. The distribution of a
species may bypass an individual country while
being reported from border countries. It may be
that insufficient collecting has occurred to pro-
duce representatives of these species in some
countries, e.g., neighboring Guatemala, Hondu-
ras, and Nicaragua have been reported to have 24
species and 13 genera not reported from El Salva-

2, nwqdla bd
Lt I)Mrmw
A1 badgu

FWdklwr a dfleinal
Doris la wa0ns

lamn abear

FULsska ybMi
Pncaiodua schumanut
PEdclrna cachl

MUqm fNruata

Fig. 1. Summary listing of first species records for
the Central American countries of this study.

dor here and in Sanborn (2001). Continued mu-
seum study and field work will no doubt result in
the identification of new species and additions to
the cicada fauna of each country.


I thank J. Brambila and D. Ziesk (FSCA), R. Brown
(UMIC), P. Cato (SDMC), R. Davidson (CMNH), M. Ep-
stein (USNM), S. Heydon (UCDC), C. Smith (UGAI),
and R. Sites (UMRM) for assistance during my visits
and Br. Leon Cook, Vince Golia, and Charles Bartlett for


BOULARD, M., AND N. M. MARTINELLI. 1996. Revision
des Fidicini, nouveau statut de la tribu, especes con-
nues et nouvelles especes (Cicadomorpha, Cicadidae,
Cicadinae). Premiere parties: Sous-tribu nouvelle des
Fidicinina. EPHE, Biol. Evol. Insectes 9: 11-81.
DISTANT, W. L. 1881. Rhynchota: Homoptera. Biologia
Centrali-Americana; contributions to the knowledge
of the fauna and flora of Mexico and Central Amer-
ica. Part 15, 1: 1-16.
DISTANT, W. L. 1883. Rhynchota: Homoptera. Biologia
Centrali-Americana: contributions to the knowledge
of the fauna and flora of Mexico and Central Amer-
ica. Part 15, 1: 17-24.
DISTANT, W. L. 1900. Rhynchota: Homoptera. Biologia
Central Americana; contributions to the knowledge
of the fauna and flora of Mexico and Central Amer-
ica. Part 15, 1: 41-43.
DISTANT, W. L. 1905. Cicadidae and Fulgoridae. Biolo-
gia Centrali Americana; contributions to the knowl-
edge of the fauna and flora of Mexico and Central
America. Part 15, 1: 140-146.

March 2006

DAVIS, W. T. 1919. Cicadas of the genus Cacama, with
descriptions of several new species. J. New York En-
tomol. Soc. 27: 68-79.
DAVIS, W. T. 1928. Cicadas belonging to the genus
Diceroprocta with descriptions of new species. J.
New York Entomol. Soc. 36: 439-458.
DAVIS, W. T. 1936. A remarkable cicada from Mexico
and other North American species. J. New York En-
tomol. Soc. 45: 101-123.
DAVIS, W. T. 1941. New cicadas from North America
with notes. J. New York Entomol. Soc. 49: 85-99.
DAVIS, W. T. 1944. The remarkable distribution of an
American cicada; a new genus, and other cicada
notes. J. New York Entomol. Soc. 52: 213-222.
DUFFELS, J. P., AND P. A. VAN DER LAAN. 1985. Cata-
logue of the Cicadoidea (Homoptera, Auchenorhyn-
cha) 1956-1980. Dr. W. Junk Publishers, Series
Entomologica 34, Dordrect. 414 pp.
METCALF, Z. P. 1963a. General Catalogue of the Ho-
moptera, Fascicle VIII. Cicadoidea. Part 1. Cica-
didae. Section I. Tibiceninae. North Carolina State
Coll. Contr. 1502: 1-585.
METCALF, Z. P. 1963b. General Catalogue of the Ho-
moptera, Fascicle VIII. Cicadoidea. Part 1. Cica-
didae. Section II. Gaeninae and Cicadinae. North
Carolina State Coll. Contr. 1502: 587-919.
METCALF, Z. P. 1963c. General Catalogue of the Ho-
moptera, Fascicle VIII. Cicadoidea. Part 2.Tibicinidae.
North Carolina State Coll. Contr. 1564: 1-492.
MOORE, T. E. 1962. Acoustical behavior of the cicada Fi-
dicina pronoe (Walker) (Homoptera: Cicadidae).
Ohio J. Sci. 62: 113-119.
MOORE, T. E. 1996. Cicadoidea, pp. 221-223 In J. E.
Llorente Bousquets, A. N. Garcia Aldrete and E.
Gonzalez Soriano [eds.], Biodiversidad, Taxonomia y
Biogeographia de Artr6podos de Mexico: Hacia Una
Sintesis de Su Conocimiento. Cd. Universtiaria,
Mexico, Instituto de Biologia, UNAM.
MOULDS, M. S. 2003. An appraisal of the cicadas of the
genus Abricta Stal and allied genera (Hemiptera:
Auchenorrhyncha: Cicadidae). Rec. Australian Mus.
55: 245-304.
SANBORN, A. F. 2001. A first contribution to a knowl-
edge of the cicada fauna of El Salvador (Homoptera:
Cicadoidea). Florida Entomol. 84: 449-450.
SANBORN, A. F. New records of cicadas from Mexico
(Hemiptera: Cicadoidea: Cicadidae). Southwest.
Nat. 51(2): in press.
LIPS. 2005. Taxonomy and biogeography of the genus
Neocicada Kato, 1932 (Hemiptera: Cicadomorpha:
Cicadidae), with descriptions of three new species.
Syst. Entomol. 30: 191-207.
SUEUR, J. 2000. Une nouvelle espece de cigale du Mex-
ique (Los Tuxtlas, Veracruz), et etude de son emission
sonore (Homoptera, Auchenorhyncha, Cicadoidea).
Bull. Soc. Entomol. France 105: 217-222.
SUEUR, J. 2002. Cicada acoustic communication: poten-
tial sound partitioning in a multispecies community
from Mexico (Hemiptera: Cicadomorpha: Cicadidae).
Biol. J. Linn. Soc. 75: 379-394.
WOLDA, H. 1984. Diversity and seasonality of Panama-
nian cicadas. Mitt. Schweizerischen Ent. Ges. 57:
WOLDA, H. 1993. Diel and seasonal patterns of mating
calls in some neotropical cicadas. Acoustic interfer-
ence? Proc. Konin. Nederlandse Akad. Wetens. Biol.,
Chem., Geol., Phys. Med. Sci. 96: 369-381.

Florida Entomologist 89(1)

Sanborn: Central American Cicadas

WOLDA, H., AND J. A. RAMOS. 1992. Cicadas in Panama,
their distribution, seasonality and diversity, pp. 271-
279 In D. Quintero and A. Aiello [eds.], The Insects of
Panama and Mesoamerica. Selected Studies. Oxford
University Press, New York.
YOUNG, A. M. 1972. Cicada ecology in a Costa Rican
tropical rain forest. Biotropica 4: 152-159.
YOUNG, A. M. 1976. Notes on the faunistic complexity of
cicadas (Homoptera: Cicadidae) in Northern Costa
Rica. Rev. Biol. Trop. 24: 267-279.

YOUNG, A. M. 1980. Habitat and seasonal relationships
of some cicadas (Homoptera: Cicadidae) in central
Costa Rica. American Midl. Nat. 103: 155-166.
YOUNG, A. M. 1981. Notes on seasonality and habitat
associations of tropical cicadas (Homoptera: Cica-
didae) in premontane and montane tropical moist
forest in Costa Rica. J. New York Entomol. Soc. 89:

Florida Entomologist 89(1)


Department of Entomology, Michigan State University, East Lansing, MI 48824

The eastern cherry fruit fly, Rhagoletis cingu-
lata (Loew), is an important late-season pest of
cherries in the eastern and Midwestern United
States (Bush 1966). Adults emerge from overwin-
tering puparia in mid-June, mate on the host
fruit, and lay eggs into cherries (Pettit & Tolles
1930; Boller & Prokopy 1976; Smith 1984).
Michigan produces approximately 75% of the
total U.S. tart cherries, Prunus cerasus L., and
12% of the U.S. sweet cherries, P. avium (L.) L.
(Anon. 2004). Zero tolerance standards for fly lar-
vae in fruit require sensitive fly monitoring sys-
tems early in the growing season as part of inte-
grated pest management (IPM) programs to pre-
vent fruit infestation. To monitor R. cingulata
flies, Pherocon AM boards are placed approxi-
mately 2.1 m (depending on the tree size) from
the ground within the tree canopy. The yellow
traps are typically baited with an ammonia and
protein hydrolysate lure (Liburd et al. 2001), pro-
viding both visual and olfactory cues to attract
flies. The first insecticide is applied after a single
fly is captured on a monitoring trap.
Steady progress has been made in developing
effective trapping systems for some important
Rhagoletis pests of temperate fruit crops. Trap
placement has been optimized for monitoring the
apple maggot fly, R. pomonella (Walsh), (Reissig
1975; Drummond et al. 1984) and the blueberry
maggot, R. mendax Curran (Liburd et al. 2000;
Teixeira & Polavarapu 2001). The optimal posi-
tion for traps to monitor R. pomonella is approxi-
mately 2.1 m above the ground (depending on the
tree size) within the apple tree canopy (Reissig
1975; Drummond et al. 1984) and 0.25-0.5 m from
fruit, while traps for R. mendax are most effective
when placed within the top of highbush blueberry
plants, Vaccinium corymbosum L., when the
bushes are 1.5 to 2.0 m high (Liburd et al. 2000;
Teixeira & Polavarapu 2001). To date, the optimal
positioning of traps for monitoring R. cingulata
has not been reported; however, based on studies
of related fruit fly species, we hypothesized that
the height of trap placement within the cherry
tree canopy would affect captures ofR. cingulata
on monitoring traps.
In 2002 and 2003, trap heights were compared
in mature, unmanaged tart cherry orchards lo-
cated in southwestern Michigan (Van Buren Co.)
with high populations of R. cingulata. The or-
chards used in these experiments contained un-
sprayed, mature trees approximately 4.6 m in
height, and planted in a 2.4-m within-row by 6.1-
m between-row spacing. This spacing allowed

sunlight to reach the entire tree, rather than the
topmost portion only. In addition, branches from
adjacent trees did not overlap.
The effect of trap height on captures of R. cin-
gulata was determined by placing unbaited
Pherocon AM traps (Trece, Inc., Adair, OK) at
three positions midway between the tree trunk
and outermost foliage of cherry trees. All traps
were placed on the southwest side of trees. Three
treatments were evaluated that included placing
traps as follows: (1) below the tree canopy (ap-
proximately 1.2 m above ground), (2) at the stan-
dard trap height (approximately 2.1 m), or (3) in
the top portion of the tree canopy (approximately
4.6 m). In order to obtain the highest trap posi-
tion, traps hung at 4.6 m were suspended from a
PVC pipe (1.4 m length and 6 mm diam.) affixed
to a tree limb such that the traps were within the
top portion of the cherry tree foliage midway be-
tween the trunk and the outermost foliage. Trees
were selected randomly and a single trap was
placed at one of the three positions in each tree.
Treatments were arranged with a distance of at
least 20 m between trees and 30 m between
blocks. Foliage surrounding all traps was re-
moved in a 0.5 m radius (Reissig 1975). Five rep-
licates of each treatment were arranged in a ran-
domized complete block design. Flies were
counted and removed from traps weekly for six
weeks in both years (17 June-30 July 2002 and 16
June-29 July 2003). To minimize position effects,
all treatments were rotated one position clock-
wise after each weekly inspection.
Total fly captures on each trap across the sea-
son in both years were subjected to analysis of
variance (ANOVA). To normalize the data, square
root-transformation (x + 0.5)1" was performed
prior to analysis. Fisher's Least Significant Dif-
ference test (LSD, SAS Institute 2000) was used
to separate mean differences among treatments
(significance level a = 0.05).
In 2002, captures ofR. cingulata were signifi-
cantly affected by trap location (F = 79.2; df= 2,8;
P < 0.05), with more R. cingulata flies caught at
4.6 m within canopies of cherry trees than on
traps placed at 2.1 m (standard trap height) or 1.2
m (Fig. 1A). Overall, more than three times the
number of flies were captured on traps placed at
4.6 m compared with traps hung at lower posi-
tions. In 2003, captures ofR. cingulata flies were
significantly affected by trap location (F = 27.0; df
= 2,8; P < 0.05). Significantly more flies were
caught on traps hung at 4.6 m compared with
traps placed at a 2.1 m or 1.2 m height (Fig. 1B).

March 2006

Scientific Notes






Trap Height (m)

Fig. 1. Number of adult R. cingulata captured (
SEM) per season on Pherocon AM boards placed at low
(1.2 m), standard (2.1 m), and high (4.6 m) positions
within cherry trees. The experiment was conducted in
2002 (A) and 2003 (B). Means with the same letter
within years are not significantly different (Fisher's
LSD Test, P < 0.05). Untransformed means are shown.

Traps in the highest canopy position caught more
than three times as many flies as those hung in
either of the two lower canopy positions. During
both years, the mean number of flies captured in
the first week on traps at the high canopy position
was greater than the 2.1- and 1.2-m canopy posi-
tions (respectively, in 2002: 49.2 3.4, 9.8 + 1.3,
1.0 + 0.28; and 2003: 10.2 + 2.4, 2.4 0.9, 0.0).
Over two growing seasons, more R. cingulata
were captured on traps placed at the highest po-
sition than on those hung at 2.1 m or 1.2 m. Sim-
ilar results were obtained with R. mendax cap-
tures within blueberry bushes, where traps
placed in the upper third of bushes captured the
greatest number of flies (Liburd et al. 2000). In
contrast, more R. pomonella were captured on
Pherocon AM traps placed at 2.1 m within the
canopy of apple trees than on traps at 1.2 or 3.0 m
(Reissig 1975; Pelz et al. unpublished). Observa-
tions of adult R. indifferens revealed that the ma-
jority of flies were present within three meters of
the ground (Frick et al. 1954). Differences in the
distribution of fruit fly species within their host
tree have also been reported for several tropical

fruit flies in the genusAnastrepha (Sivinski et al.
2004). Within their respective host trees, A. alue-
ata Stone are more abundant in the lower canopy
of trees, while A. striata Schiner are more abun-
dant in the upper canopy.
Our results suggest that R. cingulata activity
is greatest in the uppermost portion of the host
tree canopy; thus, traps placed higher in the tree
may be more visible to R. cingulata flies that are
active in the upper canopy compared with those
placed lower in the tree canopy. Direct observa-
tions of R. cingulata within cherry trees are nec-
essary to determine whether the distribution of
flies is greatest in the upper canopy and whether
that distribution changes throughout the day. In
addition, assessment of infestation rates at differ-
ent levels may also be valuable in determining
whether peak fly oviposition activity coincides
with fly population distribution, as found for R.
indifferens (Frick et al. 1954) and several Anas-
trepha spp. (Sivinski et al. 1999, 2004). In their
study, Frick et al. (1954) found that infestation of
sweet cherries by R. indifferens was greater in the
lower canopy (less than two meters) compared
with infestation high in the tree canopy (between
two and nine meters).
Finally, the sensitivity and accuracy of monitor-
ing for R. cingulata may be improved through trap
placement in a location where flies are most abun-
dant; however, because standard trapping methods
utilize ammonium acetate lures, further work
must be done to determine whether these lures will
affect fly captures on traps at different heights.


In 2002 and 2003, we compared the effect of
three trap heights on captures of R. cingulata in
Michigan cherry orchards. Overall, significantly
more flies were captured on unbaited Pherocon
AM traps hung at 4.6 m in the canopy of cherry
trees than on traps hung at 2.1 m or at a low po-
sition of 1.2 m, suggesting that R. cingulata is
more abundant in the upper portion of the host
tree canopy.


ANONYMOUS. 2004. http://www.michigan.gov/mda.
BOLLER, E. F., AND R. J. PROKOPY. 1976. Bionomics and
management of Rhagoletis. Annu. Rev. Entomol. 21:
BUSH, G. L. 1966. The taxonomy, cytology, and evolution
of the genus Rhagoletis in North America (Diptera:
Tephritidae). Bull. Mus. Comp. Zool. 134: 431-562.
Comparative efficacy and optimal positioning of
traps for monitoring apple maggot flies (Diptera: Te-
phritidae) Environ. Entomol. 13: 232-235.
Bionomics of the cherry fruit fly in eastern Washing-
ton. Washington Agric. Exp. Stn. Tech. Bull. 13.


82 Florida Entc

CASAGRANDE. 2000. Effect of trap size, placement, and
age on captures of blueberry maggot flies (Diptera: Te-
phritidae). J. Econ. Entomol. 93: 1452-1458.
THORNTON. 2001. Performance of various trap types
for monitoring populations of cherry fruit fly
(Diptera: Tephritidae) species. Environ. Entomol.
30: 82-88.
PETTIT, R. H., AND G. S. TOLLES. 1930. The cherry fruit
flies. Bull. Michigan State College Agric. Exp. Sta.
131: 1-11.
REISSIG, W. H. 1975. Performance of apple maggot traps
in various apple tree canopy positions. J. Econ. Ento-
mol. 68: 534-538.
SAS INSTITUTE. 2000. SAS/STAT User's Guide, version
6, 4th ed., vol. 1. SAS Institute, Cary, NC.


ologist 89(1) March 2006

SIVINSKI, J., M. ALUJA, AND T. HOLLER 1999. The distri-
bution of the Caribbean fruit fly, Anastrepha sus-
pense (Tephritidae) and its parasitoids
(Hymenoptera: Braconidae) within the canopies of
host trees. Florida Entomol. 82: 72-81.
Novel analysis of spatial and temporal patterns of
resource use in a group of tephritid flies of the genus
Anastrepha. Ann. Entomol. Soc. Am. 97: 504-512.
SMITH, D. C. 1984. Feeding, mating, and oviposition by
Rhagoletis cingulata (Diptera: Tephritidae) flies in
nature. Ann. Entomol. Soc. Am. 77: 702-704.
TEIXEIRA, L. A. F., AND S. POLAVARAPU. 2001. Effect of
sex, reproductive maturity stage and trap place-
ment, on attraction of the blueberry maggot fly
(Diptera: Tephritidae) to sphere and Pherocon AM
traps. Florida Entomol. 84: 363-369.

Scientific Notes


'Departamento de Entomologia Tropical, El Colegio de la Frontera Sur (ECOSUR)
Apdo. Postal 36, 30700, Tapachula, Chiapas, M6xico

2Becario COFAA. Departamento de Interacciones Planta-Insecto. Centro de Desarrollo de Productos Bi6ticos
del I. P. N. (CEPROBI). Carretera Yautepec, Jojutla, Km. 8.5, Apdo. Postal 24. San Isidro, Yautepec, Morelos, M6xico

The sapodilla bud borer, Zamagiria dixolo-
phella Dyar, has been reported attacking the sa-
podilla Manilkara zapota van Royen in Mexico
(Iruegas et al. 2002). The larvae feed on the ten-
der young shoots and fruits. Current control of
this species is based upon the use of insecticides;
however, chemical control of this pest is difficult
due to its cryptic nature. Mating disruption may
be an alternative for controlling it. Although in
Z. dixolophella the pheromone has not been iden-
tified yet, it would be worthwhile to understand
the influence of different factors in the release of
pheromone to obtain a complete picture of the fac-
tors governing the biology of the female sex pher-
omone system. Production and release of the sex
pheromone in many moths is influenced by sev-
eral biotic and abiotic factors (Landolt & Phillips
1997; Rafaeli 2002). In this study, we investigated
the possible effect of host plant and the photo-
period on the calling behavior of Z. dixolophella
under laboratory conditions as a first step to iden-
tify the sex pheromone.
Larvae of Z. dixolophella were collected in
M. zapota orchards "El Nayar" (1449'36"N and
9220'52"W at 44 masl) and "Cazanares"
(1444'40"N and 9224'20"W at 20 masl), both lo-
cated between Tapachula City and Puerto Mad-
ero, Chiapas, Mexico. Larvae were held in 3-L
clear plastic cylindrical containers (23 cm height
x 14 cm diameter), and allowed to feed upon their
host plant (tender young shoots) in controlled
conditions at 25 5C and 65 5% R H with a re-
versed photoperiod of 16: 8 h (L: D) (unless other-
wise specified). Pupae obtained were placed in
Petri dishes inside plastic cages (30 x 30 cm) and
observed constantly one or two days before emer-
gence. Most females emerged during the photo-
phase, and only these were used in the observa-
tions. The experiments started during the first
complete scotophase after emergence. Females
were observed every 10 min throughout their first
six scotophases with a red light lamp. The per-
centage of females calling daily, the daily onset of
calling time (time after lights off), and duration of
calling of each female were recorded.
The possible influence of host plant in the call-
ing behavior was investigated in two groups of
newly emerged virgin females. In the first group,
20 females were individually placed in cylindrical
containers (23 cm height x 14 cm diameter). A

fresh, tender young host plant shoot with leaves
and flowers inserted in a plastic vial with cotton
soaked in water was placed in each container. The
host plant was changed daily after each scoto-
phase. In the second group, 20 females were
placed as described above but without the pres-
ence of host plant. The opening of the containers
was covered with gauze to permit circulation of
air. A drop of natural honey was placed daily on
gauze to ensure that females had food ad libitum.
The observations were made at 25 + 5C, 65 5%
relative humidity and at 16L: 8 D photoperiod
The effect of photoperiod on the calling behav-
ior was examined under two different photoperiod
regimes: 16L: 8D and 13L: 11: D. In both cases,
larvae were collected in the field and once they
have reached the pupal stage, pupae were sexed,
and the female pupae were preconditioned under
the experimental photoperiod at which they were
to be observed. Upon emergence females were iso-
lated, placed in individual containers with host
plants at 25 5C and 65 5% relative humidity.
Twenty females were tested under each photope-
riodic regime.
The percentages of calling females were ana-
lyzed by x2 test. The data for the daily onset of
calling time and duration of calling were analyzed
by one-way repeated measures analysis of vari-
ance (ANOVA), with age as repeated measure.
Means were separated by least significant differ-
ence (LSD) at a significance level of 0.05.
Most of the females called from their first sco-
tophase independently of the presence or absence
of host plant. The mean daily onset of calling time
was not affected by the presence or absence of the
host plant, but it differed significantly with age.
The interaction between the presence of host
plant x age was not significant. Also, the presence
of host plant did not affect the length of the call-
ing period, but this parameter was influenced by
female age. The interaction between the presence
of host plant x age was not significant. In contrast
to our results, several studies have shown that
the presence of the host plant or its volatile chem-
icals stimulate the production and releasing of
the sex pheromone in several moth species (Hen-
drikse & Vos-Biinnemeyer 1987; Raina 1988;
Raina et al. 1992, 1997; Pittendrigh & Pivnick
1993). Virgin females ofHelicoverpa zea (formerly

Florida Entomologist 89(1)

Heliothis) (Boddie) (Raina et al. 1992) and Helio-
this phloxiphaga G. and R. (Raina 1988) synthe-
sized and released pheromone only in presence of
their host plants. However H. zea females reared
in laboratory for many generations did not re-
quire the host plant for the production and re-
lease the pheromone (Raina 1988). In presence of
its host plant, females of Plutella xylostella (L.)
began calling at a younger age and they spent
more time calling (Pittendrigh & Pivnick 1993).
The percentage of calling females was similar
in the two photoperiods evaluated. The mean
daily onset time of calling was significantly differ-
ent under the photoperiods tested, but this pa-
rameter was not affected by female age. The in-
teraction between age x photoperiod was signifi-
cant. In overall, females maintained at 16L: 8D
began to call earlier than females held at 13L:
11D, except in the fifth scotophase (Fig. la). The
length of the calling period differed significantly
between the photoperiods evaluated and this pa-
rameter was influenced by female age. Also, the
interaction between age x photoperiod was signif-
icant. Females held at 16L: 8D called longer than
females maintained at 13L: 11D (Fig. Ib). Our re-
sults are in agreement with the suggestion of
Haynes and Birch (1984), who proposed that pho-
toperiod would have a major influence on the call-
ing behavior of multivoltine species such as
Z. dixolophella because these species are exposed
to different photoperiod conditions at different
times of the year.

In conclusion, this study shows that the calling
behavior of Z. dixolophella is influenced by the
photoperiod, but not by the presence of host plant.
This information will be useful during the collec-
tion and identification of sex pheromone.
We thank Martha Foursali and Sergio
Gonzalez for allowing us to collect insects and
plants from their farms "El Nayar" and "Caza-
nares", respectively. We also thank Javier Valle
Mora for statistical advice, and Federico Castre-
j6n and Armando Virgen for help during the col-
lection of biological material. Economic support
for this study was provided by CONACYT (Grant
91489) and Instituto Politecnico Nacional (Grant
COTEPABE 295) through a scholarship to VRCG.


The influence of host plant and photoperiod on
calling behavior of the moth Zamagiria dixol-
ophella, a sapodilla pest in Mexico was investi-
gated under laboratory conditions. Most of the fe-
males called from their first scotophase indepen-
dently of the presence or absence of host plant.
Also, the host plant did not influence the mean
onset time of calling and the mean time spent
calling. There was an effect of photoperiod on the
mean onset time of calling and the mean time
spent calling ofZ. dixolophella.


HAYNES, K. F., AND M. C. BIRCH. 1984. The periodicity
of pheromone release and male responsiveness in
1 16i80-- -t-13L:11 the artichoke plume moth, Platyptilia carduidac-
I tyla. Physiol. Entomol. 9: 287-295.
S a a Ahost-plant stimuli in sexual behaviour of small ermine
4 -- moths (Yponomeuta). Ecol. Entomol. 12: 363-371.
Sb AND J. C. ROJAS. 2002. A new record of moth attack-
la ".b ..-ing sapodilla, with descriptions of female genitalia
b and the last instar larva. Fla. Entomol. 85: 303-308.
100 -LANDOLT, P. J., AND T. W. PHILLIPS. 1997. Host plant in-
fluences on sex pheromone behavior of phytopha-
1 2 3 4 5 0 gous insects. Annu. Rev. Entomol. 42: 371-391.
PITTENDRIGH, B. R., AND K. A. PIVNICK. 1993. Effects of
SB) the host plant Brassica juncea (L.) on calling behav-
SW iour and egg maturation in Plutella xylostella. Ento-
Sa mol. Exp. Appl. 68: 117-126.
SW00- a_ _t a RAFAELI, A. 2002. Neuroendocrine control of pheromone
Sa biosynthesis in moths. Intern. Rev. of Cytol. 213: 49-
S200W b 92.
S- ---I ..... .. -. b RAINA, A. K. 1988. Selected factors influencing neuro-
S--- hormonal regulation of sex pheromone production in
0a Heliothis species. J. Chem. Ecol. 14: 2063-2069.
1 2 3 4 6 6 RAINA, A. K., T. G. KINGAN, AND A. K. MATTO. 1992.
Scotophases Chemical signals from host plant and sexual behav-
ior in a moth. Science 255: 592-594.
Fig. 1. Calling behavior response of Z. dixolophella RAINA, A. K., D. M. JACKSON, AND R. F. SEVERSON.
at two different photoperiods under laboratory condi- 1997. Increased pheromone production in wild to-
tions (values are means SE). (A) Mean ( SE) onset bacco budworm (Lepidoptera: Noctuidae) exposed to
time of calling. (B) Mean ( SE) time spent calling. Dif- host plants and host chemicals. Environ. Entomol.
ferent letters indicate significance at P < 0.05. 26: 101-105.

March 2006

Scientific Notes


'Oregon State University, Crop and Soil Sciences, Hermiston Agricultural Research and Extension Center
P.O. Box 105, Hermiston, OR 97838

2University of Florida, Institute of Food and Agriculture, Gulf Coast Research and Education Center
14765 CR 672, Wimauma, FL 33598

3University of Florida, Institute of Food and Agriculture, Horticultural Sciences Department
P.O. Box 110690, Gainesville, FL 32611

Coccinellids (lady beetles, lady bugs or ladybird
beetles) have been used in biological control pro-
grams because of their ability to prey on economi-
cally important pests such as aphids (Hagen & Van
den Bosch 1968; Hagen 1974; Frazer 1988; Rondon
et al. 2004), whiteflies (Hoelmer et al. 1994) and
mites (Chazeau 1985; Rondon et al. 2004) on maize,
Zea mays L. (Kieckhefer & Elliot 1990), alfalfa,
Medicago sativa L. (Giles et al. 1994), and potato,
Solanum tuberosum L. (Groden et al. 1990; Hilbeck
& Kennedy 1996). Lady beetles are probably the
most visible and well known beneficial predatory in-
sects with over 450 species found in North America
(Gordon 1985). Coleomegilla maculata DeGeer (Co-
leoptera: Coccinellidae), the pink spotted lady bee-
tle, is a new world species distributed from southern
Canada, U.S. (east of the Rocky Mountains), and
Central and South America (Timberlake 1943;
Wright & Laing 1982; Gordon 1985; Munyaneza &
Obrycki 1998). Three subspecies of C. maculata
have been described based on morphological charac-
ters such as spot patterns, color, body size, genitalia,
and geographical distribution (Gordon 1985). Ac-
cording to Gordon (1985) these subspecies are C. m.
fuscilabris (Mulsant), C. m. lengi Timberlake and C.
m. strenua (Casey). Coleomegilla m. fiscilabris is
found in the southeastern U.S., including Florida,
while C. m. lengi is found from Ontario, Canada,
through northwestern Georgia. Coleomegilla m.
lengi has not been reported in Florida (Peck & Tho-
mas 1998). The criteria to determine subspecies
based only on morphological and geographical dis-
tribution has been challenged several times and
even by Darwin (1964). The assertion that "biology
should overrule taxonomy and that the term sub-
species should be referred to as species rather than
subspecies" (anonymous) is open to discussion. Nev-
ertheless, geographic distribution has been defining
in allocation of species (Odum 1950); however, the
final determination of genotypic characteristics
should be considered as definitive in insect identifi-
cation. There has not been a determination of the
actual distribution of subspecies since Gordon
(1985). To our knowledge, no further surveys have
been made to update the records.

In Florida, there has been an increasing inter-
est from the biological control industry to intro-
duce C. m. lengi (non-native), which is thought to
have a greater reproductive capability, a highly
attractive biological characteristic desired by pro-
ducers of beneficial, than C. m. fuscilabris (na-
tive) (Griffin & Yeargan 2002). However, concerns
regarding the possibility of cross genetic contam-
ination between C. m. fuscilabris and C. m. lengi
prevented the introduction (Peres 2000). Studies
by Peres & Hoy (2002) indicated that there was a
reproductive near incompatibility between the
subspecies during the first and second genera-
tions (Fl, F2); conversely, Krafsur & Obrycki
(2000) indicated that high levels of gene flow
among subspecies might be possible. Due to this
contradiction, more basic information to create a
strong argument regarding the possibility to in-
troduce C. m. lengi into Florida was needed. Thus,
the objective of this research was to compare the
development, oviposition, and feeding behavior of
C. m. fusicilabris and C. m. lengi, on the cotton
aphid, Aphis gossypii Glover (Homoptera: Aphid-
idae) as prey in the feeding behavior study, and
strawberry, Fragaria x ananassa Duchesne, as a
substrate. Strawberry plants were maintained
following Paranjpe (2003) protocols. Experiments
were conducted at the biological control labora-
tory, Protected Agricultural Project Research Sta-
tion, University of Florida, Horticultural Sciences
Department in Gainesville, FL. Both subspecies
of C. maculata were provided by Entomos
(Gainesville, FL) (voucher specimens can be
found at DPI) where they were reared on undis-
closed artificial diet.
From an initial colony (40-50 females per cage)
from Entomos, 20 egg masses of C. m. fuscilabris
and 20 of C. m. lengi were randomly selected and
isolated in individually labeled 10-cm diameter
Petri dishes and maintained at 26 + 1C, 80 5%
R.H., and 16:8 h (L:D) photoperiod. Eggs were
checked for larval eclosion every 12 h. After eclo-
sion, 20 larvae were collected randomly and isolated
in 30 ml plastic cups. Larvae were fed every second
day with an undisclosed proprietary artificial diet

Florida Entomologist 89(1)

(1.8 g), to which were added bee pollen (0.05 g) and
shrimp eggs (0.01 g). Water was provided through
wet cotton balls. Larvae were transferred to clean
plastic cups twice a week. Daily observations were
made and the number of days from instar to instar
was recorded. Instars were distinguished by the
presence of cast exuvia. After adults emerged, one
female and one male were paired (n = 20) in 30-ml
plastic cups for 48 h to facilitate mating. Gender
was determined by examining the last abdominal
sclerite under a dissecting microscope. After 48 h,
females were isolated in plastic cups (15 x 15 x 10
cm) to determine viability of eggs (% eclosion), sur-
vival (larva to adult), number of egg masses, and
number of eggs per mass produced by each female.
A small piece of gray, thick fur served as an oviposi-
tion substrate. Longevity of adults also was mea-
sured. The experiment was repeated three times
with 20 replications per treatment. The data are
presented as average (SE) over the three experi-
ments (P < 0.05). The measure of the developmental
time was analyzed by t-test for independent sam-
ples. In general, there were no significant differ-
ences between the developmental periods (egg to
adult) of C. m. fuscilabris (23 4 days) and C. m.
lengi (22 3). There were no significant differences
between C. m. fuscilabris and C. m. lengi in develop-
ment periods of their eggs (3 3; 2 1, respectively),
1st (3 2; 3 1) 2nd (4 1; 3 1), 3rd (4 1; 5 1)
and 4th instar (3 1; 3 1) larval; pre-pupal (3 2;
3 1) and pupal stages (3 2; 3 1). Female adult
longevity is significantly greater in C. m. lengi (43 +
6 days) as compared with C. m. fuscilabris (38 7).
However, there was no significant difference be-
tween male adult longevity in the two subspecies
(C. m. fuscilabris, 31 + 5 days; C. m. lengi 36 4).
The percentage of eclosion of C. m. fuscilabris eggs
to larvae (95 3) was greater than ofC. m. lengi (86
+ 4); in contrast, the percentage of survivorship
(larva to adult) was significantly greater among
C. m. lengi (73 5) than among C. m. fuscilabris (65
+ 3). The number of egg masses produced by C. m.

lengi per female per day (3 1) was not significantly
different from those produced by C. m. fuscilabris (4
+ 1). However, there were significantly more eggs
oviposited in each C. m. fuscilabris mass (11 3)
than in each C. m. lengi egg mass (8 3). Also, there
was significantly more estimated number of eggs
produced by C. m. fuscilabris (1,672 eggs) than
C. m. lengi (1,032) over the female's lifetime.
An experiment was conducted to determine
the consumption ofA. gossypii by C. m. fuscilabris
and C. m. lengi. Each experimental unit consisted
of a 10-cm diameter Petri dish, where one straw-
berry leaflet, one individual of a single predator
subspecies, and ten prey were placed. All instars
and the adults of each subspecies were evaluated.
The predators tested were starved for 8 h prior to
providing them with A. gossypii. Ten individual
prey, without a predator, per Petri dish served as
a control for the experiment. Aphids were re-
moved from infested leaves in the colony by using
a wet, fine, camel hair brush. The strawberry leaf-
lets were isolated with lanolin to confine the prey
on the upper side of the leaf relative to the Petri
dish. Petri dishes were sealed with Parafilm and
labeled. Each experiment was maintained at 21
2C, 65 5 % R.H., and 16L: 8D photoperiod.
Samples were examined under a stereo micro-
scope and the number of prey consumed at 24 h
was recorded. Each experiment was repeated
three times with five replications per treatment
in a block design. The feeding data are presented
as average number (SE) of prey consumed by a
predator at 24 h. All data were analyzed with SAS
(SAS Institute 2000). The general linear model
(GLM) procedure was used to construct analysis
of variance (ANOVA). Averaged over all five feed-
ing stages, C. m. fuscilabris consumed more
aphids (8.4 1.1) than did C. m. lengi (6.5 1.5)
(LSD, 0.05 = 1.96;F = 1.19; df = 2, 20; P > 0.09) in
24 h (Table 1). Aphid consumption by C. m. fusci-
labris 1st instar was significantly greater as com-
pared with C. m. lengi (F = 1.84; df = 4, 20; P >


Number of Aphids Consumed

Life Stage C. m. fuscilabris C. m. lengi Significance

1st instar 7 1 4 1 n.s.
2nd instar 9 1 7 2 n.s.
3rd instar 9 1 8 1 *
4th instar 9 1 7 1 *
adults 9 1 7 1 *

Total 9 1 7 1

Mean (+ SE) within subspecies. Each treatment was repeated three times with five replications in each treatment, n.s.= no sig-
nificant different; = significant different (P < 0.05)

March 2006

Scientific Notes

0.06). Also, C. m. fuscilabris 4th instar consump-
tion of aphids was greater than that of the 4th in-
star of C. m. lengi (F = 3.84; df = 4, 20; P > 0.05)
and adult C. m. fuscilabris consumed more aphids
than did adult C. m. lengi (F = 3.84; df = 4, 20; P
> 0.05). C. m. fuscilabris 2nd and 3rd instar con-
sumption of aphids was not significant different
from that of 2nd and 3rd instar of C. m. lengi (F =
3.04; df = 4, 20; P > 0.126 and F = 2.02; df = 4, 20;
P > 0.15, respectively).
Although immature and adult C. m. lengi are
larger than those ofC. m. fuscilabris (Peres 2000),
this morphological advantage does not provide
any significant benefit to C. m. fuscilabris as com-
pared with C. m. lengi. For instance, considering
total egg production as a measure of a successful
candidate for mass rearing for commercial pur-
poses, our data indicated the advantage of C. m.
fuscilabris as a mass reared subject. The 38-day
life of C. m. fuscilabris and 43-day life of C. m.
lengi adults were lower as compared with the 3
months reported by Wright & Laing (1978). We
also observed a 3-day pre-ovipositional period in
contrast to the 5 to 15 days reported by Hodek
(1973). This situation may have occurred because
our insects came from a commercial colony fed on
an artificial diet for many generations. In nature,
C. maculata spends time selecting ovipositional
sites based on availability of food such as aphids
and eggs of various species (Nault & Kennedy
2000). Our observations indicated that C. m. fus-
cilabris seems to be more aggressive than C. m.
lengi (unpublished data). C. m. fuscilabris take
only few second before starting to manipulate and
consume (handling time) the prey (Rondon et al.
2004) as compared with C. m. lengi. Although no
striking advantages emerged for one subspecies
over the other, further studies are still needed.
Results form our laboratory experiments provide
the basis to further evaluate the possible intro-
duction of C. m. lengi.
Predators were provided by Entomos (Gaines-
ville, FL). Thanks to Drs. Norm Leppla and Mar-
garet L. Smither-Kopperl of the University of
Florida, and Anna Legrand from the University of
Connecticut, for their comments and editorial
contribution. This research was funded by USDA
Special Research Grant Program, and supported
by the Florida Agricultural Experimental Station
and approved for publication as Journal Series R-


After measuring the developmental time, re-
production, and feeding of both subspecies of C.
maculata, we conclude that there were no signifi-
cant differences between subspecies in develop-
mental periods but there were different levels of
female longevity, eclosion, survival, and number
of eggs per mass. In general, C. m. fuscilabris con-

sumed more A. gossypii than C. m. lengi in 24 h
and produced more eggs per female. Further
studies are needed to conclude if the introduction
of C. m. lengi into the ecosystem of Florida would
bring additional benefits to the present predator


CHAZEAU, J. 1985. Predaceous insects, pp. 211-244 In
Predaceous mites: Their biology, natural enemies
and control. W. Helle and M.V. Sabelis (eds.).
Elsevier, N.Y. 428 pp.
DARWIN, C. 1964. On the origin of the species: a facsim-
ile of the first edition. Boston, Harvard University
Press. Pp. 346-382.
FRAZER, B. D. 1988. Coccinellidae, pp. 231-247 In A. K.
Minks and P. Harrewinj [eds.], Aphids: Their Biol-
ogy, Natural Enemies and Control. Elsevier, N.Y.
326 pp.
Prevalence of predators associated with Acyrtosi-
phum pisum (Homoptera: Aphididae) and Hypera
postica Gyllenhal (Coleoptera: Curculionidae) dur-
ing growth of the first crop of alfalfa. Biol. Control 4:
GORDON, R. D. 1985. The Coccinellidae of America and
North of Mexico. J. New York Entomol. Soc. 93: 1-912.
GRIFFIN, M. L., AND K. V. YEARGAN. 2002. Oviposition
site selection by the pink spotted lady beetle Cole-
omegilla maculata (Coleoptera: Coccinellidae):
choices among plant species. Environ. Entomol. 31:
D. L. HAYNES. 1990. Coleomegilla maculata (Cole-
optera: Coccinellidae): Its predation upon the Colo-
rado potato beetle (Coleoptera: Chrysomelidae) and
its incidence in potatoes and surrounding crops.
J. Econ. Entomol. 83: 1306-1315.
HAGEN, K. S.1974. The significance of predaceous Coc-
cinellidae in biological and integrated control of in-
sects. Entomophaga. 7: 25-44.
HAGEN, K. S., AND R. VAN DEN BOSCH. 1968. Impact of
pathogens, parasites, and predators on aphids.
Annu. Rev. Entomol. 13: 325-384.
HILBECK, A., AND G. C. KENNEDY. 1996. Predators feed-
ing on the Colorado potato beetle in insecticide-free
plots and insecticide treated commercial potato
fields in eastern North Carolina. Biol. Control 6:
HODEK, I. 1973. Biology of Coccinellidae. W. Junk. The
Hague. Pp. 294.
Interaction of the whitefly predator Delphastus pu-
sillus (Coleoptera: Coccinellidae) with parasitized
sweet potato whitefly (Homoptera: Aleyrodidae). En-
viron. Entomol. 23: 136-139.
KIECKHEFER, R. W., AND N. C. ELLIOT. 1990. A 13-year
survey of the aphidiophagous Coccinellidae in maize
fields in eastern South Dakota. Canadian Entomol.
122: 579-581.
KRAFSUR, E. S., AND J. J. OBRYCKI. 2000. Coleomegilla
maculata (Coleoptera: Coccinellidae) is a species
complex. Ann. Entomol. Soc. Am. 93: 1156-1163.
MUNYANEZA, J., AND J. J. OBRYCKI. 1998. Development
of three populations of Coleomegilla maculata (Co-
leoptera: Coccinellidae) feeding on eggs of Colorado

potato beetle (Coleoptera: Chrysomelidae). Environ.
Entomol. 27: 117-122.
NAULT, B. A., AND G. G. KENNEDY. 2000. Seasonal
changes in habitat preference by Coleomegilla mac-
ulata: implications for colorado potato beetle man-
agement in potato. Biol. Control 17: 164-173.
ODUM, E. 1950. Bird populations of the highlands
(North Carolina) plateau in relation to plant succes-
sion and avian invasion. Ecology 31: 587-605.
PARANJPE, A. 2003. Soilless media, growing containers,
plant densities, and cultivars for greenhouse straw-
berry production in Florida. M.S. Thesis. University of
Florida, Horticultural Sciences Department. 260 pp.
PECK, S. B., AND M. C. THOMAS. 1998. Arthropods of
Florida and neighboring land areas, p. 180 In A dis-
tribution Checklist of The Beetles (Coleoptera) of
Florida. V. 16. Fla. Dept. Agric. and Consumer Ser-
PERES, O. G. 2000. Reproductive incompatibility be-
tween two subspecies of Coleomegilla maculata (De-
Geer) (Coleoptera: Chrysomelidae). M.S. Thesis.
University of Florida, Department of Entomology
and Nematology. 32 pp.

March 2006

PERES, O. G., AND M. A. HOY. 2002. Reproductive in-
compatibility between two subspecies of Coleome-
gilla maculata (Coleoptera: Coccinellidae). Florida
Entomol. 85: 203-207.
The feeding behavior of the bigeyed bug, minute pi-
rate bug, and pink spotted lady beetle relative to
main strawberry pests. Environ. Entomol. 33: 1014-
SAS INSTITUTE, INC. 2000. SAS/STAT User's guide for
personal computers, version V8. SAS Institute,
Cary, NC.
TIMBERLAKE, P. H. 1943. The Coccinellidae or lady bee-
tles of the Koebele collection. Part 1. Bull. Exp. Stn.
Hawaii. Sugar Planters Assoc. Entomol. Ser. 22: 1-67.
WRIGHT, E. J., AND J. E. LAING. 1982. Stage-specific of
Coleomegilla macualta lengi Timberlake on corn in
southern Ontario. Environ. Entomol. 11: 32-37.
WRIGHT, E. J., AND J. E. LAING. 1978. The effects of tem-
perature on development, adult longevity and fecun-
dity of Coleomegilla maculata lengi and its parasite,
Perilitus conccinellae. Proc. Entomol. Soc. Ont. 109:

Florida Entomologist 89(1)

Scientific Notes


1Mississippi State University, Coastal Research & Extension Center
1815 Popps Ferry Road Biloxi, MS 39532

2The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103

Crane flies (Diptera: Tipulidae) are distributed
worldwide, ranging from fresh and marine
aquatic habitats to drier terrestrial environments
(Alexander & Byers 1981). Tipulidae is the larg-
est family in the Diptera; however, larvae have
been described for less than 10% of the named
North American species (Thompson 1990). Lar-
vae of a few Tipula species have been implicated
in damage to crops and grasslands in North
America (Hartman & Hynes 1977; Alexander &
Byers 1981; Alexander 1920). Larvae of the range
crane fly, T (Triplictipula) simplex Doane, a na-
tive species, consume roots causing damage to un-
irrigated pastureland in the San Joaquin Valley,
California (Hartman & Hynes 1977). Larvae of
other native Tipula (Serratipula) species have
been implicated in pasture damage (Alexander
1967; Gelhaus 1986). Two exotic species, the com-
mon and the European crane fly (T oleracea L.
and T paludosa Meigen, respectively) are de-
structive pests of cool-season turfgrass in the Pa-
cific Northwest, western New York (D. Peck,
Cornell University, personal communication),
and maritime provinces of Canada (Jackson &
Campbell 1975; Vittum et al. 1999; LaGasa &
Antonelli 2000).
Larvae of march flies (Diptera: Bibionidae) are
herbivores and scavengers (Hardy 1981). Larvae
of several species have been reported to damage
agronomic crops, vegetables, and grasses (Hardy
1981; Darvas et al. 2000). In the southeastern
United States, swarms of adult Plecia nearctica
Hardy, or lovebugs, are abundant in the spring
and fall. Larvae of P nearctica, however, are not a
known pest of turfgrass.
On 13-1-2004, three larvae of the subgenus
Tipula (Triplicitipula) were collected from a
group of about 30 larvae crossing pavement adja-
cent to a bermudagrass home lawn in Saucier,
Harrison County, MS. No damage to the adjacent
grass was noted. At that same site on 31-1-2004,
adult T (Triplicitipula) umbrosa Loew were col-
lected at dusk and presumed to be conspecific
with the larvae collected earlier. Females were
typically collected while at rest on a vertical sur-
face such as a building. Males were collected most
often while copulating with females. This marks
the first record of this species in Mississippi; the

species is known previously from Louisiana and
Florida (Oosterbroek 2003).
On 27-II-2004, live larvae and pupae (Tipul-
idae and Bibionidae) were collected by Wayne
Wells from under centipede grass (Eremochloa
ophiuroides [Munro] Hack.) sod which was se-
verely weakened from extensive root herbivory
and poor nutrition. The infested turf, growing on
a sod farm in Picayune, Pearl River County, MS,
was breaking apart during harvesting, indicating
damage to the roots. That same day, specimens
were submitted to the senior author for identifica-
tion. Most of the live immatures were placed into
a mixture of moistened field soil and sand in the
laboratory for rearing. A few representative lar-
vae and pupae were preserved in alcohol.
On 4-III-2004, the previously mentioned sod
farm was surveyed by the senior author. This site
contained three separate fields of centipede grass
where the root damage was such that they were
deemed unharvestable by the sod producer. Be-
cause larvae of Tipulidae and Bibionidae are not
commonly associated with damage to warm sea-
son grasses, the site was first surveyed for damage
from more common pests such as mole crickets
(Orthoptera: Gryllotalpidae, Scapteriscus spp.),
white grubs (Coleoptera: Scarabaeidae), or billbug
larvae (Coleoptera: Curculionidae). In each field,
three 1-m2 areas of damaged turf were sampled
with a soap solution for disclosing mole crickets
and adult billbug (Vittum et al. 1999). Three soil
samples, consisting of 0.1-m2 plots, were exca-
vated on each of three infested fields and the soil,
grass, and thatch examined for larvae.
Disclosing samples yielded no mole crickets or
billbugs, and only immature flies were recovered
from excavated samples. Spiny brown larvae (Bi-
bionidae) formed aggregations (about 5-10 lar-
vae) in, or just below, the thatch. Larvae in some
aggregations appeared white, not light brown,
and appeared likely infected with an ento-
mopathogen. Larger larvae and pupae (Tipul-
idae) also were located in the thatch, but solitary.
None of these larger larvae appeared infected.
Both types of larvae and the Tipula pupae were
collected for rearing. About half of the immatures
that were collected that day were packaged into
loose, moist soil and shipped overnight to the lab-

Florida Entomologist 89(1)

oratory of JKG, The Academy of Natural Sciences,
Philadelphia for rearing. The remaining imma-
tures were confined with a core of grass in a
sealed plastic container and reared in the labora-
tory at the Coastal Research & Extension Center,
Biloxi. Several crane flies emerged in early March
from pupae at both laboratory locations and were
confirmed by JKG to be Tipula umbrosa.
All bibionid larvae at both locations died be-
fore eclosion. Cadavers were firm but within a few
days became enveloped in a white fungal growth,
presumably the same pathogen noted in the field.
A fungus from these cadavers was isolated by
Dr. Charlotte Nielsen (Cornell University) and
identified by Dr. Richard Humber (USDA, ARS,
Ithaca, NY), as Evlachovaea sp. Species identifi-
cation of these larvae was not possible without an
adult specimen. However, JKG could identify
them as members of the genus Plecia, likely lar-
vae of the lovebug, P nearctica, which is abun-
dant in Mississippi in the spring and fall.
This is the first record of T umbrosa in Missis-
sippi and the first habitat record for larvae of this
species. Closely related adults and larvae (Triplic-
itipula) in the eastern United States may also de-
velop in grassy areas, particularly at edges of
woodlands; the more distantly related species in
the western United States include those that are
the pest "range crane flies" (Gelhaus 1986). Al-
though not previously recorded as pests of turf-
grass, we document that larvae of both T umbrosa
and Plecia sp. can be associated with damage to
warm season grasses, especially those with already
weakened root systems or under nutrient stress.
We thank Billy Joe Lee, Paul Jeaufreau, and
Wayne Wells (Mississippi State University, De-
partment of Plant and Soil Sciences) for collecting
specimens and assistance with the site survey.
Identifications of all larvae and adults were made
by JKG. We thank Ann Hajek, Cornell University,
for arranging for the identification of the fungal
pathogen, and George Byers, University of Kan-
sas, for comparing specimens of T umbrosa with
those he identified from Louisiana. JKG also con-
sulted an unpublished manuscript by Steven
Teale reviewing the eastern Nearctic species of
Tipula (Triplicitipula) for help in identifying
T umbrosa. Voucher specimens of T umbrosa and
Plecia sp. are on deposit in the collection of The
Academy of Natural Sciences and in the Missis-
sippi Entomology Museum, Starkville. Richard
Brown, Mark Woodrey (Mississippi State Univer-
sity), and David Boyd (USDA-ARS) provided
helpful comments on an earlier draft of this
manuscript. This paper is No. J-10723 of the Mis-
sissippi State Agricultural Experiment Station.


Larvae of march and crane flies (Bibionidae,
Plecia sp. and Tipulidae, Tipula (Triplicitipula)
umbrosa Loew) were collected from beneath dam-
aged, low maintenance centipede grass sod in Pic-
ayune, Mississippi. Larvae of both species have
not been associated previously with turf damage.
Larvae and adult T umbrosa also were found as-
sociated with turf in a residential landscape in
Saucier, MS. This is the first record of T umbrosa
for Mississippi and a new record of larval habitat.


ALEXANDER, C. P. 1920. The Crane Flies of New York,
Part II. Biology and Phylogeny. Cornell Univ. Agr.
Exp. Sta., Mem. 38: 691-1133
ALEXANDER, C. P. 1967. The Crane Flies of California.
Bull. Calif. Insect Survey 8: 1-269
ALEXANDER, C. P., AND G. W. BYERS. 1981. Tipulidae,
pp. 153-190 In J. F. McAlpine, B. V. Peterson, G. E.
Shewell, H. J. Teskey, J. R. Vockeroth, and D. M.
Wood [eds.], Manual of Nearctic Diptera. Vol. 1. Ag-
riculture Canada Research Branch Monograph 27.
ricultural dipteran pests of the Palaearctic region,
Chapter 1.15 In L. Papp and B. Darvas [eds.], Con-
tributions to a Manual of Palaearctic Diptera, Vol. 1.
Science Herald, Budapest.
GELHAUS, J. K. 1986. Larvae of the crane fly genus Tip-
ula in North America (Diptera: Tipulidae) Univ.
Kansas Science Bull. 53(3): 121-182.
HARDY, D. E. 1981. Bibionidae, pp. 217-222 In J. F.
McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey,
J. R. Vockeroth, and D. M. Wood [eds.], Manual of Ne-
arctic Diptera. Vol. 1. Agriculture Canada Research
Branch Monograph 27.
HARTMAN, M. J., AND C. D. HYNES. 1977. Biology of the
range crane fly Tipula simplex Doane (Diptera:
Tipulidae). Pan Pac Entomol 53:118-123.
JACKSON, D. M., AND R. L. CAMPBELL. 1975. Biology of
the European Crane Fly, Tipula paludosa Meigen, in
Western Washington (Tipulidae; Diptera). Wash.
State. Univ Tech. Bull. 81.
LAGASA, E. H., AND A. L. ANTONELLI. 2000. 1999 West-
ern Washington Tipula oleracea Survey (Diptera:
Tipulidae). Wash. State Dep Agric. Pub no. 034.
OOSTERBROEK, P. 2003. Catalogue of the Craneflies of
the World. Database version Oct 2003 Zoological Mu-
seum, University of Amsterdam (18,092 records)
(distributed by the author).
THOMPSON, C. F. 1990. Biosystemic information: Dip-
terists ride the third wave, p. 179-201 In M. Koszt-
arab and C. W. Schaefer [eds.], Systematics of the
North American Insects and Arachnids: Status and
Needs. Virginia Agricultural Experiment Station In-
formation Series 90-1.
Turfgrass Insects of the United States and Canada,
2nd ed. Cornell University Press, Ithaca, NY.

March 2006

Scientific Notes


1USDA, ARS, Biological Control of Pests Research Unit, Stoneville, MS

2USDA, Office of Chief Economist, Delta Research and Extension Center, Stoneville, MS

3USDA, ARS, Imported Fire Ant and Household Insect Research Unit, Gainesville, FL

The red imported fire ant (RIFA), Solenopsis in-
uicta Buren, has been encroaching on the range of
the black imported fire ant B I FA.i, .... ... .., : rich-
teri Forel to the extent that the current range of
BIFA is limited to only three states: northern
Mississippi, northwestern Alabama, and south-
ern Tennessee. In Mississippi where the two spe-
cies coexist, evidence of hybridization has been
reported (Vander Meer et al. 1985; Ross et al.
1987). These two species produce reproductively
viable Fl hybrids that were found to occupy a
broad band across the northern tier of Missis-
sippi, Alabama, and Georgia (Diffie et al. 1988).
The objective of this study was to determine the
distribution of the RIFA, BIFA, and hybrid popu-
lations in Mississippi.
Study Site: Samples of worker ants were col-
lected from field colonies in northern and central
Mississippi. Mounds were mapped with a back-
pack Trimble 124 beacon DGPS system utilizing
GIS Solo CE V3.0 software (TDS) installed on a
Compaq iPAQ Pocket PC H3900 series. A vial
sample containing 100-1000 ants was removed
from each mound, labeled for identification, and
stored on ice until frozen. Frozen samples of ma-
jor caste workers were examined and identified to
species. Ant samples identified as S. richteri were
analyzed by gas chromatography for venom alka-
loids and cuticular hydrocarbons (Vander Meer et
al. 1985). These two classes of compounds readily
distinguish BIFA and hybrid ants which are mor-
phologically identical. Field data and the results
from species and hybrid identification were en-
tered into ArcView 3.2a Geographic Information
Systems (GIS) for analysis, and for spatial pre-
sentation of the data.
The distribution of RIFA, BIFA, and hybrid
populations in Mississippi for 2001-2003 are
shown in Fig. 1. A total of 176 mounds were sur-
veyed from 52 counties in Mississippi. An earlier
report found hybrid populations in five counties
from northeastern Mississippi (Diffie et al. 1988).
In this investigation, hybrid populations were
found in twenty-seven counties extending as far
west as Bolivar County in the Mississippi Delta
(Fig. 1). Earlier reports showed BIFA populations
in eight counties of northeast Mississippi (Diffie
et al. 1988). In this investigation, BIFA popula-
tions were found in twenty-two counties extend-

Fig. 1. Spatial distribution of S. richteri, S. inuicta,
and hybrid imported fire ant colonies within Mississippi
counties, 2001 to 2003.

ing northwest to Tunica and De Soto County, and
as far south as Noxubee County in eastern Missis-
sippi. RIFA populations were found as far north
as Bolivar County in west Mississippi. However,
the northeast range of RIFA populations extends
to Kemper County. RIFA populations further
south in Mississippi extend throughout the cen-
tral and southern region of the state (Fig. 1).
Several general conclusions can be reached re-
garding the distribution of imported fire ants in
Mississippi. RIFA populations extend further
north in the west than in east Mississippi,
whereas BIFA/hybrid populations extend further
south in eastern than in western Mississippi and
can be found in several northwestern counties.

This shift in spatial distribution for RIFA,
BIFA, and hybrid populations in Mississippi sug-
gests that BIFA populations will eventually be re-
placed by RIFA and hybrid populations in
Mississippi and perhaps even in the United
States. Currently the RIFA and hybrid popula-
tions extend further north in the western part of
Mississippi, whereas in the eastern part of Mis-
sissippi the BIFA/hybrid populations extend as
far south as Noxubee County. The distribution
RIFA, BIFA, and hybrid populations can have a
significant effect on the implementation of an
areawide program to manage fire ants in Missis-
sippi. Vogt et al. (2004) listed several of these fac-
tors, including sampling, treatment thresholds,
and biological control agents. The presence of
RIFA, BIFA, and/or hybrid populations at a re-
lease site for biological control agents would prove
critical in targeting specific fire ant populations
because most biological control agents are rela-
tively host specific.
The authors express appreciation to Mr. An-
thony Pranschke and Ms. Michele Hosack for as-
sistance with this study. The use of trade
markers, firm corporation names, or mention of a
proprietary product does not constitute an en-
dorsement or recommendation for its use by the

March 2006


This study determined the distribution of the
red and black imported fire ant, and their hybrid
ant populations in Mississippi. The range of black
imported fire ant populations was found to extend
to twenty-two counties, whereas hybrid popula-
tions were found in twenty-seven counties in Mis-
sissippi. The distribution of species and hybrid
fire ant populations will be important in the de-
velopment of area-wide programs to control im-
ported fire ants in this area of the United States.

Discovery of hybrid fire ant populations in Georgia
and Alabama. J. Entomol. Sci. 23: 187-191.
AND E. I. VARGO. 1987. Biochemical phenotypic and
genetic studies of two introduced fire ants and their
hybrid (Hymenoptera:Formicidae). Evolution 41:
REZ. 1985. Biochemical evidence for hybridization in
fire ants. Florida Entomol. 68: 501-505.
CALLCOTT. 2004. Mississippi area-wide program:
Unique aspects of working with black and hybrid im-
ported fire ants. J. Agric. Urban Entomol. 20: 105-111.

Florida Entomologist 89(1)

Scientific Notes


'Universidade Federal de Alagoas, Departamento de Quimica, Laborat6rio de Quimica Entomol6gica
Campus A. C. Simoes, BR 104-Norte, kml4, Tabuleiro dos Martins, Macei6-AL, CEP 57072-970, Brazil

2Universidade Federal de Sergipe, Departamento de Engenharia Quimica, Campus Jos6 Aluisio de Campos,
Jardim Rosa Elze S/N, Sao Crist6vao-SE, CEP 49.100-000, Brazil

3Universidade de Sao Paulo, Departamento de Entomologia, Fitopatologia e Zoologia Agricola
Campus Luis de Queiroz, Caixa Postal 9, Piracicaba-SP, CEP 13.418-900, Brazil

The northeastern region of Brazil produces a
substantial amount of fruit because of its climate,
soil fertility, and good irrigation programs. How-
ever, as fruit production increases, so do tephritid
fruit flies populations. Pest fruit flies occur in
seven of the nine states that belong to this north-
eastern region, especially the following species:
Anastrepha fraterculus (Wiedemann 1830), Ana-
strepha sororcula (Zucchi 1979),Anastrepha obli-
qua (Macquart 1835), and Ceratitis capitata
(Wiedemann 1824) (Malavasi et al. 2000). In spite
of the pest status of C. capitata and Anastrepha
species in the State ofAlagoas, no publication re-
porting their occurrence in Alagoas exists (Mala-
vasi et al. 2000; Zucchi 2001).
Anastrepha fraterculus, A. obliqua, A. soror-
cula and C. capitata are of quarantine importance
in many countries, especially the last species, due
to the rigorous restrictions (Araujo et al. 2000;
Sales & Goncalves 2000). In Brazil, C. capitata
and A. fraterculus severely damage only temper-
ate fruit cultivations in the southeastern and
southern regions, respectively. Anastrepha obli-
qua and A. sororcula have been considered as sec-
ondarily important pests (Malavasi et al. 2000).
However, the expansion of fruit production in the
northeastern region may change this situation.
Therefore, a survey of tephritid populations is
necessary in order to develop control strategies
for these pests in all states of the region.
Severely infested fruits from unmanaged culti-
vars which belong to four plant species (Mangifera
indica L- var. ligata, Averrhoa carambola L., Psi-
dium guajava L. var. paloma, and Jambosia sp. L.)
were collected from three families located in six es-
tates in Alagoas (Macei6, 0939'57"S/3544'07"W;
16 m, Rio Largo, 09028'42"S/3551'12"W; 39 m,
Paripueira, 09028'30"S/3532'30"W; 5 m, Arapi-
raca, 0945'09"S/3639'40"W; 264 m, Coruripe,
10007'32"S/3610'32"W; 16 m, and Uniao dos Pal-
mares, 09009'46"S/36001'55"W; 155 m) from Feb-
ruary 2000 to July 2001. In total, thirty kilograms
of infested fruits were collected, with an average
of four larvae/fruit. The fruits were placed in con-
tainers with a layer of vermiculite as a pupation

medium, and pupae were held in plastic boxes un-
til emergence of adults. Voucher specimens were
deposited at the Departamento de Entomologia,
Fitopatologia e Zoologia Agricola, Escola Superior
de Agriculture Luiz de Queiroz (ESALQ), Univer-
sidade de Sao Paulo, Piracicaba, SP, Brazil. Iden-
tification of fruit flies was carried out by Dr.
Roberto Ant6nio Zucchi on the basis of the mor-
phological characteristics of the female ovipositor.
We report the presence of pest fruit flies for the
first time in the State of Alagoas. Anastrepha
fraterculus, A. obliqua, A. sororcula, and C. capi-
tata were identified. Anastrepha obliqua and
A. fraterculus infested all the fruits collected. In
guavas the number of A. fraterculus was higher
than that of A. obliqua. In the remaining host
fruits, A. obliqua was the predominant species.
Anastrepha sororcula was found in fruits of the
Myrtaceae family (guavas and "jambos"). Cerati-
tis capitata was reared only from starfruits.Anas-
trepha obliqua and C. capitata were recovered in
the largest numbers. In addition, a fruit fly para-
sitoid Doryctobracon areolatus Szepligeti (1911)
(Hymenoptera: Braconidae) was found (Table 1).
All parasitoids were associated with Anastrepha
We thank Leila Flavia do Nascimento Lima,
Esther Maria Gonzaga Amorim, and Regina Mar-
tins Santos (Secretaria Estadual da Agricultura,
Abastecimento e Pesca, Macei6, Alagoas) for valu-
able help during the collections in Coruripe, and
the institutions CNPq (grant no. 471828/2004-1),
FAPEAL (grant no. 20031029421-0), and CAPES
for providing financial support and scholarships.


The occurrence of Anastrepha fraterculus,
A. obliqua, A. sororcula, and Ceratitis capitata is
reported for first time in the State of Alagoas. The
specimens were obtained from starfruits A. car-
ambola, guavas P guajava, mangoes M. indica,
and "jambos" (Jambosia sp.). The parasitoid
Doryctobracon areolatus was recorded attacking
the Anastrepha species.

Florida Entomologist 89(1)

March 2006


Number of females (F)
Host family Host species Fruit fly Parasitoid and males (M)

Anacardiaceae Mangifera indica L. A. obliqua 15F; 18 M
A. fraterculus 1F
Myrtaceae Jambosia sp. A. obliqua 14F; 12M
A. fraterculus 11F; 10 M
A. sororcula 02F; 01M
Psidium guajava L. A. fraterculus 121F; 95M
A. obliqua 25F; 21M
A. sororcula 10F; 07M
D. areolatus 23F; 15M
Oxalidaceae Averrhoa carambola L. A. obliqua -174F; 169M
C. capitata -155F; 162M
A. fraterculus -04F; 02M
D. areolatus 18F; 08M


ARAiUJO, E. L., F. A. M. LIMA, AND R. A. ZUCCHI. 2000.
Rio Grande do Norte, pp. 223-226 In A. Malavasi and
R. A. Zucchi [eds.], Moscas-das-frutas de Importan-
cia Economica no Brasil. Ribeirao Preto, Holos.
geografia, pp. 93-98 In A. Malavasi and R. A. Zucchi
[eds.], Moscas-das-frutas de ImportAncia Econ6mica
no Brasil. Ribeirao Preto, Holos.

SALES, F. J. M., AND N. G. G. GONQALVES. Ceara, pp.
217-222 In A. Malavasi and R. A. Zucchi [eds.], Mos-
cas-das-frutas de Importancia Economica no Brasil.
Ribeirao Preto, Holos.
ZUCCHI, R. A. 2001. Mosca-do-mediterraneo, Ceratitis
capitata (Diptera: Tephritidae), pp. 15-22 In E. F.
Vilela, R. A. Zucchi, and F Cantor [eds.], Pragas In-
troduzidas no Brazil. Ribeirao Preto, Holos.

Scientific Notes


'Formosan Subterranean Termite Research Unit, USDA-ARS, New Orleans, LA 70124

2Insect Biocontrol Laboratory, USDA-ARS, Beltsville, MD 20705

3Center for Disease Control, Atlanta, GA 30333

4Plant Protection Research Unit, USDA-ARS, Ithaca, NY 14853

*Author for correspondence; e-mail: araina@srrc.ars.usda.gov

Most noxious weeds infesting rangeland are
exotic species (DiTomaso 2000). The genus Cen-
taurea which contains several species of knap-
weed is the most abundant group in the western
United States (Skinner et al. 2000). In their na-
tive habitat of Eurasia, natural enemies have pre-
vented knapweed from becoming an economic
problem (Keane & Crawley 2002). However, in the
United States where they were accidentally intro-
duced more than 100 years ago, these weeds, in
the absence of their natural enemies, have repro-
duced unchecked and replaced many of the more
desirable rangeland vegetation. Biological control
of weeds with imported insects and pathogens is
safe, environmentally sound, and cost effective
(McFadyen 1998), and importation and use of
highly host-specific biological control organisms
offers considerable promise for weed control. Thir-
teen insect species have been imported into the
US from Eurasia for control of knapweeds (Lang
et al. 2000). Releases of some of these species on a
Colorado grassland reduced diffuse knapweed by
77% of absolute cover (Seastedt et al. 2003). How-
ever, a major constraint to the widespread use of
these biocontrol agents is the lack of sufficient
numbers of insects for release. Story et al. (1994)
produced 20,000 adults of Agapeta zoegana on
spotted knapweed planted in field cages at a cost
of $1.32/insect. Since artificial rearing has been
used to mass-produce insects that are comparable
to wild populations, at a reasonable cost (Leyva et
al. 1995), it would be desirable to develop artificial
diets to rear some of the weed-feeding insects.
Pterolonche inspersa (Lepidoptera: Pterolon-
chidae) that feeds both internally and externally
on diffuse knapweed roots and A. zoegana (Lepi-
doptera: Tortricidae) that feeds on the roots of
spotted knapweed, were selected for this study
based on their host specificity and efficacy.
Schroeder (1977) considered P inspersa to be one
of the most promising candidates for the biologi-
cal control of diffuse knapweed in North America.
In both species, rosettes were preferred for ovipo-
sition as well as feeding by newly hatched larvae.
In northern Greece, P. inspersa was reported to be

univoltine, diapausing during the winter months
as 3rd instars (Campobasso et al. 1994; Dunn at
al. 1989). A. zoegana, which has six instars, can
complete 2-3 generations per year in Europe
(Miiller et al. 1988), whereas in British Columbia,
it is restricted to only one generation (Muir &
Harris 1987). However, there is little information
on the nature of the diapause and no descriptions
of rearing methodology for either of these species.
We report here that both species can be reared
from the egg to the adult stage on artificial diet
and that diapause can be averted or shortened
under our rearing conditions.
Adults of P. inspersa and A. zoegana were col-
lected from diffuse and spotted knapweed in west-
ern Montana during June of 1997. The moths
were placed in 500-ml cylindrical paper oviposi-
tion containers lined with wax paper, provided
with 10% honey for food, and shipped overnight to
the USDA-ARS Insect Biocontrol Laboratory,
Beltsville, MD. The containers were held at 28
2C, 55 10% RH and a photoperiodic regimen of
LD 15:9. Eggs were removed every third day and
transferred to Petri dishes lined with moist filter
paper. Newly hatched larvae were placed directly
on diet. Because there was no artificial diet avail-
able for any insect in the family Pterolonchidae,
we obtained a diet developed for the pink boll-
worm, Pectinophora gossypiella; another member
of the superfamily Gelechioidea. Similarly, for A.
zoegana, we obtained diet developed for another
member of the Tortricid family, the Eastern
spruce budworm, Choristoneura fumiferana. Both
these diets were purchased from Southland Prod-
ucts, Lake Village, AR. Recipes for the P. gossyp-
iella and C. fumiferana diets can be found in
Bartlett and Wolf (1985) and in Robertson (1985),
respectively. Roots of knapweed, obtained from
Montana, were washed, freeze-dried, and pow-
dered in a grinder. Diets were prepared with or
without 2% root powder. Following testing of the
first batch of diets, we reduced the water content
by 10% (from 930 ml recommended by Southland
Products to 835 ml per liter of diet). Three types of
containers; wax coated paper straws (7 mm in

Florida Entomologist 89(1)

diam x 10 cm long), borosilicate glass culture
tubes (10 x 75 mm) and 30-ml clear plastic cups
with paper lids (BioServ, Frenchtown, NJ) were
tested for suitability in rearing both species. All
test containers were filled with diet and infested
with either 1 or 4 larvae. The straws were placed
in desiccators containing water, and together with
tubes and cups kept in an environmental chamber
maintained at LD 15:9 and temperatures of 30+'
2C and 25 2C during the light and dark cy-
cles, respectively. Subsequently, 131 cups contain-
ing pink bollworm diet and 509 cups containing
Eastern spruce budworm diet, both with and
without 2% root powder were infested with P. in-
spersa and A. zoegana larvae, respectively. The
containers were examined every 10 days for 50
days. Some of the larvae/pupae were shipped to
Sidney, MT. Whereas adults were allowed to
emerge from the pupae, the cups with larvae were
placed in a refrigerator (3C) for 86 days to pro-
vide for diapause, after which time the cups were
returned to 28C to promote further development.
Hatch was high (> 80%) if eggs were placed on
moist filter paper in Petri dishes. Since both spe-
cies were root feeders, we first chose to test diet-
filled straws, thus providing conditions for rear-
ing that simulated the natural environment.
However, the straws became moldy even though
the diet was prepared and the straws filled in a
laminar flow hood. Culture tubes were also in-
appropriate as rearing vessels because the tubes
retained too much moisture and the larvae
invariably drowned. Only approximately 10% of

P inspersa and 2% ofA. zoegana reached the 2nd
instar in tubes, even in the presence of root pow-
der. Growing larvae in plastic cups with paper
lids was the most effective method tested. A com-
parison of results from experiments in which 1 us
4 larvae were placed in each cup of the appropri-
ate diet showed that no more than one larva sur-
vived and grew to maturity. While both species of
larvae remained alive for several weeks on diets
in which root powder was omitted, insect growth
was considerably slower than when root powder
was present in the diet. With the incorporation of
root powder, the larvae initiated feeding and tun-
neled into the diet. Whereas 6.1% of P inspersa
larvae grew to 3rd or higher instar on diet with-
out the root powder, 33.3% did so when root pow-
der was incorporated into the diet. Larvae molted
4 times and reached the 5th instar in approxi-
mately 50 days. These 5th instars contained large
amounts of fat and their mean length was 8.6 mm
(Fig. 1A). Percent survival of A. zoegana was
lower than that of P inspersa, i.e., 9.1% of the lar-
vae placed in cups containing the Eastern spruce
budworm diet with root powder survived beyond
the 3rd instar. The mean length ofA. zoegana 4th
and 5th instars was 6.0 and 9.1 mm, respectively,
and it took approximately 45 days for larvae to
reach the 5th instar (Fig. 1B). Larvae of both spe-
cies spun silk sheaths during feeding, and re-
moval from their sheaths often resulted in larval
death. Less than optimal percentages of survival
were due in large part to high mortality of first
instars. In our study, two factors appeared to con-

Fig. 1. Pterolonche inspersa andAgapeta zoegana reared on their respective artificial diets. A. 5th instar of P. in-
spersa. Note the accumulated fat in the body. B. 5th instar ofA. zoegana. C. pupa ofA. zoegana. Scale bars = 2 mm.
D. Adult A. zoegana newly closed from the pupa.

March 2006

Scientific Notes

tribute to high mortality; drowning in droplets of
moisture and lack of immediate feeding. Lower-
ing the moisture content of the diets by 10% and
allowing the diets to dry for one hour before in-
festing with larvae reduced mortality. Although
2% root powder was incorporated into the diets,
apparently it did not provide sufficient stimula-
tion for a large percentage of the newly hatched
larvae to feed. Because the young larvae of both
species mine in the rosette leaves before they tun-
nel into the roots (Miller et al. 1988), it is possible
that some chemical factor present in the leaves,
but absent in the roots, may be acting as an initial
feeding stimulant. It is suggested that in future
studies, two diets, one with root powder as used
above and a second one with 2% freeze-dried pow-
dered young leaves of corresponding plants be
prepared and poured as two layers into 22 x 52
mm plastic tubes (BioQuip, Gardena, CA) or cups.
We observed three A. zoegana pupae (Fig. 1C)
which had apparently developed without under-
going diapause. The remainingA. zoegana and all
of the P inspersa larvae entered diapause. Ap-
proximately 30 days after removing both species
of larvae from the refrigerator, the first pupa ap-
peared and within a week, the first adults began
to emerge (Fig. 1D). Twenty-three and 37 adults of
A. zoegana and P inspersa, respectively, emerged
from late April to early September. These adults
were placed in mating cups and eggs that were
laid were used to establish an F2 generation. Data
from studies performed in Europe indicate that P.
inspersa is a univoltine species and showed that
this insect enters diapause as 3rd instars
(Schroeder 1977). In our investigations, some lar-
vae developed to the adult stage without entering
diapause, indicating that it is possible to select in-
dividuals from a population and use these insects
to develop a non-diapause strain. We recognize
that this study was not a comprehensive one.
However, the basic information that we have ob-
tained can be useful in developing mass rearing
techniques for exotic biocontrol agents such as the
two lepidopterans used in this study.
We thank Jim Story and Linda White (Montana
Agricultural Experiment Station, WARC, Corval-
lis, MT) for collection and shipment of insects to
Sidney, MT. Thanks are also due to Dr. Lincoln
Smith, USDA, ARS, Albany, CA, and Dr. David Ka-
zmer, USDA, ARS, Sidney, MT for critically re-
viewing an earlier version of the manuscript.
Mention of a proprietary product is not an en-
dorsement by the US Department of Agriculture.


Laboratory rearing methods for P inspersa
and A. zoegana, introduced into North America
for the control of the exotic knapweeds, Centaurea
spp., were developed. We used known diets for the
pink bollworm and Eastern spruce budworm and

added 2% knapweed root powder to these. After
45-50 days, we obtained some 4th and 5th instars
of both species, which apparently either averted
or shortened the diapause for these individuals.


BARTLETT, A. C., AND W. W. WOLF. 1985. Pectinophora
gossypiella, pp. 415-430 In P. Singh and R. F. Moore
[eds.], Handbook of Insect Rearing Vol. II, Elsevier
Science Publishers.
TORINO, AND P. H. DUNN. 1994. Biology of Pter-
olonche inspersa (Lep.: Pterolonchidae) a biological
control agent for Centaurea diffusa and C. maculosa
in the United States. Entomophaga 39: 377-384.
DITOMASO, J. M. 2000. Invasive weeds in rangelands:
Species, impacts and management. Weed Sci. 48:
M. TAIT. 1989. Host specificity ofPterolonche inspersa
and its potential as a biological control agent for Cen-
taurea diffusa, diffuse knapweed, and C. maculosa,
spotted knapweed. Entomophaga 34: 435-446.
KEANE, R. M., AND M. J. CRAWLEY. 2002. Exotic plant
invasions and the enemy release hypothesis. Trends
Ecol. Evol. 17: 164-170.
DEL. 2000. Release and establishment of diffuse and
spotted knapweed biocontrol agents by USDA,
APHIS, PPQ in the United States. Pan-Pacific Ento-
mol. 76: 197-218.
LEYVA, K. J., K. M. CLANCY, AND P. W. PRICE. 1995. Per-
formance of wild versus laboratory populations of west-
ern spruce budworm (Lepidoptera, Tortricidae) feeding
on Douglas-fir foliage. Environ. Entomol. 24:80-87.
MCFADYEN, R. E. C. 1998. Biological control of weeds.
Annu. Rev. Entomol. 43: 369-393.
MUIR, A. D., AND P. HARRIS. 1987. Biological control of
knapweed in Britisd Columbia, 1986 Annual Report.
B.C. Ministry of Lands and Forests, Integrated Re-
search. Victoria, B.C.
apeta zoegana (L.), a suitable prospect for biological
control of spotted and diffuse knapweed, Centaurea
maculosa and C. diffusa in North America. Can. En-
tomol. 120: 109-124.
ROBERTSON, J. L. 1985. Choristoneura occidentalis and
Choristoneura fumiferana, pp. 227-236 In P. Singh
and R. F. Moore [eds.], Handbook of Insect Rearing
Vol. II, Elsevier Science Publishers.
SCHROEDER, D. 1977. Biotic agents attacking diffuse
knapweed and spotted knapweed in Europe and
their prospective suitability for biological control in
North America. Proc. Knapweed Symp. Kamloops,
B.C., Canada. pp. 108-131.
Effect of biocontrol insects on diffuse knapweed
(Centaurea diffusa) in a Colorado grassland. Weed
Sci. 51: 237-245.
SKINNER, K., L. SMITH, AND P. PRICE. 2000. Using nox-
ious weed lists to prioritize targets for developing
weed management strategies. Weed Sci. 48:640-644.
STORY, J. M., W. R. GOOD, AND L. J. WHITE. 1994. Prop-
agation ofAgapeta zoegana L (Lepidoptera: Cochyl-
idae) for biological control of spotted knapweed-
procedures and cost. Biol. Control 4: 145-148.

Florida Entomologist 89(1)

December 2005


'Museo del Instituto de Zoologia Agricola (MIZA), Facultad de Agronomia
Universidad Central de Venezuela (UCV), Maracay. 2101-A, Aragua, Venezuela

2Museo Nacional de Ciencias Naturales, CSIC. Jos6 Guti6rrez Abascal, 2. 28006, Madrid, Spain

De Leon (1959) described the genus Phyllodro-
mus with the type species, Phyllodromus leiodis De
Leon, 1959, a phytoseiid mite [Acari: Mesostig-
mata] distributed in the state of Florida, USA. This
genus remained monotypic for four decades. Moraes
et al. (1997) described a second species (P trisetatus)
from the state of Bahia, Brazil. No other species has
been described in, or assigned to, this genus.
During a recent revision of the taxonomy of
Dendrobatidae (Amphibia: Anura), we detected
that Phyllodromus De Leon is a preoccupied
name created by Jim6nez de la Espada (1875) for
a genus of frogs (Type species: Phyllodromus pul-
chellum). Hence, Phyllodromus De Leon, 1959
(Acari: Phytoseiidae) represents a junior homo-
nym ofPhyllodromus Jim6nez de la Espada, 1875
(Anura: Dendrobatidae).
We propose here Leonacarus nom. nov. for
Phyllodromus De Leon, 1959, in application of the
International Code of Zoological Nomenclature
(ICZN 2000). The name refers to "the mite of
Leon" acknowledging De Leon who described and
recognized this taxon as a separated genus. The
species currently included in Leonacarus are
L. leiodis (De Leon, 1959) comb. nov., and
L. trisetatus (Moraes & Melo, 1997) comb. nov.


Phyllodromus De Leon, 1959 (Acari: Phytosei-
idae) is a junior homonym of Phyllodromus

Jim6nez de la Espada, 1875 (Anura: Dendro-
batidae) and in application of the International
Code of Zoological Nomenclature we propose the
name Leonacarus nom. nov. for Phyllodromus
De Leon, 1959. The species currently included in
Leonacarus are L. leiodis (De Leon, 1959) comb.
nov., and L. trisetatus (Moraes & Melo, 1997)
comb. nov.
We thank J. De Marmels, L.J. Joly (Instituto
de Zoologia Agricola, Facultad de Agronomia,
UCV), I. de la Riva (MNCN), and M.A. Alonso
Zarazaga (MNCN, ICZN) for comments on the


DE LEON, D. 1959. Two new genera of Phytoseiid mites
with notes on Proprioseius meridionalis Chant
(Acarina: Phytoseiidae). Entomol. News 70: 257-262.
ICZN. 2000. International Code of Zoological Nomencla-
ture. 4th ed. International Trust for Zoological No-
menclature. London. 306 pp.
JIMENEZ DE LA ESPADA, M. 1875. Vertebrados del viaje
al Pacifico verificado de 1862 a 1865 por una
comisi6n de naturalistas enviada por el Gobierno Es-
pahol. Batracios. Madrid: Imprenta Miguel Giniesta,
ii + 208 pp. + 7 pls.
MORAES, G., E. MELO, AND G. GONDIM. 1997. Descrip-
tion of a new species of phytoseiid mite from North-
eastern Brazil and redescription of Neoseiulus
gracilis (Acari: Phytoseiidae). Florida Entomol.
80(3): 319-324.

Scientific Notes


'USDA-ARS Southern Regional Research Center, Formosan Subterranean Termite Research Unit
New Orleans, LA 70124

2Texas A&M University, Image Analysis Laboratory, College of Veterinary Medicine
College Station, TX 77843-2257

The peritrophic matrix in many insects is con-
tinuously being synthesized in the mesenteron
and excreted with the fecal matter (Wigglesworth
1972; Richards & Richards 1977; Tellam 1996;
Lehane 1997). Peritrophic matrices are classified
into 2 types according to the way they are synthe-
sized in the mesenteron. Type I is made of concen-
tric lamellae loosely attached to one another and
synthesized by the epithelial cells through the
length of the mesenteron, while type II is a single
uniform layer synthesized by a group of cells in the
anterior limit of the mesenteron (Wigglesworth
1972; Tellam 1996; Lehane 1997). The peritrophic
matrix is most commonly made of y-chitin, which
is considerably more flexible than a-chitin while in
the presence of water (Herburn 1985). Wiggles-
worth (1972) mentions that termites may possess
type II peritrophic matrices, but this has not been
confirmed. The objective of this study was to deter-
mine the type of peritrophic matrix present in
Coptotermes formosanus Shiraki.
Formosan subterranean termites were col-
lected from 2 different localities (City Park and
Gretna) around the New Orleans metropolitan
area. All the termites were brought to the labora-
tory in plastic containers. An infested pine log and
carton nest were separately transferred to a 75.7-
L plastic trash container, containing 10 L of top-
soil: sand mixture at 1:1 ratio and 3 L distilled wa-
ter. Pieces of wood (Pinus taeda, Liquidambar
styraciflua, and Carya illinoensis) were added as
a source of food. The containers were maintained
in the dark at 27 3C for 7 d to allow the termites
to settle down. Then, termites were harvested by
placing pieces of wet cardboard inside of contain-
ers for 8 h. Termites were carefully removed from
the cardboard pieces by gentle manual shaking.
Twenty termite workers were randomly se-
lected and manually placed in a fixative solution of
2% glutaraldehyde, 2% paraformaldehyde, 2% ac-
rolein, and 1.5% dimethyl sulfoxide in 0.1 M so-
dium cacodylate buffer (pH 7.4) (Kalt & Tandler
1971, modified). To facilitate penetration of the fix-
ative into the termite bodies, the selected termites
were decapitated under a stereo microscope and a
small cut at the tip of the abdomen was made.
Fixed termite abdomens were rinsed 3 times in
0.1 M sodium cacodylate buffer, post-fixed in 1%
osmium tetroxide, and embedded in Araldite/
Epon epoxy resin (Araldite 502-EMbed 812) as

described by Mollenhauer (1964). Abdomens were
cut longitudinally producing 1-pm thick sections
with an Ultracut E microtome. Five individuals
were selected for sectioning based on the quality
of fixation. Section series for each termite were
placed on glass microscope slides, allowed to dry
on a hot plate for at least 5 min, and labeled.
Sections were stained with a modification of
Humphrey and Pittman's (1974) methylene blue,
azure II, and basic fuchsin staining technique.
This technique requires 2 stain solutions. The
blue stain was prepared by mixing 0.13 g methyl-
ene blue, 0.02 g azure II, 10 ml glycerol, 10 ml me-
thyl alcohol and 80 ml distilled water. The
mixture was stirred and filtered. The red stain
was prepared by mixing 0.2 g basic fuchsin in 100
ml distilled water, and diluted 1:4 in distilled wa-
ter after stirring and filtering. The staining proce-
dure was as follows: (1) The slide was flooded with
blue stain for 15-60 seconds,(2) then 4-6 drops of
1% NaOH solution were added and spread over
the slide by tilting it for 10 seconds, (3) slides
were washed in running water and dried on a hot
plate at 80C, (4) the red stain was added for 15-
30 seconds to slides on the hot plate, and (5) slides
were finally rinsed with running water and dried.
Sections were examined with an optical com-
pound microscope (Leica DMLB, Leica Microsys-
tems, Germany) and photographed with a Leica
MPS 60 micro photographic system. Color slides
were produced on Kodak Elitechrome 160T.
Slides were digitalized at 2,000 dpi with a high
resolution scanner (Minolta Dimage Scan Multi,
Konica Minolta, Japan).
Tissue coloration in sections was sufficiently
consistent to differentiate basic tissues. The me-
senterial epithelial cells appeared dark-blue to
purple (Fig. 1A, MGE), microvilli appeared red
(Fig. 1B, MV) and peritrophic matrix pink (Fig.
1A & B, PTM). Microvilli coloration was distinc-
tive enough to make them easily identifiable. The
peritrophic matrix was located next to the mass of
microvilli (Fig. 1B). Fat body cells appeared pink
to red in color (Fig, 1A, FBC).
The peritrophic matrix of C. formosanus ap-
pears to be synthesized around the invagination
(Fig. 1A, FCI) of the stomodaeum into the me-
senteron (Fig. 1A, MSE). A group of large pink-
colored cells located around the invagination por-
tion of the stomodaeum (Fig 1A & 1B, SC) shows

Florida Entomologist 89(1)

Fig. 1. Longitudinal section of foregut and midgut junction of C. formosanus workers. (A) Lower magnification
showing the foregut invagination (FGI) into the midgut and midgut epithellium (MGE). (B) Higher magnification
of foregut invagination showing secretary cells (SC) and peritrophic matrix (PTM). A is a composite of 2 pictures
taken with a 10 x ocular and 20 x objective; B is a composite of 3 pictures taken with a 10 x ocular and 100 x objec-
tive immersed in oil.

a clear substance that migrates to the lower part
of the invagination area. These cells appear to be
secretary in nature producing a clear substance
that is easily distinguishable (Fig. 1B). At the
base of the invagination, the peritrophic matrix
(PTM) seems to originate from the clear substance
produced by the pink secretary cells and then cov-
ers the inner part of the mesenteron (Fig. 1B).
Our observations showed the presence of
structures in the anterior part of the mesenteron
resembling those described by Wigglesworth
(1972) for type II peritrophic matrix (Fig. 1). Also
the structure of the peritrophic matrix observed
in workers of C. formosanus is a continuous layer
instead of independent lamellae as described by
Wigglesworth (1972). Based on these two charac-
teristics, we describe the peritrophic matrix of
C. formosanus workers as a type II.

The rate of type II peritrophic matrix produc-
tion varies from 1 to 10 mm/h in different insect
species (Waterhouse 1954). These production rates
make type II peritrophic matrices particularly vul-
nerable to the action of chitin synthesis inhibitors.


The peritrophic matrix of workers of Coptoter-
mes formosanus Shiraki (Isoptera: Rhinotermiti-
dae) was studied from stained histological
sections. Termites were decapitated, fixed in a
mixture of paraformaldehyde-glutaraldehyde,
embedded in epoxy, sectioned at 1 pm thickness,
and stained. Our observations showed the pres-
ence of structures in the anterior part of the me-
senteron resembling those described for a type II
peritrophic matrix.

March 2006

University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs