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Bullock et al. Nematodes us. Citrus Root Weevils


MANAGEMENT OF CITRUS ROOT WEEVILS (COLEOPTERA:
CURCULIONIDAE) ON FLORIDA CITRUS WITH
SOIL-APPLIED ENTOMOPATHOGENIC NEMATODES
(NEMATODA: RHABDITIDA)

R. C. BULLOCK', R. R. PELOSI', AND E. E. KILLER2
1University of Florida IFAS Indian River Research and Education Center
2199 S. Rock Road, Ft. Pierce, FL 34945

2USDA Agricultural Research Service
2199 S. Rock Road, Ft. Pierce, FL 34945


ABSTRACT

The entomopathogenic nematodes Steinernema carpocapsae Weiser and S. rio-
bravis Cabanillas, Poiner and Raulston were compared as twice a year soil treatments
for control of the west indian sugarcane root stalk borer weevil, Diaprepes abbreviatus
(L.), and citrus root weevil, Pachnaeus litus (Germar). Additionally, four single rate
demonstrations evaluated S. riobravis and a 2- year test compared single versus dou-
ble annual treatments of S. riobravis. The 6 experiments were conducted in the Indian
River Area of Florida, U.S.A.
Two applications per year of S. carpocapsae were as effective as S. riobravis for P.
litus control but failed to control D. abbreviatus.
A single application of S. riobravis delivering 2 million nematodes per tree through
low volume irrigation systems or a tractor-mounted herbicide boom in demonstration
tests provided 64% and 89% reduction of P. litus and D. abbreviatus, respectively.
Single annual applications of 2 million S. riobravis per tree delivered with sprin-
kling cans reduced P. litus populations by 65% and D. abbreviatus populations by 85%
over a 2 year period and was statistically equal to a 2-a-year treatment of S. riobravis
that provided 80% and 95% reductions, respectively.

Key Words: Diaprepes abbreviatus, Pachnaeus litus, root weevils, citrus, Steinernema
carpocapsae, S. riobravis

RESUME

Se compare durante un aho el efecto de dos tratamientos al suelo de los nematodos
entomopat6genos Steinernema carpocapsae Weiser y S. riobravis Cabanillas, Poiner y
Raulston para el control del picudo antillano de la cana, Diaprepes abbreviatus (L.), y
el picudo de los citricos, Pachnaeus litus (Germar). Se evalu6 ademas el efecto de una
sola aplicaci6n de S. riobravis en cuatro ensayos. En un ensayo adicional realizado du-
rante 2 ahos se compare el efecto de una sola aplicaci6n por aho y de dos aplicaciones
anuales de S. riobravis. Estos 6 ensayos fueron realizados en el area de Indian River
en Florida, E.U.A.
Dos aplicaciones al aho de S. carpocapsae fueron igual de efectivas como dos apli-
caciones de S. riobravis para el control de P. litus, pero S. carpocapsae no control aD.
abbreviatus.
Una aplicaci6n de una dosis de 2 millones de nematodos por arbol de S. riobravis,
utilizando un sistema de irrigaci6n a bajo volume, o utilizando un tractor con
sistema de aplicaci6n para herbicidas, result en una reducci6n del 64% y 89% de P.
litus y D. abbreviatus, respectivamente.
Una sola aplicaci6n de 2 millones de S. riobravis por arbol utilizando regaderas
manuales dio como resultado una reducci6n del 65% de la poblaci6n de P. litus y del

















Florida Entomologist 82(1)


March, 1999


85% de D. abbreviatus durante dos afios y no difirieron estadisticamente de dos trata-
mientos por afio, lo cual di6 como resultado una reducci6n del 80% y 95% para cada
especie, respectivamente.




In the 34 years since its detection in Florida, the west indian sugarcane rootstock
borer weevil, Diaprepes abbreviatus (L.), has spread from the original find in an Apo-
pka, FL, citrus nursery (Woodruff 1968) to approximately 11,364 ha of citrus groves
distributed in 19 Florida counties (Hall 1994). Annual losses and cost of control in cit-
rus currently exceeds 72 million dollars to citrus growers (Stanley 1996).
Attempts to control D. abbreviatus have passed through failed initial eradication
efforts, suppression with chemical sprays and soil treatments (Bullock 1985, Bullock
et al. 1988, Bullock and Pelosi 1993, McCoy et al. 1995, Quintela and McCoy 1997) to
biocontrol efforts prompted by discovery of natural agents recovered in soil surveys
conducted by Beavers et al. in 1983. Some of the entomopathogens discovered have
been investigated (McCoy and Boucias 1989) and a synergism identified between
pathogens and the insecticide imidacloprid (Quintela and McCoy 1997).
The nematode component of the Beavers et al. survey generated a body of research
that identified Steinernema carpocapsae Weiser as a soil treatment for control of lar-
vae (Schroeder 1987, 1990, Downing et al. 1991, Adair 1994, Bullock and Miller 1994).
The commercial product BioVectorT, containing S. carpocapsae All strain as its active
ingredient, became available to growers. After the discovery of S. riobravis, in Texas'
Rio Grande Valley (Cabanillas et al. 1994), Schroeder (1994) compared it with S. car-
pocapsae for control of D. abbreviatus larvae under controlled conditions. He found it
superior to S. carpocapsae and its early promise has been confirmed by field tests re-
ported here.


MATERIALS AND METHODS

The research reported here was conducted during the Spring of 1994 in four citrus
groves in the Indian River Area of Florida. A fifth grove provided data from a 3-year
study. All groves were infested with D. abbreviatus and the citrus root weevil, Pach-
naeus litus (Germar).
Grove No. 1, in Indian River County, was treated 23 March. This 20-year-old block
of double bedded 'Marsh' grapefruit trees, planted at a 3.6 x 6.9m spacing, was treated
at the rate of 1.6 million nematodes per tree delivered in water at 85 GPA through a
tractor-mounted herbicide boom and applied to a pre-irrigated, surface-littered grove
floor area beneath the tree canopy. Both sides of the tree row were treated. Treat-
ments were replicated four times in 50-tree plots. Conical hardware cloth ground
traps (0.66m2 at base, 60 cm tall), capped with 0.95 liter clear plastic cups (Paradise
Plastics, Plant City, FL) attached by 15 cm long braided elastics (Goody Products,
Peachtree, GA), were placed next to the tree trunks to monitor weevil emergence from
nematode treated (n = 100) and water treated (n = 100) soil.
Grove No. 2, in Indian River County, was treated 23 March by injection of S. rio-
bravis into the microirrigation system at a rate delivering 2 million nematodes per
tree. Treatments were replicated six times in 28-tree plots. Irrigation laterals, serving
single rows of 10-year-old 'Marsh' grapefruit trees planted at a 6x9m spacing were
closed during treatment application. When nematode delivery was completed, the ir-
rigation lines were purged, laterals opened to allow water flow to control plots and the
















Bullock et al. Nematodes us. Citrus Root Weevils


block received additional irrigation provided to facilitate nematodes movement into
the soil. Ground traps were placed next to 100 trees in the treated and 100 trees in un-
treated plots.
Grove No. 3, in Indian River County, was treated 24 March by injection of S. rio-
bravis into the microirrigation system at a rate delivering 2 million nematodes per
tree. Treatments were replicated 3 times in 46-tree plots. Irrigation laterals, serving
the paired rows of 6-year-old 'Minneola' tangelo trees planted at a 3.7 x 9m spacing on
double beds, were closed during treatment application. When completed, the lines
were purged, laterals opened and the block received additional irrigation to facilitate
the nematode movement into the soil. Ground traps were placed next to 100 trees in
the treated and at 100 trees in control plots.
Grove No. 4, in St. Lucie County, was treated 14 April. A 2 quart suspension of Bio-
Vector 355, containing 1.2 billion S. riobravis, was mixed with water in a pail. The
slurry was poured into the reservoir for injection through the microirrigation system
to a block of 'Marsh' grapefruit trees planted at a 3.75 x 9m spacing on single beds.
Each tree received 2 million nematodes. Soil had been pre-moistened by 1.91 cm of
rain the day before. When treatment was completed, irrigation lines were purged and
blocked laterals that prevented water flow to beds with control trees opened. An ad-
ditional half hour of irrigation to all the blocks delivered 5 gallons of water per tree to
facilitate movement of nematodes into the soil. Ground traps were placed at the base
of 100 treated and 100 control trees and were examined weekly for presence of emerg-
ing adult weevils.
The experiment in grove No. 5 was established in a block of 7-year-old, double-bed-
ded 'Ruby' grapefruit trees, planted at a 5.4 x 7.2m spacing, at the IFAS IRREC in Ft.
Pierce. Treatments were replicated 3 times in 34-tree plots and compared two appli-
cations (spring and fall) of S. riobravis with 2 applications of S. carpocapsae annually.
One hundred untreated trees served as controls. Treatments were applied in March
and September 1994. During treatments, 2 million nematodes suspended in a gallon
of water were applied with a sprinkling can to the pre-moistened soil at the base of
each tree in each of the treatments. One hundred ground traps were placed at 100
trees in each treatment plus 100 untreated trees and were examined weekly for pres-
ence of emerging adult weevils. Data were collected for one year, 1994.In grove N. 5,
during April 1995, a single annual S. riobravis treatment was substituted for the
twice-a-year S. carpocapsae and the experiment continued for 2 years comparing sin-
gle with double applications of S. riobravis.


RESULTS AND DISCUSSION

The four short-term demonstrations conducted in the spring of 1994 provided an
over-all 89% reduction of D. abbreviatus and 64% of P. litus (Table 1). Two treatments
per year of S. riobravis provide 98% and 82% reductions of D. abbreviatus and P. litus,
respectively, compared to 7% and 53% reductions resulting from the S. carpocapsae
treatments (Table 2, 1994). However, the efficiency of the S. carpocapsae treatment,
although providing a mediocre 53% reduction, was statistically equal to that of S. ri-
obravis in reduction of P. litus populations.
The results of the 2-year study comparing one versus two applications of S. rio-
bravis (Table 2, 1995, 1996) suggest that, although two applications per year would
provide a greater average percentage reduction, the difference in performance was
not statistically significant. A total of 1,611 weevils were collected in the course of the
experiment in Grove 5. Weevil pressure remained constant through the 3 years with
control traps yielding in access of 300 adults per year.
































TABLE 1. CAPTURE OF ADULT WEEVILS IN GROUND TRAPS AT FOUR DEMONSTRATION PLOTS.

Total weevils SEM collected in 100 ground traps in following groves treated with S. riobravis:'

Grove Number


Weevil Species


abbreviatus Treated
Untreated
litus Treated
Untreated

'Separation within columns by paired t-test, 5% level.


No. 1

0 0.00 b
40 0.49a
6 0.37 b
60 0.55a


No. 2

9 0.25 a
6 0.32 a
2 0.27 b
15 0.04 a


No. 3

28 0.35 b
59 0.48 a
0 0.00 a
6 0.61 a


No. 4

2 0.28 b
249 1.10 a
62 0.15 b
113 0.40 a


Average % Reduction

89%


64%
O


D.


P.

























TABLE 2. CAPTURE OF ADULT WEEVILS IN 100 GROUND EMERGENCE TRAPS AT GROVE NO. 5.

Total adult weevils SEM caught by year:z

Nematode Species 1994x % Reduction 1995 1996 Average % Reduction r

S. carpocapsae X2'
D. abbreviatus 145 + 1.83 a 7%
P. litus 79 + 0.88 b 53%
S. riobrauis X2
D. abbreviatus 4 + 0.33 b 98% 10 + 0.49 b 19 + 0.39 b 95%
P. litus 30 + 0.51 b 82% 22 + 0.40 b 36+ 0.56 b 80%
Untreated
D. abbreviatus 165 + 1.94 a 281 + 1.05 a 267 + 2.49 a
P. litus 167 -- 1.18 a 109 -- 0.55 a 135 + 0.75 a
S. riobrauis X1
D. abbreviatus 50 + 1.26 b 31 + 0.86 b 85%

P. litus 43 0.81 b 43 + 0.54 b 65%

xSeparation of totals within columns by Duncan's Multiple Range Test, 5% level.
7X2 = applications in march and September; X1 = march application only.

















Florida Entomologist 82(1)


March, 1999


The results of these tests suggest that the success obtained in studies by Schroeder
(1994) in lab bioassay and in potted citrus can be anticipated in the field. The compar-
ison between S. carpocapsae and S. riobravis in 1994 (Table 2) indicates that the cit-
rus industry would benefit from commercialization of S. riobravis as a replacement for
S. carpocapsae in management of both D. abbreviatus and P. litus.


ACKNOWLEDGMENTS

The authors would like to thank Mike Burton, Christy Ferguson, and Danielle
Giordano for their assistance in conducting this research. Florida Agricultural Exper-
iment Station Journal Series No. R-06621.


LITERATURE CITED

ADAIR, JR., R. C. 1994. A four-year field trial of entomopathogenic nematodes for con-
trol of Diaprepes abbreviatus in flatwoods citrus grove. Proc. Fla. State Hort.
Soc. 107: 63-68.
BEAVERS, J. B., C. W. McCOY, AND D. T. KAPLAN. 1983. Natural enemies of subterra-
nean Diaprepes abbreviatus (Coleoptera: Curculionidae) larvae in Florida. En-
viron. Entomol. 12: 840-843.
BULLOCK, R. C. 1985. Potential for controlling citrus root weevil larvae and adults
with chemicals. Fla. Entomol. 68: 417-423.
BULLOCK, R. C., AND R. W. MILLER. 1994. Suppression of Pachnaeus litus and Di-
aprepes abbreviatus (Coleoptera: Curculionidae) adult emergence with Stein-
ernema carpocapsae (Rhabditida: Steinernematidae) soil drenches in field
evaluations. Proc. Fla. State Hort. Soc. 107: 90-92.
BULLOCK, R. C., C. W. MCCOY, AND J. FOJTIK. 1988. Foliar sprays to control adults of
the citrus root weevil complex in Florida. Proc. Fla. State Hort. Soc. 101: 1-5.
BULLOCK, R. C., AND R. R. PELOSI. 1993. Toxicity of imidacloprid to selected arthro-
pods in the citrus greenhouse and grove. Proc. Fla. State Hort. Soc. 106: 42-47.
CABANILLAS, R. E., G. 0. POINAR JR., AND J. R. RAULSTON. 1994. Steinernema rio-
bravis n. sp. (Rhabditida: Steinernematidae) from Texas. Fundamental and
Applied Nematology. 1: 123-131.
DOWNING, A. S., C. G. ERICKSON, AND M. J. KRAUS. 1991. Field evaluation of ento-
mopathogenic nematodes against citrus root weevils (Coleoptera: Curculion-
idae) in Florida citrus. Fla. Entomol. 74: 584-586.
HALL, D. G. 1995. A revision to the bibliography of the sugarcane rootstalk borer wee-
vil Diaprepes abbreviatus. Fla. Entomol. 78: 364-377.
MCCOY, C. W., AND D. G. BOUCIAS. 1989. Selection of Beauveria bassiana pathotypes
as potential microbial control agents of soil-inhabiting citrus weevils. Mem.
Inst. Oswaldo Cruz, Rio de Janeiro. 84, Suppl. 3: 75-80.
MCCOY, C. W., E. D. QUINTELA, S. E. SIMPSON, AND J. FOJTIK. 1995. Effect of surface-
applied and soil-incorporated insecticides for the control of neonate larvae of
Diaprepes abbreviatus in container-grown citrus. Proc. Fla. State Hort. Soc.
108: 130-136.
QUINTELA, E. C., AND C. W. MCCOY. 1997. Effects of imidacloprid on development, mo-
bility and survival of first instars of Diaprepes abbreviatus (Coleoptera: Curcu-
lionidae). J. Econ. Entomol. 90: 988-995.
SCHROEDER, W. J. 1987. Laboratory bioassay and field trails of entomogenous nema-
todes for control of Diaprepes abbreviatus (Coleoptera: Curculionidae) in citrus.
Environ. Entomol. 16: 987-989.
SCHROEDER, W. J. 1990. Suppression of Diaprepes abbreviatus (Coleoptera: Curcu-
lionidae) adult emergence with soil application of entomopathogenic nema-
todes (Nematoda: Rhabditida). Fla. Entomol. 73: 680-683.
















Scheffrahn et al.: New Amitermes 7

SCHROEDER, W. J. 1994. Comparison of two Steinernematid species for control of the
root weevil Diaprepes abbreviatus. J. Nematology. 26: 360-362.
STANLEY, D. 1996. Suppressing a serious citrus pest. Agric. Res. 44: 22.
WOODRUFF, R. E. 1968. The present status of a west indian weevil Diaprepes abbre-
viatus (L.): in Florida (Coleoptera: Curculionidae). Fla. Dept. Agric. Div. Pl. In-
dus. Entomol. Circ. No. 77.
















Scheffrahn et al.: New Amitermes


AMITERMES AMICKI, A NEW SUBTERRANEAN TERMITE
(ISOPTERA: TERMITIDAE: TERMITINAE) FROM ARUBA

RUDOLF H. SCHEFFRAHN1, NAN-YAO SU1 AND TIMOTHY G. MYLES2
1Fort Lauderdale Research and Education Center, University of Florida
Institute of Food and Agricultural Sciences, 3205 College Avenue
Fort Lauderdale, Florida, 33314, U.S.A.

2Faculty of Forestry, University of Toronto, Toronto, M5S 3B3, Canada

ABSTRACT
Amitermes amicki n. sp. is described from soldiers collected on Aruba. For compar-
ison, the soldier ofA. beaumonti is redescribed from specimens collected in Mexico,
Belize, and Cuba. Amitermes amicki is the eighth congener thus far described from
the Neotropical Region. A technique is reported to enhance the visual clarity of digi-
tized scanning electron micrographs.

Key Words: Amitermes beaumonti, Neotropics, West Indies, Netherlands Antilles,
scanning electron microscopy, digital image enhancement

RESUME

Se describe Amitermes amicki sp. n. de soldados colectados en Aruba. Para compa-
raci6n, se describe de nuevo el soldado de A. beaumonti de ejemplares colectados en
Mexico, Belice y Cuba.Amitermes amicki es el octavo cong6nero descrito de la Region
Neotropical hasta ahora. Se report una tecnica para mejorar la claridad visual de mi-
crografias electr6nicas digitizadas.




The genus Amitermes Silvestri consists of about 100 living species distributed
worldwide (Scheffrahn and Su 1987, Roisin 1989, Scheffrahn et al.1989, Constantino
1992, Sands 1992). Seven species are known from the Neotropical Region (Constan-
tino 1998), includingA. amifer Silvestri,A. aporema Constantino,A. excellent (Silves-
tri), and A. foreli Wasmann from South America; and A. beaumonti Banks, A.
















Florida Entomologist 82(1)


March, 1999


cryptodon Light, and A. ensifer Light from Central America. Of these, only A. beau-
monti has been collected from an island locality (Cuba) (Hernandez 1994, Scheffrahn
et al. 1994).
Samples of an undescribedAmitermes species were collected on two recent expe-
ditions to the island of Aruba. Aruba is located in the southern Caribbean Sea, 26 km
north of Venezuela. Herein is provided a description of the soldier caste ofA. amicki
n. sp., and a redescription of the soldier ofA. beaumonti.

MATERIALS AND METHODS
Morphometrics of specimens preserved in 85:15 (ethanol: water) were made with
a stereomicroscope fitted with a calibrated ocular micrometer. Three ratios were used
to quantify the head and mandible shape ofAmitermes soldiers, namely, the head con-
traction ratio (head width at antennae maximum head width), the marginal tooth
ratio (length of right mandible tip to marginal tooth point length of right mandible
from tip to basilateral process), and the mandible curvature ratio (arch height of right
mandible, as measured by the length of a line perpendicular to the line used to mea-
sure mandible length, mandible length). The mandible curvature ratio is similar to
the minimum mandible curvature index II as used by Light (1930) for Amitermes;
however, the former is measured to the outer margin of the right mandible and there-
fore provides a more accurate representation of mandible curvature. Light (1930)
measured minimum mandible curvature to the inner margin of the left mandible.
The holotype soldier will be deposited in the collection of the American Museum of
Natural History, New York, New York. Paratype soldiers will be deposited in the Na-
tional Museum of Natural History (Smithsonian Institution), Washington, D.C.; the
Florida State Collection of Arthropods, Florida Department of Agriculture and Con-
sumer Services, Division of Plant Industry, Gainesville, Florida; and in the first au-
thor's collection at the University of Florida Research and Education Center, Ft.
Lauderdale, Florida.
Photographic prints of scanning electron micrographs (SEMs) of Amitermes sol-
diers were scanned at 600 dots/inch and saved in .tif format. Three photo enhance-
ment software packages were used with similar results: Corel Photo House version
1.10.071, Logitech FotoTouch Color version 3.0, and Micrografx Photo Magic ver-
sion 4.0a. The scanned images were 1) deskewed to best fit the monitor screen, 2) in
a few cases antennal segments or whole antennae were copied using the freehand se-
lection tool, pasted and deskewed to achieve a more aesthetic alignment, 3) the im-
ages were then processed using the unsharp mask tool, and 4) brightness and
contrast tools were then used to improve photographic exposure. The entire image
was outlined in black, and then the background was filled in black.
After enhancing the image and blackening the background, the image was con-
verted to its negative. The line tool was used to superimpose a black line over the orig-
inal scale bar. The original bar was removed and the new bar was moved to the desired
placement and its length labeled. Individual finished images were then imported into
a Microsoft PowerPoint presentation for final layout and labeling.

Amitermes amicki Scheffrahn, new species

IMAGO. Unknown.

SOLDIER. (Figs. 1-2, Table 1). Head, in dorsal view, rectangular with slight an-
terior convergence of sides; head contraction ratio = 0.927. Head surface smooth.
Head pale yellow with lighter interior area delineating the frontal gland; vertex con-

















Scheffrahn et al.: New Amitermes


h~5~1?'' 2


1mm


r ILIIY


Figs. 1-2. Enhanced scanning electron micrographs showing the dorsal (1) and lat-
eral (2) aspects of the soldier head capsule ofAmitermes amicki n. sp.


vex in profile; mandibles orange-brown distally, concolorous with head proximally.
Head capsule with about 65-75 long bristles, sparse near posterior, more numerous
around fontanelle. Fontanelle opening on very shallow mound. Labrum roundly tri-
angulate with 10 or more long bristles; postclypeus bilobed.
Mandible curvature asymmetrical; mandibles long and evenly slender; points of
marginal teeth near middle of mandibles; marginal tooth ratio = 0.537. Posterior mar-
gin of marginal teeth projecting about 60-90 from inner surface of mandibles; anterior
margins projecting 45-60. Left mandible curving in a high arch until distal 4/5, then
curving more sharply. Right mandible also highly arched and curving more sharply at
tip, but with slight and abrupt inward bend at 1/3 length from base; mandible curvature
ratio = 0.336. Cutting edges between mandible tips and marginal teeth showing fine op-
tical serrations when viewed with light microscope, but serrations undetected on SEMs.

















Florida Entomologist 82(1)


March, 1999


TABLE 1. MEASUREMENTS OF AMITERMES AMICKI SOLDIER.

Measurements in mm
(n = 13 from 2 colonies) Range Mean + S.D. Holotype

Head length with mandibles 1.77 1.93 1.84 + 0.047 1.83
Head length to dorsal condyle 1.10 1.20 1.15 + 0.032 1.16
Head width at antennae 0.86 0.93 0.89 + 0.020 0.91
Head width, maximum 0.91 0.98 0.97 + 0.023 0.98
Head height with postmentum 0.79 0.85 0.82 + 0.020 0.81
Right mandible length,
tip to basilateral process 0.77 0.81 0.79 + 0.013 0.80
Right mandible length,
tip to tip of marginal tooth 0.41 0.44 0.42 + 0.013 0.43
Right mandible minimum curvature 0.25 0.28 0.26 + 0.008 0.26
Pronotum, maximum width 0.57 0.63 0.60 + 0.018 0.61
Pronotum, maximum length 0.31 0.37 0.35 + 0.015 0.37
Postmentum gulaa), minimum width 0.18 0.21 0.20 + 0.01 0.20
Postmentum, maximum width 0.29 0.33 0.31 + 0.01 0.31
Postmentum median length 0.65 0.72 0.68 + 0.019 0.67
Total body length 3.76 -4.90 4.51 + 0.38 4.90



Antennae with 14 articles; relative length formula 2 > 3 = 4 < 5. Postmentum
strongly inflated and bulging beneath head capsule. Pronotum pale yellow; anterior
margin orange-yellow. Pronotum margins with bristles, anterior lobe sloping steeply
upward. Apical tibial spurs 2:2:2 with numerous stout setae located above apical spines.
Comparisons. Among Neotropical species, soldiers of A. amifer, A. aporema, A.
amicki, and A. foreli have marginal teeth located near the middle of the mandibles.
Amitermes aporema from Brazil is the smallest of the group (maximum head width
0.71-0.76 mm, Constantino 1992) and has highly reduced, almost indiscernible, mar-
ginal teeth. Amitermes foreli from Panama, Columbia, and Venezuela (Constantino
1998) is the largest of the group (maximum head width 1.38-1.50 mm, Light 1932)
with long, slender marginal teeth and with mandibles sharply hooked beyond mar-
ginal teeth. Soldiers ofA. amifer from northern Argentina, Paraguay, and central Bra-
zil (Constantino 1998) are closest to A. amicki, but the former are larger (maximum
head width 1.11-1.14 mm, Light 1932). The marginal teeth oftheA. amifer soldier are
proportionally larger and more conical than the marginal teeth ofA. amicki soldiers.
The soldier ofA. amicki is nearest in size to that of the smaller A. beaumonti, but A.
amicki has a more quadrate head capsule thanA. beaumonti (head contraction index
0.927 and 0.881, respectively), more curved mandibles (mandible curvature ratio
0.336 and 0.282, respectively), medially thicker mandibles, and less barb-like and
more medially positioned marginal teeth (marginal tooth ratio 0.537 and 0.435, re-
spectively). The A. amicki soldier also lacks the fine cuticular microsculpturing found
on the soldier ofA. beaumonti (Figs. 1-4).
Type Material. HOLOTYPE soldier and 9 paratype soldiers from Palm Beach,
Aruba, (12.58 N, 70.04 W) collected 2-IX-1995 by J. A. Chase from foraging galleries
under rock (#NA003); 3 paratype soldiers from hills near Noord' (12.570 N, 70.02 W)
collected 27-IV-1995 by A. van Liempt from under plywood on soil (#NA129).
















Scheffrahn et al.: New Amitermes 11

Etymology. This species is named after Mr. Breck Amick of Amick & Son Termite
and Pest Control, St. Petersburg, Florida. Mr. Amick, along with his parents, Philip
and Toni, support basic entomological research and recognize its benefits to the pest
control industry.

Amitermes beaumonti Banks 1918; redescribed by Light (1932).

IMAGO. Snyder (1924) described winged imagos from three specimens collected
in Panama; however, specimens were collected in flight and may be of another species.
No imagos have thus far been collected in association with foragers.
SOLDIER. (Figs. 3-4, Table 2). Head, in dorsal view, ovoid in posterior with
slightly convex sides and converging moderately toward the anterior; head contrac-
tion ratio = 0.881. Head covered with fine microsculpturing visible on SEM. Head pale
yellow with lighter interior area delineating the frontal gland; mandibles orange-
brown distally, concolorous with head proximally. Vertex slightly convex in profile;
frons flat, sloping about 40 from plane of vertex. Head capsule with about 70-80 long





















1mm


Figs. 3-4. Enhanced scanning electron micrographs showing the dorsal (3) and lat-
eral (4) aspects of the soldier head capsule ofAmitermes beaumonti.

















Florida Entomologist 82(1)


March, 1999


TABLE 2. MEASUREMENTS OF AMITERMES BEAUMONTI SOLDIER

Measurements in mm
(n = 18 from 3 colonies) Range Mean + S.D.

Head length with mandibles 1.41 1.77 1.64 + 0.101
Head length to dorsal condyle 0.94 1.14 1.04 + 0.062
Head width at antennae 0.71 0.86 0.79 + 0.047
Head width, maximum 0.80 0.99 0.90 + 0.067
Head height with postmentum 0.71 0.86 0.78 + 0.048
Right mandible length,
tip to basilateral process 0.65 0.78 0.73 + 0.042
Right mandible length,
tip to tip of marginal tooth 0.28 -0.34 0.32 + 0.021
Right mandible minimum curvature 0.17 0.22 0.20 + 0.015
Pronotum, maximum width 0.47 0.55 0.52 + 0.024
Pronotum, maximum length 0.26 0.32 0.29 + 0.018
Postmentum gulaa), minimum width 0.16 0.20 0.19 + 0.013
Postmentum, maximum width 0.23 0.29 0.26 + 0.016
Postmentum median length 0.56 0.72 0.62 + 0.054
Total body length 3.86 4.75 4.22 + 0.20


bristles, sparse near posterior, more numerous around fontenelle. Fontanelle opening
on very shallow mound. Labrum lingulate with 7-8 long bristles; postclypeus bilobed.
Mandibles long; narrowing proximal to marginal teeth, then noticeably broader
until again tapering near tip; points of marginal teeth clearly distal to midpoint of
mandibles; marginal tooth ratio = 0.435. Marginal tooth barb-like; posterior margin
of marginal tooth projecting backward about 1200 from inner surface of mandible; an-
terior margin forming gradually curving confluence with mandible tip. Mandible cur-
vature symmetrical, curving evenly from base in a moderate arch, then curving more
sharply midway between marginal teeth and tips; mandible curvature ratio = 0.282.
Cutting edge between mandible tip and marginal tooth smooth. Soldiers from Quin-
tana Roo having slightly shorter and more medially angulate mandibles.
Antennae usually with 14 articles, rarely 13 or 15; relative length formula vari-
able, but 2 > 3 > 4 < 5 most common. Postmentum strongly inflated and bulging be-
neath head capsule. Pronotum pale yellow. Pronotum margins with bristles, anterior
lobe sloping upward. Apical tibial spurs 2:2:2 with several additional stout spines
near apex of middle tibia.
Comparisons. See also comparisons for A. amicki above. The soldier of A. beau-
monti is closest to those of the NearcticA. emersoni and A. silvestrianus. The mandi-
bles of A. emersoni are narrower, straighter, and proportionally longer than those of
A. amicki, while the marginal teeth on the mandibles ofA. silvestrianus are set more
basally than those ofA. beaumonti. Measurements of a single soldier from Panama in
Light's (1932) redescription of A. beaumonti were considerably larger than either
measurements in the current or the original (Banks 1918) descriptions.
Material Measured. Soldiers (6/colony) were measured from the following three
colony localities: 0.5 km S. Valladolid Nuevo, Quintana Roo, Mexico (20.93 N, 87.32
W) collected 10-XII-1997 by J. A. Chase and J. R. Mangold (#MX204); 29 km SW of Be-

















Scheffrahn et al.: New Amitermes


lize City, Belize (17.42 N, 88.44 W) collected 4-X-1994 by J. Anderson, J. A. Chase,
and J.R. Mangold (#BZ004); Bacuranao, Cuba (23.130 N, 82.24 W) collected 2-V-1974
by J. Krecek (#CU904). Additional material examined: 20 km N. Panama City, Canal
Zone, Panama, collected 15-X-1979 by J. Krecek.

BIOLOGY

The type localities ofA. amicki on Aruba are characterized by rocky soil, cacti, and
small shrubs, reflecting the arid habitat of the entire island (annual rainfall about 50
cm, mostly during November and December). Foraging groups, working within darkly
coated earthen tubes, attack wood near the soil surface. This species does not build
mounds. The foraging habit of A. amicki is typical of Amitermes, especially of the xe-
rically adapted species of the southwestern Nearctic Region (Light 1930).
Amitermes beaumonti occupies a wide range of mesic and wet habitats in Central
America and Cuba. Foraging habits of this species are similar to those of A. amicki
and other Amitermes species, and it also lacks mound nests.

DISCUSSION

The digital enhancements (Figs. 1-4) demonstrate some of the potential uses of
photo enhancement software for improving the quality and esthetic appeal of SEMs.
These enhancements can serve to heighten the features of interest by removing visual
background noise, such as SEM stubs, adhesive materials, and shadows. Structures
to be compared can be uniformly positioned and sized for easier visual interpretation.
By converting SEMs to negative images, the black background can be eliminated, re-
sulting in an image that resembles a finely stippled pen and ink drawing while retain-
ing photographic accuracy.
On the negative side, digital manipulations may also have the potential to intro-
duce error, especially if parts such as palpi and antennae are cut and repositioned for
esthetic reasons. It is, therefore, important that the types of enhancements performed
on the images be specified by authors and that digital enhancements do not alter mor-
phological elements of the specimen.

ACKNOWLEDGMENT

We are grateful to Diann Achor at the University of Florida, Lake Alfred C.R.E.C.,
for assisting with scanning electron microscopy; John De Freitas, Carmabi Founda-
tion, Curacao, for local assistance and maps; Reginaldo Constantino (Univ. Brasilia,
Brazil) for his comparison ofA. amicki withA. aporema andA. amifer; and J. A. Chase
(Terminix International, Conyers, GA), F. W. Howard, and J. Krecek (Univ. of Florida,
Ft. Lauderdale) for critically reviewing this contribution no. R-06313 of the Florida
Agricultural Experiment Station Journal Series.


REFERENCES CITED

BANKS, N. 1918. The termites of Panama and British Guiana. Bull. American Mus.
Nat. Hist. 38: 659-668.
CONSTANTINO, R. 1992. A new species ofAmitermes Silvestri fromAmapa State, Bra-
zil (Isoptera, Termitidae, Termitinae). Bol. Mus. Paraense Emilio Goeldi, ser.
Zool. 8: 337-341.
CONSTANTINO, R. 1998. Catalog of the termites of the New World (Insecta: Isoptera).
Arq. Zoologia; Sao Paulo, Brazil 35: 135-230.
















14 Florida Entomologist 82(1) March, 1999

HERNANDEZ, L. M. 1994. Una nueva especie del g6nero Incisitermes y dos nuevos reg-
istros de termites (Isoptera) para Cuba. Avicennia 1: 87-99.
LIGHT, S. F. 1930. The California species of the genusAmitermes Silvestri (Isoptera).
Univ. California Publ. Entomol. 5: 173-214.
LIGHT, S. F. 1932. Contribution toward a revision of the American species ofAmiter-
mes Silvestri. Univ. California Publ. Entomol. 5: 355-415.
ROISIN, Y. 1989. The termite genus Amitermes Silvestri in Papua New Guinea. Indo-
Malayan Zool. 6: 185-194.
SANDS, W. A. 1992. The termite genusAmitermes in Africa and the Middle East. Nat.
Res. Inst. Bull. 51. Chatham, United Kingdom. 140 pp.
SCHEFFRAHN, R. H., AND N. -Y. SU. 1987. A world list of species in the genus Amiter-
mes (Isoptera: Termitidae). Sociobiology 13: 183-190.
SCHEFFRAHN, R. H., N. -Y. SU, AND J. R. MANGOLD. 1989.Amitermes floridensis, a new
species and first record of a higher termite in the eastern United States
(Isoptera: Termitidae, Termitinae). Florida Entomol. 72: 618-625.
SCHEFFRAHN, R. H., J. P. E. C. DARLINGTON, M. S. COLLINS, J. KRECEK, AND N. -Y. Su.
1994. Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the
West Indies. Sociobiology 24: 213-238.
SNYDER, T. E. 1924. Descriptions of new species and hitherto unknown castes of ter-
mites from America and Hawaii. Proc. United States Natl. Mus. 64: 1-45.


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Florida Entomologist 82(1)


March, 1999


AULACASPIS YASUMATSUI (HEMIPTERA:
STERNORRHYNCHA: DIASPIDIDAE), A SCALE INSECT PEST
OF CYCADS RECENTLY INTRODUCED INTO FLORIDA.

FORREST W. HOWARD', AVAS HAMON2, MICHAEL MCLAUGHLIN3, THOMAS WEISSLING1
AND SI-LIN YANG1
'University of Florida, Fort Lauderdale Research & Education Center
3205 College Avenue, Fort Lauderdale, Florida 33314

2Florida Department of Agriculture & Consumer Services
Division of Plant Industry, P. 0. Box 147100, Gainesville, FL 32614

3Fairchild Tropical Garden, 11935 Old Cutler Road, Miami, FL 33156

ABSTRACT

Observations were conducted in southern Florida onAulacaspis yasumatsui Tak-
agi (Hemiptera: Sternorrhyncha: Diaspididae), a recently introduced armored scale
insect native to Southeast Asia. The insect's host plants, exclusively cycads, were
identified in the following genera: Cycas (Cycadaceae); Dioon, Encephalartos, Micro-
cycas, (Zamiaceae), and Stangeria (Stangeriaceae). Cycas spp. seemed to be preferred
over other cycad genera by this insect. Aulacaspis yasumatsui infested pinnae,
rachides, strobili, stems, and roots of various species of cycads. It generally infested

















Howard et al.:Armored Scale Insect on Cycads


all above-ground plant parts of Cycas spp. and sometimes the primary and secondary
roots to a soil depth of 60 cm. No morphological differences in the frond- and root-in-
festing forms were noted. At an ambient temperature of about 24.5C, eggs hatched
in 8-12 days. In the field, some individuals developed to second instars in 16 days and
third instars in 28 days. Third instars (mature females) laid >100 eggs. Most of the fe-
males of a generation did not live longer than 75 days. No natural enemies were ob-
served. The scale insect populations became extremely dense and killed 100% of 15
Cycas revoluta Thunburg within a year of infestation. Between 1996 and June 1998
the insect spread from a limited area in southern Miami to various sites as far as 120
km north, probably by movement of plants.

Key Words: Cycad, Hemiptera, Homoptera, Sternorrhyncha, Coccoidea, Diaspididae,
scale insect, Aulacaspis, imidacloprid


RESUME

Se llevaron a cabo observaciones en el sur de Florida sobreAulacaspis yasumatsui
Takagi (Hemiptera: Sternorrhyncha: Diaspididae), una cochinilla de armadura que
ha sido recientemente introducida desde el sudeste de Asia. Las plants hospederas
fueron exclusivamente cicas identificadas en los g6neros siguientes: Cycas (Cycada-
ceae); Dioon, Encephalartos, Microcycas (Zamiaceae); y Stangeria (Stangeriaceae).
Cycas spp. fueron preferidas en comparaci6n con las cicas de otros g6neros.Aulacaspis
yasumatsui infest6 hojas, raquises, estr6bilos, tallos, y races de varias species de ci-
cas. Generalmente infestaron todas las parties de arriba de la tierra de Cycas spp. y a
veces las races primaries y secundarias hasta una profundidad de 60 cm en el suelo.
No se observaron diferencias entire las formas que infestaron races y las que infesta-
ron parties de arriba de la tierra. Debajo de una temperature ambiental de aproxima-
damente 24.5C los huevos nacieron en 8-12 dias. En el campo, unos individuos se
desarrollaron al segundo instar en 16 dias y al tercer instar en 28 dias. Los terceros
instares (hembras maduras) ovipositaron >100 huevos. La mayoria de las hembras de
una generaci6n no vivieron mas que 75 dias. No se observaron enemigos naturales.
Las poblaciones de la cochinilla se hicieron extremadamente densas y mataron al
100% de las 15 Cycas revoluta Thunburg despues de un ano de infestaci6n. Entre 1996
yjunio de 1998, el insecto se extendi6 desde un area limitada en el sur de Miami a va-
rios sitios hasta 120 km al norte, probablemente a causa del transport de las plants.




Aulacaspis yasumatsui Takagi (Hemiptera: Sternorrhyncha: Diaspididae) was
originally described from specimens collected from a cycad, Cycas sp., in Bangkok,
Thailand, in 1972 (Takagi 1977). Cycads sensu lato are plants in the gymnosperm or-
der Cycadales, which consists of three families: Cycadaceae (one genus, Cycas), Stan-
geriaceae (one genus, Stangeria) and Zamiaceae (eight genera). Cycads are an ancient
group that predate most extant plant taxa; there are 185 species of cycads in the
world (Jones 1993).
Aulacaspis yasumatsui is considered a pest of cycads in Thailand, but is usually
maintained in low densities in that country by parasitoids (Tang et al. 1997). It was
found in Miami, Florida, on cycads grown as ornamental plants in 1996. King-sagos,
Cycas revoluta Thumberg, the most popular cycad planted in this area, were found to
be highly infested with A. yasumatsui. Fairchild Tropical Garden (FTG) and Mont-
gomery Botanical Center (MBC), both which have important world collections of
cycads, were within the initially infested area. The rampant increase and spread of
this scale insect in these collections threatened the survival of several rare and en-

















Florida Entomologist 82(1)


March, 1999


dangered species conserved in these collections. An additional concern was the threat
posed to a large concentration of nurseries in southern Florida that grow and ship
cycads throughout the U. S. and abroad. We thus initiated a research project with the
long-range objectives of elucidating the bionomics of this insect and developing pest
management methods for it.
This paper reports results on the bionomics and host relationships ofA. yasumat-
sui.


MATERIALS AND METHODS

Observations were conducted in FTG, MBC and other sites within the infested
area in southern Miami. Most observations or counts were of the scale insect itself, ne-
cessitating that the scale (i. e., the 'testa' or 'scale covering' of some authors) be re-
moved with a fine probe.
A general survey of the extent of the infestation on cycads in southern Miami was
conducted in October 1996.
On 19 November 1996, 10 pinnae from a heavily infested C. rumphii Miguel about
3 m tall growing in FTG were removed at random and examined under a microscope
and the number of males and mature females ofA. yasumatsui per 5 cm2 counted on
the abaxial and adaxial surfaces.
To determine host range data ofA. yasumatsui, all species of cycads in FTG and
plants of other families adjacent to them were examined at least weekly between Octo-
ber 1996 and June 1998. Cycads at MBC were also examined several times during this
period. Because species were not replicated and specimens were not uniform with re-
spect to environmental factors such as age, size, shading, closeness to sprinkler heads,
etc., a valid ranking of host plant species preference could not be made. However, in-
fested cycads tended to be either (1) coated with a dense population of scales so as to ap-
pear to have been snowed on; such infestations were conspicuous from a distance of 20
m or more (highly infested), or (2) infested with only a sparse population that was not
evident without close examination of the plant (lightly infested), or (3) non-infested.
During the first few months of observations, significant populations of this scale
insect infested several genera of cycads additional to Cycas. At FTG, cycads in the ge-
nus Cycas were treated with insecticides. In March 1998, observations were con-
ducted to compare the infestation levels on fronds of 14 species of cycads of other
genera represented in FTG and MBC. Only species infested withA. yasumatsui were
included. Ten of the most infested pinnae from the most infested fronds were selected
and the number of mature female scales per 5 cm2 of the abaxial surface of each pinna
determined. Cycads with infestations of stems or strobili but not fronds were noted.
An experiment was conducted to determine the developmental rate ofA. yasumat-
sui under field conditions. Five C. revoluta plants about 20 cm tall with about eight
fronds growing in plastic 16 x17 cm containers were obtained from a nursery outside
of the infested area. These were examined and determined to be free of scale insects.
On April 2, 1997, these cycads were placed on the ground in FTG. Exposure of the
plants to crawlers was maximized by (1) placing them beneath a large cycad that was
highly infested withA. yasumatsui, and (2) scattering upon them pinnae infested with
first instar crawlers ofA. yasumatsui. Observations were conducted 16, 28, 35, 41, and
75 days after initial exposure of the plants to the scale insects. On each observation
date, one pinna was removed from each plant and examined under the microscope.
Numbers of crawlers and second and third instar females were counted.
A second series of five cycads with characteristics similar to those above was
placed in FTG on August 13, 1997, and infested withA. yasumatsui as described. Ob-

















Howard et al.:Armored Scale Insect on Cycads


servations were conducted 8, 15, 21, and 35 days later with a slightly modified
method: on each observation date, two pinnae were removed from each plant and the
stages and numbers ofA. yasumatsui determined.
To determine the duration of the egg stage, the egg nearest the vulva of a female
(i. e., presumed to be newly laid) was placed on filter paper in a Petri dish. Five eggs
were observed twice daily in a laboratory at a constant temperature of about 25C un-
til each had hatched.
Barbara Judd, a horticulturist employed at the MBC nursery, informed us that she
had often foundA. yasumatsui on roots of containerized cycads. To verify that scale in-
sects were consistently associated with roots, containerized C. revoluta were lifted,
and the soil washed from the roots and examined under a stereoscopic microscope.
Ten cycads were examined in December 1997 and 10 in June 1998. In addition, in
March 1998 five mature C. rumphii and one C. wadei Merrill were selected at random
in the field. The soil was removed from roots to a depth of 60 cm and the roots exam-
ined for scale insects.
Microscope slide mounts were made of mature female specimens ofA. yasumatsui
from the roots, fronds, stems and strobili. Fifteen specimens from fronds and 25 from
roots were examined to determine whether different morphological forms were asso-
ciated with the different plant parts.
The impact of high infestations of A. yasumatsui on cycad survival was deter-
mined. Five C. revoluta plants about 0.35 m tall with about 16 fronds growing in 28 x
30 cm plastic containers and 10 plants about 20 cm tall with about 8 fronds growing
in 16 x 17 cm plastic containers were treated by soil drenches with a systemic insec-
ticide to prevent establishment of the scale insects. On 12 December 1996, the larger
cycads were drenched with five g of Merit 75%WP ( Bayer AG) (3.75 g AI of imida-
cloprid) in 500 ml of water. On 2 April 1997, the smaller cycads were drenched with
two g of this product (1.5 gAI imidacloprid) in 200 ml of water. The plants were placed
intermixed with an equal number of cycads and exposed to the scale insects as de-
scribed above. The fronds of both treated and untreated plants were observed weekly
for scale insect infestation and damage.
Distribution and host records ofA. yasumatsui are from microscope slide mounted
specimens collected by personnel of the Florida Department of Agriculture and Con-
sumer Services, Division of Plant Industry (DPI), during routine plant inspections
and by the authors.


RESULTS AND DISCUSSION

The scales of mature females of A. yasumatsui are white, 1.2 -1.6 mm long and
highly variable in form. They tend to have a pyriform shape with the exuviae at one
end, which is typical of the tribe Diaspidini (Ben-Dov 1990), but are often irregularly
circular, conforming with leaf veins, adjacent scales, and other objects. The orange
body of the female can sometimes be seen vaguely through the thin scale. The ventral
scale is extremely thin to incomplete. The scale of the male is similar to those of many
other species of Diaspididae, being 0.5-0.6 mm long, white, and tricarinate, with the
exuviae at the cephalic end.
Macroscopically in the field, the scale of the female of A. yasumatsui resembles
that of Pseudaulacaspis cockerelli (Comstock), which is also common on cycads in
Florida. However, two characteristics visible with the scale removed under a 10X
hand lens distinguish the two species: (1) The color of the body of all stages and of the
eggs of A. yasumatsui is orange, except recently molted (callow) individuals, which
are yellow. The eggs and all stages of P. cockerelli are yellow. (2)Aulacaspis yasumat-

















Florida Entomologist 82(1)


March, 1999


sui has an expanded prosoma, an unusual characteristic of the genus (Fig. 2). Finally,
scales ofA. yasumatsui are usually more numerous on the abaxial surface of pinnae,
while those of P. cockerelli are more numerous on the adaxial surface. The scale of the
female of Pinnaspis strachani (Cooley) also resembles that of A. yasumatsui, but P.
strachani is far less common on cycads than the other two scale insect species in
southern Florida.
Hundreds of cycads examined in southern Miami from October 1996-June 1998
were infested withA. yasumatsui. The scale insects were consistently more numerous
on the abaxial than on the adaxial surfaces of pinnae, and in light infestations they
occupied the abaxial surfaces exclusively. Highly infested cycads were almost com-
pletely coated with a white crust that included scales of live and dead insects (Fig. 1).
Scales of males, which are less than half the length of scales of females, were nearly
always more numerous than those of females. Crusts composed of several layers of
scales of males were especially common on rachides, where we observed a maximum
of 500 scales per cm2.
Mean numbers of live scale insects per 5 cm2 of foliar surface of a heavily infested
C. rumphii on 19 November 1996, were 43.5 (SD = 34.2) males and 10.8(SD = 12.6)
mature females on the adaxial surface, and 128.0 (SD = 87.3) males and 66.8 (SD =
26.3) females on the abaxial surface of pinnae. Examination of microscope-mounted
specimens collected from October 1996-February 1998 confirmed the presence of A.
yasumatsui on 22 species (Table 1).
In October 1996, 65 Cycas plants in FTG representing 20 species were highly in-
fested withA. yasumatsui. Light populations were observed on some species of Dioon,
Encephalartos (both Zamiaceae), and Stangeria (Stangeriaceae). Although Cycas spp.
were treated by FTG plant protection personnel several times with systemic insecti-
cides between October 1996 and January 1998, the scale insects re-infested these
hosts rapidly, and Cycas spp. continued to be more highly infested than cycads of
other genera. In early 1997, the non-Cycas genera that were hosts of A. yasumatsui
were those listed above. By March 1998, Microcyas colocoma (Zamiacae), represent-
ing an additional genus, was infested. At this time, Encephalartos ferox appeared to
be the most highly infested species of non-Cycas genera. Pinnae sampled from the
most infested fronds of E. ferox had 23.2 (SD = 9.4) scales per 5 cm2 on the abaxial sur-
face. Of the additional non-Cycas species that were infested,Encephalartos maniken-
sis, Dioon rzedowski, and D. merolae had more than 11 females and the other non-
Cycas species listed had less than three females per 5 cm2 of abaxial surface on the
most infested fronds, or infestations were confined to strobili. For comparison, a mean
of 66.8 females per 5 cm2 were observed on the abaxial surface of the highly infested
C. rumphii on 19 November 1996.
Stangeria eripus (Kunze) Baillon was only lightly infested in FTG and MGC in Oc-
tober 1996, but several plants of this species in the MBC collection were heavily in-
fested by March 1998. One specimen of this species in FTG was highly infested in
early 1997 but after several months the population on this plant diminished to a light
infestation without any discernible cause.
Although cycads of some other genera native to the Eastern and Western Hemi-
spheres were hosts, Cycas spp. were consistently more highly infested. Cycas is native
to Asia (Jones 1993) where A. yasumatsui was first collected, suggesting that it may
be the original host genus ofA. yasumatsui.
Sixteen days after containerized C. revoluta were exposed to infestations ofA. ya-
sumatsui on 2 April 1997, settled first instar crawlers were observed on the plants
(Fig. 3a). There was a mean of 69.6 (SD = 14.7) scale insects on the abaxial surface
per pinna. A mean of 84.8% (SD = 11.6) were first instars, and the remaining 15.2%













Howard et al.:Armored Scale Insect on Cycads 19


a r


,st;lm=


Fig. 1. Frond of Cycas rumphii infested withAulacaspis yasumatsui.


.


I-W
&Ib J















Florida Entomologist 82(1)


Fig. 2. Slide-mounted mature female ofAulacaspis yasumatsui, expanded prosoma
indicated.


March, 1999


























TABLE 1. CYCADS INFESTED WITH AULACASPIS YASUMATSUI IN FAIRCHILD TROPICAL GARDEN, MIAMI, OCTOBER 1996-FEBRUARY 1998.

Family Genus Species Geographic origin

Cycadaceae Cycas media R. Brown Australia & Papua New Guinea
panzhihuaensis L. Shou & S. Y. Yang China
revoluta Thunberg Japan to Ryukyu Islands
rumphii Miguel India, Southeast Asia, Oceania
seemannii A. Braun Oceania
szechuanensis W. C. Cheng & L. K. Fu China
thouarsii R. Brown ex Gaudich Africa
wadei Merrill Phillippines
Zamiaceae Dioon califanoi De Luca & Sabatori Mexico
edule Lindley Mexico
merolae De Luca Mexico
rzedowskii De Luca, Moreti, Sabatori & Vasquez Mexico
spinulosum Dyer strobilii only) Mexico
tomasellii De Luca var. sonorense Mexico





























TABLE 1. (CONTINUED) CYCADS INFESTED WITHAULACASPIS YASUMATSUI IN FAIRCHILD TROPICAL GARDEN, MIAMI, OCTOBER 1996-FEBRUARY 1998.

Family Genus Species Geographic origin

Encephalartos barteri Miguel strobilii only) Africa
ferox Bertoloni Africa
hildebrandtii nr. Lembombensis A. Braun &
Bouch6 Africa
manikensis (Gilliland) Africa
pterogonus R. A. Dyer & I. Verd Africa

whitelockii P. J. H. Hurter Africa
Microcycas colocoma (Miguel) de Candolle strobilii only) Cuba
Stangeriaceae Stangeria eriopus (Kunze) Baillon Africa

















Howard et al.:Armored Scale Insect on Cycads


were in the second instar. Thus, this species was capable of developing to second in-
star in 16 days, or possibly earlier. Presumably, the individuals in second instar were
those that had settled earliest on the plants as first instars.
Twenty-eight days after initial exposure to the scale insects, a mean of 18.1% (SD
= 11.4) of the females examined had reached third instar. Again, these were probably
individuals that had invaded and settled earliest. The remainder were first instars
(44.6%, SD = 10.2), which presumably had invaded and settled relatively recently, and
second instars (37.3%, SD = 5.58).


E 1st instar

S2nd instar

D 3rd instar


16 28 35 41 75


E 1st instar

* 2nd instar

0 3rd instar


U + -i-'-- M -r---l--- I l----l- ----
8 15 21 35
days

Fig. 3. Percentages of different instars ofAulacaspis yasumatsui on cycad host
plants at successive intervals after infestation by the scale insects. Top: April-June
1997 bottom: August-September 1997.


100-

90.

80 -

70 -
6O.

%% 50 .

40.

30,

20

10,

















Florida Entomologist 82(1)


March, 1999


After 35 days about half of the population was in second instar and about 1/3 had
reached third instar. A mean of 24.7% (SD = 10.8) of the latter had laid eggs.
After 41 days a mean of 89.1% (SD = 8.5) of the scale insects examined were third
instars. A mean of 68.2% (SD = 13.9) of these had laid eggs.
After 75 days the portion of the female population consisting of mature individuals
had fallen to 24%. Presumably, many of the females of the first generation had ex-
pended their eggs and died. A mean of 76.0% of the population were first instars, pre-
sumably of the second generation.
Later in the summer, the development of the scale insect was similar to that ob-
served in spring. Eight days after initial exposure toA. yasumatsui on August 12, the
scale insects on the cycads were all first instars. At 15 days, a small portion of them
had attained second instar; at 21 days, about 3/4 of those examined were mature fe-
males, and at 35 days all were mature females (Fig. 3b). In summary, during the warm
season, females ofA. yasumatsui developed from egg to adult within the period of one
month, and as mature females lived for about one month.
The largest egg masses observed with females beneath scales consisted of about
90-110 eggs. Usually there were in addition several empty egg shells and several
crawlers. It should be noted that this does not constitute the total number of eggs pro-
duced per female, since wedid not count the eggs developing in ovaries, or crawlers
that had already left the mother scale insect. In the laboratory at about 25C. eggs
hatched in about 8-12 days (n = 5).
Roots were infested withA. yasumatsui males and females in six of 10 C. revoluta
in containers examined in December 1997 and four of 10 examined in May 1998. The
scale insects usually were aggregated on primary roots (about 10 mm in dia) and sin-
gly or in groups of a few on secondary roots (about 2 mm in dia) near the container
sides, i. e., near the soil-air interface. Some roots that protruded from drain holes in
the containers were also infested. However, several scale insects were found on roots
at 15 cm depth and several cm inward from the container sides.
Aulacaspis yasumatsui males and females were also found on roots of four of five
C. rumphii and on one C. wadei growing in the field. They were distributed at differ-
ent depths on primary (3 cm in diam) and secondary roots in groups of a few to several
individuals from near the soil surface to a maximum depth of 60 cm (Fig. 4). Compar-
isons of 20 specimens of adult females from fronds and 25 from roots of cycads re-
vealed no morphological differences distinguishing the mature females infesting the
two different plant parts. However, the scales of females on roots often appear to be
more convex than those on fronds.
The few species of Diaspididae known to infest roots of their host plants include
Chortinaspis subterraneus Lindinger and C. iridis Balachowsky. These were found be-
low ground on the root collar of their respective hosts,Agropyrum (Graminae) and Iris
sp. (Iridaceae) (Balachowsky 1941).
Aulacaspis yasumatsui has been tentatively identified (based on field characters)
on cycads at many sites in Southeast Asia, but only in areas with a monsoon climate,
i. e. with a pronounced wet season alternating with a severe annual dry season of usu-
ally five months. It is seldom if ever on cycads in rain forest areas. Their ability to in-
fest roots is apparently not an adaptation to living on deciduous plants, for the fronds
of cycads are evergreen and scale insects commonly live on the stems. This may be an
important adaptation to surviving brush fires, which are a perennial occurrence in
these monsoon areas.
The containerized cycads that were initially exposed to A. yasumatsui on 12 De-
cember 1996 and on 2 April 1997 in each case were highly infested withA. yasumatsui
within 2 weeks. Most of the fronds of the first pp.roup, which were larger, were chlo-















Howard et al.:Armored Scale Insect on Cycads


Fig. 4.Aulacaspis yasumatsui scales on root of Cycas rumphii.

















Florida Entomologist 82(1)


March, 1999


rotic by summer and were brown and desiccated within 270 days. Most of the fronds
of the second group, which were smaller, were necrotic within 112 days after being ex-
posed. These plants never recovered and died within one year of initial infestation. In
contrast, the 15 plants treated with imidacloprid remained virtually free of scale in-
sects and their fronds were green and healthy during the entire year.
The scales of A. yasumatsui males and females are remarkably persistent. Their
presence detracts from the appearance of ornamental plants. Cycads continued to ap-
pear to be highly infested after most of the scale insects had been killed by insecti-
cides, because the scales did not readily drop off. We were unable to remove old scales
easily by mechanical means, e. g., with brushes or high pressure water sprays. Old
scales became loosened by soaking excised fronds in soapy water overnight, but sim-
ilar solutions sprayed on cycad plants did not loosen the scales. A practical treatment
that loosens old scales of armored scale insects would be a welcome development in
horticulture and plant protection.
No organized survey was conducted, but based on accumulative observations by
the Division of Plant Industry and other plant protection and horticultural personnel,
the infestation was localized in the Old Cutler Road area of Miami in 1996. We as-
sume it had been introduced into this area a few years previously. By spring of 1997,
cycads throughout southern Miami as far north as the downtown area were infested.
During the summer of 1997 we found the scale insect at several sites more than 20 km
north (North Miami, Hialeah) and east (Miami Beach) of where it is assumed to have
been introduced. By February 1998, it was found in Broward County and Palm Beach
counties at sites about 55 and 120 km north, respectively, from its assumed introduc-
tion site. Although scale insects may be spread short distances by wind dispersal of
crawlers, we surmise that the long distance spread of this scale insect was by trans-
port of infested plants.
The unusually dense populations and rapid spread of this scale insect suggests
that they were imported without their natural enemies. Indeed, we have examined
thousands of scales of this species under the microscope without observing parasitoid
exit holes. Occasionally we observed adults of ladybird beetle species (Coccinel-
lidae)long established in Florida crawling on cycads infested with A. yasumatsui.
However, we have not observed coccinellid eggs or larvae on these plants or obtained
other evidence that they feed on the scale insects. Eumaeus atala (Lycaenidae), a but-
terfly whose larvae feed on cycads, sequesters cyasin, a compound contained in
cycads, which acts as a defense against predators (Bowers & Larin 1989). Possibly,A.
yasumatsui similarly sequesters defensive chemicals from its cycad hosts, which
could be a deterrent to generalist predators.
Aulacaspis yasumatsui is an unusually severe armored scale insect pest. When not
controlled by natural enemies, which is the present situation in Florida, they are
highly damaging and often lethal to their host plants. They have a high potential to
spread to new areas via plant movement, because one to a few fecund females hidden
on the fibrous stem or on roots can easily escape detection. Once established, they are
an unusually difficult scale insect to control. The experimental treatments mentioned
above involved higher rates of imidacloprid than might be practical in the horticul-
tural industries. Furthermore, while these treatments controlled the insect on con-
tainerized cycads, we have thus far had inconsistent results with imidacloprid as well
as some other chemicals for control of this scale insect on field-grown cycads. Scale in-
sects that survive in the roots may be the source of the rapid re-infestations that we
have seen consistently when the scale insect was controlled on the above-ground por-
tions of the plant. This may also make it difficult to control the insect with natural en-
emies.

















Howard et al.:Armored Scale Insect on Cycads


Given the value of cycads as ornamentals and the endangered status of some spe-
cies of these plants, control measures for this pest are urgently needed. The results of
studies of chemical control will be reported in another publication. Biological control
and other methods are being investigated for control of this pest.
In Florida, an area where the scale insect fauna is well known, in addition toA. ya-
sumatsui, 19 species of armored scale insects (Dekle 1976) and 7 species of soft scale
insects (Coccidae) (Hamon and Williams 1984) attack cycads.Aulacaspis yasumatsui
is the only member of its genus known on cycads, thus we have coined the vernacular
name cycadd aulacaspis scale' for this species.

ACKNOWLEDGMENTS

We thank Mary Collins, Don Evans and Charles Hubbock (Fairchild Tropical Gar-
den), Barbara Judd and Terrance Walters (Montgomery Botanical Center), for assis-
tance with records, locating cycad specimens and other valuable help; Scott L. Bryan,
Jason Michewicz, and Jorge Vargas for field assistance; John Davidson (University of
Maryland) and Paris Lambdin (University of Tennessee) for reviewing the manuscript;
the Holton Nursery for supplying test plants; and the Palm Beach Palm and Cycad So-
ciety for partial financial support. The work was also partially supported by a Research
Project Enhancement Award from the University of Florida to FWH. This paper re-
ports research only. Mention of a product does not imply an endorsement or recommen-
dation. This is Agricultural Experiment Station Journal Series No. R-06369.

REFERENCES CITED

BALACHOWSKY, A. 1941. Sur un Chortinaspis Ferris nouveau d'Asie Mineure. Bull.
Soc. Entomol. de France XLVI: 9-11.
BEN-DOV, Y. 1990. Classification of diaspidoid and related Coccoidea. Pp. 97-128
D. Rosen, [ed] Armored scale insects, their biology, natural enemies and con-
trol, Vol. A. Elsevier, Amsterdam. 384 pp.
BOWERS, M. D., AND Z. LARIN. 1989. Acquired chemical defense in the Lycaenid but-
terfly, Eumaeus atala. J. Chemical Ecol. 15(4): 1133-1146.
DEKLE, G. W. 1976. Florida armored scale insects. Florida Dept. Agr. and Consumer
Serv., Div. Plant Industry, Gainesville, Florida, U. S. A. 345 pp.
HAMON, A. B., AND M. L. WILLIAMS. 1984. The Soft Scale Insects of Florida (Ho-
moptera: Coccoidea: Coccidae. Florida Dept Agr and Consumer Serv., Div.
Plant Industry, Gainesville, Florida, U. S. A. 194 p.
JONES, D. 1993. Cycads of the World. Smithsonian Institution Press, Washington, D.
C. 312 pp.
TAKAGI, S. 1977. A new species ofAulacaspis associated with a cycad in Thailand (Ho-
moptera: Cocoidea). Insecta Matsumurana New Series 11:63-72.
TANG, W., S-L YANG, AND P. VATCHARAKORN. 1997. Cycads of Thailand.: Nong Nooch
Tropical Garden and the Cycad Conservation Company, Bangkok. 34 pp.
















Florida Entomologist 82(1)


March, 1999


EFFECTS OF PESTICIDE DEPOSIT PATTERN ON RATE OF
CONTACT AND MORTALITY OF APHYTIS HOLOXANTHUS
(HYMENOPTERA: APHELINIDAE)

S. U. REHMAN, H. W. BROWNING, M. SALYANI AND H. N. NIGG
University of Florida, IFAS, 700 Experiment Station Road, Lake Alfred, FL 33850

ABSTRACT

A laboratory study was designed to determine the pesticide droplets deposition on
leaf surfaces and its effect on parasite behavior and mortality. The rate of parasite/de-
posit contact increased as the number of deposits increased, even though total volume
of deposit remained constant. The bioassay results indicated that mean percent mor-
tality ofAphytis holoxanthus in the carbaryl treatment was highest on leaf discs with
smaller and more abundant deposits than larger and fewer deposits.

Key Words: Florida red scale, Aphytis holoxanthus, deposit pattern, contact rate, par-
asite

RESUME

Se realize un studio de laboratorio para determinar el patron de deposici6n de gotas
de plaguicida en la superficie de las hojas y su efecto en la tasa de contact y la morta-
lidad de Aphytis holoxanthus. La tasa de contact de parasito/dep6sito aument6 a me-
dida que el numero de dep6sitos se increments, aun cuando el volume total de los
dep6sitos se mantuvo constant. El porcentaje de mortalidad promedio de Aphytis ho-
loxanthus en el tratamiento con carbaryl fue mas alto en los discos de hojas con dep6si-
tos mas pequenos y abundantes que en los discos con dep6sitos mas grandes y escasos.




Biological control of Florida red scale (FRS), Chrysomphalus aonidum (L.) (Ho-
moptera: Diaspididae), by the introduction ofAphytis holoxanthus DeBach has perma-
nently reduced the pest status of Florida red scale to the extent that citrus growers
generally have forgotten that during the 1950s it was one of their most serious pests
(Browning, 1994). This parasite searches actively on citrus foliage and fruit to find its
armored scale host and can be negatively affected by pesticide deposits due to high rates
of contact associated with its walking on plant surfaces (Rosen, 1973). The impact of
pesticide deposits on parasite survival may be dependent upon the pattern of deposi-
tion, i.e. size and number of deposits and on the pesticide concentration of the deposit.
Different spray techniques are used to achieve spray deposition efficiency on citrus
leaves to maximize control of target pests, but improvements in deposition may also
severely disrupt host-parasite relationships. By measuring quickly and accurately
the droplet size of released sprays, the effects of droplet size on deposit and on biolog-
ical activity of the spray in relation to a specific crop pest can be more clearly defined
(Akesson & Yates, 1989). The spray deposition efficiency of orchard airblast sprayers
generally is considered to be low and is influenced by droplet size (Reichard et al.,
1977). New leaves have less surface waxes and showed slightly better deposition pat-
tern than mature waxy leaves (Salyani, 1988). The bottom surface of leaves, having a
more rough texture, captured more droplets than the top surfaces (Salyani, 1988).

















Rehman et al.: Pesticide Deposits and Mortality


This study was designed to determine the pesticide deposition on leaf surfaces and
its effect on parasite behavior and mortality. The objectives of this study were to char-
acterize the effects of different pesticide deposit patterns on rates of parasite contact
during leaf searching, and to determine the impact of contact rate on adult parasite
mortality.


MATERIALS AND METHODS

Adult parasites were allowed to search on leaf surfaces containing various pat-
terns of pesticide deposits. The total amount of pesticide and carrier delivered was the
same among all patterns. Parasite behavior was evaluated during searching, and
rates of contact with pesticide deposits were calculated for each deposit pattern.
Bioassays were conducted according to Rehman et al. (1998), using each deposit
pattern to determine the influence of pattern on parasite mortality induced by the
pesticide residues. These experiments were conducted under laboratory conditions of
24 + 2C and 60% R.H.

A. Holoxanthus Behavior on Different Deposit Patterns

Four deposit patterns of carbaryl and untreated controls were replicated three
times. The pesticide treatment was carbaryl wettable powder (Sevin 80OS: Rhone-Pou-
lenc, Research Triangle Park, NC 27709) at 1.5 g carbaryl per liter of water (1.25 lb
per 100 gal). Water served as the control. Kaolin clay (500 mg) was added to all solu-
tions to improve the visibility of deposits. Deposit patterns of 1, 2, 5 and 10Pl1 droplets
were made with a 10 p1 _-1% micro dispenser (Drummond, USA). The leaf deposits
produced by these volumes are found in field applied deposits (Salyani and Fox, 1994).
A total of 20 1l of each solution was dispensed onto 2 x 1.7 cm'Hamlin' orange rectan-
gular leaf patches. The deposits (20 droplets, 1 il; 10 droplets, 2 il; 4 droplets, 5 il; or
2 droplets, 10 il) in each replication were air dried for 90 minutes. The actual deposit
size for each pattern was obtained by measuring the deposit through a microscope.
The length and width of the four deposits for each pattern were measured and the di-
ameter of each deposit was recorded in square millimeters (mm2).
Twenty adultAphytis holoxanthus were exposed to each deposit patterns in each
replication by confining them in a small ventilated petri dish for each single leaf
patch. Each replication was recorded for 2 h by time lapse videography at 60 frames/
sec (Panasonic, Osaka, Japan). Observations were then interpreted and recorded dur-
ing the first 10 min, the middle 10 min (from 55-65 min), and the final 10 min of the
2 h exposure period. In each case, the observations were made by analyzing the first
10 parasite searching events on each leaf patch. One searching event consisted of an
individual adult parasite traversing the leaf patch. The total time on the patch and
number of encounters with spray deposits were recorded during review of the video-
taped exposures. Rates of contact were then calculated by dividing the number of con-
tacts between the parasite and a deposit droplet, by the total time the parasite spent
on the leaf patch. These data were analyzed by PROC GLM, comparing the rates of
contact for different deposit patterns during the three time intervals monitored (first
10 min, mid 10 min, and final 10 min) and the LSD procedure (SAS Institute, 1988).

A. holoxanthus Mortality

The four deposit patterns described above for each treatment of carbaryl and con-
trol were repeated on 4.5 cm2 leaf discs punched from field collected leaves and were
















Florida Entomologist 82(1)


March, 1999


air dried for 90 minutes. Four replicates of each treatment were placed into ventilated
plastic 4.9 cm2 diameter petri dishes. The bioassays were conducted as described by
Rehman et al. (1998). After 24 h of exposure of 20 adultA. holoxanthus, mortality data
were collected from each deposit droplet pattern by detaching the petri dishes from
the ventilation system and counting the live and dead parasites in each petri dish.
The mean percent mortality was calculated and compared between deposit pattern
and presence or absence of carbaryl. The data were analyzed with ANOVA and
Fisher's least significant difference (LSD) test (SAS Institute, 1988). Abbott's corre-
tion for control mortality was not used because control mortality was below 5%.

RESULTS

A. holoxanthus Contact Rate

As the number of droplets in the deposit pattern increased, the rate of parasite/de-
posit contact increased in the controls, even though total volume of deposit remained
constant (Table 1). In the carbaryl treatment increased contact occurred with 20 drop-
lets at all times compared to the other droplet treatments (Table 1). There were es-
sentially no differences between the contact rate of other droplet numbers (Table 1).
Also, the rate of contact was highest on leaf discs with 20 deposits in the control and
treatment. Although the size of individual deposits increased with fewer droplets, the
rate of contact with larger deposits was lower than with the more numerous smaller
deposits. The droplet sizes and droplet volume for different deposit droplet numbers
are given in Table 1. According to PROC GLM (SAS Institute, 1988), the mean rate of
contact to different numbers of deposits was highly significant (F = 13.16; df = 11; P
< 0.0001 and r2 = .92), being directly proportional to the number of deposit droplets,
particularly in the control treatment. Observations of different time intervals during
the 2 h exposure indicated that the rate of contact within the different 10 min inter-
vals for pesticide and non-pesticide treatments was also significant (F = 6.77, df = 2,
P < .0107). Rates of contact for patches with 20 deposits of carbaryl were higher dur-
ing the first 10 min and decreased afterward, in comparison to the control, where
rates of contact increased during the middle and final 10 min (Table 1). Carbaryl
treatments with 10 and 20 deposit droplets showed evidence of parasite effects during
the middle 10 min and final 10 min periods. Adult parasites became sluggish and
their mobility decreased. Few sluggish or immobile parasites were observed on leaf
patches with 2 or 4 deposits of carbaryl. Control treatments showed no evidence of
negative effects on parasite activity during the 2 h exposure period. The parasites
spent most of their time walking or grooming on the leaf surface, but responded to de-
tection of deposits on the leaf surface by exhibiting antennation and probing behavior.
Occasionally, parasites were observed to stop on droplet deposits, circle around them
and tap or probe the margins of the deposits with their ovipositers. Contact with de-
posits appeared to be random. Parasites did not appear to be either attracted or re-
pelled by the deposits in either the carbaryl or control treatments.

A. holoxanthus Mortality

The bioassay indicated that mean percent mortality ofA. holoxanthus in the car-
baryl treatment was highest on leaf discs with the smallest and more abundant de-
posits. The mean percent mortality on leaf discs with 20 and 10 deposits was 53.8%
and 45.2%, respectively. There was no mortality on leaf discs with 4 deposits of car-
baryl; however, some parasites were sluggish and a few were immobile. Parasite mor-
























TABLE 1. MEAN RATE OF CONTACT OF APHYTIS HOLOXANTHUS TO DIFFERENT DEPOSIT PATTERNS.

x rate of contact (# of contact/sec) (-- SE)

Deposit diameter
Treatments Deposit numbers Drop vol (P1) (mm2) (- SE) First 10 minute Mid 10 Minute Final 10 Minute

control 2 10 5.6 (1.3) 0.07" (.04) 0.23" (.08) 0.32" (.06)
4 5 5.1 (2.0) 0.19b (.06) 0.49b (.08) 0.59b (.05)
10 2 4.3 (1.2) 0.41c (.04) 1.01/ (.18) 0.68K (.09)
20 1 3.8 (0.5) 0.75d (.13) 1.31d (.39) 1.49'(.28)

Mean 0.36 0.76 0.74

carbaryl 2 10 6.0 (.8) 0.11" (.03) 0.29" (.16) 0.09" (.03)
4 5 5.4 (0.8) 0.16" (.09) 0.39" (.14) 0.15" (.02)
10 2 4.4 (0.7) 0.29b (.07) 0.29" (.19) 0.25"b(.09)
20 1 3.9 (0.4) 1.07 c (.28) 0.69b (.11) 0.67c (.13)
Mean 0.48 0.42 0.29
Grand mean 0.42" 0.59b 0.52b

SE = Standard error of mean
Means followed by a different letter for each treatment in a column and column grand means followed by a different letter in each row are significantly different at p < 0.05 according
to LSD multiple range test.
















Florida Entomologist 82(1)


March, 1999


tality in deposit patterns without carbaryl (control) averaged 3.7% with 2 deposits
and 1.2% with 10 deposits (Fig. 1) indicating no significant effect of deposit size or
number.

DISCUSSION

According to Himel (1969), as the diameter of the average droplet size decreases,
the total number of spray droplets increases markedly. This increases the potential
for contact of the droplets with target insects. In another study Himel and Moore
(1969) mentioned that most commercial spray devices produce spray droplets whose
size (diameter) ranges from a few microns to hundreds of microns. The efficiency of
any pesticide spray and the degree of control obtained is a function of the ultimate
point of deposition of each of the pesticide spray droplets produced. In this study A.


60




50-


2 4 10 20

Number of Deposits


Fig. 1. Mean % mortality of adult Aphytis holoxanthus after 24 h exposure to
treated leaves with different numbers of deposit sizes. Error bars = standard error of
the mean.

















Rehman et al.: Pesticide Deposits and Mortality


holoxanthus did not react differentially to the carbaryl and control deposit patterns.
Random contact of A. holoxanthus with deposits in both the carbaryl and control
treatment suggested that there was no obvious attraction or repellency by these de-
posits. The lower rates of contact in the pesticide treatments of 20 deposits at the mid-
dle 10 min and final 10 min periods of the 2 h exposure were due to sluggish parasite
behavior. There was no obvious trend for other deposit sizes. In contrast, the parasites
on control deposits were very active throughout the 2 h exposure period.
Salyani and McCoy (1989) concluded that decreasing spray droplet size increased cit-
rus rust mite control. Our study showed that, for a constant amount of pesticide applied,
increased numbers of smaller droplet deposits resulted in increased contacts between ac-
tively searchingA. holoxanthus adults and static deposits. Greater mortality was expe-
rienced within treatments in which rates of contact were highest. Thus, improved
coverage of foliar pesticide applications may magnify the mortality rate of parasites.


ACKNOWLEDGMENTS

This study was supported by a Florida citrus grower box tax for research. Funds
for this project were made available from the Citrus Production Research Marketing
Order by the Division of Marketing and Development, Florida Department of Agricul-
ture and Consumer Services. Florida Agricultural Experiment Station Journal Series
No. R-04636.


REFERENCES CITED

AKESSON, N. B., AND W. E. YATES. 1989. Pesticide deposit and activity as a function
of spray atomizers and liquid formulation, pp. 2081-2090 in V. A. Dodd and
P. M. Grace (ed.), Agricultural engineering: Proceedings of the eleventh Inter-
national Congress on Agricultural Engineering, Dublin.
ALEXANDRAKIS, V. 1990. Effect of Dacus control spray, by air or ground, on the ecology
ofAspidiotus nerii Bouche (Homoptera: Diaspididae). Acta Horticul. 286: 339-
342.
BROWNING, H. W. 1994. Early classical biological control on citrus, pp 27-46 in
D. Rosen, F. D. Bennett, J. L. Capinera (eds.), Pest management in the subtrop-
ics: Biological control-a Florida perspective. Intercept Ltd., U.K.
HIMEL, C. M. 1969. The optimum size for insecticide spray droplets. J. Econ. Entomol.
62: 919-925.
HIMEL, C. M., AND A. D. MOORE. 1969. Spray droplet size in the control of spruce bud-
worm, boll weevil, bollworm, and cabbage looper. J. Econ. Entomol. 62: 916-918.
REICHARD, D. L., H. J. RETZER, L. A. LILJEDAHL, AND F. R. HALL. 1977. Spray droplet
size distributions delivered by air blast orchard sprayers. American society of
Agricultural Engineers. 20: 232-233.
REHMAN, S. U., H. W. BROWNING, H. N. NIGG, AND J. M. HARRISON. 1999. Residual ef-
fects of carbaryl and dicofol on Aphytis holoxanthus Debach (Hymenoptera:
Aphelinidae) Biol. Control. (in press).
ROSEN, D. 1973. Methodology for biological control of armored scale insects. Phytopar-
asitica 1: 47-54.
SALYANI, M. 1988. Droplet size effect on spray deposition efficiency of citrus leaves.
ASAE. 31: 1680-1684.
SALYANI, M., AND C. W. MCCOY. 1989. Spray droplet size effect on mortality of citrus
rust mite pp. 262-273 in Pesticide formulations and application system, ASTM.
Philadelphia.
SALYANI, M., AND R. D. Fox. 1994. Performance of image analysis for assessment of
simulated spray droplet distribution. Trans. ASAE 37: 1083-1089.
SAS Institute. 1988. SAS users guide: Statistics. SAS Institute, Cary, NC.
















Florida Entomologist 82(1)


March, 1999


CICADA (HOMOPTERA: CICADOIDEA) TYPE MATERIAL IN
THE COLLECTIONS OF THE AMERICAN MUSEUM OF
NATURAL HISTORY, CALIFORNIA ACADEMY OF SCIENCES,
SNOW ENTOMOLOGICAL MUSEUM, STATEN ISLAND
INSTITUTE OF ARTS AND SCIENCES, AND THE UNITED
STATES NATIONAL MUSEUM

ALLEN F. SANBORN
Barry University, School of Natural and Health Sciences, 11300 NE Second Ave.
Miami Shores, FL 33161-6695

ABSTRACT
This work is a comprehensive listing of the type material of the cicadas found in
the collections of the American Museum of Natural History, California Academy of
Sciences, Snow Entomological Museum, Staten Island Institute of Arts and Sciences,
and the United States National Museum. I have included all type material identified
in the collections. The California Academy of Sciences collection contains 4 types, 17
holotypes, 14 lectotypes, 19 allotypes, and 257 paratypes from 64 species in 14 genera.
The American Museum of Natural History contains 105 types, 2 holotypes, 8 co-types,
64 allotypes, and 149 paratypes from 136 species in 27 genera. Lectotypes and allo-
types for Tibicen linnei and T. lyricen var. engelhardti are designated. In addition, an
allotype (one of a pair) of Platypedia putnami var. lutea is designated as a paratype.
The Snow Museum has 10 types, 6 allotypes, and 351 paratypes from 35 species in 8
genera in the collection. The Staten Island Institute collection contains 15 types, 8
syntypes, 10 co-types, 5 allotypes, and 1,327 paratypes from 116 species in 24 genera.
The U.S. National Museum contains 55 types, 178 co-types, 11 allotypes, and
paratypes from 111 species of 36 genera identified. References to the original species
descriptions are provided.

Key Words: cicadas, Cicadidae, Tibicinidae, type material

RESUME
En este trabajo se present una lista comprensiva de los ejemplares tipos de las chi-
charras que se encuentran en las colecciones de las siguientes instituciones: American
Museum of Natural History, California Academy of Sciences, Snow Entomological Mu-
seum, Staten Island Institute of Arts and Sciences, y United States National Museum.
Se incluye todo material tipo identificado en las colecciones. La colecci6n en la Acade-
mia de Ciencias de California tiene 4 tipos, 17 holotipos, 14 lectotipos, 19 alotipos, y
257 paratipos de 64 species en 14 g6neros. El Museo Americano de Historia Natural
tiene 105 tipos, 2 holotipos, 8 cotipos, 64 alotipos, y 149 paratipos de 136 species en 27
g6neros. Lectotipos y alotipos de Tibicen linnei y T lyricen var. engelhardti son desig-
nados aquf. Ademas, un alotipo (uno de un par) de Platypedia putnami var. lutea es de-
signado como un paratipo. El Museo Snow de Entomologia tiene 10 tipos, 6 alotipos, y
351 paratipos de 35 species en 8 g6neros en la colecci6n. La colecci6n del Instituto de
Artes y Ciencias de Staten Island tiene 15 tipos, 8 sintipos, 10 cotipos, 5 alotipos, y
1,327 paratipos de 116 species en 24 g6neros. El Museo Nacional de Estados Unidos
tiene 55 tipos, 178 cotipos, 11 alotipos, y paratipos de 111 species en 36 g6neros iden-
tificados. Se proven referencias para las descripciones originales de las species.

















Sanborn: Cicada Type Material


Several of my current research projects have resulted in a need to locate type spec-
imens of cicadas. I have found the information available in the literature is often in-
adequate for determining the specific identity of specimens, particularly as I collect a
greater number of species. Many of the early species descriptions are vague or simply
do not provide sufficient information to allow distinct separation of species. Often,
only a few sentences were used to describe characteristics which, although sufficient
to identify a species at that time as unique, have subsequently been found in other an-
imals that were described later. Another common problem with the original descrip-
tions is the use of comparisons to other species as an integral part of the species
description. You are unable to determine if you have a particular species from these
descriptions if you do not possess examples of all the species used in the comparisons.
Similarly, the keys that have been produced for the cicadas are incomplete when try-
ing to identify a large number of species from a wide geographic range. The keys gen-
erally are restricted to the species found within a specific state, e.g. California
(Simons 1954) or Michigan (Moore 1966), and may, even then, be incomplete.
Due to these circumstances, a positive identification often requires comparing
specimens to the type material. However, the location of the type specimens is not al-
ways known. Many of the original descriptions do not state where a type was depos-
ited or simply state that the type is part of the author's collection. Over the years
these private collections have been acquired by larger institutions or parts of individ-
ual collections may have been transferred between institutions.
This work is an effort to identify type material in the large cicada depositories of
the United States. Each of these institutions has either had scientists associated with
it who described cicada species or has acquired significant collections which included
cicada type material. The California Academy of Sciences (CAS) collection contains
the collections of E. P. Van Duzee, F. H. Wymore, and B. P. Bliven. The Snow Entomo-
logical Museum of the Kansas Museum of Natural History at the University of Kan-
sas (SEMK) contains the collections ofF. H. Snow and R. H. Beamer, collaborators of
W.T. Davis. The United States National Museum (USNM) contains the P. R. Uhler col-
lection. The collections of the American Museum of Natural History (AMNH) and the
Staten Island Institute of Arts and Sciences (SIIS) share material that was originally
the Davis collection. A significant transfer of type material took place between the
SIIS and the AMNH in 1945. A large number of the cicada types designated by W. T.
Davis were transferred to the AMNH after his death (Pallister 1946a). However, there
were questions raised about the location of the type material for several species (Pal-
lister 1946b). The SIIS collection still contains the bulk of the Davis Collection includ-
ing type material for many of the species he described.
The following list identifies all type material found in the collections. Designations
of type, holotype, co-type, allotype, lectotype, and paratype were made by the original
author or by a subsequent researcher. It includes specimens that were reported to be
on loan. Labels identifying the type status of individual insects are attached to the in-
dividual specimens. Type numbers in the CAS and USNM collections are listed if a
number was attached to the specimen. Current orthography for the species is used
with higher classification following Duffels & van der Laan (1985) and Duffels (1993).

Superfamily Cicadoidea
Family Cicadidae
Tribe Platypleurinae
Subtribe Platypleuraria
Platypleura machadoi Boulard 1972: 163.
CAS one male paratype.
















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March, 1999


Platypleura fenestrata Uhler 1862a: 282.
USNM type male.

Tribe Zammarini
Borencona aguadilla Davis 1928a: 31.
AMNH type male and allotype female.
Chinaria mexicana Davis 1934: 52.
AMNH type male, allotype female, and one female paratype.
Chinaria similis Davis 1942: 178.
AMNH type male and allotype female.
SIIS two male and one female paratypes.
Chinaria vivianae Ramos 1985: 63.
USNM type male, allotype female, and one male and three female paratypes.
Odopoea cariboea Uhler 1892a: 169.
USNM type female.
Tribe Thophini
Arenopsaltria nubivena (Walker 1858: 17).
USNM co-type male.
Uhleroides hispaniolae Davis 1939: 292.
SIIS one female paratype.
USNM type male, allotype female, and one female paratype.
Uhleroides maestra Davis 1939: 291.
AMNH type male and allotype female.
SIIS two male and one female paratypes.
Uhleroides samanae Davis 1939: 294.
USNM type male.
Tribe Thophini
Thopha saccata (Fabricius 1803: 34).
USNM co-type male.
Tribe Cyclochilini

Henicopsaltria eydouxii (Guerin M6neville 1838: 181).
USNM co-type female.
Psaltoda harrisii (Leach 1814: 89).
USNM co-type male.
Psaltoda moerens (Germar 1834: 67).
USNM co-type female.
Tribe Tibicenini

Cacama californica Davis 1919a: 75.
SIIS one male and one female paratypes.
USNM type male, allotype, and three male paratypes.

















Sanborn: Cicada Type Material


Cacama carbonaria Davis 1919a: 76.
SIIS type male.
Cacama crepitans (Van Duzee 1914: 45).
CAS lectotype male (CAS Type No. 2128) and five male paratypes.
Cacama valvata (Uhler 1888: 84).
USNM type male.
Cacama variegata Davis 1919a: 73.
AMNH type male and allotype female.
CAS one male paratype.
SEMK one male paratype.
SIIS 12 male and 11 female paratypes.
Cornuplura curvispinosa (Davis 1936: 102).
AMNH one male paratype.
SIIS two male paratypes.
Cornuplura nigroalbata (Davis 1936: 104).
AMNH type female.
Diceroprocta alacris var. campechensis Davis 1938: 297.
SEMK type male and five female paratypes.
Diceroprocta albomaculata Davis 1928b: 452.
USNM type male.
Diceroprocta apache (Davis 1921: 3).
AMNH type male and allotype female.
CAS four male and one female paratypes.
SIIS 99 male and 37 female paratypes.
USNM one male and one female paratypes.
Diceroprocta apache var. ochroleuca Davis 1942: 174.
AMNH type male and allotype female.
Diceroprocta arizona (Davis 1916a: 51).
AMNH type male, allotype female, and one male paratype.
SEMK three male paratypes.
SIIS one male paratype.
USNM one male paratype.
Diceroprocta averyi Davis 1941: 93.
AMNH type male and allotype female.
SIIS seven male paratypes.
Diceroprocta bequaerti (Davis 1917b: 210).
AMNH type male.
SIIS eight male paratypes.
Diceroprocta bibbyi Davis 1928b: 453.
AMNH type male and allotype female.
CAS one male paratype.
















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SEMK -one male paratype.
SIIS seven male and one female paratypes.
USNM one male paratype.
Diceroprocta bicolor Davis 1935a: 188.
AMNH type male and allotype female.
SIIS one male paratype.
Diceroprocta biconica var. obscurior Davis 1935a: 183.
SIIS type male, allotype female, and seven male paratypes.
Diceroprocta bilaqueata (Uhler 1903a: 7).
USNM type male.
Diceroprocta canescens Davis 1935a: 179.
AMNH type male and allotype female.
CAS one male paratype.
SEMK one male paratype.
SIIS 18 male and one female paratypes.
Diceroprocta cinctifera (Uhler 1892a: 156).
USNM type male.
Diceroprocta cinctifera var. limpia Davis 1932: 246.
AMNH type male.
SIIS four male paratypes.
USNM one male paratype.
Diceroprocta cinctifera var. viridicosta Davis 1930: 60.
SEMK type male, allotype female, and 22 male and one female paratypes.
SIIS six male and one female paratypes.
Diceroprocta cleavesi Davis 1930: 61.
SIIS type male.
Diceroprocta delicate var. aurantiaca Davis 1938: 300.
SEMK type male, allotype female, and 13 male and one female paratypes.
SIIS 28 male and seven female paratypes.
USNM one male paratype.
Diceroprocta eugraphica (Davis 1916a: 52).
AMNH type male and six male paratypes.
SEMK 89 male and 21 female paratypes.
SIIS three male and one female paratypes.
USNM female allotype and five male paratypes.
Diceroprocta fraterna Davis 1935a: 187.
AMNH type male.
SIIS five male paratypes.
Diceroprocta knight (Davis 1917b: 208).
AMNH type male and one male paratype.
SIIS three male paratypes.

















Sanborn: Cicada Type Material


Diceroprocta lucida Davis 1934: 44.
AMNH type male, allotype female, and one male and one female paratypes.
SEMK one female paratype.
CAS one male paratype.
SIIS six male and three female paratypes.
USNM one male and one female paratypes.
Diceroprocta marevagans Davis 1928b: 446.
AMNH type male.
Diceroprocta oculata Davis 1935a: 189.
AMNH type male.
SIIS two male paratypes.
Diceroprocta operculabrunnea Davis 1934: 45.
AMNH type male, allotype female, and one male paratype.
USNM one male paratype.
Diceroprocta pinosensis Davis 1935a: 184.
AMNH type male.
Diceroprocta pusilla Davis 1942: 177.
AMNH type male.
SIIS three male paratypes.
Diceroprocta reperta (Uhler 1892b: 177).
USNM -type female.
Diceroprocta reticularis (Uhler 1892a: 157).
USNM type male.
Diceroprocta semicincta (Davis 1925: 41).
AMNH type male, allotype female, and two male and one female paratypes.
CAS one male and one female paratype.
SEMK three male and seven female paratypes.
SIIS 12 male and five female paratypes.
USNM six male and seven female paratypes.
Diceroprocta semicincta var. nigrans Davis 1942: 175.
SEMK type male.
Diceroprocta sordidata (Uhler 1892b: 175).
USNM type male.
Diceroprocta swalei var. davisi Metcalf 1963: 210 nom. nov. pro D. swalei castanea
Davis 1916a: 49.
AMNH type male, allotype female, and one male paratype.
SEMK one male paratype.
SIIS 11 male and two female paratypes.
USNM one female paratype.
Diceroprocta tepicana Davis 1938: 298.
AMNH type male and allotype female.
















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March, 1999


SIIS one female paratype.
Diceroprocta texana (Davis 1916a: 54).
AMNH type male and one male paratype.
SEMK three male paratypes.
SIIS 13 male paratypes.
USNM one male paratype (USNM Paratype No. 42725).
Diceroprocta texana var. lata Davis 1941: 91.
AMNH type male.
Macrotristria angularis (Germar 1834: 68).
USNM co-type male.
Macrotristria extrema (Distant 1892: 56).
USNM co-type female.
Macrotristria hieroglyphicus (Kirkaldy 1909: 391 nom. nov. pro M. hieroglyphica God-
ing and Froggatt 1904: 581).
USNM co-type female.
Macrotristria internata (Walker 1850: 98).
USNM co-type female.
Tibicen bifidus (Davis 1916a: 47).
AMNH type male, allotype female, and one male paratype.
SEMK two male and two female paratypes.
SIIS two male and two female paratypes.
Tibicen bifidus var. simplex Davis 1941: 88.
AMNH type male.
Tibicen chiricahua Davis 1923: 8.
SEMK one male paratype.
SIIS one male and one female paratypes.
Tibicen chisosensis Davis 1934: 37.
AMNH type male and allotype female.
Tibicen chloromerus var. australis (Davis 1912b: 261).
SIIS type male and type female.
Tibicen cultriformis (Davis 1915d: 239).
AMNH allotype female.
USNM type male (USNM Type No. 200068).
Tibicen davisi (Smith & Grossbeck 1907: 121).
USNM two co-type females.
Tibicen davisi var. hardeni Davis 1918: 146.
SIIS type male and one female paratype.
Tibicen dealbatus (Davis 1915c: 162).
AMNH type male and one male paratype.
SIIS nine male paratypes.

















Sanborn: Cicada Type Material


Tibicen duryi Davis 1917b: 206.
AMNH type male, allotype female, and one female paratype.
SIIS one male and two female paratypes.
Tibicen fuscus Davis 1934: 43.
USNM type male.
Tibicen hidalgoensis Davis 1941: 88.
AMNH type male.
SIIS three male paratypes.
Tibicen inauditus Davis 1917b: 204.
AMNH type male and one male paratype.
SIIS one male paratype.
Tibicen latifasciatus (Davis 1915a: 8).
AMNH type male.
SIIS 21 male paratypes.
Tibicen linnei (Smith & Grossbeck 1907: 127).
AMNH two co-type males and one co-type female.
SIIS one co-type male and one co-type female.
USNM three co-type males (all USNM Co-type No. 42730) and one co-type fe-
male (USNM Co-type No. 42730).
Tibicen longioperculus Davis 1926: 179.
USNM type male.
Tibicen lyricen var. engelhardti (Davis 1910: 458).
AMNH type male and type female.
Tibicen lyricen var. virescens Davis 1935a: 175.
AMNH type male and one female paratype.
SEMK allotype female.
SIIS two male and one female paratypes.
Tibicen minor Davis 1934: 41.
AMNH type male.
SIIS two male paratypes.
USNM three male paratypes.
Tibicen parallelus Davis 1923: 10.
AMNH type male.
Tibicen paralleloides Davis 1934: 39.
AMNH type male.
SIIS type male.
Tibicen pruinosus var. fulvus Beamer 1924: 201.
SEMK type male, allotype female, and 12 male and three female paratypes.
SIIS one male paratype and one female paratypes.
Tibicen robinsonianus Davis 1922: 41.
AMNH type male and allotype female.
















Florida Entomologist 82(1)


March, 1999


CAS two male paratypes.
SIIS 22 male and one female paratypes.
USNM two male paratypes.
Tibicen sayi (Smith & Grossbeck 1907: 121).
AMNH two male and three female co-types.
SIIS three male co-types and three female co-types.
USNM seven co-type males and five co-type females.
Tibicen similaris (Smith and Grossbeck 1907: 125).
USNM type male.
Tibicen sublaqueatus (Uhler 1903a: 9).
USNM type male.
Tibicen sugdeni Davis 1941: 89.
AMNH type male.
SIIS four male paratypes.
Tibicen texanus Metcalf 1963: 319 nom. nov. pro T. tigrinus Davis 1927: 374.
AMNH type male, allotype female, and one male paratype.
CAS one female paratype.
SIIS 15 male and seven female paratypes.
USNM one male paratype.
Tibicen townsendi (Uhler 1905: 74).
USNM type male.
Tibicen walker var. pronotalis Davis 1938: 292.
AMNH allotype female and one male paratype.
SIIS five male and one female paratypes.
USNM type male.
Tibicen winnemanna (Davis 1912a: 3).
SIIS two male and one female paratypes.
USNM type male and type female.

Tribe Cryptotympanini
Subtribe Cryptotympanaria

Cryptotympana albolineata Hayashi 1987b: 8.
USNM type male and five male and three female paratypes.
Cryptotympana distant Hayashi 1987b: 25.
USNM one male and one female paratypes.
Cryptotympana moultoni Hayashi 1987a: 206.
USNM one male paratype.
Cryptotympana praeclara Hayashi 1987a: 182.
USNM one male paratype.
Cryptotympana sibuyana Hayashi 1987b: 9.
USNM type male and two female paratypes.

















Sanborn: Cicada Type Material


Cryptotympana socialis Hayashi 1987b: 13.
USNM type male.
Cryptotympana viridicostalis Hayashi 1987b: 14.
USNM one male and one female paratypes.

Tribe Fidicinini
Beameria venosa (Uhler 1888: 82).
USNM type male.
Beameria wheeleri Davis 1934: 49.
AMNH type male and one male paratype.
SEMK five male paratypes.
SIIS three male paratypes.
USNM one male paratype.
Fidicina compostela Davis 1934: 55.
AMNH type male and one male and one female paratypes.
SIIS four male and one female paratypes.
Fidicina panamaensis Davis 1939: 288.
SIIS one male paratype.
USNM type male, allotype female, and 11 male and three female paratypes.
Ollanta caicosensis Davis 1939: 295.
USNM type male and three male paratypes.
Proarna cocosensis Davis 1935a: 191.
AMNH allotype female.
CAS type male (CAS Type No. 4220).
SIIS one male paratype.
Proarna squamigera Uhler 1895: 56.
USNM type female (USNM Co-type No. 10194).
Tympanoterpes pusilla (Berg 1879: 140).
SIIS type male and type female.

Tribe Dundubiini

Subtribe Leptopsaltriaria
Calcagninus divaricatus Bliven 1964: 98.
CAS holotype male (CAS Type No. 13802), allotype female (no CAS type num-
ber), and 16 male paratypes.

Subtribe Cosmopsaltriaria
Aceropyga corynetus ungulatus Duffels 1988: 56.
USNM one male paratype.
Cosmopsaltria aurata Duffels 1983: 81.
AMNH twelve female paratypes.
CAS 25 female paratypes.
Cosmopsaltria giganta occidentalis Duffels 1983: 72.
















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AMNH holotype male and one male and one female paratypes.
Cosmopsaltria gracilis Duffels 1983: 56.
AMNH one male paratype.
Cosmopsaltria huonensis Duffels 1983: 62.
AMNH holotype male and one female paratype.
Cosmopsaltria signata Duffels 1983: 86.
AMNH 19 female paratypes.
Diceropyga gravesteini Duffels 1977: 90.
AMNH two male paratypes.
Tribe Cicadini
Cicada obtusa Uhler 1903a: 11.
USNM type male.
Cicadatra atra appendiculata Linnavuori 1962: 31.
AMNH type male.
Cicadatra erkowitensis Linnavuori 1973: 69.
AMNH type male.
Cicadatra flavicollis Horvath 1911: 114.
USNM type female.
Cicadatra naja Dlabola 1979: 238.
USNM nine male and one female paratypes.
Cicadatra ramanensis Linnavuori 1962: 33.
AMNH type male.
Leptosemia fuscolimbata (Schumacher 1915: 111).
SIIS two type females.
Neocicada chisos (Davis 1916a: 62).
AMNH allotype female and one male paratype.
SIIS one male paratype.
USNM type male.
Tribe Psithyristriini

Subtribe Pomponariaria

Pomponia zebra Bliven 1964: 99.
CAS type male and eight male paratypes.
Tribe Moganniini
Mogannia histrionica Uhler 1862a: 283.
USNM type male.

Family Tibicinidae
Subfamily Tibicininae
Tribe Dazini

















Sanborn: Cicada Type Material


Daza nayaritensis Davis 1934: 54.
AMNH type male, allotype female, and one male and one female paratypes.
SIIS nine male and 16 female paratypes.
USNM one male and one female paratypes.

Tribe Carinetini
Carineta martiniquensis Davis 1934: 54.
AMNH type male, allotype female, and one male paratype.
SIIS four male and one female paratypes.
Carineta socia Uhler 1875: 285.
USNM type male.
Herrera laticapitata Davis 1938: 304.
AMNH allotype female.
Herrera lugubrina var. compostelensis Davis 1938: 303.
AMNH type male, allotype female, and one male and one female paratypes.
SEMK four male and one female paratypes.
SIIS 56 male and 20 female paratypes.

Tribe Tibicinini

Clidophleps astigma Davis 1917a: 7.
SIIS one male paratype.
Clidophleps beameri Davis 1936: 116.
SEMK type male and three male paratypes.
SIIS three male paratypes.
Clidophleps blaisdelli (Uhler 1892a: 163).
USNM type male.
Clidophleps distant (Van Duzee 1914: 47).
CAS lectotype male (CAS Type No. 2131) and three male paratypes.
Clidophleps distant pallidus (Van Duzee 1914: 47).
CAS lectotype female (CAS Type No. 2133) and one male paratype.
Clidophleps distant truncatus (Van Duzee 1914: 47).
CAS lectotype male (CAS Type No. 2132) and one male paratype.
Clidophleps rotundifrons (Davis 1916b: 235).
SEMK -type female.
Clidophleps tenuis Davis 1927: 382.
AMNH type male and allotype female.
CAS one female paratype.
SIIS five male paratypes.
Clidophleps vagans Davis 1925: 48.
AMNH type male.
Clidophleps wright Davis 1926: 188.
AMNH type male and allotype female.
















Florida Entomologist 82(1)


March, 1999


Okanagana albibasalis Wymore 1934a: 167.
CAS holotype male (CAS Type No. 3906), allotype female (CAS Type No. 3906),
and 27 male and three female paratypes.
SEMK one male paratype.
Okanagana annulata Davis 1935b: 304.
AMNH type male and allotype female.
SIIS nine male and two female paratypes.
Okanagana arboraria Wymore 1934a: 166.
CAS holotype male (CAS Type No. 3903), allotype female (CAS Type No. 3903),
and 29 male paratypes.
SEMK one male paratype.
SIIS one male paratype.
Okanagana arboraria crocea Wymore 1934a: 167.
CAS holotype male (CAS Type No. 3905).
SEMK one male paratype.
Okanagana arctostaphylae Van Duzee 1915: 34.
CAS lectotype male (CAS Type No. 3024), allotype female (CAS Type No. 3025),
and 1 male paratype.
USNM co-type male (USNM Co-type No. 21358).
Okanagana aurantiaca Davis 1917a: 9.
SIIS one male paratype.
Okanagana aurora Davis 1936: 107.
AMNH type male and allotype female.
SIIS one female paratype.
Okanagana balli Davis 1919b: 211.
AMNH type male, allotype female, and one male paratype.
CAS one male paratype.
SIIS six male and one female paratypes.
USNM five male and one female paratypes.
Okanagana bella Davis 1919b: 198.
AMNH type male and allotype female.
SIIS 36 male and 14 female paratypes.
Okanagana canescens Van Duzee 1915: 37.
CAS lectotype male (CAS Type No. 3028), and allotype female (CAS Type No.
3029).
Okanagana cruentifera (Uhler 1892a: 161).
USNM type male.
Okanagana davisi Simons 1953: 191.
CAS holotype male (CAS Type No. 7155) and 4 male paratypes.
Okanagana ferrugomaculata Davis 1936: 110.
SIIS one male paratype.
USNM type male.

















Sanborn: Cicada Type Material


Okanagana formosa Davis 1926: 181.
AMNH type male and allotype female.
Okanagana fratercula Davis 1915b: 20.
AMNH type male.
Okanagana fumipennis Davis 1932: 251.
AMNH type male and allotype female.
SEMK one male and one female paratypes.
SIIS 13 male and four female paratypes.
Okanagana gibbera Davis 1927: 377.
AMNH type male, allotype female and one male and one female paratypes.
CAS two female paratypes.
SIIS four male and 11 female paratypes.
USNM one male paratype.
Okanagana hesperia (Uhler 1876: 342).
USNM type male.
Okanagana hirsuta Davis 1915b: 13.
AMNH type female.
Okanagana hirsuta var. catalina Davis 1936: 114.
SIIS type male.
USNM allotype female.
Okanagana lurida Davis 1919b: 192.
USNM type male.
Okanagana luteobasalis Davis 1935b: 302.
AMNH type male, allotype female, and one male and one female paratypes.
CAS two male and two female paratypes.
SEMK one male and one female paratypes.
SIIS 45 male and 12 female paratypes.
USNM one male and one female paratypes.
Okanagana magnifica Davis 1919b: 189.
AMNH type male, allotype female, and one male and one female paratypes.
SEMK 12 male and 11 female paratypes.
CAS one male and one female paratypes.
SIIS 62 male and 94 female paratypes.
USNM seven male and seven female paratypes.
Okanagana mariposa Davis 1915b: 12.
AMNH type male.
Okanagana mariposa var. oregonensis Davis 1939: 300.
AMNH type male and allotype female.
Okanagana napa Davis 1919b: 194.
USNM type male.
Okanagana nigriviridis Davis 1921: 9.
















Florida Entomologist 82(1)


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AMNH type male and allotype female.
SIIS one male paratype.
USNM one male paratype.
Okanagana nigrodorsata Davis 1923: 13.
AMNH type male and allotype female.
CAS three male paratypes.
SIIS two male paratypes.
USNM one male and two female paratypes.
Okanagana opacipennis Davis 1926: 186.
AMNH type female.
Okanagana oregona Davis 1916b: 233.
AMNH type male and one male paratype.
CAS allotype female (CAS Type No. 2130) and one male paratype.
SIIS three male and four female paratypes.
Okanagana ornata Van Duzee 1915: 33.
SIIS type male and co-type male.
Okanagana orithyia Bliven 1964: 97.
CAS holotype male (CAS Type No. 13803), allotype female (no CAS type num-
ber), and 11 male and seven female paratypes.
Okanagana ornata Van Duzee 1915: 33.
CAS lectotype female (CAS Type No. 3032).
Okanagana pallidula Davis 1917b: 213.
AMNH one male paratype.
CAS one male paratype.
SIIS type male and seven male paratypes.
Okanagana pallidula var. nigra Davis 1938: 308.
AMNH type male.
SIIS five male paratypes.
Okanagana pernix Bliven 1964: 93.
CAS holotype male (CAS Type No. 13804) and one male paratype.
Okanagana rhadine Bliven 1964: 96.
CAS holotype male (CAS Type No. 13805) and one female paratype.
Okanagana rubrocaudata Davis 1925: 46.
AMNH type male and allotype female.
Okanagana rubrovenosa Davis 1915b: 11.
AMNH type male.
CAS one male paratype.
Okanagana rubrovenosa var. rubida Davis 1936: 113.
SEMK type male, allotype female, and 23 male and two female paratypes.
SIIS three male paratypes.
Okanagana salicicola Bliven 1964: 92.

















Sanborn: Cicada Type Material


CAS holotype male (CAS Type No. 13806), allotype female (no CAS type num-
ber), and five male paratypes.
Okanagana schaefferi Davis 1915b: 11.
USNM type male.
Okanagana sequoia Bliven 1964: 94.
CAS holotype male (CAS Type No. 13807), allotype female (no CAS type num-
ber), and seven female paratypes.
Okanagana simulata Davis 1921: 12.
AMNH one male paratype.
CAS one male paratype.
SIIS type male and one male paratype.
Okanagana sperata Van Duzee 1935: 25.
CAS holotype male (CAS Type No. 4077).
Okanagana striatipes var. beameri Davis 1930: 68.
CAS two male paratypes.
SEMK type male, allotype female, and 26 male paratypes.
SIIS 18 male and one female paratypes.
USNM five male paratypes.
Okanagana sugdeni Davis 1938: 306.
AMNH allotype female.
SIIS one female paratype.
Okanagana synodica var. nigra Davis 1944: 220.
AMNH type male and allotype female.
Okanagana tanneri Davis 1930: 64.
AMNH one male paratype.
SEMK type male.
Okanagana triangulata Davis 1915b: 14.
AMNH type male.
Okanagana triangulata croncina Wymore 1934b: 174.
CAS holotype male (CAS Type No. 3915) and three male paratypes.
SEMK one male paratype.
Okanagana tristis Van Duzee 1915: 35.
CAS lectotype male (CAS Type No. 3026), allotype female (CAS Type No. 3027),
and one male paratype.
SIIS co-type female.
Okanagana tristis var. rubrobasalis Davis 1926: 184.
AMNH type male and allotype female.
Okanagana uncinata Van Duzee 1915: 41.
CAS lectotype male (CAS Type No. 2129).
Okanagana utahensis Davis 1919b: 216.
AMNH type male, allotype female, and one male and one female paratypes.
















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SIIS 22 male and four female paratypes.
USNM three male paratypes.
Okanagana vandykei Van Duzee 1915: 38.
CAS lectotype male (CAS Type No. 3030), allotype female (CAS Type No. 3031),
and two male paratypes.
Okanagana venusta Davis 1935b: 299.
AMNH type male and allotype female.
SIIS two male and three female paratypes.
Okanagana villosa Davis 1941: 95.
CAS type male (CAS Type No. 4980).
Okanagana viridis Davis 1918: 153.
AMNH type male.
Okanagana vocalis Bliven 1964: 90.
CAS holotype male (CAS Type No. 13808), and allotype female (CAS Type No.
13808).
Okanagana wymorei Davis 1935b: 305.
AMNH type male.
SIIS one male paratype.
Okanagana yakimaensis Davis 1939: 299.
SIIS one male paratype.
Okanagodes gracilis Davis 1919b: 221.
AMNH type male, allotype female, and one male paratype.
SIIS one male and two female paratypes.
USNM 13 male and four female paratypes.
Okanagodes gracilis var. pallida Davis 1932: 256.
AMNH type male and allotype female.
SEMK one male and one female paratypes.
SIIS 25 male and 15 female paratypes.
Okanagodes gracilis var. viridis Davis 1934: 57.
AMNH type male, allotype female, and one male paratype.
SIIS 103 male and 12 female paratypes.
USNM one male paratype.
Okanagodes terlingua Davis 1932: 257.
AMNH type male.
SIIS one male paratype.
Tibicinoides cupreosparsus (Uhler 1889: 43).
USNM type female.
Tibicinoides mercedita (Davis 1915b: 16).
AMNH type male and one male and one female paratypes.
CAS one male and one female paratypes.
SIIS two male and one female paratypes.

















Sanborn: Cicada Type Material


USNM one male paratype (USNM Paratype No. 42728).
Tibicinoides minute (Davis 1915b: 17).
AMNH type male and one male paratype.
CAS two male paratypes.
SIIS three male paratypes.
USNM one male paratype (USNM Paratype No. 42729).
Tribe Parnisini
Adeniana planiceps (Horvath 1917: 7).
USNM type male and allotype female.
Tribe Taphurini
Psallodia espinii Uhler 1903b: 18.
USNM -two co-type males.
Selymbria ahyetios Ramos & Wolda 1985: 178.
USNM type male, allotype female, and two female paratypes.
Selymbria pluvialis Ramos & Wolda 1985: 177.
USNM type male, allotype female, and two female paratypes.

Tribe Chlorocystini
Papuapsaltria brassi de Boer 1995b: 17.
AMNH one male and one female paratypes.
Papuapsaltria dioedes de Boer 1995b: 37.
AMNH one male and two female paratypes.
Papuapsaltria woodlarkensis de Boer 1995b: 39.
AMNH one male and two female paratypes.
Thaumastopsaltria lanceola de Boer 1992: 38.
AMNH three male and three female paratypes.
Thaumastopsaltria pneumatica de Boer 1992: 29.
AMNH two male and two female paratypes.

Tribe Gymnotympanini

Baeturia colossea de Boer 1994b: 144.
AMNH one male paratype.
Baeturia digitata Blote 1960: 78.
AMNH one male paratype.
Baeturia gressitti De Boer 1989: 26.
CAS one male and two female paratypes.
Baeturia guttulinervis Blote 1960: 67.
AMNH five female paratypes.
Baeturia marmorata Blote 1960: 67.
AMNH two male paratypes.
Baeturia nana (Jacobi 1903: 13).
















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SIIS type male and type female.
Baeturia nasuta Blote 1960: 61.
AMNH four male and 17 female paratypes.
Baeturia retracta de Boer 1994a: 170.
AMNH one female paratype.
Baeturia toxopeusi Blote 1960: 76.
AMNH three male and five female paratypes.
Baeturia versicolor de Boer 1994b: 150.
AMNH one male paratype.
Gymnotympana nenians Jacobi 1903: 14.
SIIS type female.
Gymnotympana stenocephalis de Boer 1995a: 58.
AMNH one male paratype.

Tribe Cicadettini
Cicadetta calliope var. floridensis (Davis 1920: 131).
AMNH type male and allotype female.
Cicadetta camerona (Davis 1920: 134).
USNM type male (USNM Type No. 42721), allotype female (USNM Type No.
42732), and one male paratype.
Cicadetta hackeri (Distant 1915: 51).
USNM co-type male.
Cicadetta kansa (Davis 1919c: 340).
AMNH type male and allotype female.
SIIS one male paratype.
Cicadetta pellosoma (Uhler 1862a: 283).
USNM type female.
Cicadetta rubea (Goding & Froggatt 1904: 651).
USNM type female (USNM Type No. 27411).
Cicadetta stradbrokensis (Distant 1915: 50).
USNM co-type male.
Cicadetta texana (Davis 1936: 105).
AMNH type male.
Cicadetta toowoombae (Distant 1915: 52).
USNM co-type male.
Cicadetta tympanistria Kirkaldy 1907: 18.
USNM type male.
Leptosalta radiator (Uhler 1897: 276).
USNM type male (USNM Type No. 3144) and six male co-types.
Pauropsalta mneme (Walker 1850: 181).
USNM co-type male.
















Sanborn: Cicada Type Material


Subfamily Platypediidae

Tribe Platypediini
Neoplatypedia ampliata (Van Duzee 1915: 29).
CAS lectotype male (CAS Type No. 3017) and one female paratype.
Neoplatypedia constricta Davis 1920: 123.
SIIS type male, allotype female, one male and one female paratypes.
USNM four male and one female paratypes.
Platypedia affinis Davis 1939: 301.
USNM type male.
Platypedia aperta Van Duzee 1915: 29.
CAS lectotype male (CAS Type No. 2134), allotype female (CAS Type No. 2135),
and 5 male paratypes.
Platypedia areolata (Uhler 1862b: 285).
USNM type female.
Platypedia australis Davis 1941: 95.
AMNH type female.
Platypedia balli Davis 1936: 118.
AMNH type male.
SIIS three male paratypes.
Platypedia barbata Davis 1920: 120.
SIIS type male and allotype female.
Platypedia bernardinoensis Davis 1932: 259.
AMNH type male and allotype female.
CAS 1 male paratype and 2 female paratypes.
SEMK one male and one female paratypes.
SIIS three male and five female paratypes.
Platypedia falcata Davis 1920: 113.
SIIS type male.
Platypedia intermedia Van Duzee 1915: 30.
CAS lectotype male (CAS Type No. 3018) and allotype female (CAS Type No.
3019).
Platypedia laticapitata Davis 1921: 14.
SIIS type male and allotype female.
Platypedia latipennis Davis 1921: 14.
AMNH type male.
Platypedia mariposa Davis 1935b: 307.
AMNH type male and allotype female.
SIIS one male paratype and two female paratypes.
Platypedia middlekauffi Simons 1953: 194.
CAS holotype male (CAS Type No. 7156), allotype female (no CAS type num-
ber), and one male and two female paratypes.
















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Platypedia minor Uhler 1888: 81.
USNM type male.
CAS two male and four female paratypes.
Platypedia mohavensis Davis 1920: 100.
AMNH type male and allotype female.
SIIS 23 male and 23 female paratypes.
USNM one male and one female paratypes.
Platypedia mohavensis var. rufescens Davis 1932: 258.
AMNH type male and allotype female.
CAS two male paratypes.
SEMK 54 male and 3 female paratypes.
SIIS 29 male and 37 female paratypes.
Platypedia putnami (Uhler 1877: 455).
USNM type male.
Platypedia putnami var. keddiensis Davis 1920: 108.
AMNH type male and allotype female.
SIIS one male paratype and one female paratype.
Platypedia putnami var. lutea Davis 1920: 106.
AMNH type male, two allotype females, and one male paratype.
CAS one male and one female paratypes.
USNM ten male and seven female paratypes.
Platypedia putnami var. occidentalis Davis 1920: 106.
CAS lectotype male (CAS Type No. 3020) and allotype female (CAS Type No.
3020).
SIIS one male and one female paratypes.
USNM one female paratype.
Platypedia rufipes Davis 1920: 101.
SIIS two female paratypes.
USNM type male, allotype female, and three female paratypes.
Platypedia rufipes var. angustipennis Davis 1932: 260.
AMNH type male and allotype female.
SEMK one male and one female paratypes.
SIIS 16 male and 18 female paratypes.
Platypedia scotti Davis 1935b: 308.
SIIS type male, allotype female, and three male and four female paratypes.
Platypedia sierra Wymore 1935: 143.
CAS holotype male (CAS Type No. 4104), allotype female (no CAS type num-
ber), and 19 male and 12 female paratypes.
SIIS two male paratypes.
Platypedia similis Davis 1920: 112.
AMNH type male and allotype female.
















Sanborn: Cicada Type Material


CAS one male paratype.
SIIS one male and one female paratypes.
Platypedia sylvesteri Simons 1953: 193.
CAS holotype male (CAS Type No. 7157).
Platypedia tomentosa Davis 1942: 182.
AMNH type male.
SIIS one male paratype.
Platypedia usingeri Simons 1953: 192.
CAS holotype male (CAS Type No. 7158), allotype female (no CAS type num-
ber), and two male paratypes.
Platypedia vanduzeei Davis 1920: 115.
CAS type male (CAS Type No. 3022) and allotype female (CAS Type No. 3023).
SIIS three male and one female paratypes.

I am designating the male and female co-types of Tibicen linnei (Smith & Gross-
beck) from West Farms in the AMNH collection as the lectotype and allotype of the
species. Both the male and female specimens of T lyricen var. engelhardti (Davis) in
the AMNH collection are labeled as type. Pallister (1946a) identified the male as the
type and the female as an allotype. The male, therefore, should be labeled as a lecto-
type and the female as the allotype for the species since Davis did not make a desig-
nation when he described the species (Davis 1910). I would also change the
designation of one of the two female specimens of Platypedia putnami var. lutea Davis
from allotype to paratype. Both were collected at the same location on the same date.
No change to the co-type designation of the T. sayi specimens is made because T. sayi
has been synonymized with T chloromerus (Walker) (Davis 1923).
This list adds type material from 93 species to the material identified in Hennes-
sey (1990) in the SIIS collection. I have identified an additional type specimen for
Okanagana simulata, a co-type of 0. ornata, and an allotype for Neoplatypedia con-
stricta. Hennessey (1990) listed the specimens of Tibicen latifasciatus and T winne-
manna as syntypes although they are labeled as paratypes. He states there is, and I
found, a T. latifasciatus specimen labeled as "type" in the AMNH. I found both a male
and female T winnemanna specimen labeled "type" in the USNM. The specimens in
the SIIS collection should be considered paratypes since Davis identified the speci-
mens as such when he described the species (labels attached to specimens and Davis
1912a). Davis (1912a) also designated the male specimen in the USNM as the type
and the female as the allotype for the species.
Pallister (1946b) identified 15 species whose types and/or allotypes were either de-
posited in the Davis Collection or probably in the Davis Collection. Of the specimens
he lists as part of the Davis collection I have been unable to find the allotypes of Tibi-
cen davisi var. hardeni and Okanagana simulata. I found the type of Clidophleps ro-
tundifrons (Davis 1916b: 235) in the SEMK. Of the specimens he lists as probably in
the Davis Collection I found the type and allotype of Tibicen winnemanna in the
USNM, the type and allotype of Okanagana rubrovenosa var. rubida at the SEMK,
and Platypedia scotti in the SIIS. The remaining specimens Pallister (1946b) identi-
fied are currently part of the SIIS collection.
I can answer some of the questions raised by Pallister (1946b) with this work. This
list adds type material from 48 Davis species to the material identified in Pallister
(1946a). I have identified additional types from 18 species, allotypes from 11 species,
and paratypes from 33 species. There are two specimens identified in Pallister (1946a)

















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March, 1999


as being transferred to the AMNH collection that I was unable to locate. I found the
type male of Okanagana tanneri not in the AMNH collection but in the SEMK. I have
not located the allotype of O. viridis.
There is one specimen of Okanagana striatipes (Haldeman) and one specimen of
Okanagana triangulata Davis that are labeled respectively as "Type" and "Holotype
male" in the general CAS collection. The specimen of 0. striatipes, however, was col-
lected in 1929 in Sedona, Arizona, while Haldeman (1852: 359) described the species
from a specimen collected on an expedition to the Great Salt Lake Valley of Utah. I
found the type of 0O. triangulata designated by a label in Davis' handwriting with a lo-
cation label matching the type location given in the original description (Davis 1915b:
14) in the AMNH. These two specimens in the CAS collection are therefore incorrectly
labeled as types.
Finally, there are 43 male and 17 female specimens identified as paratypes of
Platypedia plumbea Van Duzee (manuscript name) in the CAS collection. The descrip-
tion for P. plumbea has not been published.


ACKNOWLEDGMENTS

I would like to thank Edward Johnson at the SIIS, Randall Schuh and Donna En-
glund at the AMNH, Norman Penny at the CAS, Rob Brooks, George Byers and Karla
Segelquist at the SEMK, and Marc Epstien and Richard Froeschner at the USNM for
their assistance in accessing the collections. P. K. Phillips read an early draft of the
manuscript and assisted at the CAS. This work was supported financially in part
through an AMNH Collection Study Grant.


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HORVATH, G. 1911. Hemiptbres recolt6s par M. le Dr. W. Innes Bey en Egypte. Bull.
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HORVATH, G. 1917. Description d'une nouvelle Cigale d'Egypt. Bull. Soc. d'Entomol.
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JACOBI, A. 1903. Singzikaden von Ost-Neuguinea. Gesell. f. Naturf. Freunde Berlin
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KIRKALDY, G. W. 1907. Leaf-hoppers supplement. (Hemiptera.) Hawaiian Sugar
Planters' Assoc. Div. Entomol. Bull. 3: 1-186.

















Sanborn: Cicada Type Material


KIRKALDY, G. W. 1909. Hemiptera, old and new, No. 2. Canadian Entomol. 41: 388-
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LEACH, W. E. 1814. The zoological miscellany; being descriptions of new, or interest-
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60 Florida Entomologist 82(1) March, 1999

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4444444444444444444444444444444444444444444444444444















Florida Entomologist 82(1)


March, 1999


USE OF AN ACOUSTIC EMISSION DETECTOR FOR LOCATING
FORMOSAN SUBTERRANEAN TERMITE (ISOPTERA:
RHINOTERMITIDAE) FEEDING ACTIVITYWHEN INSTALLING
AND INSPECTING ABOVEGROUND TERMITE BAIT STATIONS
CONTAINING HEXAFLUMURON

THOMAS J. WEISSLING'AND ELLEN M. THOMS2
1University of Florida, Ft. Lauderdale Research & Education Center
3205 College Ave., Ft. Lauderdale, FL 33314

2Dow AgroSciences, Tampa, FL 33629

ABSTRACT

Soft- and hard-style aboveground bait stations containing 0.5% hexaflumuron in a
cellulose matrix (Recruit" AG), were installed indoors in two condominium buildings
and in one home. Bait stations were affixed to wood where Formosan subterranean
termites [FST (Coptotermes formosanus Shiraki)] were observed in foraging tubes
and subsurface galleries and adjacent to these locations where termite feeding was
detected using an acoustic emission detector (AED). Bait stations were inspected
monthly for the presence of termites, bait matrix consumption, and acoustic emission
(AE) counts on adjacent wood. Bait stations were added as needed. Six to fourteen
aboveground bait stations were installed in each structure at 3 or 6 placement sites.
Termites fed in a mean of 73.3% of the stations installed to consume a total of 42-149

















Weissling & Thoms:Above Ground Termite Baiting


g of bait matrix. FST at the home had ground contact and consumed 2- 3.5 times more
bait matrix than aerial infestations of FST at the condominiums. Acoustic emission
counts generally declined from pretreatment levels at all monitoring locations follow-
ing installation of the bait stations, with the exception of one condominium where AE
counts peaked 6 weeks following installation. At all sites, bait matrix consumption in
stations peaked during the first two months following installation of the aboveground
bait stations. Baiting eliminated detectable FST activity in all structures, as indi-
cated by lack of visual signs of termites and AED activity for at least two months.
Elimination of detectable activity followed two or more months of bait matrix con-
sumption by termites. Mean time to elimination of detectable activity for FST popu-
lations was 3.3 months (range 2-4 months).

Key Words: Acoustic Emission Detector, Formosan Subterranean Termite, Recruit
AG, Aboveground Termite Bait Station, hexaflumuron, Coptotermes formosanus


RESUME

Se instalaron dentro de una casa y dos condominios estaciones de cebo de dos esti-
los, suaves y duras, que contenman un 0.5% de hexaflumuron en una matriz de celulosa
(Recruit" AG). Las estaciones de cebo se fijaron sobre madera en la que se habian ob-
servado termitas subterraneas formosas ("FST", Coptotermes formosanus Shiraki) en
tuneles de alimentaci6n y en galerfas bajo la superficie, y tambien se colocaron adya-
centes a estos sitios donde alimentaci6n por termitas fu6 detectada utilizando un de-
tector de emisi6n acustica ("AED"). Las estaciones de cebo fueron inspeccionadas
mensualmente en busca de la presencia de termitas, consumo de la matriz del cebo, y
del numero de emisiones acusticas en madera adyacente. Se anadieron estaciones de
cebo cuando fue necesario. Se instalaron entire seis y catorce estaciones de cebo en cada
estructura en tres a seis lugares. Las termitas se alimentaron en un promedio del
73.3% de las estaciones de cebo instaladas, consumiendo un total de 42 a 149 g de la
matriz del cebo. La FST en la casa tuvo contact con el suelo y consumi6 de 2 a 3.5 mas
de la matriz del cebo que la FST en las infestaciones areas en los condominios. El nu-
mero de las emisiones acusticas generalmente decline de los niveles anteriores al tra-
tamiento en todas las localidadaes despues de instalar las estaciones de cebo, con la
excepci6n de un condominio en que el numero de las emisiones acusticas alcanz6 su
maximo seis semanas despues de la instalaci6n de las estaciones. En todos los sitios,
el consumo de la matriz del cebo alcanz6 su maximo en los dos primeros meses despues
de la instalaci6n de las estaciones de cebo arriba del piso. El uso del cebo elimin6 la ac-
tividad detectable de la FST en todas las estructuras, que fu6 indicado por la ausencia
de signos visuales de las termitas y de la actividadAED de por lo menos dos meses. Se
elimin6 la actividad detectable despues de dos meses mas de consumo de la matriz del
cebo por las termitas. El tiempo promedio para la eliminaci6n de toda actividad detec-
table de las poblaciones de FST fue de 3.3 meses (rango de 2 a 4 meses).




Field evaluations have documented the elimination of subterranean termite activ-
ity by application of RecruitR(DowElanco, Indianapolis, IN), a bait matrix containing
the chitin synthesis inhibitor hexaflumuron applied in-ground (Su et al. 1995, De-
Mark et al. 1995, Grace et al. 1996) and aboveground (Su et al. 1997). In these field
trials, mark-recapture methods were used to delineate termite foraging territories
and to estimate foraging population sizes before and after bait application. Untreated
wooden blocks in monitoring stations were used to measure wood consumption rates
by termite populations and to determine foraging activity (DeMark et al. 1995, Su et
al. 1995, Su et al. 1997).
















Florida Entomologist 82(1)


March, 1999


Delineating termite foraging territories and determining foraging activity of
aboveground termite populations can be very difficult. Mark-recapture methods are
disruptive and labor intensive. Subterranean termites foraging above-ground, com-
pared to those in soil, are more likely to abandon foraging areas after disturbance (N.-
Y. Su, Ft. Lauderdale Research and Education Center, University of Florida, personal
communication). Su et al. (1997) also documented that aboveground monitoring and
baiting stations, compared to in-ground stations, had a lower acceptance rate for bait
feeding by subterranean termites.
Acoustic emission (AE) detection has been successfully used to delineate above-
ground foraging territories and foraging activity for drywood termites (Scheffrahn et
al. 1997). Scheffrahn et al. (1993) verified that the acoustic emission detector (AED)
quantitatively records high frequency sound as wood is fed upon by termites. Schef-
frahn et al. (1997) then utilized acoustic emissions as a simple, non-disruptive method
to quantify drywood termite activity in structural wood before and after application
of localized chemical treatments.
The AED has not been previously used to determine subterranean termite activity
in structural wood as a measure of treatment efficacy. Therefore, the purpose of this
study was to evaluate the AED for delineating and measuring subterranean termite
feeding activity in structural wood as criteria to position installation of aboveground
bait stations containing hexaflumuron. In addition, the AED was evaluated as a tool
to measure subterranean termite foraging activity before and after consumption of
bait containing hexaflumuron.

MATERIALS AND METHODS

Termite Bait Stations.

Two types of aboveground bait stations, hard- and soft-style, containing 0.5%
hexaflumuron in a cellulose matrix (RecruitRAG) were evaluated. The hard-style sta-
tion consisted of a rigid, plastic container (10 cm by 10 cm by 4 cm), with a snap-on
cover containing 25 g of matrix (Fig. 1). The soft-style station consisted of a flexible
laminated foil pouch (15 cm by 15 cm) and contained 15 g of matrix (Fig. 2). On the
front of the soft-style station a cover flap sealed a removable inspection flap (7.5 cm
dia.) which covered the matrix. The back of the soft-style station had a removable ac-
cess flap (7 cm by 7 cm) and flexible adhesive used to affix the station to the target site.

Study Sites.

The aboveground bait stations were evaluated within three structures in Broward
County, Florida; two multi-story condominiums (Condol and Condo2) and one single-
story, single family residence (Home). All structures were infested with Formosan
subterranean termites (FST), Coptotermes formosanus Shiraki. Formosan subterra-
nean termite infestations in both condominium buildings were aerial; with no detect-
able ground contact as indicated by the presence of FST on the uppermost story of
each building (16th and 6th stories for Condo 1 and Condo2, respectively) with no activ-
ity on lower stories. The ability of FST to establish aerial infestations on flat rooftops
of high rise (4-14 storied) buildings similar to Condol and Condo2 in Hallandale, FL
was documented by Su et al. (1989). Formosan subterranean termites infesting the
home were in contact with the ground.
In Condol, FST had infested wooden baseboards and door trim in the hallway
(HALL) and the master bedrooms (BR) of two condominium units adjacent to the el-















Weissling & Thoms:Above Ground Termite Baiting


Station




















Cover

Bait
matrix

Fig. 1. Hard-style aboveground bait station containing hexaflumuron, showing
snap-on cover.


evator (EL) on the top (16th) floor (Fig. 3). Formosan subterranean termites were no
longer active in the master bedroom of one condominium when this study was initi-
ated, so aboveground bait stations were installed in the hallway and master bedroom
of the other condominium as indicated by Fig. 3. Infested baseboards and door trim
had been replaced 6 months prior to testing in the condominium unit of Condol.
These baseboards and trim were subsequently reinfested and were used as sites for
installation of aboveground bait stations. In Condo2, FST infested 2x4 wood framing
in an elevator machinery (EL)/storage room (SR) on the roof-top of this 6-story build-
ing (Fig. 4). No localized insecticide treatments had been applied to control these
aerial FST infestations.
In the Home, FST had infested doorframes in the recreation room (REC; Fig. 5).
The Home was on low-lying, continuously water-saturated soil adjacent to a canal.
The foundation was grade-beam construction comprised of an interior grid of load-
bearing footings poured beneath the slab. Two professional pest control companies
were treating the Home for subterranean termites. One company repeatedly applied
soil termiticides around the perimeter foundation and sub-slab. The water-saturated
soil and grade-beam construction may have prevented the establishment of a contin-
uous termiticide soil barrier beneath the structure, because FST continued to reinfest
the Home after termiticide applications. Drilling through the slab revealed that FST
had constructed extensive carton in the honey-combed chambers beneath the slab.















Florida Entomologist 82(1)


March, 1999


Cover
Flap


Removable
Bait flap
Bait
Matrix



Removable -
inspection
flap Flexible
adhesive
Front Back

Fig. 2. Soft-style aboveground bait station containing hexaflumuron, showing
cover flap and removable inspection flap on station front and removable flap and flex-
ible adhesive on station back.


The other pest control company installed 20 in-ground SentriconR System (Dow-
Elanco, Indianapolis, IN) stations on June 6, 1995. These stations, containing moni-
toring devices, were inspected monthly (x = 37.5 days) and 23 additional in-ground
stations were added during subsequent inspections. The installation and inspection
procedures followed label and manufacturer guidelines. Nonetheless, termites were
never detected in the in-ground monitoring stations. Water-saturated soil and exten-
sive termiticide perimeter treatments appeared to deter FST from foraging outside of
the foundation to sites where in-ground monitoring stations were installed.

AED Monitoring.
Infested wood members were located initially by the observations of building occu-
pants. Wood members were visually surveyed for signs of FST infestation and were
monitored using the AED (LocatorR) currently under development by DowElanco (In-
dianapolis, IN). The AED is a hand-held, battery-powered instrument designed to de-
tect low frequency ultrasonic sound emitted as wood fibers are damaged by insect
chewing. It consists of a main processor unit connected by two coaxial cables to two
identical sensors. The sensors are attached with putty to the wood members to be
monitored for termite activity.
One AED monitoring location was established adjacent to each bait placement
site, even if no termite feeding activity was detected using the AED. These locations
were monitored once for 60 seconds during each visit, and marked for monitoring on
subsequent visits. In areas where there was no visible termite activity (e.g. subsurface
galleries, feeding damage or emergence holes) but termite feeding activity was de-
tected using the AED, 2.3 mm diameter holes were drilled through the wood surface
to intersect termite galleries. These holes were located within a 50 cm2 area to fit
within the access area on the back of aboveground stations.














Weissling & Thoms:Above Ground Termite Baiting 65

S ____1 I-^-- I I- r----l|






HALL
k7C C BAIC]


C LR
BR BA K

I i



Fig. 3. Floor plan of top (16th) floor of Condol. Shaded areas are condominiums in-
fested by Formosan subterranean termites prior to application of aboveground bait
stations. Stars show placement sites for bait stations on baseboards in hallway
(HALL) and master bedroom (BR) and on door trim of closet (C) of master bedroom;
K = Kitchen, LR = Living room, BA = bathroom.


Installation of Aboveground Bait Stations.

Bait stations were affixed to wood where termites were visually observed in forag-
ing tubes and subsurface galleries and/or the AED detected termite feeding. Direc-
tions on the Recruit AG label (DowElanco 1996) were followed. For soft-style stations,
the adhesive backing and back access flap were removed and the exposed matrix was
aligned with termite gallery openings, exit holes, drill holes, or mud foraging tubes.
Soft-style stations were affixed to baseboards and door trim in Condol (13 June, 1996)
and the Home (28 October, 1996) using a staple gun or hot glue, in addition to the ad-
hesive on the station back. Hard-style stations were affixed to 2 x 4's in Condo2 (13
June, 1996) using screws and hot glue. All stations were located indoors. Thirty to
forty ml of water was added to the matrix of each station after installation, and sta-
tions were resealed with covers.

Inspection of Aboveground Bait Stations.
Bait stations were initially inspected every 2-6 weeks. After carefully removing
bait station covers, the presence or absence of termites in stations was noted, and the
percent of bait matrix consumed was estimated. In addition, AED readings were
taken during each inspection and if additional termite activity, indicated by damage,
mud tubes, and/or AE counts, was observed at new locations in wood, additional sta-
tions were installed at these sites. If less than 50% of the bait matrix was consumed
at the time of inspection, it was re-moistened with water and the station cover was put
back in place. If termites had consumed 50% or more of the matrix, a second station

















Florida Entomologist 82(1)


March, 1999


Fig. 4. Floor plan of roof top of six-story Condo2. Shaded area is elevator machin-
ery (EL)/storage room (SR) infested with Formosan subterranean termites prior to ap-
plication of aboveground bait stations. Stars show placement sites of bait stations on
2 x 4 wood framing in machinery/storage room.

was stacked on top the original, per Recruit AG label directions (DowElanco 1996).
The cover of the original station was removed prior to stacking and the matrix in the
station was moistened, if necessary. Subsequent hard-style stations were affixed us-
ing screws. Subsequent soft-style stations were affixed using a staple gun or hot glue,
in addition to the sealant on the back of the station. Two hard-style stations or three
soft-style stations were the maximum number of stations that could be stacked.
Thirty to forty ml of water was added to the matrix of each new station after installa-
tion. The percent of matrix consumed in stacked stations was based on visual inspec-
tion of the outer most layer at each bait placement site on subsequent inspections.
Additional consumption of matrix in underlying stations could not be determined un-
til all stations were removed at the end of the trial to calculate the total amount of ma-
trix consumed for each site. Trials were completed and stations were removed when
no termite activity, as determined by visual inspection, AE monitoring, and destruc-
tive sampling of previously infested wood, and sound structural wood, was observed
for two or more months following at least two months of bait consumption. Condol
and Condo2 were inspected approximately 7 and 19 months after completion of bait-
ing for signs of FST activity. Inspection consisted of visual survey, destructive sam-
pling, and monitoring using the AED. Home could not be inspected after bait removal
because of a change in ownership.


............. 11 .............. .... ... . . . . . . . .















Weissling & Thoms:Above Ground Termite Baiting 67



BA C









REC
BAt












Fig. 5. Floor plan of single-story Home. Stars show placement sites of bait stations
on door trim of the bathroom (BA) and closet (C)in the recreation room (REC).


RESULTS AND DISCUSSION

Six to fourteen aboveground bait stations were installed in each structure at 3 or
6 placement sites (Table 1). In all structures, bait stations were stacked, and at
Condo2 and the Home, new bait station placement sites were added after the initial
installation. Termites fed at 14 of 15 (93.3%) of the bait station placement sites, con-
suming an estimated total of 42-149 g matrix at each structure and feeding on a mean
of 73.3% of the bait stations installed in a structure. With the exception of one bait sta-
tion placement site at Condo2, bait stations that were not fed upon had been installed
late in the baiting program, when termite populations and subsequent feeding had al-
ready begun to decline. Su et al. (1997) reported that FST only fed in a mean of 17.6%
of the aboveground bait stations applied. To compensate for lack of termite activity in
stations, Su et al. (1997) installed more bait stations per structure compared to the
number installed in this study (X = 18 vs. 10, respectively).
Ground-based FST at the Home, compared to aerial FST infestations at Condol
and Condo2, consumed 2 and 3.5 times more matrix, respectively (Table 1). Su et al.
(1997) obtained similar results; LES, a ground-based FST colony, consumed 710.7 mg
of hexaflumuron compared to 747 mg consumed in this study by the Home FST colony.
Baiting eliminated all detectable signs of FST foraging activity at the three study
structures, as indicated by the absence of live termites in structures and bait stations,
no new termite damage or foraging tubes, and no AE counts. In addition, no dispersal
















Florida Entomologist 82(1)


Total g Matrix Consumed

Mean AE Count


Condo 1








, ,, S


I S 8


.200
t-
- 150
0
.100 0
LU
-50






350
300oo
250 C
200 o
150 mu
100
50
0





. 400

- 300
C
0
- 200 0
UJ
-100

0


Inspection Date


Fig. 6. Foraging activity (Total g Bait Matrix After Previous Inspection) and mean
AE counts ( SD) per minute per monitoring location before, during, and after appli-
cation of aboveground bait stations containing hexaflumuron; Condol, Condo2, and
Home, Broward County, Florida, 1996-1997.


March, 1999

















Weissling & Thons:Above Ground Termite Baiting


flights were observed nineteen months after baiting was completed at the condomin-
iums. Mean time to elimination of detectable activity was 3.3 months for FST popula-
tions in this study, compared to 5.8 months for FST populations by Su et al. (1997).
Comparisons of matrix consumption and AE counts for Condol, Condo2 and the
Home are shown in Fig. 6. Table 1 compared to Fig. 6 depicts six more g of matrix con-
sumed (149 vs. 143 g, respectively) for the Home, because additional consumption of
matrix in underlying stations could not be determined until all stations were removed
and carefully inspected at the end of the trial.
After installation of aboveground bait stations at all sites, AE counts generally de-
clined below pretreatment levels except at Condo2 where counts peaked 6 weeks fol-
lowing bait station installation. At all sites, matrix consumption and number of
termites in stations peaked during the first two months following installation of bait
stations. No termites were found in stations during the last two inspections at Condo 1
and Condo2, and during the last inspection at the Home. The absence of termites co-
incided with the absence of AE counts at monitoring locations. In addition, when the
doorframes of the Home were replaced in March, 1997, only dead FST were found in
the damaged wood.
In this study, a higher percentage of aboveground bait stations containing hexaflu-
muron were fed upon by FST and fewer bait stations were required to eliminate de-
tectable termite activity more rapidly than previously documented by Su et al. (1997).
Recruit AG has a different cellulose matrix (nonparticulate) compared to the particu-
late matrix containing hexaflumuron evaluated by Su et al. (1997). Laboratory and
field testing (E. King, pers. comm.) have documented that subterranean termites pref-
erentially consume the matrix of Recruit AG compared to that evaluated by Su et al.
(1997). Therefore, the improved matrix of Recruit AG could explain part of the im-
proved performance observed in this study. Nonetheless, researchers installing Re-
cruit AG without using an AED to determine where termites were feeding in wood
had a lower percentage of aboveground stations fed upon compared to that in this
study. A similar study was conducted at six structures in Texas using the same above-
ground termite bait stations and protocol, with the exception that an AED was not
available to determine bait placement sites (T. Atkinson, pers. comm.). In that study,
subterranean termites fed upon only 38 out of 86 placement sites (44%) at the six
structures. Subterranean termite activity was, however, eliminated at these six sites
in Texas. These studies suggest that aboveground bait stations containing hexaflumu-
ron can eliminate detectable subterranean termite activity in structures. These stud-
ies also indicate that the AED has the potential to be used to determine optimal bait
placement in order to reduce the number of stations needed to achieve control.
Because of the potential for bait avoidance, absence of termites in baiting stations,
by itself, is not considered to be a reliable method of measuring the efficacy of toxic
baits on subterranean termites (Su & Scheffrahn 1996). Rather, foraging activity is
considered to be a more reliable evaluation variable (Su & Scheffrahn 1996). The ab-
sence of termites in bait stations in this study coincided with the lack of AE counts in
previously-infested wood. In addition, no termite activity could be detected with the
AED in sound wood adjacent to previously infested wood indicating that termites had
not moved away from baits to other locations. These observations suggest that the
AED can be used to reliably evaluate the efficacy of control procedures for subterra-
nean termites in aboveground wood by detecting foraging activity.
This study confirmed the findings of previous research (Su et al. 1997) that baits
containing hexaflumuron applied in aboveground stations eliminate detectable activ-
ity of aerial and ground-based populations of subterranean termites within struc-
tures. Baits such as Recruit AG will be useful where conditions prevent structural
































TABLE 1. SUMMARY OF BAITING USING ABOVEGROUND BAIT STATIONS CONTAINING HEXAFLUMURON FOR FORMOSAN SUBTERRANEAN TERMITES IN-
FESTING THREE STRUCTURES IN BROWARD COUNTY, FLORIDA.

Total No. Total No. g dry wt Months to No
Station Stations Used Matrix mg AI Termite Activity
Study Site Station Type FST Colony Placement Sites (Fed Upon) Consumed Consumed After Instal

Condol Soft-style Aerial 3 6(3) 42 210 2
Condo2 Hard-style Aerial 6 10(7) 75 375 4
Home Soft-style Ground 6 14(12) 149 747 4 00

















Weissling & Thoms:Above Ground Termite Baiting


infestations of subterranean termites from foraging to in-ground bait stations (e.g.
aerial infestations, previous soil termiticide treatment, water-saturated soil), or at lo-
cations where soil access for installation of in-ground bait stations is limited by as-
phalt or concrete pavement.

ACKNOWLEDGMENTS

The authors appreciate the contributions of Rudolf Scheffrahn who reviewed and
improved the manuscript. This manuscript is Florida Agricultural Experiment Sta-
tion Journal Series R-06026.

REFERENCES CITED

DEMARK, J. J., E. P. BENSON, P. A. ZUNGOLI, AND B. M. KARD. 1995. Evaluation of
hexaflumuron for termite control in the southeast U.S. Down To Earth 50(1):
20-26.
DOWELANCO. 1996. Recruit AG label, for Experimental Use Only, DowElanco, India-
napolis, IN, 11 pp.
GRACE, J. K., C. H. M. TOME, T. G. SHELTON, R. J. OSHIRO, AND J. R. YATES III. 1996.
Baiting studies and consideration with Coptotermes formosanus (Isoptera: Rhi-
notermitidae) in Hawaii. Sociobiol. 28: 511-520.
SCHEFFRAHN, R. H., N. -Y. SU, AND P. BUSEY. 1997. Laboratory and field evaluations
of selected chemical treatments for control of drywood termites (Isoptera: Kal-
otermitidae). J. Econ. Entomol. 90: 492-502.
SCHEFFRAHN, R. H., W. P. ROBBINS, P. BUSEY, N.-Y. SU, AND R. K. MUELLER. 1993.
Evaluation of a novel, hand-held acoustic emissions detector to monitor ter-
mites (Isoptera: Kalotermitidae, Rhinotermitidae) in wood. J. Econ. Entomol.
86: 1720-1729.
Su, N.Y., AND R. H. SCHEFFRAHN. 1996. A review of the evaluation criteria for bait-
toxicant efficacy against field colonies of subterranean termites (Isoptera). So-
ciobiol. 28: 521-530.
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) in high-rise buildings. J. Econ. Entomol. 82: 1643-1645.
Su, N.-Y., E. M. THOMS, P. M. BAN, AND R. H. SCHEFFRAHN. 1995. A monitoring/bait-
ing station to detect and eliminate foraging populations of subterranean ter-
mites (Isoptera: Rhinotermitidae) near structures. J. Econ. Entomol. 88: 932-
936.
Su, N.-Y., P. M. BAN, AND R. H. SCHEFFRAHN. 1997. Remedial baiting with hexaflu-
muron in above-ground stations to control structure-infesting populations of
the Formosan subterranean termite (Isoptera: Rhinotermitidae). J. Econ. En-
tomol. 90: 809-817.
















Florida Entomologist 82(1)


March, 1999


THE DISTRIBUTIONS OF THE CARIBBEAN FRUIT FLY,
ANASTREPHA SUSPENSE (TEPHRITIDAE) AND ITS
PARASITOIDS (HYMENOPTERA: BRACONIDAE) WITHIN THE
CANOPIES OF HOST TREES

JOHN SIVINSKI', MARTIN ALUJA2 AND TIM HOLLER3
IUSDA-ARS, CMAVE, P.O. Box 14565, Gainesville, Florida, USA

2Instituto de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico

3USDA-APHIS-PPQ, Caribbean Fruit Fly Station, 1911 SW 34th St.
Gainesville, Florida, USA


ABSTRACT

In the area of LaBelle, Florida (Hendry County), the Caribbean fruit fly, Anas-
trepha suspense (Loew), is commonly attacked by three braconid parasitoids, Doryc-
tobracon areolatus (Szepligeti), Diachasmimorpha longicaudata (Ashmead), and
Utetes anastrephae (Viereck). Fruits from fifteen individual trees of four species, Suri-
nam Cherry (Eugenia uniflora L.), Cattley guava (Psidium cattleianum Sabine),
guava (P. guajava L.), and loquat (Eriobotryajaponica [Thunb.]), were systematically
sampled in order to determine the distribution of A. suspense and its parasitoids
within the tree's canopies. Fruits infested byA. suspense were lighter than infested
ones in P. guajava. This may be due to the presence of the larvae. There was no evi-
dence that A. suspense preferred to oviposit in fruits at particular heights above
ground or distances from canopy edges. Fruits containing larvae parasitized by U.
anastrephae were significantly lighter than those containing larvae parasitized by ei-
ther D. areolatus or D. longicaudata, and it was not present in P. guajava, the species
with the heaviest fruits. There were no differences, either overall or within host tree
species, among the heights above ground or distances from canopy edges of fruits con-
taining larvae parasitized by any of the three braconids. Niche similarity inD. areola-
tus and D. longicaudata may be due to the absence of a shared evolutionary history.
Both are recent introductions to Florida, but while D. areolatus is a neotropical spe-
cies, D. longicaudata is from the Indo-Philippine region. Thus, there has been little
opportunity for divergence. Knowledge of the distributions within tree canopies of the
pest-fly, and of its natural enemies, may lead to improvements in its biological control.

Key words: Diachasmimorpha longicaudata, Doryctobracon areolatus, Utetes anas-
trephae, biological control, parasitoids, fruit flies

RESUME

En el area de LaBelle, Florida (condado de Hendry), la mosca del Caribe,Anastre-
pha suspense (Loew), es atacada comunmente por tres parasitoides brac6nidos, Do-
ryctobracon areolatus (Szepligeti), Diachasmimorpha longicaudata (Ashmead), y
Utetes anastrephae (Viereck). Para determinar la distribuci6n deA. suspense y sus pa-
rasitoides dentro de las copas de los arboles se colectaron sistematicamente frutas de
quince arboles individuals de cuatro species, Eugenia uniflora L., Psidium cattlei-
anum Sabine, P. guajava L., y Eriobotrya japonica [Thunb.]. Se document que las
guayabas infestadas porA. suspense eran mas ligeras que las que no estaban infesta-
das. Esto puede ser debido a la presencia de las larvas. No hubo evidencia de que A.
suspense prefiriera ovipositar en frutas en particulares alturas o distancias de los bor-
des de las copas dentro de los arboles. Se not6 que las frutas con larvas parasitadas

















Sivinski et al.: Parasitoid distributions 73

por U. anastrephae eran significativamente mas ligeras que aquellas con larvas para-
sitadas ya sea por D. areolatus o D. longicaudata; U. anastrephae no estuvo present
en P. guajava, la especie con los frutos mas pesados. No se detectaron diferencias, ya
sea en forma general o entire species de hospederos, entire las alturas y las distancias
de los bordes de las copas dentro de los arboles, y entire las frutas con larvas parasita-
das por cualquiera de los tres brac6nidos. Es possible que la similitud de los nichos eco-
16gicos de D. areolatus y de D. longicaudata sea debida a la ausencia de una historic
compartida de evoluci6n. Las dos species fueron introducidas recientemente a Flo-
rida, y mientras que D. areolatus es una especie neotropical, D. iongicaudata es de la
region Indo-Filipina, asi que la oportunidad para una divergencia no ha sido mayor.
El conocimiento sobre la distribuci6n dentro de las copas de los arboles de esta mosca
plaga, y tambien la de sus enemigos naturales, nos puede ser util para mejorar su con-
trol biol6gico.






The Caribbean fruit fly,Anastrepha suspense (Loew), was sporadically captured in
Florida throughout the first half of the century (Baranowski et al. 1993). These immi-
grants from the Greater Antilles failed to establish until 1965, when a population in-
troduced into the Miami area expanded over the peninsular portion of the state
(Weems 1966).Anastrepha suspense attacks over 90 species of fruits in Florida (Norr-
bom & Kim 1988) and has restricted the movement of citrus fruits into California,
Texas, and Japan. To protect citrus exports, the Florida Department of Agriculture
has organized a system of fly-free zones based on negative-trapping (Simpson 1993).
Shortly after the establishment ofA. suspense, parasitoids were introduced for its
control. The first of these was the opiine braconid Doryctobracon areolatus (Szepligeti)
(= Parachasma cereus) (Baranowski & Swanson 1970, 1971). Doryctobracon areolatus
is a widespread species, ranging from Mexico to Argentina (Wharton & Marsh 1978).
It is a parasitoid of several pestiferous Anastrepha species, including the Mexican
fruit fly, A. ludens (Loew), and finds its hosts in both native and commercial, exotic
fruits (e.g., Aluja et al. 1990, Hernandez-Ortiz et al. 1994). While initially abundant
in south Florida, it was soon replaced at the original site of introduction by a second
opiine braconid, Diachasmimorpha longicaudata (Ashmead) (Baranowski et al.
1993). This Indo-Philippine species was originally recovered from Bactrocera spp. but
proved to attack a number of tephritids and was widely disseminated (Clausen 1978).
At present, D. areolatus is common only in the northern portion of the Caribbean fruit
fly's range, while D. longicaudata predominates in the south (Eitam 1998). There is a
relatively narrow region, just south-west of Lake Okechobee, where both species are
abundant. A third opiine species, Utetes anastrephae (Viereck), is native to Florida,
where it historically parasitized Anastrepha spp. present in the extreme southern
portion of the state and the Florida Keys (Baranowski et al. 1993). The spread of A.
suspense has allowed it to increase its own range into the middle of the peninsula
(Eitam 1998). In addition to Florida, U. anastrephae occurs throughout Latin Amer-
ica, reaching as far south as Argentina (Wharton and Marsh 1978). It has a noticeably
shorter ovipositor than either D. areolatus or D. longicaudata.
Augmented releases of D. longicaudata have been employed to help maintain Flor-
ida's fly-free zones (see Sivinski et al. 1996). It would be useful if the environments fa-
vored by D. areolatus, D. longicadata and U. anastrephae were better understood. In this
way, augmentative releases of one or all species could be tailored to particular conditions.
Should there be "gaps" in the foraging behaviors of the three species, new natural ene-
















Florida Entomologist 82(1)


March, 1999


mies that flourish in these specific microhabitats might be identified and imported. One
context in which the species might differ is their distributions within host tree canopies.
Characteristics of the canopy structure affect, or are suspected to affect, the forag-
ing of a number of parasitic Hymenoptera species (see Godfray 1994 and cit.). Differ-
ences within the canopy in light, heat, humidity, and risk of predation are among the
factors that could select for both nonrandom distributions of hosts and specialized for-
aging tactics by natural enemies. For example, the red scale, Aonidiella aurantii
(Mask.), is 27 fold more vulnerable to attack by aphelinid parasitoids when feeding on
the periphery of citrus trees rather than in their interior (Murdoch et al. 1989). Closer
to the present topic, the opiine braconid Psyttalia (= Opius) concolor (Szepligeti) ap-
pears to parasitize a greater proportion of tephritids in the upper halves of large
Greek olive trees (Kapatos et al. 1977). Darby (1933) suggested that the opiine Doryc-
tobracon crawfordi (Viereck) is more apt to attackA. ludens larvae in mangos than in
sweet limes because the relatively open foliage of the lime provides less refuge from
heat and low humidities. In southern Mexico, D. areolatus is more likely than, U.
anastrephae to parasitize Anastrepha obliqua (Macquart) in fruits near the margins
of certain tree canopies (Sivinski et al. 1997).
In this study, we examined the spatial distributions ofA. suspense and its three lo-
cal parasitoids in the canopies of four species of host trees. We also examined the size,
as estimated by weights, of fruits that escaped infestation byA. suspense, that con-
tainedA. suspense but not parasitoids, and those that contained bothA. suspense and
its various parasitoids. Information on the temporal distribution of D. areolatus and
D. longicaudata in Florida can be found in Sivinski et al. (1998).


METHODS

Fruits were collected from preferred host tree species in the LaBelle area, Hendry
Co., Florida from 1992 to1994. These included six Surinam cherry trees (Eugenia uni-
flora L.), three loquat (Eriobotrya japonica (Thunb.), one guava (Psidium guajava L.),
and five Cattley guava (Psidium cattleianum Sabine) (Table 1). Fruit sampling was
done systematically, i.e., fruits from every part of the trees were collected every week.
Fruits were obtained by gently shaking the branch next to their petioles. If they fell,
they were assumed to have nearly completed their tenure on the tree. Thus, the fly
larvae inside the fruits had been exposed to attack by parasitoids for a "typical" period
in that location. Prior to shaking fruits, distance from the ground and distance from
the edge of the foliage was determined, as was compass direction measured in de-
grees. After removal, fruits were weighed and then kept individually for one week on
damp vermiculite at ca. 27C. and ambient humidity. Fruit weight provides a useful
estimate of fruit size (Sivinski 1991). Pupae were held for an additional four weeks,
after which the emerging adults were identified. Voucher specimens are in the collec-
tion of J. Sivinski at USDA-CMAVE, Gainesville, FL.
Statistical comparisons of the weights and canopy-locations of fruits containingA.
suspense and its various parasitoids were made by multivariate ANOVA, with the
various means distinguished through the Waller-Duncan k-ratio t-test ("proc GLM",
SAS Inst. Raleigh, N.C. 27605). In the initial analysis of all fruits from all trees of all
species, the error term used to derive the F value was taken from the interaction of all
trees of all species and the species of insect present in the fruits. The relationships
among compass direction, infestation, and parasitism were examined by dividing the
canopy into six sections, each encompassing 60 degrees. Comparisons of the distribu-
tions ofA. suspense and its various parasitoids among these sections were by two-way
crosstabulation chi-square tests (SAS Inst. Raleigh, N.C. 27605).
















Sivinski et al.: Parasitoid distributions 75

TABLE 1. THE NUMBERS OF THE VARIOUS SPECIES OF HOST TREES AND THE NUMBERS OF
FRUITS SAMPLED. ALSO INCLUDED ARE THE LEVELS OF INFESTATION (A. SUS-
PENSA PUPAE RECOVERED / G OF PICKED FRUIT HELD IN THE LABORATORY)
AND THE MEAN PERCENT PARASITISM (PARASITOIDS / PARASITOIDS + ADULTA.
SUSPENSE) BY THE DIFFERENT PARASITOIDS IN THE VARIOUS HOST TREES
(UTETES ANASTREPHAE = UA; DORYCTOBRACONAREOLATUS = DA; DIACHAS-
MIMORPHA LONGICAUDATA = DL). FRUITS OF THE VARIOUS HOST TREE SPE-
CIES ARE CHARACTERIZED BY MEAN SIZE (WEIGHT IN GRAMS) AND LOCATION
WITHIN THE CANOPIES (MEAN HEIGHT FROM THE GROUND [CM] AND DISTANCE
FROM THE EDGE OF THE CANOPY [CM]). IN ALL APPROPRIATE INSTANCES STAN-
DARD ERRORS ARE ENCLOSED IN PARENTHESES FOLLOWING THE MEANS.

Surinam cherry loquat Cattley guava guava

N of trees 6 3 5 1
N of fruits 1336 336 656 315
pupae / gram 0.49 (0.01) 0.08 (0.005) 0.96 (0.02) 0.061 (0.006)
Ua parasitism 2 (0.4) 0.3 (0.3) 0.4 (0.001) 0
Da parasitism 26 (1.0) 16 (3.0) 18 (1.0) 0.5 (0.3)
Dl parasitism 14 (1.0) 0 13 (1.0) 0.4 (0.3)
weight (grams) 3.68 (0.04) 15.3 (0.21) 7.41 (0.12) 47.17 (0.10)
height (cm) 195.9 (2.68) 289.9 (6.44) 185.0 (3.01) 213.7 (4.35)
distance (cm) 12.10 (0.45) 24.05 (3.11) 8.75 (0.28) 10.51 (0.54)


RESULTS

Anastrepha suspense was the only tephritid reared from the samples, and D. are-
olatus and D. longicaudata were the predominate parasitoids. U. anastrephae was
present in relatively small numbers and was not recovered at all from guava, the spe-
cies with the largest fruits. In Table 1 the various species of trees are described in
terms of numbers of trees sampled, numbers of fruits sampled, mean infestations (A.
suspense larvae / g), mean percent parasitism (parasitoids / parasitoids +A. suspense)
by the three species of parasitoids, mean fruit weights, mean heights of fruits above
ground, and the mean distances of fruits from the edge of the canopies. There were
significant differences among the mean heights and weights of the various species of
host fruits, however, there were no differences in the mean distances fruits hung from
the various canopy edges (Table 2).
Overall, fruits that containedA. suspense larvae and uninfested fruits did not dif-
fer in locations within the canopies; i.e., they were similar in terms of heights above
ground and distances from canopy edges (Table 2). There were significant differences
in the weights of infested and uninfested fruits, and there was a significant interac-
tion between species of host fruit and the pattern of weight difference in infested and
uninfested fruits (Table 2). When host species were examined individually, only
guava, whose infested fruits were lighter, displayed any significant weight difference
(Table 3).
Fruits from all trees of all species of hosts that contained unparasitized larvae and
those containing larvae parasitized by U. anastrephae, D. areolatus, and D. longicau-
data did not differ in heights above ground or distances from the edges of canopies
(Table 4). However, fruits with unparasitized larvae were significantly heavier than
those containing parasitized larvae, and those fruits containing larvae parasitized by

















Florida Entomologist 82(1)


March, 1999


TABLE 2. ABOVE-THE WEIGHTS (G) AND WITHIN CANOPY LOCATIONS (HEIGHTS ABOVE
GROUND [CM] AND DISTANCES FROM CANOPY MARGINS [CM]) OF FRUITS FROM
ALL TREES OF ALL HOST SPECIES THAT WERE EITHER INFESTED WITH A. SUS-
PENSA OR WERE UNINFESTED. STANDARD ERRORS ARE IN PARENTHESES FOL-
LOWING MEANS. BELOW-THE RESULT OF A MULIVARIATE ANOVA THAT
COMPARES THE MEAN WEIGHTS, HEIGHTS ABOVE GROUND, AND DISTANCES
FROM CANOPY MARGINS OF A) FRUITS THAT WERE EITHER INFESTED WITH A.
SUSPENSE OR UNINFESTED, B) WEIGHTS, HEIGHTS, AND DISTANCES OF FRUITS
OF DIFFERENT SPECIES, AND C) THE INTERACTION BETWEEN PATTERNS OF IN-
FESTATION REGARDING FRUIT WEIGHT, HEIGHT, AND DISTANCE AND HOST SPE-
CIES.


Uninfested n = 516


Infested n = 2152


Weight
Height
Distance


Infestation


21.96 (1.04)
217.23 (4.37)
15.15 (1.42)


Weight
Height
Distance


Host Species Weight
Height
Distance
Infest* Host Weight
Height
Distance


8.61 (0.24)
204.74 (2.15)
12.03 (0.48)


F

14.12
.026
0.06
1329.8
6.3
0.69
27.17
1.06
3.03


p
0.0037*
0.62
0.81
0.0001*
0.0009*
0.58
0.0001*
0.41
0.08


TABLE 3. A COMPARISON OF THE WEIGHTS (G) OF FRUITS THAT DO AND DO NOT CONTAIN
LARVAE OF A. SUSPENSE IN ALL OF THE VARIOUS HOST TREE SPECIES. STAN-
DARD ERRORS ARE IN PARENTHESES FOLLOWING THE MEANS, AND SAMPLE
SIZES FOLLOW STANDARD ERRORS.

Uninfested Infested df F p

Surinam 3.7 (0.11) 3.7 (0.04) 1 0.61 0.47
Cherry n = 195 n = 1168
Loquat 15.3 (0.32) 15.3 (0.28) 1 0.06 0.83
n = 130 n = 205
Cattley 7.3 (0.87) 7.4 (0.12) 1 4.47 0.13
Guava n = 24 n = 631
Guava 50.5 (1.61) 43.4 (1.12) 1 12.61 0.0004*
n = 167 n = 148

















Sivinski et al.: Parasitoid distributions 77

TABLE 4. ABOVE- THE WEIGHTS (G) AND WITHIN CANOPY LOCATIONS (HEIGHTS ABOVE
GROUND [CM] AND DISTANCES FROM CANOPY MARGINS [CM]) OF FRUITS FROM
ALL TREES OF ALL HOST SPECIES THAT EITHER CONTAINED UNPARASITIZED
LARVAE OF A. SUSPENSE (= AS), OR CONTAINED LARVAE PARASITIZED BY THE
VARIOUS PARASITOIDS (UTETES ANASTREPHAE = UA, DORYCTOBRACON ARE-
OLATUS = DA, DIACHASMIMORPHA LONGICAUDATA = DL). STANDARD ERRORS
IN PARENTHESES FOLLOW MEANS, AND MEANS SHARING THE LETTER FOLLOW-
ING THE STANDARD ERROR ARE NOT SIGNIFICANTLY DIFFERENT. BELOW-THE
RESULT OF A MULIVARIATE ANOVA THAT COMPARES THE MEAN WEIGHTS,
HEIGHTS, AND DISTANCES FROM CANOPY MARGINS OF A) FRUITS THAT CONTAIN
UNPARASITIZED A. SUSPENSE LARVAE AND THOSE THAT CONTAIN LARVAE PAR-
ASITIZED BY THE VARIOUS PARASITOIDS, B) WEIGHTS, HEIGHTS, AND DIS-
TANCES OF FRUITS OF THE VARIOUS HOST SPECIES THAT CONTAIN A.
SUSPENSE LARVAE, AND C) THE INTERACTION BETWEEN THE PATTERNS OF
PARASITISM REGARDING FRUIT WEIGHT, HEIGHT, AND DISTANCE AND HOST
TREE SPECIES.

As Ua Da Dl

Weight 11.14 (0.45) a 4.78 (0.50) c 6.0 (0.19) b 6.3 (0.2) b
Height 211.3 (3.3) a 218.4 (19.4) a 204.2 (4.1) a 181.9 (4.0) a
Distance 11.78 (0.71) a 16.76 (6.62) a 11.62 (0.86) a 11.40 (0.71) a

df F p

Parasitoid Weight 3 2.56 0.091
Height 3 0.55 0.66
Distance 3 2.59 0.08
Host Species Weight 3 58.0 0.0001*
Height 3 0.40 0.76
Distance 3 0.01 0.99
Parasitoid* Host Weight 7 2.37 0.07
Height 7 0.53 0.80
Distance 7 2.35 0.07

'A Waller-Duncan K-ratio t-test found some means to be significantly different.


U. anastrephae were significantly lighter than those containing larvae parasitized by
D. areolatus or D. longicaudata (Table 4). In the later two parasitoid species, there
were significant negative correlations between the mean weights of fruits from the
various host species and the mean percent parasitism of the fruit fly larvae they con-
tained (Fig. 1). However, within species there were no relationships between fruit
weights and percent parasitism (Table 5).
There were no significant differences between the compass locations of infested
and uninfested fruits, either overall or in any of the individual host fruit species. Nei-
ther were there any differences among the compass location distributions of the var-
ious parasitoid species.

DISCUSSION
Anastrepha suspense larvae were evenly distributed within the canopies of host
trees. Guava fruits containing fly larvae were lighter in weight than those that did

















Florida Entomologist 82(1)


March, 1999


30

2 0 U. anastrephae r-0.65, p=0.35
25 D. areolatus r=-0.97, p=0.03
] D. longicaudatus r=-0.99, p=0.007

20





10 -
5




0



-5 ....--
0 1 10 20 30 40 50
Sc Cg L Mean fruit weight (g) G


Fig. 1. The relationships between the mean fruit weights of the various host trees
(Sc = Surinam cherry, L = loquat, Cg = Cattley guava, G = guava) and the percent par-
asitism ofA. suspense larvae by the various parasitoids (Utetes anastrephae, Dorycto-
bracon areolatus, and Diachasmimorpha longicaudata). The drawings of abdomens
near the different lines illustrate the relative lengths of the ovipositor in the various
parasitoid species. The data point for D. longicaudata on loquat was omitted from the
analysis because this parasitoid is rarely abundant during the winter months when
loquat is fruiting.



not, and similarly, various fruits in southern Mexico, containing differentAnastrepha
species, are often lighter than uninfested fruits (Sivinski et al. 1997). Some tephritids
oviposit in small, green fruits and increase the rate of maturation and, by unknown
means, decrease the size of the fruits they occupy, e.g., the papaya fruit fly, Toxotry-
pana curvicauda Gerstaeker (Landolt 1985). Since female A. suspense often oviposit
in immature guava fruits (e.g. Burk 1983), lighter infested fruits may be due to the ac-
tion of the larvae. Alternative explanations are that females prefer to lay eggs, or lar-
vae are better able to survive, in smaller fruits.
Doryctobracon areolatus parasitized A. suspense larvae in all the fruit species
sampled, while D. longicadata was obtained from larvae in all but loquat fruits. Utetes
anastrephae was absent in the large fruits of guava, and common only in the smallest
fruits, those of Surinam cherry. In southern Mexico, U. anastrephae is also restricted
to smaller fruits, and its limited foraging range is presumably reflected in its shorter
ovipositor (Sivinski et al. 1997). In general, and in D. areolatus and D. longicaudata
in the present study, mean percent parasitism is significantly higher in host tree spe-
cies with smaller fruits (e.g., Sivinski et al 1997). This may be due to a preference for
smaller fruits (Sivinski 1991), or the greater access parasitoids have to fly larvae liv-

















Sivinski et al.: Parasitoid distributions 79

TABLE 5. COMPARISONS OF THE WEIGHTS (G) OF FRUITS THAT CONTAINED UNPARASIT-
IZED A. SUSPENSE LARVAE AND THOSE THAT CONTAINED LARVAE PARASITIZED
BY THE VARIOUS PARASITOIDS (UTETES ANASTREPHAE = UA, DORYCTOBRA-
CON AREOLATUS = DA, AND DIACHASMIMORPHA LONGICAUDATA = DA). STAN-
DARD ERRORS ARE IN PARENTHESES FOLLOWING MEANS AND SAMPLE SIZES
FOLLOW STANDARD ERRORS. MEANS SHARING A LETTER ARE NOT SIGNIFI-
CANTLY DIFFERENT. THE SINGLE SPECIMEN OF UA RECOVERED FROM LOQUAT
WAS DISCARDED FROM THE CALCULATIONS.

As Ua Da Dl F p

Surinam 3.67 (0.07) 4.24 (0.38) 3.62 (0.09) 4.04 (0.09) 0.41 0.75
Cherry 538 a 18 a 316 a 189 a
Loquat 15.6 (0.33) 15.2 (0.63) 41.86 0.11
141 a 33 a
Cattley 6.75 (0.20) 9.6 (0.20) 7.46 (0.19) 8.08 (0.23) 2.05 0.19
Guava 196 a 2 a 219 a 201 a
Guava 43.3 (1.22) 38.8 (1.56) 41.0 (1.45) 0.24 0.79
132 a 4 a 2 a



ing in the shallow pulp of smaller fruits. The absence ofD. longicaudata from loquats
may be due to the tree's late winter fruiting season. Diachasmimorpha longicaudata
is relatively rare throughout the winter months in the LaBelle area of Florida, regard-
less of the host tree (Sivinski et al. 1998).
There were no distinct microhabitat preferences among the three parasitoid spe-
cies, either in terms of heights within the canopies or distances from the canopy edges.
Fruits containing larvae parasitized by U. anastrephae were farther from the edges of
the host tree canopies than those containing larvae parasitized by D. areolatus or D.
longicaudata, albeit not significantly so. However, in southern Mexico U. anastrephae
often parasitized larvae in fruits that were deeper, on average, in the canopies of host
trees than those attacked by D. areolatus (Sivinski et al. 1997)
One difference in the foraging of D. areolatus and D. longicaudata, not addressed
in the present study, is the latter species' propensity to forage over fallen fruits (e.g.,
Purrcell et al., 1996, see however, Sivinski et al. 1998). In a Mexican study, D. areola-
tus attacked larvae in fruits on the ground under tree canopies, but to a lesser extant
than D. longicaudata (Sivinski et al. 1997).
It is possible that the substantial overlap in the spatial niches of D. areolatus and
D. longicaudata in tree canopies is due to a lack of shared history over evolutionary
time. Competition between the two long-ovipositor species may not yet have led to se-
lection for divergence in morphology and behavior. There is similar overlap between
the within-canopy distributions of D. longicaudata introduced into Mexico and yet an-
other neotropical Doryctobracon species, D. crawfordi (Veireck) (Sivinski et al 1997).
Competition between D. areolatus and D. longicaudata is also the most likely rea-
son for their generally distinct geographical distributions in Florida. It appears that
only the ability ofD. areolatus to forage in habitats with relatively low host diversity
and /or survive in climates characterized by low winter temperatures has allowed it
to resist displacement by D. longicaudata (Eitam 1998; Sivinski et al. 1998). Interest-
ingly, U. anastrephae with its short ovipositor and specialization on only smaller
fruits, is present in the Florida ranges of both long ovipositor species.

















Florida Entomologist 82(1)


March, 1999


There are other examples of the coexistence of native and introduced parasitoid
species with different ovipositor lengths, and the displacement of native species by in-
troduced species with similar ovipositor lengths; e.g., both occurred among the Ich-
neumonidae attacking the Swaine jack pine sawfly, Neodiprion swainei Middleton
(Price 1972). In this instance, ovipositor length is correlated to the host stage at-
tacked. Short ovipositor species lay eggs in exposed larvae while long ovipositor spe-
cies reach down to pupae buried in leaf litter. The significance of ovipositor length in
braconid fruit fly parasitoids, and how it influences interspecific competition, is less
obvious. As far as is known all three of the braconids examined in Florida attack fruit
fly larvae, not eggs or pupae, inside fruits (Eitam 1998; J. Sivinski and M. Aluja, per-
sonal observation). Since longer ovipositors appear to be able to reach both shallow
and deeply placed hosts, i.e., long ovipositor species are present in the entire size
range of host fruits, the adaptive significance of U. anastrephae's short ovipositor re-
mains to be determined.


ACKNOWLEDGMENT

Avi Eitam, Michael Hennessy, and Regina Sugayama suggested numerous im-
provements to an earlier draft of the manuscript. Debbie Smith and Janet Voelker col-
lected and watched over a great deal of fruit. Nothing could have been done without
them. Gina Posey was a great help in the production of the tables and Valerie Malcolm
prepared the manuscript. Victor Chew was of inestimable assistance in analyzing the
data. Kevina Vulinec drew the parasitoid abdomens included in the figure. This study
was performed in part with funds from USDA-OICD International Collaborative Re-
search Project 60-43YK-0016.


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ALUJA, M., J. GUILLEN, P. LIEDO, M. CABRERA, E. RIos, G. DE LA ROSA, H. CELE-
DONIO, AND D. MOTA. 1990. Fruit infesting tephritids (Dipt.: Tephritidae) and
associated parasites in Chiapis, Mexico. Entomophaga 35: 39-38.
BARANOWSKI, R., AND R. W. SWANSON. 1970. Introduction of Parachasma (= Opius)
cereus (Hymenoptera: Braconidae) into Florida as a parasite ofAnastrepha sus-
pensa (Diptera: Tephritidae). Florida Entomol. 53: 161-162.
BARANOWSKI, R., AND R. W. SWANSON. 1971. The utilization of Parachasma cereus
(Hymenoptera: Braconidae) as a means of suppressing Anastrepha suspense
populations. Proc. Tall Timbers Conf. Biol. Cont. by Habit. Manag. No. 3: 249-
252.
BARANOWSKI, R., H. GLENN, AND J. SIVINSKI. 1993. Biological control of the Caribbean
fruit fly, Anastrepha suspense (Loew). Florida Entomol. 76: 245-250.
BURK, T. 1983. Behavioral ecology of mating in Caribbean fruit flies, Anastrepha sus-
pensa (Loew). Florida Entomol. 66: 330-344.
CLAUSEN, C. P. 1978. Introduced Parasites and Predators of Arthropod Pests and
Weeds: a World Review. USDA-ARS Agr. Handbook No. 480.
DARBY, H. H. 1933. Insects and microclimate. Nature. 131: 839.
EITAM, A. 1998. Biogeography of Braconid Parasitoids of the Caribbean Fruit Fly,
Anastrepha suspense (Loew) (Diptera: Tephritidae), in Florida. Dissertation,
University of Florida.
GODFRAY, H. C. J. 1994. Parasitoids Behavioral Ecology and Evolutionary Ecology.
Princeton Univ. Press, Princeton N. J.
HERNANDEZ-ORTIZ, V., R. PERE-ALONSO, AND R. WHARTON. 1994. Native parasitoids
associated with the genus Anastrepha (Diptera: Tephritidae) in Los Tuxtlas,
Veracruz, Mexico. Entomophaga 39: 171-178.

















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KAPATOS, E., B. S. FLETCHER, S. PAPPAS, AND Y. LAUDEHO. 1977. The release of Opius
concolor and 0. concolor var. siculus (Hym.: Braconidae) against the spring
generation of Dacus oleae (Diptera: Tephritidae) on Corfu. Entomophaga 22:
265-270.
LANDOLT, P. 1985. Papaya fruit fly eggs and larvae (Diptera: Tephritidae) in field-col-
lected papaya fruit. Florida Entomol. 68: 354-356.
MURDOCH, W. W., R. F. LUCK, S. J. WALDE, J. D. REEVE, AND D. S. YU. 1989. A refuge
for red scale under control by Aphytis: structural aspects. Ecology 70:1707-
1714.
NORRBOM, A. L., AND K. C. KIM. 1988. A list of the reported host plants of the species
ofAnastrepha (Diptera: Tephritidae). USDA-APHIS-PPQ Pub No. 81-52.
PRICE, P. W. 1972. Parasitoids utilizing the same host: adaptive nature of differences
in size and form. Ecology 53: 190-195.
PURCELL, M., C. JACKSON, J. LONG, AND M. BATCHELOR. 1994. Influence of guava rip-
ening on parasitism of the oriental fruit fly, Bactrocera dorsalis (Hendel)
(Diptera: Tephritidae), by Diachasmimorpha longicaudata (Ashmead) (Hy-
menoptera: Braconidae) and other parasitoids. Biol. Cont. 4: 396-403.
SIMPSON, S. E. 1993. Caribbean fruit fly-free zone certification protocol in Florida
(Diptera: Tephritidae). Florida Entomol. 76: 228-232.
SIVINSKI, J. 1991. The influence of host fruit morphology on parasitism rates in the
Caribbean fruit fly (Anastrepha suspense [Loew]). Entomophaga 36: 447-455.
SIVINSKI, J., C. 0. CALKINS, R. BARANOWSKI, D. HARRIS, J. BROMBILA, J. DIAZ,
R. BURNS, T. HOLLER, AND G. DODSON. 1996. Suppression of a Caribbean fruit
fly (Anastrepha suspense (Loew): Tephritidae) population through augmented
releases of the parasitoid Diachasmimorpha longicaudata (Ashmead) (Bra-
conidae). Biol. Cont. 6: 177-185.
SIVINSKI, J., M. ALUJA, AND M. LOPEZ. 1997. The spatial and temporal distributions
of parasitoids (Hymenoptera) of MexicanAnastrepha species in the canopies of
host trees. An. Entomol. Soc. America 90: 604-618.
SIVINSKI, J. M. ALUJA, T. HOLLER, AND A. EITAM. 1998. Phenological comparison of
two braconid parasitoids of the Caribbean fruit fly (Diptera: Tephritidae). En-
viron. Entomol. 27: 360-365.
WEEMS, H. 1966. The Caribbean fruit fly in Florida. Proc. Florida Sta. Hort. Soc. 79:
401-405.
WHARTON, R., AND P. M. MARSH. 1978. New World opiine (Hymenoptera: Braconidae)
parasitic on Tephritidae (Diptera). J. Washington Acad. Sci. 68: 147-167.
















Florida Entomologist 82(1)


March, 1999


SEMI-ARTIFICIAL REARING OF THE LARVAE OF
ANASTREPHA OBLIQUA (DIPTERA: TEPHRITIDAE) IN
MANAUS, AMAZONAS-BRAZIL

LILIAN A. SALDANHA1 AND NELITON M. SILVA2
1Department of Entomology, University of Nebraska-Lincoln
Lincoln, NE 68583-0816

2Universidade do Amazonas, Manaus, Amazonas, Brazil.

ABSTRACT

Larvae of the West Indian fruit fly, Anastrepha obliqua (Macquart), were success-
fully reared on a meridic diet of: agar, wheat flour, sucrose, nipagin, antibiotic, sodium
benzoate, yeast, water and different amounts from 10 to 50 percent of powdered de-
hydrated araca-boi (Eugenia stipitata) (Myrtaceae). For each evaluated diet, 100 lar-
vae were reared. Anastrepha obliqua F, were successfully reared. The meridic diets
with 20 to 40 percent of powdered dehydrated araca-boi were observed to provide the
optimum growth medium for larval development. Under these conditions, 38% of the
larvae completed development at 26 + 1C, 88 + 6% RH, and a photoperiod of 12:12
(L:D). The average life cycle was 43.3 days (eggs, 4 days, larvae, 26.3 days, pupae, 13
days). These results provide a basis for rearingA. obliqua in semi-artificial conditions.
However more studies are needed to better understand pupal humidity requirements.

Key Words: West Indian fruit fly, larvae rearing, meridic diet, nutrition, diet selection,
Insecta

RESUME

As moscas-das-frutas sao conhecidas mundialmente como pragas pelos danos que
causam a fruticultura. Para a realizagao de testes de m6todos de control 6 necessario
o conhecimento da biologia da praga. Para tanto foi desenvolvido o present trabalho
de criacao de larvas de Anastrepha obliqua em condigoes semi-artificiais utilizando
dieta semi-artificial constituida de: dieta basica (agar, amido, sucrose, 1Avedo, ben-
zoato de s6dio, antibi6tico, nipagin e agua destilada) acrescida de quantidades cres-
cente de fruto de araca-boi (Eugenia stipitata) (Myrtacea) liofilizado, e polpa fresca
nas percentagens que variaram de 10 a 50. As dietas ap6s preparadas eram armaze-
nadas em tubos de ensaio, nos quais colocava-se uma larva rec6m eclodida. 0 conjunto
tubo-larva era mantido em temperature ambiente (26C + 1C) umidade de 88 + 6% e
fotoperiodo de 12 horas, at6 atingir o period de pupa. Esta era entao, ap6s 24 horas,
pesada e transferida para tubo de ensaio contend areia peneirada autoclavada e le-
vemente umidecida e observada at6 emergencia do adulto. Foram criadas para cada
dieta testada 100 larvas, num total de 1100 larvas. As dietas contend 20, 30 e 40%
de fruto liofilizado proporcionaram um desenvolvimento adequado das larvas, com
tempo medio de desenvolvimento de 27.7 + 3.68 dias; 24.7 + 3.6 dias e 24.3 + 4.5 dias,
respectivamente. Pela primeira vez, para a esp6cie em estudo, obteve-se F, reproduti-
vamente viavel, com tempo de desenvolvimento de 43.3 dias (ovos = 4; larvas = 26.3;
pupas = 13 dias). A utilizagao de fruto de araca-boi liofilizado revelou-se adequada
para confeccao de dieta de criacao de larvas deAnastrepha obliqua. Entretanto maio-
res investigacoes sobre a umidade do substrato para pupa devem ser realizadas.

















Saldanha and Silva: Rearing of West Indian fruit fly


Throughout the world, various fruit flies are economically important pests. The fe-
males oviposit into the fruits and the larvae develop inside, feeding on pulp or seed.
Many commercial fruits are affected making them inappropriate for commercializa-
tion and industrial purposes. The life cycle is completed in the soil, where the larvae
pupate and the imagos emerge and start a new cycle (Morgante 1991).
According to Foote et al. (1993), the West Indian fruit flyAnastrepha obliqua (Mac-
quart) occurs throughout the Greater and Lesser Antilles.Anastrepha obliqua is also
found in more parts of world than any of the other Anastrepha species. It has been re-
corded in Central America (Belize, Costa Rica, Guatemala, Honduras, Mexico, Nica-
ragua, Panama), North America (USA), and South America (Argentina, Brazil,
Colombia, Ecuador, Guyana, Peru, Suriname, Venezuela) (White and Elson-Harris
1992).
Thirty species of fruit flies occur in the Brazilian Amazon (Silva et al. 1996).Anas-
trepha obliqua is the most important economic fruit fly in the State of Amazon, dam-
aging a variety of fruits (Silva 1993, Silva et al. 1996, Zucchi et al. 1996, Ronchi-Teles
and Silva 1996). The amount of fruit production is increasing in this area. With the in-
crease of host plants, there is a possibility that the number ofA. obliqua will increase,
as will their potential to damage the crops. Damage affects fruit exportation, in-
creases the costs per acre to produce the crops and overall decreases fruit yield.
Zucoloto et al. (1979), Polloni (1981) and Jorge (1987) cultured A. obliqua in the
laboratory on an artificial diet. Continuous rearing ofA. obliqua has come up against
two major obstacles. The first obstacle concerns the poor nutritional balance of the lar-
val diet. The second obstacle is the unusual behavior of the emerged adults and their
inability to reproduce.
This study was conducted to evaluate a variety of meridic diets for rearing larvae
ofA. obliqua. Meridic diet can be defined as a diet composed of defined chemicals but
can contain crude components (Reinecke 1985). If found, the optimum medium can be
used to study A. obliqua biology, including the basic life cycle and rearing require-
ments. This knowledge is important in developing management strategies for this
pest.


MATERIALS AND METHODS

Meridic diets were evaluated. Diets were prepared following standard laboratory
procedures, however the diet was not autoclaved. Microbial development was inhib-
ited by the addition of anti-fungal (nipagin) and antibiotic ingredients. The amounts
of antibiotic and nipagin were determined by trial and error. The trials were based on
the nutritional requirement ranges for insects determined by Parra (1979). The
amounts are important to provide the correct balance of the microbial flora in the me-
dium.
The basic composition of the diet was similar to the diet developed by Zucoloto et
al. (1979) and evaluated by Jorge (1987). Adjustments were made to their original
protocol in an attempt to reduce microbial contamination and to increase the nutri-
tional value of the diet. The diets used in this experiment were prepared using the ba-
sic ingredients (Table 1) as well as additional ingredients, which varied with each
trial. The varied ingredients were powdered dehydrated araga-boi, varying in concen-
tration from 10 to 50 percent, and araga-boi pulp. Araga-boi fruit was used, because it
is the primary host ofA. obliqua. The use of dehydrated fruit was proposed to improve
larval development by reducing contamination caused by bacterial and fungal devel-
opment, presented when fresh pulp was used, and also prevent first instar larvae
drowning. Thus, eleven different diets were evaluated for rearing maggots (Table 2).

















Florida Entomologist 82(1)


March, 1999


TABLE 1. INGREDIENTS FOR ARTIFICIAL DIET FOR ANASTREPHA OBLIQUA.

Basic diet

Ingredients Quantity

Agar 4.0 g
Yeast 9.0 g
Wheat flour 8.0 g
Sucrose 12.0 g
Sodium benzoate 0.05 g
Antibiotic 0.05 g
Nipagin 2.0 ml
Water 220 ml



The chemical ingredients were obtained from a local store. The powdered dehydrated
araca-boi was prepared at INPA-Food production laboratory by freezing a mixture of
the fruit at each stage of maturity. The concentration of the nutrients in the fruit was
determined following standard procedures of food analysis. Araca-boi, dry matter has
8-10.75% protein, 5-6.5% fiber, 69.98-71.63% carbohydrate and contains the following
micronutrients: phosphate, potassium, calcium, and magnesium. Furthermore, one
hundred grams of fresh pulp contain 7.75mg of Vitamin A, 9.85mg of Vitamin B,, and
7.68mg of Vitamin C (Pinedo et al. 1981, Cavalcante 1991).
The medium was prepared by mixing water, sucrose, wheat flour, and agar in a
beaker. The mixture was heated for 5 minutes. The solution was placed in a vertical
hood to cool for 3 minutes. The remaining ingredients were added, stirring constantly
with a glass rod. The pH was checking using pH paper and adjusted to pH 3 using hy-


TABLE 2. PERFORMANCE OF ANASTREPHA OBLIQUA LARVAE IN DIFFERENT DIETS.

Diet number Description Performance'

1 Basic diet (Table 1) Poor
2 Basic diet + 10% dehydrated fruit Poor
3 Basic diet + 20% dehydrated fruit Moderate
4 Basic diet + 30% dehydrated fruit Moderate
5 Basic diet + 40% dehydrated fruit Moderate
6 Basic diet + 50% dehydrated fruit Low
7 Basic diet + 10% araca-boi pulp Poor
8 Basic diet + 20% araca-boi pulp Poor
9 Basic diet + 30% araga-boi pulp Poor
10 Basic diet + 40% araca-boi pulp Poor
11 Basic diet + 50% araca-boi pulp Poor

'The diet performance was evaluated based on the larval development.

















Saldanha and Silva: Rearing of West Indian fruit fly


drochloric acid. The mixture was poured into glass tubes (9.8 x 1.2 cm), stopped with
a cotton ball and placed at a 300 angle to prevent the maggots from drowning. The
tubes were stored at 4C until use.
The larvae used in this experiment were the offspring of adults reared in labora-
tory conditions of 26 + 1C, 88 + 6% RH and a photoperiod of 12:12(L:D). All of these
adults were fed with two different solutions consisting of: 1) sugar cane hydrolysate
(50ml) mixed with brown sugar (100g); 2) honey (10%). Two cotton balls on Petri
dishes were placed in each cage. Each cotton ball was saturated with one of the above
mentioned solutions.
Initially, artificial gel-forming compounds were placed in the cages to stimulate
the females to lay eggs. However, these females did not use the structures provided,
instead they laid eggs throughout the cage. The cages consisted of a wire skeleton cov-
ered with nylon fabric. Using a camel hair paint brush, the eggs were collected every
morning and afternoon. The eggs were placed into Petri plates with 1% sodium ben-
zoate solution until the larvae hatched. The larvae were washed in a new 1% sodium
benzoate solution to reduce the microbial contamination on their bodies. One larva
was placed in a glass tube with meridic diet. The tubes were closed with cotton and
placed at 26 + 1C, 88 + 6% RH and a photoperiod of 12:12 (L:D). The tubes were ob-
served daily thereafter and pupation was recorded. Pupae were removed after 24
hours, their weights were recorded, and they were placed on moistened sand in glass
tubes covered with cotton to prevent desiccation. They were incubated at 26 + 1C, 88
+ 6%RH and a photoperiod of 12:12(L:D). Adults that emerged were confined in cages,
fed as described previously, and observed daily. Their reproductive capability was
evaluated.


RESULTS AND DISCUSSION

Only the diets with powdered dehydrated araca-boi allowed the development of
larvae to the adult stage (Table 2: Diets 3, 4, 5, and 6). The percentages of larvae that
completed development and the developmental periods of West Indian fruit fly mag-
gots with various amounts of powdered dehydrated araca-boi are shown in Table 3.
The results of this experiment demonstrated that the amount of powdered dehy-
drated araca-boi improved the rearing conditions and the reproductive capability of
the emerged adults, because it increased the nutritional value of the diet. The use of
powdered dehydrated araca-boi is equally important because it can be used, not only
during the fruit season, but also throughout the year. Additionally, the use of dehy-
drated fruit reduced microbial development and death of first instar larvae, when
compared to the use of fresh pulp.
The percentage of pupation and the rate of growth in this experiment were consis-
tent with previous results observed by Jorge (1987). However, emerged adults in this
experiment showed reproductive capability when compared to the results given by
Jorge (1987). Another result that differs from Jorge (1987) is the number of emerged
adults. In this experiment more adults emerged than in Jorge's experiment. Using a
protocol similar to the one developed by Zucoloto et al. (1979) we could not reproduce
the 100% pupation that they observed. We could not compare emergence results be-
cause the authors did not report these results. Meanwhile, Moreno et al. (1997) found
that diets containing casein is the best for pupal development. The authors measured
the flight quality of the flies, while in this work reproductive ability was measured.
The different concentrations of powdered dehydrated araca-boi in the diets influ-
enced the numbers of larvae attaining maturity (pupation) and their rate of develop-
ment. The percentages of larvae completing development varied from 0 to 38.

















Florida Entomologist 82(1)


March, 1999


TABLE 3. COMPARISON OF MERIDIC DIET WITH DEHYDRATED ARAQA-BOI AT 26C, 88%
RH AND A PHOTOPERIOD OF 12:12 (L:D) FOR REARING ANASTREPHA OBLIQUA.

Diet No No No Duration (Days) % %
number pH Larvae Pupae Adults Larvae Pupae Pupation Emergence

1 3 100 0 -
2 3 100 0 -
3 3 100 38 12 27.7 12.1 38 31.5
4 3 100 28 11 24.7 14.2 28 39.3
5 3 100 30 12 24.3 13.6 30 40.
6 3 100 24 4 25 12.8 24 16.7


Optimum development was obtained using powdered dehydrated araga-boi at concen-
trations ranging from 20 to 40 percent (Table 3). Concentrations higher or lower than
the optimal range yielded substantially fewer pupae or adults. The rate of growth
(time to pupation) in this experiment (13.1 days, at 26 + 1C, 88 + 6% RH, and a pho-
toperiod of 12:12(L:D) was the same as observed by Jorge (1987) at 26.5C (13.6 days).
However, these results differ from those observed by Polloni (1981), who reported 18.2
days at 22.5C, which was probably a result of lower temperature.
Instead of Petri dishes, we used glass tubes to hold the media for larval develop-
ment, because we noted that larvae were drowning in the Petri dishes.
West Indian fruit fly larvae can be reared in semi-artificial conditions on meridic
diets with powdered dehydrated araga-boi from 20 to 40 percent. However, the meth-
odology described here is only a beginning and further refinements are necessary. In
particular, optimum conditions for pupae need additional investigation followed by
further evaluation of more generations.

ACKNOWLEDGMENTS

Sincere thanks to Drs. Jos6 R. P. Parra, Mohamed Habib and, Sandra M. Darwich,
for their suggestions and encouragement. Also, special thanks to Dr. J. Morgante for
providing the sugar cane hydrolyzate used in this research.

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ZUCCHI, R. A., N. M. SILVA, AND S. SILVEIRA NETO. 1996. Anastrepha species from the
Brasilian Amazon: Distribution, Hosts, and lectotypes designations, pp. 259-
264 in B. A. McPheron and G. J. Steck, [eds.] Fruit fly pests. St. Lucie Press,
Delray Beach, Fl.
ZUCOLOTO, F. S., S. PUSCHE, AND C. M. MESSAGE. 1979. Valor nutritivo de dietas ar-
tificiais para Anastrepha obliqua (Diptera: Tephritidae). Bolm. Zool., Univ. S.
Paulo 4: 75-80.


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Robacker: Attraction of Mexican Fruit Fly to Synthetic Lures 87


ATTRACTION OF WILD AND LABORATORY-STRAIN MEXICAN
FRUIT FLIES (DIPTERA: TEPHRITIDAE) TO TWO SYNTHETIC
LURES IN A WIND TUNNEL

DAVID C. ROBACKER
Crop Quality and Fruit Insects Research, USDA, Agricultural Research Service 2301
South International Blvd., Weslaco, TX 78596

ABSTRACT

Attraction of laboratory-strain Mexican fruit flies,Anastrepha ludens (Loew), and
wild-type flies to two synthetic lures was evaluated in a wind-tunnel. The lures were
BioLure (ammonium acetate and putrescine) and AMPu (ammonium carbonate,
methylamine HC1, and putrescine). In one experiment, wild-type flies from the state

















Florida Entomologist 82(1)


March, 1999


of Nuevo Leon, Mexico, were evaluated against laboratory-strain flies that originated
in Nuevo Leon. Yellow panels containing AMPu attracted >2.5 times more females
and >3.5 times more males of both fly strains than panels containing BioLure. In an-
other experiment, wild-type flies from the state of Chiapas, Mexico, were evaluated
against the Nuevo Leon laboratory strain. Results of this experiment were similar to
the first except the differences in attractiveness between AMPu and BioLure to flies
of both strains were less pronounced. The difference in relative attractiveness of
AMPu and BioLure in the two experiments was related to the time of year when the
experiments were conducted rather than to inherent differences between the fly
strains. In both experiments, BioLure was about two times more attractive to fe-
males than to males whereas AMPu was only slightly more attractive to females. Both
lures were more attractive to laboratory-strain flies than to wild-type flies from either
region of Mexico.

Key Words: Anastrepha ludens, attractants, ammonia, ammonium acetate, pu-
trescine, methylamine

RESUME

La atracci6n de las moscas mexicanas de la fruta (Anastrepha ludens [Loew]) sil-
vestres y unas de laboratorio a dos cebos atrayentes sint6ticos fue evaluada en un tui-
nel de viento. Los cebos utilizados fueron BioLure (acetato de amonia y putrecina) y
AMPu (carbonato de amonia, hidrocloro de metilamina, y putrecina). En un experi-
mento, moscas silvestres del estado de Nuevo Le6n, M6xico, fueron evaluadas en com-
paraci6n con moscas del laboratorio criadas originalmente de moscas que se
obtuvieron en Nuevo Le6n. Paneles amarillos con AMPu atrajeron >2.5 de veces mas
de moscas hembra y > 3.5 de veces mas de moscas macho de las dos lines de moscas
que panels con BioLure. En otro experiment, moscas silvestres del estado de Chia-
pas, M6xico, fueron evaluadas en comparaci6n con las del laboratorio de origen de
Nuevo Le6n. Los resultados de este experiment fueron similares al del primero, con
la excepci6n de que las diferencias de atracci6n entire AMPu y BioLure para las mos-
cas de las dos cepas fue menos pronunciada. En los experiments, la diferencia de la
atracci6n relative de AMPu y BioLure estuvo mas relacionada a la 6poca del aflo en
que los experiments se llevaron a cabo mas que a diferencias inherentes entire las dos
lines de moscas. En los dos experiments, BioLure fue aproximadamente dos veces
mis atractivo para las hembras que para los machos, mientras que AMPu fue ligera-
mente mas atractivo para las hembras. Los dos cebos fueron mas atractivos para las
moscas de la linea del laboratorio que para las silvestres de cualquiera de las dos re-
giones de M6xico.




Robacker & Warfield (1993) developed and Robacker (1995) modified an attractant
for the Mexican fruit fly,Anastrepha ludens (Loew), called AMPu consisting of ammo-
nium carbonate, methylamine HC1 and putrescine. Later, Heath et al. (1995) devel-
oped an attractant for the Mediterranean fruit fly, Ceratitis capitata (Wiedemann),
consisting of ammonium acetate and putrescine. The latter attractant, commercially
available as BioLure, was improved by addition of trimethylamine (Heath et al.
1997). Field tests showed that the original two-component BioLure also attracted
the Mexican fruit fly (Heath et al. 1995) and that addition of trimethylamine did not
affect attractiveness to the Mexican fruit fly (Heath et al. 1997).
Wind-tunnel bioassays demonstrated that AMPu was about two times more at-
tractive than BioLure to laboratory-strain Mexican fruit flies in experiments de-
signed to test effects of hunger and gamma irradiation of flies on their responses to

















Robacker: Attraction of Mexican Fruit Fly to Synthetic Lures 89

the two lures (Robacker 1998). One-day experiments conducted in citrus orchards in
south Texas with fresh AMPu lures each day and BioLure lures ranging in age from
0-24 days confirmed the wind-tunnel bioassays. In these field tests in which the lures
were tested on sticky traps, AMPu was about 2.5 times more attractive than Bi-
oLure to sterile, laboratory-strain Mexican fruit flies.
The purpose of this research was to test the hypothesis that wild-type Mexican
fruit flies, or flies that originated from different populations, may not respond to these
two lures in the same way as the laboratory-strain flies. Wild-type flies from both
northeastern and southern Mexico were tested against the laboratory strain. The ex-
periments were conducted using the wind-tunnel bioassay described in Robacker
(1998).

MATERIALS AND METHODS

Insects and Laboratory Conditions

Laboratory-strain flies were from a culture that originated from fruit of chapote
amarillo, Sargentia greggii S. Wats., a native host of the Mexican fruit fly, collected in
Nuevo Leon, Mexico, in 1987. The culture has been maintained on laboratory diet
since establishment. Wild-type flies were from the Montemorelos area of Nuevo Leon
in northeastern Mexico and the Tapachula area of Chiapas in southern Mexico. Nuevo
Leon flies were obtained from larvae that egressed from either grapefruit or chapote
amarillo collected during the spring of 1997. Chiapas flies were obtained from larvae
that egressed from either sweet or sour orange collected during the winters of 1997
and 1998. Adult flies were maintained on sugar and water (provided separately) be-
cause previous work showed that this feeding regime maximized responses to both
types of lures used in this work (Robacker 1998). Laboratory conditions for holding
flies were 22 + 2C, 50 + 20% relative humidity and photophase from 0630 to 1930 h
provided by fluorescent lights.

Lures

BioLure lures were obtained from Consep, Inc. (Bend, Oregon). They consisted of
an ammonium acetate packet and a putrescine packet. The 2 packets were taped to-
gether with their membrane openings unobstructed and separated from each other
for use in the wind tunnel bioassay. BioLure lures were used each day for 5 days fol-
lowing removal from refrigeration. The BioLure lures produce stable emissions of
their components for at least 4 weeks in the laboratory (Heath et al. 1995). Laboratory
tests to measure emissions of BioLure components indicated that the lures emit (at
laboratory temperature 23C) about 300 ig/h of ammonia (Heath et al. 1997). Emis-
sion of acetic acid is probably about 3-12 ig/h. These rates for acetic acid were re-
ported by Heath et al. (1995) for a similar ammonium acetate packet (Consep, Inc.)
that emitted 100-500 ig/h of ammonia. Putrescine emission has not been determined
for BioLure.
AMPu was used in an agar formulation in 1.9 ml polypropylene microcentrifuge
tubes (A. Daigger & Company, Inc., Wheeling, Illinois) (Robacker 1995, 1998). AMPu/
agar lures were prepared by mixing equal volumes of hot agar solution (Bacto Agar,
Difco Laboratories, Detroit, Michigan) and aqueous AMPu containing 120, 200, and
20 mg/ml, respectively, of ammonium carbonate (ACS Reagent quality, Aldrich Chem-
ical Co., Inc., Milwaukee, Wisconsin), methylamine hydrochloride (99%, Sigma Chem-
ical Co., St. Louis, Missouri) and putrescine (98%, Aldrich). Final concentrations in

















Florida Entomologist 82(1)


March, 1999


AMPu/agar tubes were 60, 100, and 10 mg/ml of the three chemicals and 1% agar in
a final volume of 1.7 ml. The pH of the AMPu/agar formulation was 8.7-8.9. AMPu
tubes were capped and stored in a refrigerator. They were used in tests for 1 day after
removal from refrigeration. Lures were discarded after 1 day because this agar for-
mulation was developed only for short-term delivery of the AMPu components. Previ-
ous laboratory tests to measure emissions of AMPu components indicated that these
lures emit (at 35C) about 300 ig/h of ammonia, 40 ig/h of methylamine, 17 ng/h of
putrescine, and 20 ng/h of 1-pyrroline, a chemical that forms spontaneously in the
lures (Robacker and Bartelt 1996).


WIND-TUNNEL BIOASSAY

Bioassays were conducted in a plexiglass wind tunnel with dimensions 0.3 x 0.3 x
1.2 m. Wind tunnels of similar dimensions have been used successfully for bioassays of
fruit fly semiochemicals (Landolt et al. 1992, Epsky et al. 1997). The bioassay method
used for this work was modeled after that used by Landolt et al. (1992) for the Medi-
terranean fruit fly and was used previously with the Mexican fruit fly (Robacker 1998).
Each end of the wind tunnel was screened to allow airflow. The downwind end con-
tained a baffle system to create a uniform airflow through the chamber. Air was pulled
through the chamber at 0.4 m/sec by an exhaust fan connected to the downwind end.
Air leaving the chamber was vented from the room by a ceiling exhaust fan. The top
of the chamber had 2 circular service openings (12.8 cm diam) with plexiglass covers,
one located near each end of the chamber to allow easy access to the chamber interior.
A 100 W "soft white" light bulb (General Electric Co., Cleveland, Ohio) in a reflecting
lamp was positioned 17 cm above the downwind end of the chamber. The purpose of
this light was to hold flies non-responsive to lure odors in the downwind end by posi-
tive phototaxis and thus minimize random flying into the upwind end of the chamber.
Overhead lighting was provided by 2 banks of 4 fluorescent "cool white" lights each
(F40CW, General Electric).
For each bioassay one AMPu tube or BioLure lure was attached to the side of a
yellow plastic panel (10 by 13 cm). The panel was suspended in a fixed position from
the service opening at the upwind end of the chamber so as to provide a broad visual
stimulus to responding flies downwind and with the lure on the upwind side of the
panel away from responding flies. In this configuration the panel was 21 cm away
from the upwind end and nearly in the center of the air stream (midway between top
and bottom and the sides).
Flies were introduced one at a time into the downwind end of the chamber in clear
plastic vials (7 cm by 3 cm diam) placed on top of a beaker on the bottom of the cham-
ber directly below the downwind service opening. In this configuration, the top of the
vial (where the fly would emerge) was located in the center of the air stream. Each fly
was allowed 5 min to leave the vial (fly or walk off of the vial). Flies that did not leave
in 5 min were not included in the data. Once a fly left the vial, the fly was allowed 5
min to fly or walk upwind or contact the panel. Upwind movement was scored if flies
passed a point 2/3 of the distance from the release vial to the panel.
As a control, each test of a wild fly was followed by a test of attraction of a labora-
tory-strain fly to the same lure. Also, tests were conducted in two identical chambers.
One chamber was used for AMPu and the other for BioLure for a series of 40-50
tests. Chambers then were washed with soapy water (all test chemicals are water sol-
uble) and the lures were tested in the other chambers for a series. Comparison of data
obtained in the two chambers indicated that fly responses were not affected by cham-
ber.

















Robacker: Attraction of Mexican Fruit Fly to Synthetic Lures 91

Statistical Analyses

Effects of fly type, sex, lure type, time of year when bioassays were conducted, and
various factor interactions on responses of flies in the wind-tunnel bioassay were
tested by Chi-square using the Loglinear Model procedure of SYSTAT 7.0 (SYSTAT
1997). The effect of fly age on response was tested using option Cochran (Cochran's
test of linear trend) of the XTAB procedure of SYSTAT 7.0.

RESULTS AND DISCUSSION

Wild-Type vs Laboratory-Strain Flies from Nuevo Leon

Loglinear models containing the factors fly type, sex, and lure type and all inter-
actions of these factors with response (response or not) were constructed. Pearson X2
was not significant for either upwind movement ()2 = 6.4; df = 5; P = 0.27) or contact
with panels (x' = 3.9; df = 5; P = 0.57) indicating that the complete models fit the ob-
served response frequencies.
Fly type had significant effects in models for both upwind movement and contact
with panels. Wild-type flies from Nuevo Leon moved upwind (x' = 163.6 for both lures
combined; df = 1; P < 0.001) and contacted (Z = 175.3 for both lures combined; df = 1;
P < 0.001) panels at much lower rates than laboratory-strain flies that originated
from Nuevo Leon (Table 1). Wild-type flies whose larvae had egressed from chapote
amarillo did not respond differently to the lures than flies from grapefruit.
Sex of flies had little effect on upwind movement of either strain toward either lure
or on contact with panels with AMPu. Females of both strains contacted panels with
BioLure at higher rates than did males (Z' = 4.9 from a reduced model containing
data for BioLure only; df = 1; P < 0.05). Although sex had less effect on responses to
AMPu than to BioLure, the interactionof sex and lure type was not significant.
Lure type also had significant effects. More flies of both strains moved upwind (x'
= 45.9 for both strains combined; df = 1; P < 0.001) and contacted (x' = 57.9 for both
strains combined; df = 1; P < 0.001) panels with AMPu than panels with BioLure.
Lure type also had significant effects when only wild-type flies were included in the
analysis. More wild-type flies moved upwind ()2 = 13.0; df = 1; P < 0.001) and con-


TABLE 1. UPWIND MOVEMENT AND CONTACT WITH PANELS WITH SYNTHETIC LURES IN A
WIND TUNNEL BY LABORATORY-STRAIN AND WILD-TYPE MEXICAN FRUIT FLIES,
BOTH STRAINS ORIGINATING FROM NUEVO LEON, MEXICO.

Upwind Movement Contact with Source

BioLure AMPu BioLure AMPu

laboratory Males 22.8 47.1 9.8 36.6
strain
Females 27.7 51.1 17.5 44.9
wild-type Males 5.2 15.7 0.5 5.2
Females 5.6 10.2 1.7 4.5

Values are percentages with n's per cell: male flies with BioLure, 193; females with BioLure, 177; males
with AMPu, 191; females with AMPu, 176.

















Florida Entomologist 82(1)


March, 1999


tacted (Z2 = 9.3; df = 1; P < 0.01) panels with AMPu than panels with BioLure. Rel-
ative responses of wild-type flies and laboratory-strain flies to the two lures differed
as indicated by significant interactions of lure type by fly type for both upwind move-
ment (Z2 = 3.9; df = 1; P < 0.05) and contact with panels (Z2 = 7.2; df = 1; P < 0.01).
These interactions indicated that responses to BioLure by wild-type flies were
slightly less than expected based on main effects of lure type and fly type.


Wild-Type Flies from Chiapas vs Laboratory-Strain Flies

Loglinear models like those used to analyze the Nuevo Leon fly responses were
constructed. Pearson xZwas not significant for either upwind movement (Z2 = 3.0; df =
5;P = 0.70) or contact with panels (Z' = 1.8; df = 5;P = 0.88).
Wild-type flies from Chiapas moved upwind (x2 = 16.4 for both lures combined; df
= 1;P < 0.001) and contacted (x2 = 33.0 for both lures combined; df= 1;P < 0.001) pan-
els with lures at lower rates than the laboratory-strain flies that originated in Nuevo
Leon (Table 2). Effects of larval host of the wild-type flies on responses of the flies to
the lures could not be determined because the pupae from the two hosts were mixed
together before eclosion.
Females of both strains moved upwind (x = 10.5 for both strains combined; df = 1;
P < 0.01) and contacted (x2 = 16.3 for both strains combined; df = 1;P < 0.001) panels
with lures at higher rates than did males. As with wild-type flies from Nuevo Leon,
the effect was more pronounced for response to BioLure than for response to AMPu
although interactions of sex and lure were not significant for either upwind movement
or contact with panels.
More flies of both strains moved upwind (Z2 = 10.2 for both strains combined; df =
1;P < 0.01) and contacted (xZ = 13.1 for both strains combined; df= 1; P < 0.001) panels
with AMPu than panels with BioLure AMPu was not significantly more attractive
than BioLure when only wild-type flies were included in the analysis. Interactions
of fly type with lure type were not significant for either upwind movement or contact
with panels.



TABLE 2. UPWIND MOVEMENT AND CONTACT WITH PANELS WITH SYNTHETIC LURES IN A
WIND TUNNEL BY WILD-TYPE MEXICAN FRUIT FLIES FROM CHIAPAS, MEXICO,
AND BY LABORATORY-STRAIN FLIES ORIGINATING FROM NUEVO LEON, MEXICO.

Upwind Movement Contact with Source

BioLure AMPu BioLure AMPu

laboratory Males 15.6 23.8 4.8 17.0
strain
Females 18.3 29.8 12.5 22.3
wild-type Males 10.2 12.2 2.0 4.1
flies
Females 9.2 18.2 4.2 5.8

Values are percentages with n's per cell: male flies, 147; females with BioLure, 120; females with AMPu,
121.

















Robacker: Attraction of Mexican Fruit Fly to Synthetic Lures 93

Effect of Fly Sex

Data from the current work and from Robacker (1998) indicated that BioLure
was much more attractive to females than to males (females/males = 1.2 to 6.1 in var-
ious experiments). Corresponding ratios for attraction to AMPu were smaller (fe-
males/males = 1.1 to 2.8). I conclude that AMPu and BioLure differ in their
propensities to attract males and females.


Effect of Fly Age

Bioassays were conducted with wild-type flies from both Nuevo Leon and Chiapas
ranging in age from 3-60 days post eclosion. The relationship between age of flies and
their responses to the two lures was investigated.
Table 3 shows the percentages of wild-type flies of 3 age groups that contacted pan-
els with lures. The percentage of males responding to BioLure appeared to increase
with age of flies. Cochran's test of linear trend indicated the effect was not significant.
The percentages of older males (>18 days) that responded to both lures were
higher than the percentages of 3-7 day old males that responded whereas the opposite
was true for females (Table 3). A loglinear model containing the factors age and sex
and the interaction o f age, sex, and response (contact or not) was constructed to test
whether age affected responses of males and females differently. Pearson x was not
significant (x2 = 3.5; df = 4; P = 0.48). The interaction of sex and age was significant
(Z = 4.3; df = 1; P < 0.05) indicating that age affected responses of the sexes differ-
ently.
More flies contacted panels with AMPu than those with BioLure at all fly ages
(Table 3). Relative attractiveness of the lures at different fly ages was not significantly
different.
Previous work showed that age of laboratory-strain flies between 6 and 17 days old
had little effect on attraction of the flies to AMPu and BioLure (Robacker 1998). The
analysis conducted here indicates that age also did not have great effects on responses of
wild-type flies to these two lures. I do not conclude that age would have little or no effect
on responses to these lures or similar semiochemicals in nature where conditions differ.



TABLE 3. CONTACTS WITH PANELS WITH SYNTHETIC LURES IN A WIND TUNNEL BY WILD-
TYPE FLIES (NUEVO LEON AND CHIAPAS FLIES COMBINED) OF DIFFERENT AGES.

3-7 days 8-18 days >18 days

BioLure 0.0 0.6 2.0
males
AMPu 3.0 5.2 4.8
BioLure 5.1 1.4 3.4
females
AMPu 5.4 5.5 4.2

Values are percentages with n's per cell: 0-7 days old, 33-39; 8-18 days old, 144-155; males >18 days old, 145-
148; females >18 days old, 118-119.

















Florida Entomologist 82(1)


March, 1999


Effect of Time of Year

AMPu was much more attractive than BioLure to wild-type flies from Nuevo
Leon but only slightly more attractive than BioLure to wild-type flies from Chiapas.
The following analysis indicates that this difference is not due to inherent differences
between flies from Nuevo Leon and Chiapas, but more likely is due to the time of year
when the bioassays were conducted.
Bioassays with wild-type flies from Nuevo Leon were conducted from June to Sep-
tember, 1997. Bioassays with wild-type flies from Chiapas were conducted during
April and May, 1997, and during March and April, 1998. During each experiment, bio-
assays with laboratory-strain flies were conducted as controls. Laboratory-strain flies
tested during June to September along with wild-type flies from Nuevo Leon re-
sponded like the wild-type flies in that their responses to AMPu were much higher
than to BioLure (Table 1). Likewise, laboratory-strain flies tested during March to
May along with wild-type flies from Chiapas responded like the wild-type flies in that
their responses to AMPu were only slightly higher than to BioLure(Table 2). Thus,
responses of wild-type flies from the two regions of Mexico were similar to responses
of the laboratory-strain flies in each experiment.
Because the laboratory strain was the same in both experiments, the effect of time
of year on responses of the laboratory strain was analyzed to determine if time of year
may have affected the relative attractiveness of AMPu and BioLure to the two wild-
type strains. Table 4 shows responses in wind-tunnel bioassays of 4-17 day old, sugar-
fed, protein-deprived laboratory-strain flies to the two lures during spring (February-
May) and summer (June-September). Table 4 includes data from the current work
and from experiments conducted previously to test effects of food deprivation and
gamma irradiation(Robacker 1998).
AMPu was about four times more attractive than BioLure to males during both
test periods (Table 4). However, the relative responses of females to AMPu and Bi-
oLure changed. AMPu was only about 30% more attractive than BioLure during
spring bioassays but 2.5 times more attractive during summer bioassays.
A loglinear model containing the factors lure and time of year and all interactions
of these factors with response (response or not) was constructed to test the hypothesis
that laboratory-strain females responded differentially to the 2 lures at different
times of the year. Pearson x was not significant (Z2 = 0.9; df = 2; P = 0.64). The inter-
action of lure and time of year was significa nt (Z = 5.9; df = 1;P < 0.05) indicating that


TABLE 4. CONTACTS WITH PANELS WITH SYNTHETIC LURES IN A WIND TUNNEL BY LAB-
ORATORY-STRAIN MEXICAN FRUIT FLIES AT 2 TIMES OF THE YEAR.

BioLure AMPu

spring 7.0 27.8
males
summer 8.9 39.0
spring 20.0 26.4
females
summer 16.9 43.6

Values are percentages with n's per cell: male flies during spring, 227-229; males during summer, 210-213; fe-
males during spring, 200-201; females during summer, 195.

















Robacker: Attraction of Mexican Fruit Fly to Synthetic Lures 95

the relative attractiveness to laboratory-strain females of AMPu compared with Bi-
oLure changed from spring to summer. This analysis indicates that the difference in
relative attractiveness of AMPu and BioLure to wild-type females from Nuevo Leon
compared with those from Chiapas probably is not due to inherent differences be-
tween flies from Nuevo Leon and Chiapas. The factors that caused the time-of-year ef-
fect are not known but they exerted their effects equally on laboratory-strain and
wild-type flies.

Wind-Tunnel Responses in the Absence of Lures

Control bioassays were not conducted to determine how many wild-type flies
would move upwind and contact panels when no lures were present. Previous re-
search with laboratory-strain flies in this bioassay system showed that about 9% of
males and 4% of females moved upwind and 0.6% of both males and females contacted
panels that did not contain lures (Robacker 1998). Responses by wild-type females but
not males to BioLure were higher than responses by laboratory-strain flies when no
lures were present (Tables 1 & 2), however, it was not possible to determine from this
comparison if wild-type females actually responded to BioLure. Positive responses
to AMPu can be inferred because responses of wild-type flies (Nuevo Leon & Chiapas
combined) to AMPu were significantly greater than responses to BioLure (Z' = 8.3
for upwind movements by males, df = 1, P < 0.01; x2 = 7.5 for contacts by males, df =
1, P < 0.01; x2 = 6.6 for upwind movements by females, df = 1, P = 0.01; x = 2.2 for con-
tacts by females, df = 1, P = 0.14).

Implications for Attractants Research

Laboratory-strain flies (not irradiated) responded to the lures at much higher
rates (21.5%) (summed over both lures and both experiments) than wild-type flies
(3.4%) in this work, and nonirradiated flies responded at higher rates (29.4%) than ir-
radiated flies (14.4%) in previous work (Robacker 1998). However, attractiveness of
the 2 lures relative to each other varied little with fly type: i.e. AMPu/BioLure re-
sponse (contacts) ratios were similar for wild-type flies (2.6) (summed over Nuevo
Leon and Chiapas flies) vs laboratory-strain control flies from the experiments with
wild flies (2.8) in this work, and for irradiated (2.8) vs nonirradiated laboratory-strain
flies (2.4) in previous work. These data indicate that semiochemical research con-
ducted with laboratory-strain Mexican fruit flies, irradiated or nonirradiated, should
not differ qualitatively from that done with wild-type flies.
As discussed above, laboratory-strain flies responded quantitatively much differ-
ently than wild-type flies to both lures. It is not known if this result is due to morpho-
logical or physiological differences in neural apparatus between the fly types, or to
differences in the way each fly type reacts to laboratory conditions.

Implications for Trapping Programs

Results in this work and in Robacker (1998) in which AMPu was consistently more
attractive than BioLure to both laboratory and wild-type Mexican fruit flies suggest
that AMPu should be a better lure than BioLure in trapping programs. However,
other factors such as local climatic conditions and the type of trap used with the lures
could have profound effects on the relative attractiveness of the lures. Thus, results of
these laboratory bioassays and even the field experiments conducted with irradiated
flies in south Texas (Robacker 1998) may be poor predictors of field-trapping experi-
ments with wild flies in localities other than Texas.
















Florida Entomologist 82(1)


March, 1999


Direct field comparisons of the two lures in populations of wild flies have not been
conducted because a formulation of AMPu proven to function properly for more than
2 days in the field has not been developed. Obviously, a long-lasting formulation is
needed before AMPu could even be considered for use in trapping programs.

ACKNOWLEDGMENTS

I thank Maura Rodriguez and Connie Chavez for technical assistance, Don Tho-
mas and Daniel Moreno for obtaining the wild flies, and Bob Heath (USDA-ARS,
Gainesville, Florida) and Peter Landolt (USDA-ARS, Wapato, Washington) for the
wind tunnels. Use of a product brand in this work does not constitute an endorsement
by the USDA.

REFERENCES CITED

EPSKY, N. D., B. D. DUEBEN, R. R. HEATH, C. R. LAUZON, AND R. J. PROKOPY. 1997. At-
traction ofAnastrepha suspense (Diptera: Tephritidae) to volatiles from avian
fecal material. Fla. Entomol. 80: 270-277.
HEATH, R. R., N. D. EPSKY, B. D. DUEBEN, J. RIZZO, AND F. JERONIMO. 1997. Adding
methyl-substituted ammonia derivatives to a food-based synthetic attractant
on capture of the Mediterranean and Mexican fruit flies (Diptera: Tephritidae).
J. Econ. Entomol. 90: 1584-1589.
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