Title: Florida Entomologist
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Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1994
Copyright Date: 1917
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Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
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Braman et al.: Zoysiagrass Susceptibility to S. vicinus


POTENTIAL RESISTANCE IN ZOYSIAGRASSES
TO TAWNY MOLE CRICKETS (ORTHOPTERA:
GRYLLOTALPIDAE)

S.K. BRAMAN', A.F. PENDLEY', R.N. CARROW2
AND M.C. ENGELKE3
Department of Entomology and 2 Department of Crop &
Soil Science, University of Georgia,
College of Agriculture and Environmental Sciences
Experiment Stations, Georgia Station, Griffin, GA 30223

STexas A&M University, Research & Extension Ctr.,
17360 Coit Road., Dallas, TX 75252

ABSTRACT

Reduction in growth by tawny mole crickets, Scapteriscus vicinus Scudder, at den
sities equivalent to 15 adults per 0.09 m2 varied among nine experimental and three
commercially available zoysiagrass (Zoysia Willd.) cultivars. Reductions in root dry
weights after a four week infestation period were similar among all cultivars, and av
eraged 27.1% less than non-infested controls. Shoot dry weight reduction was most
severe for DALZ 8516, DALZ 9006, and Meyer zoysia. The cultivars that retained the
highest percentage of their normal growth were DALZ 8502, DALZ 8514, DALZ 8701,
and Emerald zoysia. Crickets fed on cultivars that were least damaged usually pro
duced the least number of eggs. However, when the most severely injured selection
(DALZ 8516) served as the host, a similarly low number of eggs was observed. Crick
ets fed on Meyer zoysia and DALZ 8508 produced the greatest number of eggs.

Key Words: Host plant resistance, Scapteriscus spp., turfgrass

RESUME

La reducci6n del crecimiento de los grillotopos aleonados, Scapteriscus vicinus
Scudder, a densidades equivalentes a 15 adults por 0.09 m2, vari6 en nueve culti
vares experimentales y tres comerciales de hierba zoysia (Zoysia Willd.). Las reduc
ciones en el peso seco de las raices fueron similares en todos los cultivares, luego de
un period de 4 semanas infestaci6n, y promediaron un 27.1% menos que los testigos
no infestados. La reducci6n en el peso seco de los brotes fue mas several en las zoysias
DALZ 8516, DALZ 9006 Y Meyer. Los cultivares que retuvieron el mas alto porcentaje
de su crecimiento normal fueron DALZ 8502, DALZ 8516, DALZ 8701 y Emerald. Los
grills que se alimentaron de los cultivares menos danados, usualmente produjeron
menor numero de huevos. Sin embargo, cuando el clon mas severamente danado
(DALZ 8516) sirvi6 como hospedante, fue observado un numero de huevos similar
ente bajo. Los grills alimentados de las zoysias Meyer y DALZ 8508 prdujeron el
mayor numero de huevos.





Mole crickets in the genus Scapteriscus have become the most serious pests of turf
grasses in the southeastern United States since their entry into this country, probably
via the ballast of ships at approximately 1900 (Walker & Nickle 1981, Nickle & Cast
ner 1984, Walker 1984). The tawny mole cricket, Scapteriscus vicinus Scudder, is the


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Florida Entomologist 77(3) September, 1994


more damaging of the two species common in Georgia. Management of these pests has
involved chemical, cultural, and classical biological control efforts (Walker 1984, Hud
son et al. 1988).
Laboratory and field screening has identified turfgrass genotypes that are rela
tively resistant, tolerant, or less preferred by various insects and mites (see reviews
by Reinert 1982, Quisenberry 1990). Limited research has focused on resistance to
mole crickets and white grubs (Potter & Braman 1991). Resistant cultivars are
needed to provide a safe, economical control strategy for these serious turf pests.
Herein, we report the results of a greenhouse study that evaluated nine experimental
and three commercially available zoysiagrass (Zoysia Willd.) cultivars for their sus
ceptibility to mole cricket injury and suitability as oviposition substrates.


MATERIALS AND METHODS

Zoysiagrass plugs (4.6 x 4.6 x 6.3 cm) of 12 cultivars were transplanted into gran
ular calcinated clay (Turface, Applied Industrial Materials, Corp., Deerfield, IL) in
plastic pots (15 cm diam) in a greenhouse. Pots were watered daily and fertilized once
per week with a solution containing 250 ppm NPK (Peters" 20-20-20). Milorganite
was applied (2.25 gm per pot) once per month. Turf was cut weekly to a height of 5 cm
Experimental cultivars included a range of leaf textures, colors, and growth rates
(Carrow 1992). Six months after transplanting, plugs were transferred to PVC tubes
(38 cm tall; 15 cm diam) containing fine sand (children's play sand). These tubes were
covered at the bottom with plastic petri dish lids and were placed into wooden box
frames and equipped with drip irrigation. Watering and fertilization regimes were
maintained as before for one month before infestation with adult crickets.
Adult S. vicinus were collected in Tifton, GA during April, 1992 using a standard
acoustic trap similar to that described by Walker (1982). Twenty-four treatments (12
cultivars each infested with mole crickets and the same 12 cultivars non-infested)
were arranged in a randomized complete block design with 7 replications (168 total
tubes). Two female and one male mole cricket were introduced into each of the 84
tubes designated as infested treatments (252 total crickets were used). Each con
trainer was covered with 32-mesh saran screen (Chicopee Manufacturing Co., Gaines
ville, GA) to prevent escape of crickets. Non-infested cages were also covered with
screens to ensure equivalent light, temperature and humidity conditions. Water and
fertilizer regimes were maintained as described previously. Greenhouse microplots
were destructively sampled after four weeks of exposure to crickets.
Top growth was clipped to a height of 5 cm two weeks after crickets were intro
duced into the PVC containers. Clippings were placed in paper bags, oven dried for 7
days, and then weighed. Top growth dry weight was also recorded four weeks after
cricket infestation. Numbers of green shoots per 18 cm2, selected at random from the
182 cm2 surface of each tube, and root dry weights were determined at the terminal
tion of the experiment.
Sand from each experimental PVC microplot (1343 kg total for all plots) was sifted
to recover mole cricket adults and eggs. Adult survival and numbers of eggs were re
corded and compared among cultivars using the GLM procedure (SAS 1985). Num
bers of eggs per cultivar were also regressed against injury. Means were separated
following a significant analysis of variance by a least significant difference test (Sokal
& Rohlf 1981). Growth reductions of roots, shoots, and shoot density (expressed as a
percentage of non-infested plants for each cultivar) were transformed using an arc
sine square root of the proportion before being subjected to analysis of variance and
mean separation using a least significant difference test.









Braman et al.: Zoysiagrass Susceptibility to S. vicinus 303

RESULTS AND DISCUSSION

Reduction in top growth two weeks after infestation was statistically similar
among all cultivars (F 1.50; df=11,66; P>0.05; Table 1). Growth of cricket-infested
plants averaged 67.9% of non-infested plants at that time. Reduction in shoot dry
weights differed (F=4.17; df=11,66; P 0.0001) among cultivars four weeks after infes
station. DALZ 8516 achieved only 10.1% of its normal growth when infested with mole
crickets. DALZ 8502, however, maintained 55% of its normal growth even under this
high infestation level. The pest density used in this study was equivalent to 15 adult
crickets per 0.09 m2 ( 1 ft2). This represents more than 15 times the pest density re
quiring chemical intervention to protect turf. Total reduction in top growth for the en
tire four week period ranged from 35.7 to 74.8% (inverse of the extremes presented in
Table 1).
Shoot density followed a pattern similar to that of clipped dry weights. Root weight
reduction, however, was similar (F=1.45; df=11,66; P>0.05) among all cultivars eval
uated and averaged 72.9% of non-infested controls (Table 1). The majority of the dam
age observed was confined to the crown of each infested zoysiagrass plug. Dead turf
was first visible at the center of each plug and expanded outward with increasing time
of exposure.
Adult survival at the termination of the four week exposure period was not signif
icantly affected by cultivar (F 1.58; df=11,66; P>0.05) and averaged 70.0% on all cul
tivars (data not given). Egg production during this time period, however, differed
(F=2.63; df=11,66; P 0.003) among cultivars (Fig.l). Mean numbers of eggs per culti
var ranged from 4.5 to 35.4. In general, tubes containing cultivars that were least
damaged (DALZ 8502, DALZ 8514, Emerald, DALZ 8701, and DALZ 8507) also con
trained lower numbers of eggs, except for DALZ 8516, which was the most severely
damaged, but contained relatively few eggs. Meyer zoysia and DALZ 8508 supported
the greatest egg production. Regression of number of eggs per cultivar against injury
was significant (F 16.9; df=11,82; P 0.0001; r2 0.2), however, the low r2 value indi
cated that plant injury explained little of the variation for numbers of eggs laid.

TABLE 1. ZOYSIAGRASS RESPONSE TO ADULT S. VICINUS INDUCED INJURY.

Mean + s.e. % of Noninfested Controls (n=7)1

Shoot Dry Weight Root Dry Shoot Density

Cultivar 2 wk 4 wk Total Weight per 18 cm2

DALZ8502 87.7 0.2 55.4 0.1a 64.3 + 0.1a 72.0+0.1 49.8 + 0.1b
DALZ8514 73.0+0.1 41.6 +0.lab 55.0+0. lab 97.8+0.1 43.9 0.1bc
Emerald 100.3+0.5 41.0 + 0.1ab 54.7 + 0.lab 72.5+0.1 41.0 + 0.1bc
DALZ8701 72.4+0.1 42.2 + 0.lab 52.9 + 0.lab 72.5+0.1 80.9 + 0.2a
DALZ8507 75.5+0.2 34.1+0.1abc 50.7+0.1ab 75.9+0.1 30.9 0.1abc
DALZ8508 75.6+0.1 23.0 + 0.1bc 41.6 + 0.1abc 69.1+0.1 23.0 + 0.lab
ElToro 59.4+0.1 27.6 + 0.1bc 41.3 + 0.1abc 72.3+0.1 43.5 + 0.2bc
DALZ8512 60.6+0.1 25.1 + 0.bc 38.8 + 0.bc 76.2+0.1 20.2 0.1cd
DALZ8501 58.2+0.1 21.5 + 0.1bc 37.4 + 0.1bc 69.5 +0.1 28.7 + 0.1bc
Meyer 55.5 0.1 19.1 + 0.1bc 34.9 0.1bc 62.9 0.1 23.1 + 0.1bcd
DALZ9006 48.6 0.1 25.1 + 0.1bc 34.7 + 0.1bc 60.5 +0.1 34.7 + 0.1bcd
DALZ8516 49.0+0.1 10.1 + 0.1bc 25.2 + 0.1c 72.1 0.1 9.0 + 0.1d
1Means within a column followed by no letter or the same letter are not significantly different (P>0.05;LSD test)
































8507 El Toro
8508 8512


September, 1994


8501 9006
Meyer 8516


Zoysiagrass Cultivar
Fig. 1. Mean numbers of eggs laid during a four week period on 12 zoysiagrass cul
tivars. Bars with different letters were significantly different (P<0.05;LSD).

Results of this no-choice evaluation of 12 cultivars of zoysiagrass under intense
mole cricket pressure revealed distinct differences in susceptibility to injury and suit
ability for egg production by the tawny mole cricket. Reinert & Busey (1984) showed
that mole crickets preferred the finer textured varieties within a species of grass.
They discussed the need to define the relative contribution of nonpreference and host
plant tolerance. The results reported in the present study suggest a similar tolerance
of injury and reduced suitability for oviposition among fine textured (DALZ 8502) and
wider bladed (DALZ 8514) zoysiagrass selections. In large monocultures, such as golf
courses, parks, commercial properties, etc., nonpreference in the absence of other re
distance mechanisms has a limited value. The potential for resistance in zoysiagrass
to damage by tawny mole crickets demonstrated in this study offers characteristics in
addition to nonpreference that should permit selected cultivars to be planted with re
duced risk of mole cricket injury.

ACKNOWLEDGMENT

Appreciation is extended to W. G. Hudson for helpful discussion and collection of
crickets. The Georgia Golf Course Superintendent's Association provided partial sup
port for this project.

REFERENCES CITED

CARROW, R.N. 1992. Zoysiagrass performance, water use, and rooting as affected by
traffic and nitrogen, p. 9 in Anonymous. 1992 Turf. Res. Summary. United
States Golf Assn., Far Hill, NJ.


Florida Entomologist 77(3)


304

40


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w 30
0)

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0
5 20
.0
E
=3
C
� 10
a,



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A

AB







B



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I I


8502 Emerald
8514 8701









Braman et al.: Zoysiagrass Susceptibility to S. vicinus 305

HUDSON, W. G., J. H. FRANK, AND J. L. CASTNER 1988. Biological control of Scap
teriscus spp. mole crickets (Orthoptera: Gryllotalpidae) in Florida. Bull. Ento
mol. Soc. America 34: 192-198.
NICKLE, D.A., AND J. L. CASTNER. 1984. Introduced species of mole crickets in the
United States, Puerto Rico, and the Virgin Islands (Orthoptera: Gryllotalpi
dae). Ann. Entomol. Soc. America 77: 450-465.
POTTER, D. A., AND S. K. BRAMAN. 1991. Ecology and management of turfgrass in
sects. Annu. Rev. Entomol. 36: 383-406.
QUISENBERRY, S. S. 1990. Plant resistance to insects and mites in forage and turf
grasses. Florida Entomol. 73: 411-421.
REINERT, J. A. 1982. A review of host resistance in turfgrasses to insects and acarines
with emphasis on the southern chinch bug, p. 3-12 in H.D. Niemczyk and B. G.
Joyner [eds.]. Advances in turfgrass entomology. Hammer Graphics, Piqua,
OH, 150 pp.
REINERT, J. A., AND P. BUSEY. 1984. Resistant varieties, p. 35-40 in T. J. Walker [ed.].
Mole crickets in Florida. Florida Agric. Exp. Stn. Bull. 846.
SAS INSTITUTE INC. 1985. SAS Users Guide: Statistics, version 5 edition, Cary, NC,
956 pp.
SOKAL, R. R., AND F. J. ROHLF. 1981. Biometry Second edition. W. H. Freeman & Co.,
San Francisco.
WALKER, T. J. 1982. Sound traps for sampling mole cricket flights (Orthoptera: Gryl
lotalpidae: Scapteriscus). Florida Entomol. 65:105-109.
WALKER, T. J. 1984. Mole crickets in Florida. Florida Agric. Exp. Stn. Bull. 846. 54 pp.
WALKER, T.J., AND D.A. NICKLE. 1981. Introduction and spread of pest mole crickets:
Scapteriscus vicinus and Scapteriscus acletus reexamined. Ann. Entomol. Soc.
America 76:507-517.


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MATING FREQUENCY OF THE PAPAYA FRUIT FLY
(DIPTERA: TEPHRITIDAE) WITH AND WITHOUT HOST FRUIT


PETER J. LANDOLT
Insect Attractants, Behavior, and Basic Biology
Research Laboratory, Agricultural Research Service
U.S. Department of Agriculture, Gainesville, FL 32604


ABSTRACT

Frequency of mating of female papaya fruit flies, Toxotrypana curvicauda Gers
taecker, was affected by the availability of host fruit for oviposition. Mature females
held for 5 days with males in cages in the laboratory without host fruit nearly always
mated once. Most mature females held in cages for 5 days with both males and imma
ture papaya fruit mated more than once. Remating by females held with fruit oc
curred occasionally before oviposition, indicating that an increased propensity to
remate may not be due to sperm depletion. Mature males kept for 5 days with females
mated up to 10 times (x=5.6+0.6).

Key Words: Insecta, Toxotrypana curvicauda, oviposition, pheromone, kairomone, re
mating


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Florida Entomologist 77(3) September, 1994


RESUME

La frecuencia de apareamiento de hembras de la mosca frutera de la papaya, Tox
otrypana curvicauda Gerstaecker, fue afectada por la disponibilidad del fruto hos
pedante para la ovoposici6n. En el laboratorio las hembras maduras mantenidas en
jaulas durante 5 dias, con machos y sin el fruto hospedante, casi siempre se aparearon
una sola vez. La mayoria de las hembras maduras mantenidas enjaulas durante 5
dias con machos y frutos no maduros de papaya se apare6 mas de una vez. Nuevos
apareamientos de hembras mantenidas con frutos tuvieron lugar ocasionalmente an
tes de la ovoposici6n, indicando que el aumento en la propensi6n al reapareamiento no
debe ser el resultado del agotamiento del esperma. Los machos maduros mantenidos
durante 5 dias con hembras se apareron hasta 10 veces (x=5.6+0.6)

Key Words: Insecta, oviposition, pheromone, kairomone, remating





The frequency of mating in tephritid fruit flies is an important aspect of their sex
ual behavior. It is relevant to the development of those pest control programs based
in part on sexual interactions. For example, sex attractants developed for females
may be more effective for species that remate frequently and may then repeatedly re
spond to male sex pheromone. Also, female mating frequency directly affects the effi
cacy of the sterile male insect technique for control and eradication of tephritid fruit
flies.
Frequency of mating varies among females of species of fruit-infesting tephritids.
Females of the apple maggot fly, Rhagoletis pomonella (Walsh), mate frequently, as of
ten as weekly, possibly to maintain fertility levels (Prokopy & Roitberg 1984). Tropical
species of fruit flies that lek, such as Ceratitis capitata Wiedemann (Nakagawa et al
1971), and Anastrepha suspense (Loew) (Sivinski & Heath 1988), are thought to mate
usually once, with rematings due either to insemination failures at first matings or to
sperm depletion following extensive oviposition. Landolt & Hendrichs (1983) reported
that female papaya fruit flies, Toxotrypana curvicauda Gerstaecker, usually mate
once, based on observations of flies in a field cage. This might be expected because of
their relatively short reproductive lifespan and limited egg production.
Studies of female papaya fruit fly responses to male sex pheromone, however, dem
onstrated that females are attracted to males even after mating (Landolt & Heath
1988, 1990). Such sex attraction by mated females may be to locate a potential second
mate or may be a strategy to locate host fruit, since males in papaya fields normally
call from immature fruit (Landolt et al. 1992). Because of this apparent discrepancy
and the limited current data on T curvicauda mating frequency, the subject was re
addressed. We report here the frequency of mating of both sexes of the papaya fruit
fly in the laboratory, and effects of the presence of papaya fruit for oviposition on fe
male mating frequency.


MATERIALS AND METHODS

Papaya fruit flies were obtained as mature larvae in infested fruit collected in com
mercial papaya plantings in the Redlands area of Dade County and in Sarasota
County, Florida. Larvae emerging from fruit pupated in sterilized potting soil. Paper
cannisters (250 ml) of pupae in potting soil were kept in the laboratory in screened
cages for adult emergence. Newly emerged flies were sorted by sex daily and were









Landolt: Papaya Fruit Fly Mating Frequency


kept in screened cages with cups of sugar water and inverted water bottles on the cage
tops. The laboratory was at 22� C and 50 + 10% RH, with overhead fluorescent light
ing. Lights were on a 14:10 (L:D) cycle, with lights on at 0600 hours (E.S.T.) and off
at 2000 hours (E.S.T).
Mating frequency was determined for mature female papaya flies held with imma
ture papaya fruit (preferred oviposition sites), for mature female flies without fruit,
and for mature male flies without fruit. For the first experiment, reproductively ma
ture females (> 6 days old, Landolt 1984) were held one per cage from 0730 hours on
a Monday to 1700 hours on the following Friday. Each morning at 0800 hours, a male
was placed in each cage and observations were made at 30 min intervals until 1700
hours when the male was removed. Because matings on average take 90 min (Landolt
& Hendrichs 1983), it was assumed that none would be missed if observations were
made every 30 min. At each 30 min observation time, records were made of mating
and oviposition (for females held with fruit). Females without fruit generally do not
attempt to oviposit (Landolt & Reed 1990) and no oviposition attempts were seen in
this study by females held without fruit (egg deposition or probing with the ovipositor
on the cage). After flies mated, the male was replaced. Each cage included a small cup
of sugar water on cotton within the cage and an inverted water bottle on the cage top.
Twenty-six females were tested in this way without fruit; 10 during one week, 10 dur
ing a second week and 6 during a third week. This experiment, using the same proto
col, was also conducted for 34 mature females held with males and papaya fruit (as 2
sets of 6, 1 set of 12, and 1 set of 10 during 4 different weeks). Papaya fruit used were
green immature fruit 5 -7 cm in diam and 7 -11 cm in length. During the replicates
conducted to determine mating frequency of females without fruit, papaya was also
excluded from the laboratory to avoid host odor effects on mating behavior.
To assess mating frequency of males, a similar protocol was followed. For five con
secutive days, at 0800 hours, a mature unmated female was placed in each cage con
training one male. Observations were made each 30 minutes, until 1700 hours, when
the female was removed. The female was replaced after each copulation, with another
mature unmated female. This was done for 21 mature (>4 days old) males, all held
without papaya fruit (as 2 sets of 6 and 1 set of 5 during 3 different weeks).
Mean mating frequencies for females held with and without papaya fruit were
compared using Student's t-test.


RESULTS

Nearly all female papaya fruit flies held in cages without fruit mated only once
during the 5-day test period (Fig. 1). Mean number of matings per female (L SE) was
1.00 + 0.06. Females held in cages with papaya fruit mated up to 4 times, with most
females mating more than once (Fig. 1). Mean number of matings per female was 1.82
+ 0.15. Females without fruit mated significantly less than did females with fruit
(t=4.63, df=58, p<0.0001). Males mated up to 10 times over the course of the 5-day test
period, with nearly all mating multiple times (Fig. 2). Mean number of matings per
male was 5.60 + 0.6.
Matings by females held without fruit were predominantly during the morning
hours, while matings of females held with fruit were more dispersed throughout the
day (Fig. 3). Also, 24 of the 26 females held with males and without fruit mated on the
first day of the experiment, and 31 of the 33 females held with males and with fruit
mated on the first day of the experiment. The pattern of ovipositions observed during
these tests appeared to be bimodal, with peaks of activity in early morning and again
in late afternoon (Fig. 3).







Florida Entomologist 77(3)


100


80


60


40


20


0


0 1 2 3 4 5 6
NUMBER OF MATINGS


100


80


60


40


20 X i;



0 1 2 3 4 5 6
NUMBER OF MATINGS
Fig. 1. Percent of mature female papaya fruit flies that mated from 0 to 6 times
over 5 days, when (A) held in cages without immature papaya fruit or (B) held with
immature papaya fruit for oviposition.


September, 1994








Landolt: Papaya Fruit Fly Mating Frequency


50


II
z 40

%)
CO
w
-J 30



O 20
l-



0.


0 1 2 3 4 5 6 7 8 9 10

NUMBER OF MATINGS
Fig. 2. Percent of male papaya fruit flies held without fruit that mated from 0 to
10 times over 5 days.

Of the 33 females held in cages with both males and papaya fruit, 23 remated at
least once (Fig. 1). Of the 23 that remated, 13 (57%) mated the second time before ovi
position. Of the 13 that remated after oviposition, only 6 did so within 4 h after ovipo
sition.

DISCUSSION

These data indicate that the papaya fruit fly female typically mates more than
once if provided access to host fruit for oviposition. Although the remating rate for fe
males held with males and without fruit was very low (4%), a much higher percentage
(72%) of those females held with males and fruit remated. This finding appears to be
similar to that known for other species of tropical frugivorous Tephritidae. Sivinski &
Heath (1989) reported only 10% of female A. suspense remated when held the first
week with males and no oviposition site, but 60% of those females held the first week
with both males and oviposition sites remated. Sixty percent of Mediterranean fruit
fly females held in cages with fruit for oviposition remated (Nakagawa et al. 1971) and
fifty percent of female Anastrepha ludens (Loew) provided wax oviposition domes re
mated (Robacker et al. 1985).
It is not yet known why a high percentage of papaya fruit fly females remated in
this study. Multiple mating in tephritid fruit flies is thought to be due to 1) forced mat
ings by males controlling access to oviposition sites, as in Rhagoletis pomonella
(Prokopy & Roitberg 1984), 2) poor sperm transfer in initial matings, as suspected in
part for C. capitata (Mazomenos et al 1977), or 3) sperm depletion following extensive
oviposition, as in A. suspense (Sivinski & Heath 1988). The possibility of transfer of
nutrients with male ejaculate was tested for A. suspense (Sivinski & Smittle 1987),









Florida Entomologist 77(3)


40 -


20



10




8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16 16-17
TIME OF DAY


8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16 16-17
TIME OF DAY
Fig. 3. Percent of matings by females held without fruit (crosshatched bars) or
with fruit (open bars) at hourly intervals throughout the day (A), and percent of ovi
positions observed at hourly intervals throughout the day (B).


September, 1994









Landolt: Papaya Fruit Fly Mating Frequency


but with no evidence of a significant male contribution. In the case of the papaya fruit
fly, there is no clear indication which of these explanations may account for the ob
served multiple matings.
The male papaya fruit fly in the field defends fruit from other males and courts ar
riving females, conceivably controlling female access to oviposition sites. However, fe
males appear capable of resisting courting males, making it unlikely that multiple
matings by females are the result of forced copulations by males. The observed in
creased remating rates of females held with fruit, compared to those held without
fruit, indicate that poor sperm transfer during first matings was not a problem con
tributing to multiple mating. Insemination failures at first matings would be expected
to result in rematings with or without available oviposition sites. In other studies of
tephritid mating frequency, remating appeared to clearly relate to extended periods of
oviposition and was attributed to sperm depletion (Sivinski & Heath 1988; Cunning
ham et al 1971; Robacker et al 1985, Tzanakakis et al 1968). Remating by female pa
paya fruit flies clearly was associated with access to oviposition sites (immature
papaya fruit) and oviposition activity, suggesting a similar need to replace depleted
sperm stocks used during oviposition. However, papaya fruit fly females are relatively
short lived (Landolt & Hendrichs 1983) and deposit limited numbers of eggs (Knab &
Others 1914) compared to other frugivorous fruit flies. It is not known if males are
similarly limited in numbers of sperm they can deliver during insemination, thus re
quiring female remating following oviposition. An additional possibility is that female
papaya fruit flies arriving at good oviposition sites occupied by males may mate or re
mate as an expression of mate choice, since a male at the site has demonstrated suc
cess both in host location and territorial defense. Such success may involve heritable
traits in the male that are beneficial to the female's offspring.
Mated female papaya fruit flies may respond to male pheromone (Landolt & Heath
1988, 1990) both to locate potential oviposition sites and prospective mates. Previous
studies indicated that enhanced attraction by mated females occurred to a combine
tion of host odor and male pheromone (Landolt & Reed 1990, Landolt et al 1992). It
was suggested that such behavior may aid females attempting to locate host fruit, by
orienting to males calling from such fruit. If ovipositing female T curvicauda need to
remate following oviposition, such a strategy ensures the availability of both fruit and
a mate simultaneously for an arriving female.
Male papaya fruit flies in this study mated three times more often than females
during the same 5-day time period. This is not a measure of male lifetime fertility,
since males may live for considerably longer periods of time, but is an indication of
their potential relative to females for polygamy. These results are similar to those re
ported for male C. capitata by Nakagawa et al (1971), with males mating every sev
eral days at every opportunity provided.

ACKNOWLEDGMENTS

Technical assistance was provided by K. M. Davis-Hernandez. J. Burns and S.
Sapp kindly provided access to their papaya groves for collection of fruit and papaya
fruit flies.

REFERENCES CITED

CUNNINGHAM, R. T., G. F. FARIAS, S. NAKAGAWA, AND D. L. CHAMBERS. 1971. Repro
duction in the Mediterranean fruit fly: Depletion of stored sperm in females.
Ann. Entomol. Soc. America 64:312-313.









Florida Entomologist 77(3) September, 1994


KNAB, F., AND W. W. YOTHERS. 1914. The papaya fruit fly. J. Agric. Res. 2: 447-453.
LANDOLT, P. J. 1984. Reproductive maturation and premating period of the papaya
fruit fly, Toxotrypana curvicauda (Diptera: Tephritidae). Florida Entomol. 67:
240-244.
LANDOLT, P. J., AND R. R. HEATH. 1988. Effects of age, mating and time of day on be
havioral responses of female papaya fruit fly, Toxotrypana curvicauda Gers
taecker (Diptera: Tephritidae) to synthetic pheromone. Environ. Entomol. 17:
47-51.
LANDOLT, P. J., AND R. R. HEATH. 1990. Effects of pheromone release rate and time of
day on catches of male and female papaya fruit flies on pheromone-baited fruit
model traps. J. Econ. Entomol. 83: 2040-2043.
LANDOLT, P. J., AND J. HENDRICHS. 1983. Reproductive behavior of the papaya fruit
fly, Toxotrypana curvicauda Gerstaecker (Diptera: Tephritidae). Ann. Entomol.
Soc. America 76: 413-417.
LANDOLT, P. J., AND H. C. REED. 1990. Behavior of the papaya fruit fly (Diptera: Te
phritidae): host finding and oviposition. Environ. Entomol. 19: 1305-1310.
LANDOLT, P. J., H. C. REED, AND R. R. HEATH. 1992. Attraction of female papaya fruit
fly (Diptera: Tephritidae) to male pheromone and host fruit. Environ. Entomol.
21: 1154-1159.
MAZOMENOS, B., J. L. NATION, W. J. COLEMAN, K. C. DENNIS, AND R. ESPONDA. 1977.
Reproduction in Caribbean fruit flies: Comparisons between a laboratory
strain and a wild strain. Florida Entomol. 60: 139-144.
NAKAGAWA, S., G. J. FARIAS, D. SUDA, R. T. CUNNINGHAM, AND D. L. CHAMBERS. 1971.
Reproduction in the Mediterranean fruit fly: frequency of mating in the labor
tory. Ann. Entomol. Soc. America 64: 949-950.
PROKOPY, R. J., AND B. D. ROITBERG. 1984. Foraging behavior of true fruit flies. Amer
ican Scientist. 72: 4149.
ROBACKER, D. C., S. J. INGLE, AND W. G. HART. 1985. Mating frequency and response
to male-produced pheromone by virgin and mated females of the Mexican fruit
fly Southwest. Entomol. 10: 215-221.
SIVINSKI, J., AND R. R. HEATH. 1988. Effects of oviposition on relating, response to
pheromones and longevity in the female Caribbean fruit fly, Anastrepha sus
pensa (Diptera: Tephritidae). Ann. Entomol. Soc. America 81: 1021-1024.
SIVINSKI, J., AND B. SMITTLE. 1987. Male transfer of materials to mates in the Carib
bean fruit fly, Anastrepha suspense (Diptera: Tephritidae). Florida Entomol.
70:233-238.
TZANAKAKIS, M. E., J. A. TSITSIPIS, AND A. P. ECONOMOPOULOS. 1968. Frequency of
mating in females of the olive fruit fly under laboratory conditions. J. Econ. En
tomol. 61:1309-1312.









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida313

ANNOTATED CHECKLIST
OF THE BARK AND AMBROSIA BEETLES
(COLEOPTERA: PLATYPODIDAE AND SCOLYTIDAE)
OF TROPICAL SOUTHERN FLORIDA

THOMAS H. ATKINSON'3AND STEWART B. PECK2
'Entomology and Nematology Department, University of Florida
Gainesville, FL USA 32611

2Department of Biology, Carleton University
Ottawa, Ontario, Canada K1S 5B6

ABSTRACT

The fauna of Scolytidae and Platypodidae is reviewed for tropical southern Florida
(Collier, Broward, Dade, and Monroe Counties). The family Platypodidae is repre
sented by 3 species, all in the genus Platypus. The family Scolytidae includes 83 spe
cies in 37 genera in the region. This total includes 20 species considered immigrants
to the area. Three species previously reported from the region, Cryptocarenus spatu
latus Wood, Xyleborus xylographus (Say), and Araptus politus (Blandford), probably
do not occur there. Feeding habits, mating systems, hosts, and distributions are sum
marized for all species included.

Key Words: Taxonomy, distributions, ecology, hosts, introduced species

RESUME

Se revisa la fauna de las families Scolytidae y Platypodidae de la region tropical
del sur de Florida (condados de Collier, Broward, Dade y Monroe). La familiar Platypo
didae esta representada por 3 species, todas en el genero Platypus. La familiar Scoly
tidae esta representada por 83 species en 37 generous. Este total incluye 20 species
las cuales se consideran inmigrantes al area. Tres species previamente reportadas de
la region, Cryptopcarenus spatulatus Wood, Xyleborus xylographus (Say) y Araptus
politus (Blandford), probablemente no existen alli. Habitos alimenticios, sistemas de
apareamiento y distribuciones se resume para todas las species incluidas.





Bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) constitute a
diverse group of beetles that bore in a variety of woody tissues and are well repre
sented in most temperate and tropical forests. Best known are species of Dendrocto
nus, Ips, and Scolytus that are primary pests of conifers in high-latitude forests,
although most of these are atypical of the group as a whole. All species breed in live,
stressed, or newly-killed host material. Adults excavate a system of galleries within
which eggs are laid and immature development occurs. The greater part of the life cy
cle takes place within host tissues, except for dispersal between hosts. A fascinating
range of patterns of host plant utilization (tissues consumed, relationships with fun
gal symbionts, and host specificity) and mating behaviors exists within this basic life

SCurrent address: Dow Elanco, 13355 Noel Rd., Suite 1045 Dallas, TX 75240-6604



This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
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Florida Entomologist 77(3) September, 1994


history paradigm. Most species leave a persistent physical record of their life history
and activities in the woody tissues of their hosts that can be interpreted by a student
of the group. As a consequence, one can easily tabulate several important biological
parameters for a large proportion of the species in an area, including those that have
not been studied in detail.
The taxonomy of the group is well-known (Wood 1982, Wood & Bright 1993) for the
United States as a whole, but there is little information available from southern Flor
ida, largely due to its remoteness. Some collecting was done near the turn of the cen
tury by E. A. Schwarz and A.D. Hopkins. S. L. Wood collected there in the 1950's.
Besides ourselves, more recent collectors of note include D.E. Bright, M. A. Deyrup, R.
H. Turnbow, E. G. Riley and R. A. Anderson. Bright's (1985) checklist of the Caribbean
highlighted the paucity of available information from that region. No student of the
group has ever collected extensively on any of the Greater Antilles.
The physical, historical, and floristic geography of southern Florida has been re
viewed by Peck (1989). The area is geologically recent and has a subtropical climate.
It has had direct and continuous land contact with temperate areas, but has never
had any land connections with any tropical areas that would serve as source areas for
its current biota. Most of the Neotropical flora of southern Florida has apparently dis
persed there by wind, water, or birds from the Bahamas and Greater Antilles (Tom
linson 1980).
This checklist represents a convergence of 2 separate projects, a faunal study of
the entire group for the southeastern United States by the first author and a biogeo
graphic study of the insect fauna of tropical southern Florida by the second author
(Peck 1989). The purpose of this paper is to document the scolytid and platypodid
fauna of tropical southern Florida for a subsequent ecological and biogeographical
analysis.


METHODS

The second author has collected extensively in the region and accumulated large
numbers of specimens of bark and ambrosia beetles as part of a large scale project on
the biogeography of the insect fauna of tropical southern Florida (Peck 1989). Sam
pling was done with flight intercept traps that were left in place over extended periods
for 5 years in areas of natural vegetation on the Keys and mainland (sites described
in Peck 1989). Over 2,500 specimens of Scolytidae and Platypodidae were collected.
Most specimens and representatives of all species collected by the second author were
deposited in the Canadian National Collection, the Canadian Museum of Nature, and
the S.B. Peck collection (all in Ottawa, Ontario). Some representative specimens were
deposited in the Florida State Collection ofArthropods, Gainesville, Florida.
The first author has been involved in a faunal study of the Scolytidae and Platy
podidae of the southeastern United States for several years. A total of 1,250 speci
mens (other than those collected by the second author) were examined from southern
Florida, as well as an additional 3,000 specimens of the same species from other parts
of the United States, Canada, the Caribbean, and Latin America. Scolytidae and
Platypodidae were examined by the first author from the following collections (abbre
viations used are from Arnett & Samuelson (1986)): Archbold Biological Station, Lake
Placid, Florida (ABSC); Canadian Museum of Nature Collection, Ottawa, Ontario
(CMNC); E. G. Riley private collection, College Station, Texas (EGRC), Department of
Forestry and Environmental Science, State University of New York, Syracuse, New
York (DFEC), Florida State Collection of Arthropods, Gainesville, Florida (FSCA),
Henry and Ann Howden collection, Ottawa, Ontario, (CHAH), Museum of Compara









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida315

tive Zoology, Cambridge, Massachusetts (MCZC), R.H. Turnbow private collection, Ft.
Rucker, Alabama (RHTC), S.L. Wood private collection, Provo, Utah (SLWC); T.H. At
kinson private collection, Riverside, California (THAC), University of Georgia at Ath
ens Georgia (UGCA), and U. S.. National Museum of Natural History, Washington,
D.C. (USNM).
In addition to data taken from museum specimens, the relevant literature on the
Scolytidae and Platypodidae was critically reviewed by the first author for specific dis
tribution records in southern Florida, as well as host records and distribution records
for the entire ranges of the species included.


ADVENTIVE SPECIES

Exactly what constitutes an adventive species and unambiguous criteria by which
one might judge particular cases are matters of considerable debate (Frank & McCoy
1990, 1992, Whitehead & Wheeler 1990). Whitehead & Wheeler (1990) argued that as
a general criterion one should consider species to be adventive if newly reported (or
detected) in the United States. They recognized that southern Florida had not been
well-collected historically and that new records for Antillean species did not necessary
ily imply that these species were non-native. Frank & McCoy followed this criterion
literally in their recent list of new immigrants into Florida (1992) (i.e., new record
means new arrival), although they also acknowledged the lack of complete historical
information on the insect fauna of tropical southern Florida.
Part of the problem is semantic (Frank & McCoy 1990). Many terms have been
used inconsistently in the literature to indicate natural occurrence, occurrence
through human intervention (accidental or not), and various types of restricted dis
tributions. Frank & McCoy (1990) restricted the term "introduced" to mean species
deliberately brought into a new area. They used the term "immigrant" to refer to spe
cies that have arrived and become established without intentional human activities.
Their definition is deficient in an important sense in that it does not distinguish be
tween the accidental establishment of species through human agency and the move-
ment and establishment of species into historically new areas by natural means of
dispersal, passive or active. To further complicate matters, there is no reason to sup
pose that the process of accumulation of the biota of southern Florida through "natu
ral means" has come to a stop.
This distinction is not trivial for southern Florida. Given that southern peninsular
Florida (including the Keys) has only become emergent in Wisconsonian and recent
geological periods (Tomlinson 1980, Peck 1989) and has never had any land connec
tions to the Greater Antilles, virtually any species with an Antillean or Neotropical
distribution currently found in Florida is immigrant by this definition. Nonetheless,
most of these occur there naturally and should be considered native. These would not
be considered "indigenous" as defined by Frank & McCoy (1990). The same would also
be true of any Neartic or Holarctic species occurring naturally in the Florida penin
sula as well, although it has been available for colonization by terrestrial organisms
since the mid-late Tertiary and Pleistocene. Some Nearctic and Holartic species have
only recently been reported from the peninsula (e.g., Deyrup & Atkinson 1987, Atkin
son et al. 1991), yet no one would think to call these "immigrants".
To avoid confusion, we consistently use the terms adventivee" and "immigrant" to
refer to species that do not naturally occur in southern Florida. None of the adventive
species known from the area have been deliberately introduced in the sense of Frank
& McCoy (1990). We use the term "native" to refer to any species which we believe oc
curs naturally in southern Florida whether "indigenous" (sensu Frank & McCoy 1990)









Florida Entomologist 77(3) September, 1994


or "immigrant" in the sense of having dispersed to Florida as part of a natural dis
persal process not mediated by deliberate or accidental human activity.
There is no evidence that any Neotropical species of Scolytidae or Platypodidae
has been introduced, intentionally or accidentally, into Florida in modern times. Such
an event might have occurred more than 150 years ago and passed unnoticed (i.e., be
fore the publication dates of works treating the area). Neotropical adventives may be
recognized by any combination of the following criteria: (1) the main part of the spe
cies' historical distribution does not include the Caribbean; (2) strict association with
a non-native host plant; (3) clear association with disturbed habitats and / or absence
from natural communities; and (4) documented invasion and / or subsequent spread.
Several species that originated in the Old World tropics have become established in
this century in the United States and have shown an historical pattern of distribution
consistent with spread from a single point, even when the original invasion and colo
nization passed unremarked. Examples include Xylosandrus germanus (Blandford)
(Bright 1968, Chapin & Oliver 1986, Staines 1984, Weber & MacPherson 1982), X.
compactus (Eichhoff) (Dixon & Woodruff 1982, Ngoan et al. 1976), X. crassiusculus
(Motschulsky) (Anderson 1974, Chapin & Oliver 1986, Deyrup & Atkinson 1987, Ko
vach & Gorsuch 1985, Atkinson et al. 1991), and Xyleborus atratus (Eichhoff) (Atkin
son et al. 1990, 1991). In all of these cases, either the historical pattern of spread can
be documented from collection records (Xylosandrus spp.) or else a "novel species"
known from another continent is detected over a wide area almost simultaneously
(Xyleborus atratus).
We maintain that any species that is currently found in southern Florida and in
the Bahamas and / or the Greater Antilles should be considered native (although al
most certainly immigrant) unless there is evidence to the contrary. There are 24 spe
cies in our area that have this distribution pattern. By the same token, any
Neotropical species whose modern range includes the Gulf Coast of the southeastern
United States and Texas and is continuously distributed into Mexico and Central
America should also be considered native (12 species, some of which are also widely
distributed in the eastern United States and / or the Caribbean).
By our restricted criteria, we consider 20 species of Scolytidae found in southern
Florida to be immigrants, mostly from Africa or Asia. Frank & McCoy (1992), by a me
chanical application of the rule of "recent record means new immigrant ", listed 3
other species as immigrants (Pseudothysanoes securigerus (Blackman), Theoborus
solitariceps Schedl, and Trischidias exigua Wood) that we consider natives. Wood
(1977, 1982) listed an additional 14 Neotropical species as immigrants with distribu
tions that fit one of the 2 patterns described above. Cryptocarenus '. ... I i . .1. i.
C. seriatus I --. i I Xyleborus volvulus (F), Xylosandrus zimmermanni (Hopkins),
and Corthylus spinifer Schwarz have ranges that include the Antilles and South Flor
ida. Hypothenemus eruditus Westwood, H. seriatus (Eichhoff), Xyleborus affinis Eich
hoff, and X. ferrugineus (F) are continuously distributed along the Gulf Coast into
Mexico and Central America. In contrast, other Neotropical species with similar dis
tributions were treated as native. Dendrosinus bourreriae Schwarz, Chaetophloeus
insularis (Blackman), Pycnarthrum hispidum I i n i Scolytodes schwarzi (Hop
kins), and Ambrosiodmus lecontei Hopkins are found in the Antilles and South Flor
ida. Cnesinus strigicollis LeConte is continuously distributed along the Gulf Coast
through Texas and into Mexico. In neither publication were the criteria explained by
which this conclusion was reached. All of these putative "Neotropical exotics" are host
generalists (polyphagous) and many reproduce by inbred polygyny. As noted by Wood
(1977), this is true of many species recently established in North America, but it does
not constitute a priori evidence that a particular species is non-native.









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida317

EXCLUDED SPECIES

Three species previously reported from southern Florida, Cryptocarenus spatula
tus Wood, Xyleborus xylographus (Say), and Araptus politus (Blandford), probably do
not occur there. Cryptocarenus spatulatus and Araptus politus were intercepted com-
ing into Miami from Latin America (Wood 1982), but no specimens of either species
have ever been collected from local populations. Araptus politus is a host specialist
and breeds in seeds of Mucuna spp. (Leguminosae) which are not known to occur in
Florida. Xyleborus xylographus was cited by Bright (1968) and Wood (1982) from Key
West (presumably from the same specimen(s) in the Canadian National Collection).
This species breeds in oaks and no other authentic specimens have ever been found in
peninsular Florida despite extensive collecting by the first author in northern Florida.
The only species of oak that occurs in southern Florida is Quercus virginiana L., found
only in Dade County near Miami and on Key Largo (Long & Lakela 1971, Tomlinson
1980). The "Key West" material most likely represents an error in labelling or an in
perception.


ORGANIZATION OF CHECKLIST

The sequence of subfamilies and tribes follows Wood & Bright (1992). Species are
arranged alphabetically within genera, and genera alphabetically within tribes. Ab
breviations for feeding habits (host tissues consumed), degree of host specificity, and
mating system are listed in parenthesis for each species. Host information and overall
distribution patterns are also included. Abbreviations are used to indicate: (1) Feed
ingHabits: ph, phloem feeding (= true bark beetles) (phloeophagy); xm, feeding on ec
tosymbiotic fungi ( ambrosia beetles) (xylomycetophagy); xy, direct feeding on
sapwood (xylophagy); my, pith of twigs and branches (myelophagy); sp, seeds (sper
matophagy); myc, non-ambrosial fungi (mycophagy); (2) Host Specificity: mo, re
stricted to hosts of a single species or genus (monophagy); ol, restricted to hosts of a
single family (or to a very limited number of hosts) (oligophagy); po, not host specific
(polyphagy); (3) Mating Systems: mg, monogyny; hpg, harem polygyny (including big
amy); ipg, inbred polygyny (mating terminology follows Kirkendall 1983). A question
mark (?) after any abbreviation indicates an absence of data.
Collection localities are listed by county for the four southern Florida counties in
the following sequence: Collier, Broward, Dade and Monroe. Each locality is followed
by a parenthetical reference to the collections) in which specimens are deposited (ab
breviations listed in Methods) or a literature reference. Complete collection data are
not included because of space limitations, but are available from the authors. Speci
mens were examined by the first author for all localities for which a collection is des
ignated.


ANNOTATED CHECKLIST OF THE PLATYPODIDAE AND SCOLYTIDAE
OF SOUTHERN FLORIDA


Family Platypodidae

Platypus compositus Say. (xm, po, mg). Found in large trunks and stumps of most
hardwoods within its range, also bald cypress, Taxodium distichum (L.) Rich.. Widely
distributed in the Neotropics, apparently adventive to Africa. Southeastern U.S. from









Florida Entomologist 77(3) September, 1994


Texas eastward. Collier: Royal Palm Hammock St. Park (USNM); Dade: Paradise
Key (USNM); Monroe: Big Pine Key (CMNC); Sugarloaf Key (CMNC).
Platypus flavicornis (F.). (xm, mo, mg). Found in trunks and stumps of all species
of Pinus (Pinaceae) within its range. Southeastern U.S., from eastern Texas. Dade:
Homestead (FSCA); Opa-locka (FSCA).
Platypus parallelus (F). (xm, po, mg). Breeds in large diameter host material of
virtually any woody plant within its range. It has also been reported from palm
trunks. Widely distributed in the Neotropics. In the U.S. found only in southern Flor
ida and southern Texas. Broward: Fort Lauderdale(FSCA); Collier: Collier Semi
nole St. Park (FSCA); Marco Island (FSCA); 8 mi SE Naples, (USNM); Dade:
Everglades Natl. Park, Royal Palm Hammock (CMNC); Homestead (FSCA); 6 mi N
Homestead, (USNM); Miami (FSCA); Miami Beach (FSCA); Perrine (FSCA); West Mi
ami (FSCA); Monroe: Big Pine Key (EGRC); Big Torch Key (CMNC); Cape Sable
(FSCA); John Pennekamp St. Park (ABSC); Key Largo (USNM, RHTC); Key West
(FSCA); No Name Key (EGRC); Plantation Key (FSCA); Stock Island (FSCA, USNM).


Family Scolytidae
Subfamily Hylesininae
Tribe Hylastini

Hylastes salebrosus Eichhoff. (ph, mo, mg). Found in roots, stumps, or branches in
contact with the soil of most Pines within its range. Southeastern U.S., from Texas
eastward. Dade: "Biscayne Bay" (Wood 1982).
Hylastes tenuis Eichhoff. (ph, mo, mg). Found in roots, stumps or branches in con
tact with the soil of most Pines within its range. Southeastern U.S., from Texas east
ward, also known from Hispaniola. Southwestern U.S. to central Mexico in montane
regions. Monroe: Key West (USNM).


Tribe Tomicini

Dendroctonus terebrans (Olivier). (ph, mo, mg). Found in stumps, large roots, and
the lower portion of large pines. Southeastern U.S. from eastern Texas to Atlantic sea
board. Broward: Fort Lauderdale (Wood 1982); Dade: Kendall (FSCA); Opa-locka
(FSCA).


Tribe Bothrosternini

Cnesinus strigicollis LeConte. (my, po, mg). Very polyphagous, breeding in twigs of
a wide variety of plant species. Southeastern U.S. from eastern Texas to lower Atlan
tic seaboard. In Mexico along Gulf Coast to Yucatan Peninsula. Dade: Elliot Key
(ABSC); Miami (FSCA); Deering Estate (CMNC); Matheson Hammock (CMNC);
Monroe: Key Largo (UGCA, CMNC, Wood 1982); N. Key Largo (ABSC); Plantation
Key (FSCA, ABSC).
Pagiocerus frontalis (F.) (sp, mo, mg). Breeds in fleshy seeds of several genera of
Lauraceae, notably Persea. Widely distributed in lowland Neotropical areas. In U.S.
found along lower Gulf Coast and lower Atlantic seaboard to North Carolina. It has
not been collected in the Keys but native and exotic species of Persea (avocado) do oc
cur there (Long & Lakela 1971, Tomlinson 1980). Collier: Monroe Sta. (ABSC);
Dade: Chekika St. Rec. Area, 50 km SW Miami (CMNC); Miami, Deering Estate
(CMNC).









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida319

Tribe Phloeotribini

Phloeotribus texanus Schaeffer. (ph, mo, mg). Breeds in branches of Celtis spp (Ul
maceae). Southeastern U.S. from Texas eastward; lowland regions of Mexico. Collier:
Collier Seminole St. Park (ABSC); Dade: "Biscayne" (Wood 1982).

Tribe Phloeosinini

Dendrosinus bourreriae Schwarz (xy, po, mg). Reported from branches of several
unrelated families of tropical hardwoods. Known only from southern Florida and
Greater Antilles. This species has not been collected from the mainland although
some of its reported hosts are found in tropical hardwood hammocks there (Long &
Lakela 1971, Tomlinson 1980). Monroe: Key Largo (FSCA); Stock Island (FSCA).
Phloeosinus taxodii taxodii Blackman. (ph, mo, mg). Breeds in branches of bald cy
press, Taxodium distichum (Taxodiaceae). Southeastern U.S. from eastern Texas. A
different subspecies, P taxodii taxodiicolens Wood, is found throughout Mexico in
Montezuma cypress, Taxodium mucronatum Ten. Collier: Naples, 8 mi SE (USNM).

Tribe Hypoborini

Chaetophloeus insularis (Blackman). (ph, mo, mg). Breeds in branches of native
and exotic Manilkara spp. (Sapotacae). Southern Florida and Greater Antilles. This
species has not been collected from the mainland although its native host, Manilkara
bahamensis (Baker) Lam. & Meeuse, is found in there and an exotic host, M zapota
(L.) Royen is grown widely in southern Florida (Long & Lakela 1971, Tomlinson
1980). Monroe: Big Pine Key, Cactus Hammock (CMNC); Big Torch Key (CMNC);
Cudjoe Key (CMNC); Fat Deer Key (CMNC); Key West (Wood 1982); Lower Mate
cumbe Key (Wood 1982); No Name Key (CMNC); Sugarloaf Key (CMNC, Wood 1982).

Subfamily Scolytinae
Tribe Scolytini

Cnemonyxficus (Schwarz). (ph, mo?, mg). The type series was reportedly collected
from Ficus (Moraceae), probably due to an error in identification. The true host is the
manchineel tree, Hippomane mancinella L. (Euphorbiaceae) (Atkinson 1993). This
species breeds in larger branches (> 3 cm diameter) and trunks of its host tree. The
host occurs throughout the Keys and on the mainland near Cape Sable, although it is
not abundant (Long & Lakela 1971, Tomlinson 1980). Southern Florida, Bahamas,
Virgin Islands. Monroe: Big Pine Key (CMNC, THAC); Key West (MCZC, Wood
1982).
Cnemonyx vagabundus Wood. (ph, mo?, mg). The type series was reportedly col
elected from Piscidia piscipula (Leguminosae), probably due to an error in identifica
tion. The true host is the manchineel tree, Hippomane mancinella L. (Euphorbiaceae)
(Atkinson 1993). This species breeds in smaller branches (< 3 cm diameter) and
trunks of its host tree. Southern Florida, Puerto Rico and Panama. Monroe: Big Pine
Key (CMNC, THAC); Key West (Wood 1982).

Tribe Ctenophorini

Pycnarthrum hispidum (Ferrari). (ph, mo, mg). Branches of native and exotic figs
(Ficus spp.) (Moraceae). Widely distributed in lowland Neotropical areas. In the U.S.









Florida Entomologist 77(3) September, 1994


known only from southern Florida and southern Texas. Broward: 1 mi N Andytown,
(RHTC); Dade: "Biscayne" (MCZC); Coconut Grove (MCZC); Homestead (SLWC); Mi
ami (FSCA); Monroe: Key Largo (UGCA); Plantation Key (FSCA); Stock Isl. (FSCA);
Sugarloaf Key (SLWC).
Scolytodes schwarzi (Hopkins). (ph, mo, hpg). Breeds in shaded-out branches of
living Ficus spp. (Moraceae). Based on personal experience (THA), this species does
not breed in cut branches. Southern Florida, Greater Antilles, lowland regions of Mex
ico. Broward: 1 mi N Andytown (RHTC); Collier: Corkscrew Swamp Sanctuary
(ABSC); Dade: "Biscayne" (MCZC); Deering Estate Park (CMNC); Matheson Ham
mock (ABSC); Miami (Wood 1982); Monroe: Big Pine Key (CMNC); Everglades Natl.
Park (Wood 1982); Key Largo (ABSC, Wood 1982); Plantation Key (Wood 1982); Sug
arloaf Key (SLWC).


Tribe Micracini

Micracis swainei Blackman. (xy, po, hpg). Reported from branches of a variety of
hardwood species. It is commonly found in willow (Salix) in Florida. Southeastern
U.S. from eastern Texas, lowland regions of Mexico and Central America. Dade: 6 mi
N. Homestead (USNM).
Micracisella nanula (LeConte). (my, po, mg). Breeds in twigs of a variety of trees,
shrubs, and vines. Southeastern U.S. from Texas eastward. A closely related species,
M. opacithorax Schedl (possibly conspecific) is found in southern Texas and the Gulf
Coast of Mexico. Dade: "Biscayne" (DFEC, MCZC); Homestead (Wood 1982); Miami
(Wood 1982); Monroe: Big Pine Key (ABSC, Wood 1982); Key Largo (Wood 1982); Key
Vaca, Marathon (Wood 1982); Key West (Wood 1982, Blackman 1928); Missouri Key
(Wood 1982); N. Key Largo (ABSC); Sugarloaf Key (CMNC, Wood 1982).
Pseudothysanoes securigerus (Blackman). (?,?,?). Nothing is known about the hosts
or feeding habits of this species. Species of Pseudothysanoes may be either phloem or
sap-wood borers. All studied species are bigynous. Southern Florida, Hispaniola.
Dade: Everglades Natl. Park, Long Pine Key (CMNC); Monroe: Big Pine Key
(CMNC); No Name Key (CMNC).
Thysanoes fimbricornis LeConte. (xy, po, hp). Breeds in branches of a variety of
hardwood species. Southeastern U.S. from eastern Texas. Found in lowland areas of
eastern and southeastern Mexico. Dade: "Biscayne" (Blackman 1928).


Tribe Ipini

Ips avulsus (Eichhoff). (ph, mo, hp). Breeds in branches and crowns of all pines
within its range. Southeastern U.S. from eastern Texas. Dade: "Biscayne" (MCZC);
Miami (MCZC).
Ips calligraphus (Germar). (ph, mo, hp). Breeds in trunks and large branches of all
pines within its range. Eastern U.S. and southeastern Canada, Montane regions of
the southwestern U.S., Mexico, Guatemala and Honduras. Cuba and Hispaniola.
Dade: Everglades Natl. Park, Paradise Key (Hopping 1965b); Monroe: Key Largo
(Hopping 1965b); Key West (Hopping 1965b).
Ips grandicollis (Eichhoff). (ph, mo, hpg). Breeds in branches and trunks of all
pines within its range. Eastern U.S. and southeastern Canada, Montane regions of
the southwestern U.S., Mexico, Guatemala and Honduras. Cuba and Hispaniola.
Dade: Biscayne (MCZC); Everglades Natl. Park, Long Pine Key (CMNC); Everglades
Natl. Park, Paradise Key (Hopping 1965a); Homestead (Hopping 1965a); Monroe:
Big Pine Key (CMNC).









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida321

Tribe Dryocoetini

Coccotrypes advena (Blandford). (sp, po, ipg). Breeds in seeds of a variety of plants.
Widely distributed in Old World tropics, adventive in New World. Dade: Coral Gables
(FSCA). This species was intercepted in imported seeds and may not actually be es
tablished in Florida.
Coccotrypes carpophagus (Hornung). (sp, ol, ipg). Breeds in seeds of palms, espe
cially Sabal palmetto (Walt.) Lodd. ex Schultes and Washingtonia robusta Wendl.
Widely distributed in Old World tropics, adventive in New World. Dade: Coconut
Grove (Wood 1982); Deering Estate Park (CMNC); Everglades Natl. Park, Royal Palm
Hammock (CMNC); Homestead (ABSC); Matheson Hammock (CMNC); Monroe: Big
Pine Key (CMNC); Big Torch Key (CMNC); Cudjoe Key (CMNC); Fat Deer Key
(CMNC); Grassy Key (ABSC); Key Largo (CMNC); Key Vaca, Marathon (CMNC,
MCZC); No Name Key (CMNC); Plantation Key (ABSC); Sugarloaf Key (CMNC).
Coccotrypes cyperi (Beeson). (sp-ph, po, ipg). Extremely polyphagous. Breeds in
phloem and seeds of many hosts. Widely distributed in Old World tropics, adventive
in New World. Dade: Deering Estate Park (CMNC); Homestead (ABSC); Miami
(FSCA); Monroe: Big Pine Key (CMNC); Fat Deer Key (CMNC).
Coccotrypes dactyliperda (F). (sp, ol, ipg). Breeds in seeds of palms, especially
Phoenixspp. Widely distributed in Old World tropics, adventive in New World. Mon-
roe: Sugarloaf Key (CMNC).
Coccotrypes distinctus (Motschulsky). (sp, ol, ipg). Breeds in seeds of palms, espe
cially Phoenixspp., Sabal palmetto, and Washingtonia robusta. Widely distributed in
Old World tropics, adventive in New World. Collier: Collier Seminole St. Park
(ABSC); Dade: Chekika State Rec. Area (CMNC); Coconut Grove (Wood 1982); Deer
ing Estate Park (CMNC); Everglades Natl. Park, Long Pine Key (CMNC); Fairchild
Tropical Gardens (FSCA); Homestead (ABSC); Matheson Hammock (CMNC); Miami
(FSCA); Miami Beach (Wood 1982); Old Cutler Hammock (CMNC); Monroe: Planta
tion Key (ABSC); Stock Isl. (ABSC, CHAH, CMNC).
Coccotrypes rhizophorae (Hopkins). (sp, ol, ipg). Hosts: (Rhizophoraceae) Breeds in
expanding prop roots and propagules of its host, Rhizophora mangle L. All other spe
cies of Coccotrypes found in the New World are believed to be immigrants from Africa
or Asia. C. rhizophorae is a special case and possible exception because it may have
immigrated to the New World without human aid or intervention. Coccotrypes rhizo
phorae is also known from southeastern Asia in Rhizophora spp. Given that this bee
tle breeds in the fruits and developing seedlings of red mangrove which are capable
of floating long distances in salt water, it is entirely plausible that this insect was dis
persed to the New World by infested host propagules,just as its host presumably dis
persed to the New World by ocean currents (Tomlinson 1980). Collier: Everglades
City (USNM); Dade: Homestead (Wood 1982); Miami (MCZC, THAC); Miami, Deer
ing Estate (CMNC); Monroe: Key Largo (ABSC, USNM); Key West (Wood 1982).
Coccotrypes robustus Eichhoff (sp, ?, ipg). Reported from seeds of Euterpe sp. (Pal
mae). No host records from Florida. This species is known only from southern Florida
and the Greater Antilles. It is presumably of Old World origin as are all other species
in this genus. Dade: Deering Estate Park (CMNC); Monroe: Big Pine Key (CMNC);
Big Torch Key (CMNC); Cudjoe Key (CMNC); Fat Deer Key (CMNC); Key Vaca
(CMNC); No Name Key (CMNC); Sugarloaf Key (THAC).
Coccotrypes vulgaris (Eggers). (sp, po, ipg). Breeds in seeds of many hosts. Widely
distributed in Old World tropics, immigrant in New World. Dade: Everglades Natl.
Park, Long Pine Key (CMNC).
Dendrocranulus carbonarius (Hopkins). (my, ol?, mo?). All species of Dendrocran
ulus with known habits breed in the stems of cucurbit vines. Presumably D. carbon









Florida Entomologist 77(3) September, 1994


arius does so as well. The original host record from Annona is almost certainly due to
an error in labelling. Found in southern Florida and Caribbean Islands. This species
has not been collected in Florida since its original description. Dade: "Biscayne Bay"
(Wood 1982).

Tribe Crypturgini

Crypturgus alutaceus Schwarz. (ph, mo, mg). Breeds in phloem of dead or dying
pines (Pinaceae). Southeastern U.S. from eastern Texas to Atlantic seaboard. Collier:
Collier Seminole St. Park (Wood 1982); Monroe: Big Pine Key (Wood 1982).

Tribe Xyleborini

Ambrosiodmus devexulus (Wood). (xm, po, ipg). In wide variety of hosts. Dade:
Homestead (Wood 1982).
Ambrosiodmus lecontei Hopkins. (xm, po, ipg). Wide variety of hosts. Usually
found in branches or other small diameter material. Peninsular Florida and Carib
bean. Dade: "Biscayne" (MCZC); Everglades Natl. Park, Long Pine Key (CMNC); Ev
erglades Natl. Park, Royal Palm Hammock (CMNC); Homestead (FSCA, Wood 1982);
Miami (FSCA, Wood 1982); North Miami (FSCA); Monroe: Key Largo (CMNC, Wood
1982); Key Vaca (CMNC); Key West (Wood 1982); No Name Key (CMNC); Sugarloaf
Key (CMNC).
Ambrosiodmus obliquus (LeConte). (xm, po, ipg). Wide variety of hosts. Widely dis
tribute in lowland Neotropical areas. Southeastern U.S. Dade: Everglades Natl.
Park, Long Pine Key (CMNC); Everglades Natl. Park, Royal Palm Hammock
(CMNC); Homestead (Bright 1968); Monroe: Cudjoe Key (CMNC); No Name Key
(CMNC); Sugarloaf Key (CMNC).
Dryocoetoides sp. (xm, po?, ipg). Hosts unknown. A single specimen was collected
in a flight-intercept trap. It may represent an undescribed species because it does not
agree with any species of this genus previously reported from the Caribbean or Me
soamerica and is not of any species found in the USNM or SLWC. Monroe: Big Torch
Key (CMNC).
Premnobius cavipennis Eichhoff. (xm, po, ipg). Breeds in trunks of a wide variety
of hosts. Native to Africa, widely distributed in lowland Neotropical areas. In the U.S.,
found in southern Florida only. Collier: Collier Semenole St. Park (Bright 1968);
Dade: Chekika State Rec. Area (CMNC); Everglades Natl. Park, Long Pine Key
(CMNC); Everglades Natl. Park, Royal Palm Hammock (CMNC); Homestead (Bright
1968); Miami (Wood 1982); Monroe: Big Pine Key (CMNC); Flamingo Camp, Ever
glades (ABSC); Key Largo (CMNC); No Name Key (CMNC); Sugarloaf Key (CMNC).
Theoborus solitariceps Schedl. (xm, po, ipg). Found in branches of a wide variety
of hosts. Widely distributed in lowland Neotropical areas including the Caribbean.
Known only from a single collection in Florida. Dade: Deering Estate Park (CMNC).
Xyleborinus gracilis (Eichhoff). (xm, po, ipg). Breeds in a wide variety of hosts.
Lowland Neotropical areas. In U.S. known from Florida, Louisiana and North Caro
lina (Bright 1987). Dade: Biscayne Bay (Wood 1982); Everglades Natl. Park, Long
Pine Key (CMNC).
Xyleborinus saxeseni (Ratzeburg). (xm, po, ipg). Breeds in virtually any woody
plant within its range, including conifers. Eurasian species, immigrant in the New
World. Dade: Deering Estate Park (CMNC); Everglades Natl. Park, Long Pine Key
(CMNC).









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida323

Xyleborus affinis (Eichhoff). (xm, po, ipg). Breeds in trunks of most woody plants
within its range. Widely distributed in lowland Neotropical areas, southeastern U.S.
Dade: Chekika State Rec. Area (CMNC); Deering Estate Park (CMNC); Everglades
Natl. Park, Long Pine Key (CMNC); Everglades Natl. Park, Royal Palm Hammock
(CMNC); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Cudjoe Key
(CMNC); Key Largo (CMNC, RHTC); Key West (MCZC); Sugarloaf Key (CMNC).
Xyleborus ferrugineus (F.). (xm, po, ipg). Breeds in trunks of most woody plants in
its range. Widely distributed in lowland neotropical areas, eastern U.S., southeastern
Canada. Dade: Chekika State Rec. Area (CMNC); Coconut Grove (MCZC); Deering
Estate Park (CMNC); Everglades Natl. Park, Long Pine Key (CMNC); Everglades
Natl. Park, Royal Palm Hammock (CMNC); Monroe: Big Pine Key (CMNC); Big
Torch Key (CMNC); Cudjoe Key (CMNC); Fat Deer Key (CMNC); Key Largo (CMNC,
RHTC); Key Vaca (CMNC); No Name Key (CMNC); Sugarloaf Key (CMNC).
Xyleborus pubescens (Zimmermann). (xm, mo, ipg). Breeds in trunks of pines (Pi
naceae). This is one of the few species of this genus that is host specific. Southeastern
U.S. Collier: Collier Seminole St. Park (Wood 1982); Dade: Chekika State Rec. Area
(CMNC); Dade Co. (Wood 1982); Everglades Natl. Park, Long Pine Key (CMNC); Par
adise Key (Wood 1982); Monroe: Big Pine Key (CMNC); Key Largo (UGCA); Key
West (Bright 1968).
Xyleborus volvulus (F.). (xm, po, ipg). Breeds in trunks of most woody plants in its
range. Widely distributed in lowland Neotropical areas. In U.S. only in southern Flor
ida and southern Texas. Broward: Ft. Lauderdale (FSCA); Collier: Naples (FSCA,
Bright 1968); Dade: Biscayne Bay (Bright 1968); Chekika State Rec. Area (CMNC);
Coconut Grove (MCZC); Coral Gables (FSCA); Deering Estate Park (CMNC); Ever
glades Natl. Park, Long Pine Key (CMNC); Everglades Natl. Park, Royal Palm Ham
mock (CMNC); Homestead (FSCA); Miami (FSCA); Miami Beach (FSCA); Perrine
(FSCA); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Cudjoe Key
(CMNC); Fat Deer Key (CMNC); Key Largo (CMNC, FSCA, RHTC); Key Vaca
(CMNC); Key West (Bright 1968); Loggerhead Key (FSCA); No Name Key (CMNC);
Plantation Key (FSCA); Sugarloaf Key (CMNC).
Xyleborus xylographus (Say). (xm, mo, ipg). This species breeds in Oaks (Quercus
sp.) (Fagaceae). The record from Key West (Brightl968, Wood 1982) is either based on
a misidentification or an interception since this species is not known to breed any
where in peninsular Florida. (T.H.A., unpublished).
Xylosandrus compactus (Eichhoff). (xm, mo, ipg). Breeds in twigs of a wide variety
of living trees, shrubs, and vines resulting in the death of the twig. This species sel
dom breeds in cut material. Native to southeastern Asia. Found along Gulf Coast to
Texas. Broward: Ft. Lauderdale (Wood 1982); Collier: Collier Seminole St. Park
(ABSC); Copeland (ABSC); Dade: Chekika State Rec. Area (CMNC); Coconut Grove
(Wood 1982); Deering Estate Park (CMNC); Everglades Natl. Park, Long Pine Key
(CMNC); Everglades Natl. Park, Royal Palm Hammock (CMNC); Miami (Wood 1982);
Naranja (FSCA); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Cudjoe
Key (CMNC); Key Largo (CMNC, Wood 1982); N Key Largo (ABSC); No Name Key
(CMNC); Plantation Key (ABSC); Sugarloaf Key (CMNC).
Xylosandrus crassiusculus (Motschulsky). (xm, po, ipg). Breeds in stems of a wide
variety of trees and shrubs. This species frequently breeds in living stems, especially
near ground level. Secondary fungal invasions of old galleries may result in death of
the plant. Native to Asia. First detected in coastal South Carolina, now found west to
Texas (Atkinson et al. 1991). Collier: Collier Seminole St. Park (Deyrup & Atkinson
1987); Dade: Chekika State Rec. Area (CMNC); Deering Estate Park (CMNC); Ever









Florida Entomologist 77(3) September, 1994


glades Natl. Park, Long Pine Key (CMNC); Everglades Natl. Park, Royal Palm Ham
mock (CMNC); Old Cutler Hammock (CMNC).
Xylosandrus zimmermanni (Hopkins). (xm, po, ipg). Breeds in small diameter ma
trial. Not known to attack living hosts. Lowland Neotropical areas, southern Florida.
Dade: Biscayne Bay (Bright 1968); Everglades Natl. Park, Long Pine Key (CMNC);
Everglades Natl. Park, Long Pine Key (CMNC).

Tribe Cryphalini

Cryptocarenus heveae (Hagedorn). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Widely distributed in lowland Neotropical areas, southern Florida.
Collier: Collier Seminole St. Park (ABSC, Wood 1982); Dade: Everglades Natl. Park,
Long Pine Key (CMNC).
Cryptocarenus seriatus Eggers. (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Widely distributed in lowland Neotropical areas, peninsular Florida.
Collier: Collier Seminole St. Park (ABSC, Wood 1982)); Copeland (ABSC); Faka
hatchee Strand (ABSC); Ochopee (Wood 1982); Dade: "Biscayne" (Wood 1982); Deer
ing Estate Park (CMNC); Entrance Everglades Natl. Park (RHTC); Everglades Natl.
Park, Long Pine Key (CMNC); Hialeah (FSCA); Miami (FSCA); Paradise Key (Wood
1982); Monroe: Big Pine Key (Wood 1982); Big Torch Key (CMNC); Fat Deer Key
(CMNC); Grassy Key (Wood 1982); Key Largo (Wood 1982); Key Vaca (Wood 1982);
Key West (Wood 1982); Lower Matecumbe Key (Wood 1982); Stock Island (FSCA).
Cryptocarenus spatulatus Wood. (xm, po, ipg). The record from Miami (Wood 1982)
is based on an interception from Peru. There is no indication at present that it is ac
tually established in Florida.
Hypocryphalus mangiferae Schedl. (ph, mo, mg). Breeds in branches of mango
(Mangifera indica L.) (Anacardiaceae). Native to southern Asia, widely established in
tropical areas of the world where its host is grown, including southern Florida. Dade:
Homestead (Wood 1982); Perrine (Wood 1954).
Hypothenemus areccae (Hornung). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Immigrant from Old World tropics. Dade: Everglades Natl. Park,
Long Pine Key (CMNC); Homestead (FSCA).
Hypothenemus birmanus (Eichhoff). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Immigrant from Old World Tropics. Broward: Andytown, 1 mi N
(RHTC); Hollywood (ABSC); Collier: Collier Seminole St. Park (ABSC, Wood 1982);
Fakahatchee Strand (ABSC); Dade: Chekika State Rec. Area (CMNC); Deering Es
tate Park (CMNC); Everglades Natl. Park, Long Pine Key (CMNC); Everglades Natl.
Park, Royal Palm Hammock (CMNC); Homestead (ABSC, Wood 1982); Matheson
Hammock (THAC, Wood 1982); Miami (Wood 1982); Perrine (Wood 1982); Monroe:
Big Pine Key (CMNC); Cudjoe Key (CMNC); Key Largo (UGCA, CMNC, Wood 1982);
Key Vaca (CMNC); Lower Matecumbe Key (ABSC); N. Key Largo (ABSC); No Name
Key (CMNC); Sugarloaf Key (CMNC).
Hypothenemus brunneus (Hopkins). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Immigrant from Old World tropics. Collier: Copeland (ABSC);
Dade: "Biscayne" (MCZC); Everglades Natl. Park, Long Pine Key (CMN
C); Homestead (Wood 1982); Miami (Wood 1982); Monroe: Key Largo (Wood
1982); Key West (Wood 1982); Matecumbe Key (Wood 1982); Plantation Key (ABSC,
FSCA); Sugarloaf Key (Wood 1982).
Hypothenemus californicus Hopkins. (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Immigrant from Old World tropics. Dade: Homestead (Wood 1982);









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida325

Perrine (Wood 1982); Monroe: Key Largo (Wood 1982); Key Vaca (Wood 1982); Key
West (Wood 1982); Matecumbe Key (Wood 1982); Plantation Key (Wood 1982).
Hypothenemus columbi Hopkins. (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Immigrant from Old World tropics. Dade: Homestead (Wood 1982);
Perrine (Wood 1982).
Hypothenemus crudiae (Panzer). (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Immigrant from Old World tropics. Broward: Davie (FSCA); Collier:
Collier Seminole St. Park (ABSC); Monroe Sta. (ABSC); Dade: Chekika State Rec.
Area (CMNC); Elliot Key (ABSC); Everglades Natl. Park, Long Pine Key (CMNC); Hi
aleah (FSCA); Homestead (FSCA); Monroe: Big Pine Key (CMNC); Big Torch Key
(CMNC); Cudjoe Key (CMNC); Fat Deer Key (CMNC); Key Largo (CMNC, ABSC); No
Name Key (ABSC); Sugarloaf Key (CMNC).
Hypothenemus dissimilis (Zimmermann). (my, po, ipg). Breeds in pith of twigs of
a wide variety of hosts. Eastern U.S. Dade: "Biscayne Bay" (Wood 1982).
Hypothenemus eruditus Westwood. (ph-my, po, ipg). Breeds in pith or phloem of
twigs and branches of a wide variety of hosts. Widely distributed in lowland Neotro
pical areas, eastern U.S. Broward: Andytown, 1 mi N (RHTC); Collier: Collier Sem-
inole St. Park (ABSC); Copeland (ABSC); Dade: Chekika State Rec. Area (CMNC);
Deering Estate Park (CMNC); Elliot Key (ABSC); Everglades Natl. Park, Long Pine
Key (CMNC); Everglades Natl. Park, Royal Palm Hammock (CMNC); Homestead
(ABSC); Old Cutler Hammock (CMNC); Monroe: Big Pine Key (CMNC); Big Torch
Key (CMNC); Cudjoe Key (CMNC); Fat Deer Key (CMNC); Islamorada (UGCA); Key
Largo (CMNC); Key Vaca (CMNC); No Name Key (CMNC); Upper Matecumbe Key
(CMNC).
Hypothenemus_. , (Hopkins). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Mexico, Cuba, southern Florida. Dade: Everglades Natl. Park, Long
Pine Key (CMNC, THAC); Homestead (Wood 1982); Monroe: Key Largo (Wood 1982);
Key West (Wood 1982); Long Key (Wood 1982); Matecumbe Key (Wood 1982); Planta
tion Key (Wood 1982).
Hypothenemus hirsutus (Wood). (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Known from southern Florida only. Dade: Elliott Key (ABSC); Mon-
roe: Big Pine Key (CMNC, Wood 1982); Big Torch Key (CMNC); Cudjoe Key (CMNC);
Fat Deer Key (CMNC); Grassy Key (Wood 1982); Key Largo (Wood 1982); Key Vaca
(Wood 1982); Key West (Wood 1982); Matecumbe Key (Wood 1982); No Name Key
(ABSC, CMNC); Plantation Key (Wood 1982); Sugarloaf Key (CMNC, Wood 1982).
Hypothenemus interstitialis (Hopkins). (my, po, ipg). Breeds in pith of twigs of a
wide variety of hosts. Widely distributed in lowland Neotropical areas, southeastern
U.S. Collier: Collier Seminole St. Park (ABSC); Fakahatchee Strand (ABSC); Mon
roe Station (ABSC); Dade: Deering Estate Park (CMNC); Everglades Natl. Park,
Long Pine Key (CMNC); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Key
Largo (ABSC, CMNC, UGCA); Key Vaca (CMNC); No Name Key (ABSC); Sugarloaf
Key (CMNC).
Hypothenemusjavanus (Eggers). (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Immigrant from Old World. Broward: Delray Beach (Wood 1982); Hol
lywood (ABSC); Collier: Collier Seminole St. Park (Wood 1982); Dade: Chekika State
Rec. Area (CMNC); Coconut Grove (Wood 1982); Deering Estate Park (CMNC); Ever
glades Natl. Park, Long Pine Key (CMNC); Everglades Natl. Park, Royal Palm Ham
mock (CMNC); Homestead (Wood 1982); Miami (Wood 1982); Paradise Key (Wood
1982); Perrine (Wood 1982); Monroe: Key Largo (CMNC, Wood 1982); Key Vaca
(CMNC); N. Key Largo (ABSC); No Name Key (CMNC); Sugarloaf Key (CMNC).









Florida Entomologist 77(3) September, 1994


Hypothenemus miles (LeConte). (my?, ?, ipg). Hosts unknown. Gulf Coast from
Florida to Texas. Dade: Everglades Natl. Park, Long Pine Key (CMNC); Monroe:
Sugarloaf Key (CMNC).
Hypothenemus obscurus (F.). (my, po, ipg). Breeds in pith of twigs of a wide variety
of hosts. Immigrant from Old World. Dade: Homestead (FSCA); Miami (Wood 1982);
Monroe: Fat Deer Key (CMNC); Key Vaca (CMNC); No Name Key (CMNC); Sugar
loaf Key (CMNC).
Hypothenemus pubescens (Hopkins). (my, ol, ipg). Breeds in stems of grasses. Mex
ico, Puerto Rico, southern Florida, southern Texas. Dade: Everglades Natl. Park,
Long Pine Key (CMNC); Monroe: Cudjoe Key (CMNC); Fat Deer Key (CMNC); Key
Largo, Pennekamp St. Park (CMNC); Key Vaca (Wood 1982); Key West (Wood 1982);
Missouri Key (Wood 1982).
Hypothenemus seriatus (Eichhoff). (my, po, ipg). Breeds in pith of twigs of a wide
variety of hosts. Widely distributed on lowland Neotropical areas, eastern U.S. Bro-
ward: Davie (FSCA); Ft. Lauderdale (USNM); Dade: Everglades Natl. Park, Long
Pine Key (CMNC); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Cudjoe
Key (CMNC, FSCA); Fat Deer Key (CMNC); Islamorada (UGCA); Key Largo (UGCA,
CMNC); Key West (FSCA); No Name Key (CMNC); Stock Isl. (CMNC, FSCA); Sugar
loaf Key (CMNC); Windley Key (CHAH).
Hypothenemus setosus (Eichhoff). (my, po, ipg). Breeds in pith of twigs of a wide va
riety of hosts. Native to Old World, southern Florida. Dade: Miami (Wood 1982).
Hypothenemus squamosus (Hopkins). (my, po, ipg). Breeds in pith of twigs of a
wide variety of hosts. Lowland areas of Mexico, Central America, and the Antilles.
Southern Florida. Monroe: Key Largo (FSCA, Wood 1982); Matecumbe Key (Wood
1982).
Hypothenemus sp. (my, po, ipg). Breeds in pith of twigs of a wide variety of hosts.
Southern Florida, Cuba. Collier: 1 mi W Ochopee (THAC); Dade: Everglades Natl.
Park, Long Pine Key (CMNC); Homestead (ABSC); Monroe: Big Pine Key (CMNC);
Key Largo (CMNC); Key Vaca (CMNC). This species does not match any included for
North and Central America (Wood1982) or from the Caribbean (Bright 1985). It was
compared to all available types in the U.S. National Museum of Natural History (in
cluding types of Hopkins) and the S.L. Wood collection. It may be a previously unde
tected and undescribed native or Antillean species or may have been introduced from
the Old World.
Scolytogenes knabi (Hopkins). (ph-my, po?, mg). Breeds in pith of twigs of a wide
variety of hosts. Lowland Neotropical areas, Peninsular Florida. Collier: Collier
Seminole St. Park (ABSC, ABSC); Dade: Miami (Wood 1982); Chekika State Rec.
Area (CMNC); Monroe: Crawl Key (RHTC); Fat Deer Key (CMNC); Plantation Key
(Wood 1982); Sugarloaf Key (Wood 1982).
Trischidias atoma (Hopkins). (myc, ?, ipg). Breeds in fungus-infested twigs and
branches. Southeastern U.S. Collier: Copeland (ABSC); Dade: Everglades Natl.
Park, Long Pine Key (CMNC); Monroe: No Name Key (CMNC); Sugarloaf Key
(CMNC).
Trischidias exigua Wood. (myc, ?, ipg). Breeds in carbonaceous fruiting bodies of
ascomycete fungi on branches (Deyrup 1987). Known from southern Florida and the
Yucatan Peninsula. Dade: Everglades Natl. Park, Long Pine Key (CMNC); Old Cut
ler Hammock (CMNC); Monroe: Big Pine Key (CMNC); Big Torch Key (CMNC); Key
Largo (CMNC); Sugarloaf Key (CMNC).
Trischidias minutissima Wood. (myc, ?, ipg). Found in fungus pustules on aerial
roots of red mangrove. Known only from Keys. Monroe: Sugarloaf Key (SLWC).









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida327

Trischidias striatus Atkinson. (?, ?, ?). Hosts unknown. Known only from Keys.
Dade: Everglades Natl. Park, Long Pine Key(CMNC); Monroe: Big Pine Key
(CMNC); Big Torch Key (CMNC); Cudjoe Key (CMNC); Key Largo (CMNC); No Name
Key (CMNC); Sugarloaf Key (CMNC).

Tribe Corthylini

Araptus dentifrons Wood. (my, mo, mg). In stems of milkweed vine (Sarcostemma
clausum (Jacq.) R. & S.) (Asclepiadaceae). Known from several localities in Mexico,
southern Texas and southern Florida. Collier: Fakahatchee Strand (ABSC); Monroe
Station (ABSC).
Araptus politus (Blandford). This species was intercepted in Miami in "bird seed"
but there is no evidence that it has become established (Wood 1982). It breeds in the
large seeds of Mucuna spp. (Leguminosae) which are not known to occur in Florida.
Corthylus papulans Eichhoff (= spinifer Schwarz). (xm, po, mg). Breeds in
branches of a wide variety of hosts. Widely distributed in lowland neotropical areas,
peninsular Florida. Broward: Andytown, 1 mi N (RHTC); Collier: Copeland (ABSC);
Ochopee Trail Lake campsite (UGCA); Dade: Everglades Natl. Park, Long Pine Key
(CMNC); Miami (FSCA, Wood 1982); Paradise Key (Wood 1982); Monroe: Key Largo
(UGCA); Ramrod Key (Wood 1982); Sugarloaf Key (CMNC).
Monarthrum mali (Fitch). (xm, po, hpg). Breeds in trunks of a variety of hardwood
species. Widely distributed in eastern North America. Collier: Collier Seminole St.
Park (Wood 1982).
Pityoborus comatus (Zimmermann). (xm, mo, mg). In shaded-out branches of liv
ing pines. Southeastern U.S., Bahamas. Dade: "Biscayne Bay" (Wood 1982); Ever
glades Natl. Park, Long Pine Key (UGCA); Monroe: Big Pine Key (Wood 1982).
Pityophthorus annectens LeConte. (ph, mo, hpg). In branches of pines. Southeastern
U.S. Montane areas of southwestern U.S., Mexico, Central America, and Caribbean.
Dade: Homestead (Bright 1981); Monroe: Big Pine Key (Bright 1981).
Pityophthorus borrichiae Wood. (ph, mo, hpg). Stems of Borrichia arborescens (L.)
DC., B. frutescens (L.) DC. (Compositae). Known only from the Keys, although both
hosts are found on the mainland as well (Long & Lakela 1971, Tomlinson 1980). Mon-
roe: Key Largo (SLWC).
Pityophthorus concentralis Eichhoff. (ph, mo, hpg). Branches and trunks of Me-
topium toxiferum (L.) Krug & Urban (Anacardiaceae). Southern Florida, Cuba. Col-
lier: Royal Palm Hammock St. Park (Bright 1981); Dade: "Biscayne Bay" (MCZC,
Wood 1982); Deering Estate Park (CMNC); Everglades Natl. Park, Long Pine Key
(CMNC); Everglades Natl. Park, Royal Palm Hammock (CMNC); Paradise Key
(Bright 1981); Monroe: Big Pine Key (CMNC, Bright 1981); Big Torch Key (CMNC);
Fat Deer Key (CMNC); Everglades Natl. Pk., Flamingo (RHTC); Key Largo (THAC,
UGCA); Key Vaca (Wood 1982, Bright 1981); Key West (Wood 1982); No Name Key
(ABSC, CMNC); Plantation Key (Wood 1982, Bright 1981); Stock Isl. (Wood 1982);
Sugarloaf Key (CMNC, Bright 1981, Wood 1982).
Pityophthorus crinalis Blackman. (ph, mo, hpg). Toxicodendron radicans (L.)
Kuntze (Anacardiaceae). This species has not been collected in the Keys although its
host is found there (Long & Lakela 1971). Southeastern U.S. Collier: Collier Semi
nole St. Park (ABSC); Dade: S. Miami, Deering Estate Park (CMNC).
Pityophthorus lautus Eichhoff. (ph, mo, hpg). Toxicodendron radicans (Anacardi
aceae). Widely distributed in eastern North America, possibly a species complex.
Specimens referred to Pityophtorus lautus have been collected in several unrelated









Florida Entomologist 77(3) September, 1994


hosts in the eastern United States including Cercis canadensis L. (Leguminosae),
Acer saccharinum L. (Aceraceae), Sassafras albidum (Nutt.) Nees. (Lauraceae), Ul
mus rubra Muhl. (Ulmaceae), Pinus strobus L. (Pinaceae), as well as several hosts in
the Anacardiaceae. At first glance this might appear to be a polyphagous species ex
cept that it does not normally occur in any hosts other than those mentioned above
and their close relatives. All specimens referable to this species collected in peninsu
lar Florida have only been collected in poison ivy, despite the fact that some of these
other hosts also occur in the same areas (TH. A., unpublished data). It may occur in
the Keys as well as on the mainland since poison ivy is found on the islands (Long &
Lakela 1971. Collier: Copeland (ABSC).
Pityophthorus pulicarius (Zimmermann). (ph-my, mo, hpg). In phloem and pith of
branches and twigs of pines (Pinaceae). Southeastern U.S. Collier: Royal Palm Ham
mock St. Park (Bright 1981); Dade: Biscayne (MCZC); Everglades Natl. Park, Long
Pine Key (CMNC); Homestead (Bright 1981); Paradise Key (Bright 1981); Monroe:
Big Pine Key (ABSC, CMNC, Bright 1981).
Pityopthorus pecki Atkinson. (ph?, ?, hpg?). Hosts unknown. Known only from the
Keys. Monroe: Big Pine Key (CMNC); No Name Key (CMNC).


ACKNOWLEDGMENTS

Travel by the first author to visit the Museum of Comparative Zoology was sup
ported by an Ernst Mayr Grant from that institution and to Provo, Utah to visit the
S. L. Wood collection by a grant from the American Philosophical Society. Field work
of S. Peck in southern Florida was partially supported by research grants to S. B.
Peck from the Natural Sciences and Engineering Research Council of Canada. Jaram
ila Peck greatly aided in making collections under most unpleasant conditions of heat,
humidity, and mosquitoes in the hardwood hammock forests. This is Florida Agricul
tural Experiment Station Journal Series No. R-03178.


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United States (Coleoptera, Scolytidae). U.S. Dept. Agric. Coop. Econ. Ins. Rept.
24(45-48): 863-864.
ARNETT, R., AND P.A. SAMUELSON. 1986. The Insect and Spider Collections of the
World. Brill/ Flora & Fauna Publications. Gainesville.
ATKINSON, T.H. 1993. Rediscovery of 2 neotropical bark beetles (Coleoptera: Sco
lytidae) from southern Florida. Coleopterists Bull. (in press).
ATKINSON, T.H., R.J. RABAGLIA, AND D.E. BRIGHT. 1990. Newly detected exotic spe
cies of Xyleborus (Scolytidae) in eastern North America, with a revised key to
species. Canadian Entomol. 122: 93-104.
ATKINSON, T. H., R. J. RABAGLIA, S. J. PECK, AND J. L. FOLTZ. 1991. New records of
Scolytidae and Platypodidae from the U.S. and Bahamas. Coleopterists Bull.
45: 152-164.
BLACKMAN, M. W. 1928. Notes on Micracinae with description of twelve new species.
New York St. Coll. For., Syracuse, Tech. Pub. 25: 185-208.
BRIGHT, D.E. 1968. Review of the tribe Xyleborini in America north of Mexico (Co
leoptera: Scolytidae). Canadian Entomol. 100: 1288-1323.
BRIGHT, D.E. 1981. A taxonomic monograph of the genus Pityophthorus Eichhoff in
North America (Coleoptera: Scolytidae). Entomol. Soc. Canada Mem. 118: 1
378.
BRIGHT, D.E. 1985. Studies on West Indian Scolytidae (Coleoptera) 3. Checklist of Sc
olytidae of the West Indies, with descriptions of new species and taxonomic
notes. Entomol. Arb. Mus. Frey 33/34: 169-187.









Atkinson & Peck:Bark and Ambrosia Beetles of South Florida329

BRIGHT, D.E. 1987. Notes on the occurrence of Xyleborinus gracilis (Eichhoff) in the
United States (Coleoptera: Scolytidae). Coleopterists Bull. 41: 338.
CHAPIN, J. B., AND A. D. OLIVER. 1986. New records for Xylosandrus and Xyleborus
species (Coleoptera: Scolytidae). Proc. Entomol. Soc. Washington 88: 680-683.
DEYRUP, M.A. 1987. Trischidias exiguaWood, new to the Untied States, with notes on
the biology of the genus (Coleoptera: Scolytidae). Coleopterists Bull. 41: 339
343.
DEYRUP, M.A., AND T.H. ATKINSON. 1987. New records of Scolytidae (Coleoptera) from
Indiana and Florida. Great Lakes Entomol. 20: 67-68.
DIXON, W. N., AND R. E. WOODRUFF. 1982. The black twig borer, Xylosandrus compac
tus (Eichhoff) (Coleoptera: Scolytidae). Fla. Dept. Agric. Consumer Serv., Div.
Plant Indus. Entomol. Circ. No. 250, 2 p.
FRANK, J.H., AND E.D. MCCOY. 1990. Endemics and epidemics of shibboleths and
other things causing chaos. Florida Entomol. 73: 19.
FRANK, J.H., AND E.D. MCOY. 1992. The immigration of insects to Florida, with a tab
ulation of records published since 1970. Florida Entomol. 75: 128.
HOPPING, G.R. 1965a. The North American species in Group IX of Ips DeGeer (Co
leoptera: Scolytidae). Canadian Entomol. 97: 422-434.
HOPPING, G.R.1965b. The North American species in Group X of Ips DeGeer (Co
leoptera: Scolytidae). Canadian Entomol. 97: 803-808.
KIRKENDALL, L.R. 1983. The evolution of mating systems in bark and ambrosia bee
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KOVACH, J., AND C. S. GORSUCH. 1985. Survey of ambrosia beetle species infesting
South Carolina peach orchards and a taxonomic key for the most common spe
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LONG, R.W., AND O. LAKELA. 1971. A flora of tropical Florida: a manual of the seed
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bles, Fla.
NGOAN, N. D., R. C. WILKINSON, D. E. SHORT, C. S. MOSES, AND J. R. MANGOLD. 1976.
Biology of an introduced ambrosia beetle, Xylosandrus compactus, in Florida.
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manus in America north of Mexico (Coleoptera: Scolytidae). Great Lakes Ento
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Univ. Kansas Sci. Bull. 36: 959-1089.
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umes).









Florida Entomologist 77(3) September, 1994


USE OF A BIOTINYLATED DNA PROBE FOR DETECTION OF
THE ASTER YELLOWS MYCOPLASMALIKE ORGANISM IN
DALBULUS MAIDIS AND MACROSTELES FASCIFRONS
(HOMOPTERA: CICADELLIDAE).


FERNANDO E. VEGA,'2 ROBERT E. DAVIS,3 ELLEN L. DALLY,3 PEDRO BARBOSA',
ALEXANDER H. PURCELL,4 AND ING-MING LEE3
'Department of Entomology, University of Maryland, College Park, MD 20742.

Present address: Subtropical Insects Research Unit, U. S. Horticultural Research
Laboratory, USDA, ARS, 2120 Camden Road, Orlando, FL 32803.

3Molecular Plant Pathology Laboratory, USDA, ARS, Beltsville, MD 20705.

4Department of Environmental Science, Policy and Management, Entomology
Program, University of California, 201 Wellman Hall, Berkeley, CA 94720.


ABSTRACT

A DNA probe was used to detect aster yellows mycoplasmalike organism acquisi
tion by the corn leafhopper, Dalbulus maidis (DeLong & Wolcott), an insect that does
not transmit aster yellows mycoplasmalike organism, as well as by Macrosteles fasci
frons (Stal), a vector insect. Results show the effectiveness of the probe for pathogen
detection in both the non-vector and the vector insect.


RESUME

Se utilize una prueba de DNA para detectar la adquisicidn del organismo de mico
plasma de las estrias amarillas (AYMLO) por la chicharrita del maiz, Dalbulus mai
dis (DeLong & Wolcott), un insecto que no transmite AYMLO, asi como de Macrosteles
fascifrons (Stal) un insecto vector. Los resultados muestran la efectividad de la prueba
para detectar el pat6geno en el insecto no vector y en el insecto vector.





Positive and negative effects have been reported in vector and non-vector insects
after feeding on diseased plants (Vega 1992). In at least one case, an expansion of host
plant range occurred when a non-vector insect species fed upon a diseased host (Ma
ramorosch 1958). This system involved the aster yellows mycoplasmalike organism
-an aster leafhopper [Macrosteles fascifrons (Stal); Homoptera: Cicadellidae] (So
rensen & Sawyer 1989) transmitted a plant pathogen with host plants in over 40 dif
ferent families (McCoy et al. 1989) -and the corn leafhopper, Dalbulus maidis
(DeLong & Wolcott), (Homoptera: Cicadellidae).
D. maidis is a Zea spp. specialist (Nault & DeLong 1980) that cannot normally sur
vive on China asters [Callistephus chinensis (L.) Nees] (Maramorosch 1958) nor does
it transmit the pathogen causing aster yellows (Maramorosch 1952). However, if D.
maidis feeds on asters infected with the aster yellows mycoplasmalike organism, not
only can it survive on the plant, but after a minimum of three days on the plant it can
survive on healthy asters and other non-host plants, such as carrots and rye, on which


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Vega et al.: Use of a Biotinylated DNA probe


it typically would not survive (Maramorosch 1960; Purcell 1988). This phenomenon is
called conditioning (Maramorosch 1958).
Recently, Vega et al. (1993) confirmed the presence of the aster yellows mycoplas
malike organism in D. maidis by using the polymerase chain reaction (PCR). The ob
jective of this study was to determine whether a biotinylated DNA probe could be used
to detect the mycoplasmalike organism in a vector and a non-vector insect.

MATERIALS AND METHODS

Healthy and aster yellow-infected aster plants, as well as healthy and infected M.
fascifrons, were provided by the fifth author. The non-vector insects (i.e., Dalbulus
maidis) had been maintained in colonies at the University of Maryland greenhouse
for 15-20 generations.

Pathogen Acquisition

Vector and non-vector insects were placed on asters infected with the 'Severe
(SAY) strain of the aster yellows mycoplasmalike organism for various acquisition ac
cess periods (AAP). The AAP was followed by various incubation periods, during
which insects were held on uninfected asters in the case of M. fascifrons, or on unin
fected corn (cv. Aristogold Bantam Evergreen) in the case of D. maidis. When D. mai
dis was allowed to feed on healthy aster plants, the insect died within 4 days
(Maramorosch 1958). To reduce mortality of the non-vector insect after the AAP on
SAY infected asters, we held D. maidis on corn for the incubation period. D. maidis
was given an AAP of 7 days followed by incubation periods of 0, 7 or 14 days and an
AAP of 14 days followed by incubation periods of 0 or 7 days. M. fascifrons was given
an AAP of 7 days followed by incubation periods of 0, 7, or 14 days and an AAP of 14
days followed by an incubation period of 0, 7, or 14 days. Negative controls consisted
of insects that fed on corn (D. maidis) or on aster (M. fascifrons). M. fascifrons AAP
and incubation periods were conducted at the University of California, Berkeley. D.
maidis AAP and incubation periods were conducted at the University of Maryland
greenhouse. Insects were frozen (-86 C) after each treatment combination until DNA
analysis.

DNA Extraction

Batches of five insects were separately placed in 12 ml glass tissue homogenizers
(Bellco Biotechnology, Vineland, N.J.) and macerated several times in 400 Pl of extrac
tion buffer (100 mM Tris, 50 mM EDTA, 500 mM NaCI, pH 8.0), 2 pl of 13mercaptoeth
anol, and 20 pl of 20% sodium dodecyl sulfate (SDS). The solution was transferred to
a 1.5 ml Eppendorf tube and re-macerated with a mini-pestle. After clarifying three
times by centrifugation, nucleic acids were extracted using chloroform-isoamyl alco
hol and TE-saturated phenol. To precipitate nucleic acids, 2.5 volumes of cold absolute
ethanol was added to samples followed by 30 min at 86 C, and 20 min of centrifuga
tion at 14,000 rpm. The DNA pellet was resuspended in 100 pl 6x saline sodium cit
rate (SSC; lx= 0.9 M NaCI, 0.09 M sodium citrate, pH 7.0).

Filter Preparation

Resuspended DNA was denatured by adding to each 50 pl SSC, 3 p1 of 2N NaOH,
boiling for 10 min, cooling in ice and neutralizing with 3 pl of 2M TRIS and 1 pl 1.5









Florida Entomologist 77(3) September, 1994


M NaAc. Three microliters of undiluted and serial dilutions were spotted on nitrocel
lulose paper and baked at 80 C for 2 h under vacuum.


Filter Pre-hybridization

Filters were pre-hybridized with gentle rotation at 42 C for 2-4 h in presence of
50% formamide, 5x SSC, 5X Denhardt's solution (0.5 g Ficoll, 0.5 g polyvinylpyrroli
done, 0.5 g bovine serum albumin), 25 mM sodium phosphate and 0.5 mg/ml of dena
tured salmon sperm DNA.


Probe Hybridization and Visualization

Hybridization was conducted at 42 C using 5% Dextran sulfate, 45% formamide,
5X SSC, 1X Denhardt's solution, 20 mM sodium phosphate (pH 6.5), 0.2 mg pl of de
natured salmon sperm DNA, and approximately 0.3 mg of biotinylated DNA probe
per ml. The probe was prepared by cloning aster yellows mycoplasmalike organism
DNA fragments into plasmid vectors pSP64 or pSP65 which were then amplified in
Escherichia coli strain JM83 (for a full description see Lee & Davis 1988). After 12-16
h incubation, filters were rinsed twice for 3 min in 2X SSC with 0.1% SDS, 2 rinses of
0.2X SSC with 0.1% SDS (both rinses at room temperature), and 2 rinses at 50 C for
15 min using 0.16X SSC with 0.1% SDS. For signal detection, filters were rinsed for
1 min in buffer 1 (0.1 M Tris-HC1, 0.15 M NaCI, pH 7.5), and blocked for 1 h at 64 C
in buffer 2 (3% bovine serum albumin in buffer 1). Filters were incubated with gentle
rotation for 10 min using streptavidin-alkaline phosphatase diluted in buffer 1, fol
lowed by two 15 min rinses in buffer 1 and one 10 min rinse in buffer 3 (0.1 M Tris
HCl, 0.1 M NaCI, 0.05 M MgC1,, pH 9.5). To visualize reactions, filters were incubated
for about 30 min in the dark using nitroblue tetrazolium and 5-bromo-4-chloro-in
dolylphosphate diluted in buffer 3. To stop reactions, filters were washed in terminal
tion buffer (20 mM Tris, 0.5 mM EDTA, pH 7.5). A bluish color indicated a positive
reaction, i.e., presence of the pathogen's DNA.


RESULTS AND DISCUSSION

For D. maidis given a 7-day AAP, higher amounts of DNA were detected as the in
cubation period increased from 0 to 7, and then to 14 day (Table 1). There was a re
duction in the SAY DNA detected as the incubation period increased from 0 to 7 days
in insects given a 14-day AAP (Table 1). Similar reductions in aster yellows concern
tration have been reported in the alimentary canal of M. fascifrons by Sinha &
Chiykowski (1967). SAY DNA was detected in M. fascifrons at all dilutions and in all
treatments except for the control and for insects given a 7 day AAP and no incubation
period (Table 1).
Our results (Table 1) indicate that a biotinylated DNA probe can be used for the
detection of SAY MLO in vector and non-vector insects. Even though sensitive tech
niques such as PCR are now routinely used for pathogen detection, biotinylated DNA
probes are a sound alternative in cases where PCR technology is not available. In this
study, the use of a biotinylated DNA probe yielded information that suggests an in
crease in pathogen titer as the incubation period increased for non-vector insects with
a 7 days AAP
If widespread, the conditioning phenomenon could play an important role in the
expansion of insects host plant range. The availability of different pathogen detection









Vega et al.: Use of a Biotinylated DNA probe


TABLE 1. SAY MLO DETECTION IN D. MAIDIS AND M. FASCIFRONS USING A BIOTINY
LATED DNA PROBE.

Dalbulus maidis

Nucleic acid dilutions

AAP' IP2 1X3 1/2X 1/4X 1/8X

0 0
7 0 +5
7 7 + + +
7 14 + + +
14 0 + + +
14 7 + +

Macrosteles fascifrons

Nucleic acid dilutions

AAP IP 1X 1/2X 1/4X 1/8X

0 0
7 0
7 7 + + + +
7 14 + + +
14 0 + + +
14 7 + + + +
14 14 + + + +

1AAP, acquisition access period, in days.
2IP, incubation period following the AAP, in days.
'lx = undiluted DNA; 1/2X = DNA sample diluted in half, etc.
'Negative reaction, i. e. no hybridization signal.
'Positive hybridization reaction.

techniques will allow research aimed at understanding the mechanisms) responsible
for conditioning.

ACKNOWLEDGMENTS

This work was performed while the first author was visiting researcher in the Mo
lecular Plant Pathology Laboratory (USDA, ARS). This is Scientific Article No. 8792,
Contribution No. A-6580 of the Maryland Agricultural Experiment Station.

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Florida Entomologist 77(3) September, 1994


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Florida Entomologist 77(3)


LEPIDOPTEROUS PESTS OF COTTON AND THEIR
PARASITOIDS
IN A DOUBLE-CROPPING ENVIRONMENT

F. C. TINGLE, E. R. MITCHELL, AND J. R. MCLAUGHLIN
Insect Attractants, Behavior, and Basic Biology
Research Laboratory, Agricultural Research Service
U.S. Department of Agriculture, Gainesville, FL 32604


ABSTRACT

Seasonal populations of Spodoptera frugiperda, S. exigua, S. eridania, Heliothis
virescens, Helicoverpa zea, Pseudoplusia includes, and their parasitoids, were mon
itored on late-season cotton in northcentral Florida in 1992. The cotton was planted
as a second crop, following corn, in one field and as a first crop in a second field in
which cotton has not been grown for many years. At least twelve species of parasitoids
emerged from lepidopterous larvae collected from the cotton plants. The most com
mon parasitoids were Cotesia marginiventris, Meteorus autographae, Cardiochiles ni
griceps, Netelia sayi, and Copidosoma truncatellum. The native parasitoid that
showed the most potential as a biological control agent was C. marginiventris, be
cause it attacks a broad range of pests including the Heliothis/Helicoverpa complex,
Spodoptera spp., and Pseudoplusia includes.


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.


September, 1994









Tingle et al.: Cotton Pests and Parasitoids in Florida


Key Words: Cotton, Heliothis/Helicoverpa, Spodoptera, parasitoids

RESUME

Las poblaciones estacionales de Spodoptera frugipereda, S. exigua, S. eridania,
Heliothis virescens, Helicoverpa zea, Pseudoplusia includes, y sus parasitoides, fu
eron muestreadas periodicamente en algod6n tardio en la region noroccidental de
Florida en 1992. El algod6n fue plantado en un campo como segundo cultivo, despues
de maiz, y en otra area donde no se habia sembrado algod6n durante muchos anos. Al
menos doce species de parasitoides emergieron de las larvas de lepidopteros colecta
das de las plants de algod6n. Los parasitoides mas comunes fueron Cotesia margin
iventris, Meteorus autographae, Cardiochiles nigriceps, Netelia sayi, y Copidosoma
trucatellum. El parasitoide native que mostr6 el mayor potential como agent de con
trol biologico fu6 C. marginiventris, porque ataca un amplio rango de plagas, incluy
endo el complejo Heliothis/Spodoptera, Spodoptera spp., y Pseudoplusia includes.





Cotton had not been grown commercially in northcentral Florida for over 50 years
until a 60-ha field was planted following an early corn crop in Gilchrist County in
1991. This crop was successful and more acreage was planted in 1992. Because cotton
had not been planted in this area for many years and particularly because it was
planted as a second crop in a double-cropping environment, we chose to monitor pop
ulations of cotton pests and their parasitoids in cotton in 1992. Information on the
parasitoids was needed to define the natural populations and to determine their po
tential value in integrated pest management programs.


MATERIALS AND METHODS

The cooperating grower planted cotton in a 18-ha dryland field on June 6 as the
first crop of the year and in a nearby 152-ha irrigated field July 11-18 following an
early corn crop that was harvested for silage. Two cotton varieties, Delta Pine 90 and
HS 46, were planted in each of the two fields, which were separated by a highway and
tree line. Information on insect pest infestations and damage levels was provided to
the grower who applied conventional insecticide treatments at his discretion. Approx
imately 52% of the insecticide applications involved synthetic pyrethroids. Other pes
ticide groups applied included carbamates, chlorinated bicyclic sulfites, organo
phosphates, and Bacillus thuringiensis. The dryland cotton received 12 applications
of pesticide, and 10 applications were made in the irrigated field. Although some irri
gation was used, it was not a major factor that influenced the outcome of the study be
cause rainfall was sufficient throughout the growing season.
Cotton plants (stems, leaves, squares, blooms, and bolls) were checked for damage
and lepidopterous eggs and larvae were counted twice weekly in each of the 2 variety
ies at 6 preselected locations in the dryland field and at 12 locations in the irrigated
field (25 randomly selected plants per location). Different plants were checked on each
sample date. The eggs and larvae were collected by hand, returned to the laboratory,
and reared individually in 33-ml plastic cups on artificial diet (Guy et al. 1985) for
emergence of adults or adult parasitoids. Larval and egg collections were made from
June 30 through October 16 in the dryland cotton. The larval and egg collections in
the irrigated field began on July 21 and continued through November 20. Parasitiza
tion levels were determined throughout the season based on the emergence of moths









Florida Entomologist 77(3) September, 1994


or parasitoids from collected eggs and larvae. Identification of pest and parasitoid spe
cies was verified by comparison with preserved specimens.

RESULTS AND DISCUSSION

The insect surveys and the emergence of adults from eggs and larvae collected
from the late-season cotton enabled us to monitor field populations (Fig. 1 3) of the fol
lowing pest species: fall armyworm (FAW), Spodoptera frugiperda (J.E. Smith); beet
armyworm (BAW), S. exigua (Hiibner); southern armyworm (SAW), S. eridania
(Cramer); tobacco budworm (TBW), Heliothis virescens (F); corn earworm (CEW),
Helicoverpa zea (Boddie); and soybean looper (SBL), Pseudoplusia includes
(Walker).
Prior to September 1, the predominant pest species in the dryland cotton, as indi
cated by the number of adults that emerged from collected larvae and eggs, was FAW
(Figures 2A and 3A, respectively). Parasitoids could have been a major factor in pre
venting the emergence of adults from the TBW and CEW larvae found on the cotton
plants (Figure 1B), because 50 to 100% of the field-collected TBW and CEW larvae
were parasitized during August. Also during August, 65 to 75% of SBL larvae were
parasitized (Fig. 1C), whereas less than 40% of the FAW and BAW larvae were para
sitized (Figure 1A). Parasitism of FAW and BAW larvae increased rapidly, reaching
100% during the first week of September. At this time SBL became the most common
pest (Figures 1C and 2B) until the cotton matured. However, very few SBL eggs were
found on the cotton plants in either field, and no parasitoids emerged from any of the
field-collected lepidopteran eggs. Predation of larvae and eggs by beneficial insects
was not assessed.
The predominant pest species in the later-planted cotton in the irrigated field was
TBW, as indicated by adult emergence from larval and egg collections (Fig. 2D and
3D), until outnumbered by FAW in late August (Fig. 2C and 3C). Subsequently, BAW
larvae predominated in September. As in the dryland cotton, parasitoids apparently
prevented a substantial number of TBW and CEW larvae from reaching the adult
stage. Parasitization of TBW and CEW larvae peaked at 80% during the first week of
September and again at almost 70% during the first week of October (Fig. 1E). Para
sitization of FAW and BAW larvae did not exceed 50% throughout the season (Fig.
1D).
SBL larval populations in the irrigated cotton increased in mid-September and
peaked two weeks later (Fig. 1F and 2D). Parasitism levels of the SBL larvae ranged
from 20 to 50% during September and October. More FAW than BAW eggs were col
elected from cotton plants during October (Fig. 3C), but by late October more BAW
adults emerged from the field-collected larvae than any other pest species (Fig. 2C).
No SAW larvae were found in either field, but some SAW egg masses were collected
from plants in the irrigated field in late October (Fig. 3C). Differences between the lar
val and egg collection data likely result from protection of the eggs and small larvae
in the laboratory from insecticide applications, predators, parasitoids, heavy rains,
and other environmental hazards. Most of the field-collected larvae of each pest spe
cies were half grown or smaller.
At least 12 species of parasitoids emerged from the lepidopterous larvae collected
from the cotton plants. These included the following Hymenoptera: Cardiochiles nig
riceps (Viereck) [Braconidae], Chelonus insularis Cresson [Braconidae], Copidosoma
truncatellum (Dalman) [Encyrtidae], Cotesia marginiventris (Cresson) [Braconidae],
Meteorus autographae Muesebeck [Braconidae], Netelia sayi (Cushman) [Ichneu
monidae], Ophion sp. [Ichneumonidae], and Pristomerus spinator (F.) [Ichneu









Tingle et al.: Cotton Pests and Parasitoids in Florida


DRY LAND


1 15 29 12 26 0 23 7 21 4 18
JUL AUG 8EP OCT NOV


IRRIGATED
FAW and BAW


1 15 29 12 26 9 2 7 21 4 18
JUL AUG 8EP OCT NOV


Fig. 1. Number of Spodoptera (FAW and BAW), Heliothis/Helicoverpa (TBW and
CEW), and P includes (SBL) counted per 100 cotton plants and percentage parasit
ism of field-collected larvae in each field.

monidae]. Four species of Diptera, all Tachinidae, were identified: Archytas
marmoratus (Townsend), Eucelatoria rubentis (Coquillett), Lespesia archippivora (Ri
ley), and Winthemia rufopicta (Bigot).


5.0
4.5
4.0
3.5
3.0
S2.5
,20
1.56
1.0
-0.5
o0.0

& so
46
40
35
30

L 20
UJ 15
'10






S40
z so
wS
m80


BBL









338 Florida Entomologist 77(3) September, 1994


DRY LAND IRRIGATED

10 FAW *BAW A FAW S BAW C 100
100
90 90
80 80
70 70
60 60
50 50
40 40

20 20






80 80
S70 70
0 60
50 50
40 40
30 30
20 \ 20
10 10

1 15 29 12 26 9 237 21 418 1 15 29 12 26 9237 21 418
JUL AUG SEP OCT NOV JUL AUG SEP OCT NOV
Fig. 2. Percent of each pest species as determined by emergence of adults from lar-
vae collected from cotton plants in each field (FAW = fall armyworm; BAW = beet ar-
myworm; TBW = tobacco budworm; CEW = corn earworm; SBL = soybean looper).


Average parasitism of FAW and BAW combined was 23% in each field although the
average numbers of larvae counted per week were almost 5 times greater in the irri-
gated field than in the dryland cotton. The number of FAW and BAW larvae per 100
plants averaged 0.7 0.3 (June 30-October 16) in the dryland cotton and 3.4 + 0.9
(July 21-November 20) in the irrigated cotton that followed an early corn crop. Most
of the parasitoids that emerged from the FAW and BAW larvae were C. marginiven-
tris. This species made up 66% and 83%, respectively, of the parasitoids that emerged
from larvae collected in the dryland and irrigated fields. The second most common
parasitoid in each field was M. autographae (10 and 15% of the parasitism in the dry-
land and irrigated fields, respectively).
Parasitism of the TBW and CEW larvae combined averaged 65% per week
throughout the season in the dryland cotton but only 29% in the later-planted irri-
gated cotton. Also, the TBW and CEW larval population was almost 4 times greater
in the dryland cotton [5.9(x) 2.9(SEM) larvae per 100 plants] than in the irrigated
field (1.5 0.5 larvae per 100 plants).









Tingle et al.: Cotton Pests and Parasitoids in Florida


7

Co.'

w 4
Z 3
15 2



150
52 35
120

IDS
W a
C Igoo

-5 60

Sso
z 45
30
15
n


DRY LAND


o TBW E CEW


IRRIGATED


339




100
90

70 m
80
750 f
6So
so m
z
40 1
30 0
20 n
10 m


100


Co
5O
70 -
rni
60
50
40 2
40
30
20
10
0


1 15 29 12 26 9 23 7 21 4 18 1 15 29 12 26 9 23 7 21 4 18
JUL AUG SEP OCT NOV JUL AUG 8EP OCT NOV
Fig. 3. Number of Spodoptera (FAW, BAW, anf SAW) egg masses and Heliothis/
Helicoverpa (TBW and CEW) eggs counted per 100 cotton plants and percent of each
species as determined by emergence of adults from egg collections in each field (FAW
fall armyworm; BAW = beet armyworm; SAW = southern armyworm; TBW = tobacco
budworm; CEW = corn earworm).


Almost one half (48%) of the parasitoids that emerged from the TBW and CEW
collected from plants in the irrigated field were C. marginiventris, which was also the
predominant parasitoid species emerging from FAW and BAW larvae from both fields.
Only 23% of the parasitoids that emerged from TBW and CEW larvae in the dryland
cotton were C. marginiventris. The most common parasitoid that emerged from the
TBW and CEW larvae from the dryland field was N. sayi, comprising 47% of the total
parasitoids that emerged from these species. N sayi made up 20% of all parasitoids
from TBW and CEW larvae collected in the irrigated field.
Another important parasitoid that emerged from the collections of TBW and CEW
larvae was C. nigriceps. Although TBW and CEW were not identified to species before
emergence of adults, C. nigriceps is restricted to TBW (Marsh 1978). This parasitoid
made up about 25% of the total parasitoid species that emerged from larvae collected
from cotton in each field. In a previous 3-year study in an adjoining county, C. nigri
ceps was the predominant parasitoid species reared from TBW larvae collected from
tobacco (Tingle & Mitchell 1982). N. sayi, which was the most prevalent parasitoid of


o TOW a CEW


F.









Florida Entomologist 77(3) September, 1994


TBW in postharvest tobacco (Tingle & Mitchell 1982), also emerged from TBW larvae
collected from cotton in the present study.
The weekly average percentage parasitism of SBL larvae collected from cotton
during the season was 49% in the dryland field and 39% in the irrigated field. The av
erage number of SBL larvae counted per 100 plants per week was [9.9(x) 4.6(SEM)]
in the dry-land cotton and 4.2 + 1.0 in the irrigated field. Although C. marginiventris
emerged from 58% of the parasitized SBL larvae collected in the irrigated field, 92%
of parasitized SBL in the earlier planted cotton contained C. truncatellum. This spe
cies also emerged from 32% of the parasitized SBL larvae collected from cotton in the
irrigated field.
C. truncatellum, a polyembryonic egg-larval parasitoid of several insect pests, has
a high reproductive potential and has been reported as the most prevalent parasitoid
of SBL (Burleigh 1971). However, this parasitoid may not be a promising candidate in
an integrated control program for SBL because Hunter & Stoner (1975) found that
SBL larvae parasitized by this species consumed 35% more food than unparasitized
larvae. The increased food consumption of parasitized larvae could outweigh benefits
resulting from reduced SBL populations.
C. marginiventris, however, has potential as a biological control agent because it
attacks a broad range of lepidopterous pests such as the Heliothis Helicoverpa group,
Spodoptera spp., and Pseudoplusia includes. This parasitoid frequently causes high
mortality among these pests (e.g., Tingle et al. 1978; McCutcheon & Turnipseed 1981,
Pair et al. 1982, 1986). Ashley (1979) reported that C. marginiventris is one of the
most frequently recovered parasitoids from FAW larval collections. It is an important
natural enemy of FAW because it parasitizes first and second-instar larvae that die
when they reach the fourth instar (Ashley et al. 1982).
Growers in northcentral Florida are experimenting with various crops and produce
tion schemes in an attempt to increase profitability on existing acreage. The concept
of producing a cash crop such as cotton following a crop of silage corn has received at
tention and appears promising. Of particular concern to growers pursuing this prac
tice has been the mix of insect pests encountered in cotton planted so late in the
growing season, and also how the pest complex might be exacerbated by the early (i.e.,
first) crop of corn.
This study indicates that the species of lepidopterous pests in dryland (early sea
son) and irrigated (late season, 2nd crop) cotton were virtually the same as was the
parasitoid complex associated with these pests. Although several species of parasi
toids were recorded, C. marginiventris was the most prevalent and had the largest im
pact because of its propensity to attack a broad range of pests including the Heliothis/
Helicoverpa complex, Spodoptera spp., and Pseudoplusia includes.
The data presented here suggest that parasitoids in general, and C. marginiven
tris in particular, could play a major role in control of most lepidopterous pests on cot
ton. Pest control strategies should foster the preservation of these and other natural
enemies via the selection and judicious use of pesticides that have minimal impact on
the natural enemies complex and, where possible, provide suitable habitats in and
around cotton fields to encourage parasitoid development and survival.


ACKNOWLEDGMENTS

We gratefully acknowledge the technical assistance of W. Copeland, J. Leach, J.
Rye, R. Navasero-Ward, F Adams, C. Green, and R. Hemphill of this laboratory. Also,
we thank J.R. Ruberson (Dept. Entomol., Coastal Exp. Stn., Tifton, GA) and person
nel of the Division of plant Industry (DPI), Fla. Dept. of Agric., Gainesville, FL, for









Tingle et al.: Cotton Pests and Parasitoids in Florida


providing preserved specimens to aid in identification of parasitoids. Mention of a pro
prietary product does not constitute an endorsement or recommendation for its use by
USDA.


REFERENCES CITED

ASHLEY, T. R. 1979. Classification and distribution of fall armyworm parasites. Flor
ida Entomol. 62: 114-123.
ASHLEY, T. R., V. H. WADDILL, E. R. MITCHELL, AND J. RYE, 1982. Impact of native
parasites on the fall armyworm, Spodoptera frugiperida (Lepidoptera: Noctu
idae), in south Florida and the release of the exotic parasite, Eiphosoma vitti
cole (Hymenoptera: Ichneumonidae). Environ. Entomol. 11:833-837.
BURLEIGH, J. B. 1971. Parasites reared from the soybean looper in Louisiana 1968-9.
J. Econ. Entomol. 64: 1550-1551.
GUY, R. H., N. C. LEPPLA, J. R. RYE, C. W. GREEN, S. L. BARRETTE, AND K. A. HOLLIEN.
1985. Trichoplusia ni, pp. 487-494, in P. Singh and R. F. Moore [eds.] Handbook
of Insect Rearing, Vol. 2. Elsevier Science Publishers B.V, Amsterdam.
HUNTER, K. W., JR., AND A. STONER. 1975. Copidosoma truncatellum: Effect of para
sitization on food consumption of larval Trichoplusia ni. Environ. Entomol. 4:
381-382.
MARSH, P. M. 1978. The Braconid parasites (Hymenoptera) of Heliothis species (Lep
idoptera: Noctuidae). Proc. Entomol. Soc. Washington 80: 1536.
MCCUTCHEON, G. S., AND S. G. TURNIPSEED. 1981. Parasites of lepidopterous larvae
in insect resistant and susceptible soybeans in South Carolina. Environ. Ento
mol. 10: 69-74.
PAIR, S. D., M. L. LASTER, AND D. F. MARTIN. 1982. Parasitoids of Heliothis spp. (Lep
idoptera: Noctuidae) larvae in Mississippi associated with sesame interplant
ings in cotton, 1971-1974: Implications of host habitat interaction. Environ.
Entomol. 11: 509-512.
PAIR, S. D., J. R. RAULSTON, A. N. SPARKS, AND P. B. MARTIN. 1986. Fall armyworm
(Lepidoptera: Noctuidae) parasitoids: Differential spring distribution and inci
dence on corn and sorghum in the Southern United States and Northeastern
Mexico. Environ. Entomol. 15: 342-348.
TINGLE, F. C., T. R. ASHLEY, AND E. R. MITCHELL. 1978. Parasites of Spodoptera ex
igua, S. eridania (Lepidoptera: Noctuidae) and Herpetogramma bipunctalis
(Lepidoptera:Pyralidae) collected from Amaranthus hybridus in field corn. En
tomophaga 23: 343-347.
TINGLE, F. C., AND E. R. MITCHELL. 1982. Effect of synthetic pheromone on parasiti
zation of Heliothis virescens (F.) (Lepidoptera: Noctuidae) in tobacco. Environ.
Entomol. 11: 913-916.









Florida Entomologist 77(3) September, 1994


EGG MORPHOLOGY OF ANASTREPHA OBLIQUA AND SOME
COMPARATIVE ASPECTS WITH EGGS OF ANASTREPHA
FRATERCUL US (DIPTERA:TEPHRITIDAE)

TERESITA MURILLO' AND LUIS FERNANDO JIRON2.
Laboratorio de Control de Calidad, Corporaci6n PIPASA,
San Antonio de Bel6n, Heredia, Costa Rica.

SMuseo de Insectos, Facultad de Agronomia,
Universidad de Costa Rica, San Jose, Costa Rica.

ABSTRACT

The egg morphology of Anastrepha obliqua (Macquart) is described utilizing both
light and scanning electron microscopy. Other observations include oviposition depth
by A. obliqua on mango and comparative analysis with A. fraterculus (Wiedemann)
egg morphology. A. obliqua eggs bear a conspicuous structure at the anterior pole,
which is called a respiratory horn. A discussion of this term and definitions given by
other authors for similar structures found in other fruit fly eggs is given. We suggest
that the occurrence of the respiratory horn outside the mango fruit peel by A. obliqua
may be a useful character for post-harvest inspection and mango fruit disinfestation.

Key Words: Mango, fruit flies, insect egg anatomy, insect respiration

RESUME

La morfologia de Anastrepha obliqua (Macquart) es descrita por medio de micros
copia de luz y electr6nica de barrido. Otras observaciones incluyen la profundidad de
ovoposici6n por A. obliqua en mangos y el analysis comparative con la morfologia del
huevo de A. fraterculus (Wiedemann). Los huevos de A. obliqua poseen una notable es
tructura en el polo anterior, que es llamada cuerno respiratorio. Se ofrece una discu
sion acerca de este termino y de definiciones dadas por otros autores para estructuras
similares en otros huevos de moscas fruteras. Sugerimos que la aparici6n del cuerno
respiratorio fuera de la cascara del mango en A. obliqua puede ser una caracteristica
until para la inspecci6n post cosecha y la desinfecci6n de los frutos.





The taxonomy of Anastrepha fruit flies is based primarily on adult forms; however,
egg morphology also has taxonomic value, and more descriptive work needs to be done
(Norrbom & Foote 1989). Emmart (1933) published descriptions of the eggs of
A.ludens (Loew), A. obliqua (Macquart) (confused with A. fraterculus (Wiedemann)),
A. striata Schiner and A. serpentina (Wiedemann). Sein (1933) reported that a portion
of the egg of A. obliqua remains outside of the fruit peel after oviposition. Later,
Lawrence (1979) described the eggs of A. suspense (Loew). More recently, Norrbom
(1985) described the general characters of the eggs of eight Anastrepha species, in
cluding A. obliqua. Additional descriptions of the eggs of A. bistrigata Bezzi (Steck &
Malavasi 1988), A. grandis (Macquart) (Steck & Wharton 1988) and A. ludens (Car
roll & Wharton 1989) have appeared since then.
Based on the above mentioned descriptions, there is some confusion concerning
the morphology of A. obliqua eggs. Emmart (1933) described a twisted anterior end of


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Murillo & Jiron: Anastrepha obliqua Egg Morphology 343

the egg, hairs around the micropyle and on the posterior end, and a sculptured surface
on the anterior (micropylar) end. Sein (1933) described a smooth surface over the en
tire egg. Norrbom (1985) described A. obliqua eggs as having an anterior lobe covered
by pore-like openings and a reticulated surface at the base of this lobe. The latter au
thor made these observations on ova dissected from preserved females and expressed
some reservations about the appearance of the egg after oviposition.
Here, we describe the morphology of the eggs of A. obliqua with light and scanning
electron microscopy (SEM). The position of the egg after oviposition into mango fruit
is also described with SEM. We also make a comparative description with A. fratercu
lus egg morphology.


MATERIALS AND METHODS

Adults of A. obliqua were reared at the laboratories of Organismo Internacional
Regional de Sanidad Agropecuaria (OIRSA) in Guadalupe (San Jose Prov, elev. 1200
m.) obtained from field infested mangoes collected at Orotina (Alajuela Prov, elev. 200
m.). About 300 of these adults (both sexes) were held in a 1 m3screened cage and al
lowed to oviposit into an artificial medium consisting of a 10 cm diam agar hemi
sphere coated with a beeswax-paraffin mixture. Thirty eggs were retrieved and placed
in glycerin on depression slides and measured under a compound light microscope.
General features were observed with light microscopy by mounting another thirty
eggs in Hoyers temporary mounting medium. Adults of A. fraterculus were obtained
from a laboratory colony (generation F23), and eggs were obtained by placing a piece
of black cloth externally on the upper wall of the colony cage for oviposition. They
were treated as described above.
Eggs of both Anastrepha species were fixed for SEM in 2.5% glutaraldehyde and
2.0% paraformaldehyde, in phosphate buffer (0.1 M, pH 7.4), post-fixed in 1% osmium
tetraoxide, dehydrated in graded alcohol, and critically point dried. Stub mounted
specimens were examined with a scanning electron microscope (Hitachi S-570). Prep
aration for cryofracture was carried out, as an intermediate step after alcoholic dehy
dration, using a substitution graded mixture up to 1:1 alcohol: DMSO (dimethyl
sulfoxide). An aluminum surface frozen by liquid nitrogen was the support for the
sample, which was cryofractured by a stroke with a razor blade. After the SEM prep
aration process, we observed some problems keeping the egg in turgid form; however,
the anterior pole maintained its shape, thus facilitating its observation.
Five mature mango fruits of Yellow, Mora and Tommy Atkins varieties were ex
posed to oviposition by about 300 A. obliqua adults (both sexes) in a 1 m3 screened
cage for twelve hours. Thirty 1 cm3 pieces of fruit rind, each containing one egg, were
taken from each variety for microscopic examination and measurement of the exposed
portion of the egg with a dissecting microscope. Oviposition depth into mango fruit by
A. obliqua was calculated as the difference between the average egg length and the
length of each of the exposed respiratory horns. Ten pieces of each mango variety, each
containing one egg, were similarly treated for SEM as described above to examine the
exposed egg horn after oviposition.


RESULTS AND DISCUSSION

Anastrepha obliqua and A. fraterculus are closely related species belonging to the
fraterculus subgroup (Norrbom 1985); however, their egg morphologies are clearly dif
ferent.









Florida Entomologist 77(3) September, 1994


Anastrepha obliqua eggs observed by light microscopy are creamy white with the
anterior end drawn into a respiratory horn and the posterior end bluntly rounded
(Fig. 1). Average measurements of thirty A. obliqua eggs were: total length 1.447 mm
(1.181-1.584 mm), greatest width 0.225 mm (0.202-0.259 mm) and respiratory horn
length 0.142 mm (0.086-0.245 mm).
The respiratory horn is a projection of the chorion and aeropyles are grouped on
the apex; this structure obviously functions in gaseous exchange. It has been observed
that eggs hatch after being in bubbling water for 48 hours (for mass rearing purposes)
(Aagesen, personal communication). Thus, it seems possible that the egg's respiratory
system can function as a plastron. At the base of the respiratory horn, we observed a
micropyle (Fig. 3). No hair-like structures were found as Emmart (1933) previously
reported.
Our use of the term respiratory horn is made in the same sense as that of Hinton
(1981) who used it to describe structures on eggs which are used in atmospheric res
piration. In the case of A. obliqua, this structure seems obviously to function in respi
ration, and the term respiratory horn seems preferable to other terms commonly used
to describe similar structures on tephritid eggs such as pedicell" (Headrick & Goeden
1990), "stalk" (Benjamin 1934, Cavender & Goeden 1984), or "lobe" and "appendage"
(Norrbom 1985). Pedicel, for example, is the diminutive of the latin ped or pedis and
means "little foot"; stalk refers to a lengthened part on which an organ grows or is
supported; thus, neither of the former terms is appropriate to the structure seen here,
and other terms are less descriptive of its apparent function.
Most of the egg's surface is smooth with no obvious structures or "sculpturing" on
it. However, it is reticulated near both ends and these areas have a granular texture
(Fig. 4). The reticulation at the respiratory horn's base is more strongly defined, but
less polygonal than at the posterior end.
The entire chorion as well as the respiratory horn is cavernous; thus, there is a
continuous network of open spaces extending from the respiratory horn to the rest of
the chorion over the entire egg body. The internal structures of the chorion and respi
ratory horn are shown in Fig. 5 and 6, respectively.
Bacteria were observed on recently oviposited eggs of A. obliqua. Previous obser
vations by Murillo et al. (1990) showed that some bacterial flora, which they called
"indigenous microflora", are associated with A. obliqua eggs just after oviposition.
These bacterial species belong to the Enterobacteriaceae group, which also have been
reported on the eggs of other tephritid species (Drew & Lloyd 1989, Howard et al.
1985).
Anastrepha fraterculus eggs are elongated, creamy white, with a very short chori
onic extension anteriorly and bluntly rounded posteriorly (Fig. 2). The micropyle is
more conspicuous than that observed in A. obliqua, and it is located almost at the an
terior apex of the egg (Fig. 8). The eggs of A. fraterculus bear polygonal reticulations
over the anterior pole and subdistally to the posterior pole, and they are more clearly
defined than those of A. obliqua eggs. There are aeropyles on the borders of the retic
ulations on the anterior pole, surrounding the micropyle (Fig. 9).
Anastrepha obliqua lay their eggs individually with the respiratory horn remain
ing at the fruit's surface from the micropyle upward (Fig. 7). This suggests that the A.
obliqua egg needs unobstructed air exchange with the atmosphere before the larva
hatches. The angle of the respiratory horn relative to the fruit surface may change,
but it remains exposed. Average oviposition depth was 1.34 mm in the three mango
varieties studied, and there were no statistically significant differences among them
(1.347 mm oviposition depth for Yellow and Tommy Atkins, and 1.334 mm for Mora;
SE 0.022, 0.032 and 0.040, respectively; P< 0.05).








Murillo & Jiron: Anastrepha obliqua Egg Morphology 345


Figs. 1-4.
(1) The egg of Anastrepha obliqua (Bar = 120 pm).
(2) The egg of Anastrepha fraterculus (Bar = 115 pm).
(3) Respiratory horn of the egg of A. obliqua, aeropyles at the apex and micropyle at
the base (arrow) (SEM) (Bar =14 pm).
(4) Granular surface of the anterior reticulated area of A. obliqua egg (SEM) (Bar = 3
pm).


wo









Florida Entomologist 77(3)


Figs. 5-9.
(5) Cryofracture of the chorion of the A. obliqua egg (SEM) (Bar = 1 pm).
(6) Cryofracture of the respiratory horn of the A. obliqua egg (SEM) (Bar = pm).
(7) Respiratory horn of A. obliqua egg emerging from the mango peel (SEM) (Bar 50 m).
(8) Micropyle at the anterior apex of A. fraterculus egg (SEM) (Bar = 5 pm).
(9) Aeropyles on the borders of reticulations in the area surrounding the micropyle of
A. fraterculus egg (SEM) (Bar = 5 pm).


September, 1994









Murillo & Jiron: Anastrepha obliqua Egg Morphology 347

Plastron-bearing respiratory horns have been reported in species belonging to
seven dipteran families (Dryomyzidae, Sepsidae, Coelopidae, Sphaeroceridae, Droso
philidae, Cordiluridae and Muscidae) (Hinton 1981). However, so far there has been
no report of its occurrence in Tephritidae. Mouzaki & Margaritis (1987) suggested
that the respiratory structures present in the family Drosophilidae have been re
placed in Tephritidae by: a) an area for respiration on the main body of the egg, b)
later by the whole egg's surface, and c) very limited areas on the main body and the
anterior cup of the egg. However, at least in A. obliqua a respiratory horn is present,
and the structures found in A. nigrifascia (Stone), A. pittieri (Caravallo) (Norrbom
1985), Paracantha gentilis (Hering) (Headrick & Goeden 1990), P cultaris (Coquillett)
(Cavender & Goeden 1984), P culta (Wiedemann), P forficula (Benjamin), Toxotry
pana curvicauda Gerstaecker, and Zonosemata electa (Say) (Benjamin 1934) eggs also
may serve a similar function.
In addition to a respiratory function, the egg horn of A. obliqua may play a role in
the recognition of infested hosts (fruits) by gravid females. Takasu and Hirose (1988)
found that Ooencyrtus nezarae Ishii (Encyrtidae: Hymenoptera) utilizes the egg stalk
(as they named it) as an external marker, which permits the female to recognize hosts
which are already infested. It is possible that A. obliqua females, before oviposition,
also check suitable fruit for a physical marker (exposed horn), and as a parasitism in
dictator complementary to oviposition deterring pheromones (chemical marker)
(Prokopy & Roitberg 1984). However, this function needs to be demonstrated.
After the larva hatches it migrates into internal fruit tissues. The respiratory horn
maintains its physical integrity attached to the fruit peel, unless it is broken off by
rubbing or handling.
Perhaps the exposure of the A. obliqua respiratory horn, outside the mango fruit
peel, could be an aspect to consider when planning post-harvest technology for fresh
fruits. There may be value in developing a chemical, physical or biological method to
detect the presence of the respiratory horn. These methods could be developed also for
use in fruit disinfestation, e.g., addition of a surfactant to a fruit bath may allow water
to enter the chorion interior and limit egg respiration.

ACKNOWLEDGMENTS

The authors thank Dr. Mario Vargas and Dr. Francisco Hernandez who collabo
rated in this study, and Dr. William Ramirez for his suggestions in early manuscripts.
Dr. Allen Norrbom gave valuable suggestions and manuscript reviewing. We thank
Unidad de Microscopia Electr6nica (UME) of the Universidad de Costa Rica for all fa
cilities they gave for this study, and for the training course we received in that Center;
also, Organismo Internacional Regional de Sanidad Agropecuaria (OIRSA) for speci
mens, and Jorge Ramirez for his comments on the final manuscript draft. Finally, the
authors thank the Consejo Nacional de Investigaciones Cientificas (CONICIT) for eco
nomic support for the publication of this paper.

REFERENCES CITED

BENJAMIN, F. H. 1934. Descriptions of some native trypetid flies with notes on their
habits. United States Dept. Agric. Technical Bulletin 401.
CARROLL, L. E., AND R. A. WHARTON. 1989. Morphology of the immature stages of
Anastrepha ludens (Diptera: Tephritidae). Ann. Entomol. Soc. America 82:
201-214.
CAVENDER, G. L., AND R. D. GOEDEN. 1984. The life history of Paracantha cultaris
(Coquillet) on wild sunflower, Helianthus annus L. ssp. lenticularis (Douglas)









Florida Entomologist 77(3) September, 1994


Cockerell, in southern California (Diptera: Tephritidae). Pan-Pacific. Entomol.
60:213-218.
DREW, R. A. I., AND A. C. LLOYD. 1989. Bacteria associated with fruit flies and their
host plants, pp. 131-140 in A.S. Robinson and G. Hooper, [eds], World Crop
Pests. Fruit Flies, their biology, natural enemies and control. Amsterdam,
Elsevier Science. v. 3A.
EMMART, E. W. 1933. The eggs of four species of the fruit flies of the genus Anas
trepha. Proc. Entomol. Soc. Washington 35: 184-191.
HEADRICK, D., AND R. D. GOEDEN. 1990. Description of the immature stages of Para
cantha gentilis (Diptera: Tephritidae). Ann. Entomol. Soc. America 83: 220-229.
HINTON, H. E. 1981. Biology of insect eggs. Pergamon Press, Oxford. v. I, 316 p.
HOWARD, D. J., G. L. BUSH, AND J. A. BREZNAK. 1985. The evolutionary significance
of bacteria associated with Rhagoletis. Evolution 39:405-417.
LAWRENCE, P. O. 1979. Immature stages of the Caribbean fruit fly Anastrepha sus
pensa. Florida Entomol. 62: 214-219.
MOUZAKI, D. G., AND L. H. MARGARITIS. 1987. Comparative structural study of the
egg-shell (chorion) in Dacus oleae, Rhagoletis cerasi, Ceratitis capitata, and Eu
rytoma amygdali. Fruit Flies Procs. II International Symposium, Crete, Sept.
1986. p. 79-87.
MURILLO, T., P. RIVERA, F. HERNANDEZ, AND L. F. JIRON. 1990. Indigenous microflora
of the West Indies fruit fly, Anastrepha obliqua (Diptera Tephritidae). Fruits
(France) 45: 629-631.
NORRBOM, A. L. 1985. Phylogenetic analysis and taxonomy of the cryptostrepha, dac
iformis, robusta and schausi species groups of Anastrepha Schiner (Diptera: Te
phritidae). Ph.D. Thesis. Pennsylvania Sta. Univ. 359 p.
NORRBOM, A. L., AND R. H. FOOTE. 1989. The taxonomy and zoogeography of the ge
nus Anastrepha (Diptera: Tephritidae), pp. 15-26 in A.S. Robinson and G.
Hooper, [eds.], World Crop Pests. Fruit Flies, their biology, natural enemies
and control. Amsterdam, Elsevier Science. v. 3A.
PROKOPY, R. J., AND B. ROITBERG. 1984. Foraging behavior of true fruit flies. Ameri
can Scientist 72: 4149.
SEIN, F. 1933. Anastrepha (Trypetidae, Diptera) fruit flies in Puerto Rico. J. Dept. Ag
ric. Puerto Rico 17:183-196.
STECK, G. J., AND A. MALAVASI. 1988. Description of immature stages of Anastrepha
bistrigata (Diptera Tephritidae). Ann. Entomol. Soc. America 81: 1004-1009.
STECK, G. J., AND R. A. WHARTON. 1988. Description of immature stages of Anas
trepha interrupta, A. limae, A. grandis (Diptera: Tephritidae). Ann. Entomol.
Soc. America 81: 994-1003.
TAKASU, K., AND Y. HIROSE. 1988. Host discrimination in the parasitoid Ooencyrtus
nezarae: the role of the egg stalk as an external marker. Entom. Exper. & Appl.
47:45-48.









Childers: Copper Formulations and Rust Mite Control 349

EFFECT OF DIFFERENT COPPER FORMULATIONS TANK-
MIXED WITH FENBUTATIN-OXIDE
FOR CONTROL OF CITRUS RUST MITES (ACARI:
ERIOPHYIDAE) ON FLORIDA CITRUS

CARL C. CHILDERS
University of Florida, IFAS
Citrus Research and Education Center
700 Experiment Station Road
Lake Alfred, FL 33850

ABSTRACT

One or more rates of fenbutatin-oxide (= Vendex") 4 L applied alone were compared
with the same rates of Vendex tank-mixed with one or more formulations of copper
and an untreated check for control of citrus rust mite, Phyllocoptruta oleivora (Ash
mead) (Acari: Eriophyidae), in 4 citrus grove sites between 1988 and 1991. The copper
formulations corresponded to recommended rates in the Florida Citrus Spray Guide
and included: 99% GC basic copper sulfate (pentahydrate) with 53% metallic copper,
a 77% WP copper hydroxide formulation containing 50% metallic copper and a 61.4%
copper hydroxide DF formulation containing 40% metallic copper. Reduced effective
ness of Vendex occurred when combined with all 3 copper formulations. Both copper
hydroxide formulations were more disruptive in reducing the effectiveness of Vendex
in controlling citrus rust mite compared with the basic copper sulfate formulation. In
creases in citrus rust mite numbers occurred in the copper-only treatments in 2 of the
3 field experiments. In both instances, population increases in the copper-only treat
ments occurred at the same time as those in the untreated check trees, suggesting a
stimulatory effect by the copper compounds.

Key Words: Chemical control, Phyllocoptruta oleivora, miticides, spray tank-mixes

RESUME

Varias dosis de oxido de fenbutatina (Vendex) 4 L aplicadas solas fueron compare
das con mezclas en tanque de diferentes formulaciones de cobre mas fenbutatina alas
mismas dosis y con un testigo no tratado, para el control del acaro tostador, Phyllo
coptruta oleivora (Ashmead) (Acari: Eriophyidae), en cuatro localidades con huertas
de citricos entire 1988 y 1991. Las formulaciones de cobre correspondieron a las reco
mendadas en la Guia de Aspersiones de los Citricos de la Florida e incluyeron: 99%
GC de sulfato de basico de cobre (pentahidratado) con 53% de cobre metalico, una for
mulaci6n de hidroxido de cobre al 77% WP conteniendo 50% de cobre metalico y una
formulaci6n del 61.4% de hidroxido de cobre DF con un 40% de cobre metalico. La efec
tividad del Vendex se redujo cuando este fue combinado con las tres formulaciones de
cobre. Las formulaciones de hidroxido de cobre redujeron mas fuertemente la efecti
vidad del Vendex que el sulfato basico de cobre en el control del acaro tostador. En dos
de los tres campos experimentales se observaron aumentos en la mortalidad de los
acaros tostadores. En ambos casos, la poblaci6n en los tratamientos de cobre solo au
ment6 al mismo tiempo que en los arboles no tratados, sugiriendo un efecto estimula
dor por parte de los compuestos de cobre.





Copper compounds are recommended for use on Florida citrus to control several
fungal diseases including greasy spot (Mycosphaerella citri Whiteside), melanose (Di

This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Florida Entomologist 77(3) September, 1994


aporthe citri Wolf) and citrus scab (Elsinoe fawcettii Bitancourt & Jenkins) (Timmer
& McGovern 1993, Whiteside 1988). Several formulations of copper, that contain me
tallic copper, including copper sulfate and copper hydroxide, have been used by Flor
ida citrus growers for many years. The materials are relatively inexpensive while
providing fungicidal activity. Many citrus growers apply copper in the postbloom and
summer sprays, in combination with a miticide or insecticide, for broad spectrum pest
control and to minimize application costs.
The citrus rust mite, Phyllocoptruta oleivora (Ashmead), and the spider mites, Eu
tetranychus banks (McGregor) and Panonychus citri (McGregor), can be serious pests
during the postbloom and summer periods and thus may require a miticide applica
tion during one or both times, depending upon citrus variety, mite presence and risk
of population increase (Childers 1987).
Thompson (1939) found that P citri increased following copper sprays on citrus
and Holloway et al. (1942) reported that this spider mite was more abundant follow
ing sprays containing compounds of copper, zinc and lime compared with untreated
trees. Griffith & Fisher (1949) observed that Phyllocoptruta oleivora and Panonychus
citri increased abnormally on citrus in Florida following the use of sprays containing
copper, zinc, or lime or exposure to road dust or fertilizer residues on fruit and leaves.
Johnson (1960a) evaluated 5 copper formulations, including basic copper sulfate, cop
per oxide, copper hydroxysulfate, copper sulfate, and tetra copper calcium oxychlo
ride, in combination with zineb for control of Phyllocoptruta oleivora. All of the copper
compounds and rates tested reduced the effectiveness of zineb in the summer spray
but not in the postbloom spray. This supported earlier work by Johnson et al. (1957)
that showed zineb provided less control of P oleivora when combined with fixed cop
per compared with zineb alone.
Residual control by chlorobenzilate or dicofol against the citrus rust mite was re
duced when copper or methidathion was tank-mixed for use on citrus in Texas (Dean
1979). Densities of Brevipalpus phoenicis (Geijskes) (Acari: Tenuipalpidae) increased
after frequent copper oxychloride applications on tea in Indonesia (Oomen 1982).
Eger et al. (1985) evaluated tank mixtures of copper with chlorpyrifos or ethion ap
plied in 2.338 or 4.676 kiloliters of water per hectare using an airblast sprayer. They
found that residual control of the citrus rust mite was reduced with the copper + or
ganophosphate combinations compared with the organophosphate compounds ap
plied alone. Hirsutella thompsonii Fisher, a fungus pathogenic to citrus rust mite, was
suspected of being adversely affected by the copper.
Vendex is not affected by differences in water pH because the organotin compound
is extremely stable. The formulation forms a suspension, not a solution, in water (Sea
man & Riedl 1986). This would suggest that a coating of the miticide by petroleum oil
would reduce the efficacy of the contact miticide (Childers & Selhime 1983).
Grower complaints of poor citrus rust mite control with fenbutatin-oxide + copper
tank mixes prompted the field evaluations reported in this paper. The residual activ
ity of one or more copper formulations in combination with one or more rates of Ven
dex was evaluated between 1987 and 1991 and compared with the miticide applied
alone at the same rates. Results of these studies are reported here.

MATERIALS AND METHODS

One or more rates of Vendex, 4 pounds active ingredient per gallon liquid formu
nation (4 L) (479 g AI per liter) (E. I. DuPont de Nemours, Wilmington, DE), applied
alone were compared with the same rates of Vendex tank-mixed with one or more for









Childers: Copper Formulations and Rust Mite Control 351

mulations of copper for citrus rust mite control and an untreated control in 4 citrus
grove sites between 1988 and 1991 (Table 1). The rates of copper corresponded to rec
ommended rates in the Florida Citrus Spray Guide. The copper formulations included
99% basic copper sulfate (pentahydrate) granular crystals (GC) with 53% metallic
copper (Tennessee Chemical Co., Copperhill, TN), a formulation of 77% copper hy
droxide WP containing 50% metallic copper (Kocide Chemical Corp., Houston, TX)
and a 61.4% copper hydroxide dry flowable (DF) formulation containing 40% metallic
copper (Griffin Corp., Valdosta, GA).


Experimental Sites

At grove site 1, plots consisted of 9 trees arranged in a 3 by 3 grid (Table 1). At sites
2 and 3, treatments were assigned to plots of 25 trees arranged in a 5 by 5 grid and at
site 4, 36 trees arranged in a 6 by 6 grid. All treatments were arranged in a random
ized complete block design based on motile numbers of citrus rust mites per tree taken
during the pretreatment counts and replicated 5 times for each site (Childers & Sel
hime 1983).
The center tree was sampled from each plot in all of the sites except site 2 where
2 center trees were sampled from each plot. Each sample tree was a vigorous, healthy
tree representative of the block and each plot was separated from adjacent plots by 2
trees within the row and 2 rows between treatments. Foliar spray treatments were
applied in the listed volumes per ha with properly calibrated airblast sprayers on the
dates listed in Table 1.

TABLE 1. LOCATIONS, CHARACTERISTICS OF CITRUS PLANTINGS, DATES OF MITICIDE
APPLICATION, SPRAYERS, GALLONAGES AND TRACTOR SPEEDS.

Citrus Grove Site

Characteristic 1 2 3 4

Location Baseball City Mineola vic. Lake Placid vic. Ft. Meade vic.
Polk County Lake County Highlands Polk County
County
Cultivar Valencia orange Hamlin orange Valencia orange Hamlin orange
Tree height
(meters) 4.57-5.49 m 2.13-3.35 m 4.27-5.18m 2.44-3.66 m
Spacing 7.62 x 7.62 m 7.62 x9.14 m 9.14 x 9.14m 4.57x 7.62 m
(meters) ( 173 trees/ ( 240 trees/ ( 119 trees/ ( 287 trees/hect
hectare) hectare) hectare) are)
Application May 26-27, June 20-22, June 25-26, July 8-10,
dates 1988 1989 1991 1991
Type sprayer FMC 4000 CP Southwind pto FMC 970 Swanson
Durand pto
Liters per
hectare 1,169 1,169 1,403 720
Spray pressure 160 psi 200 psi 180 psi 100 psi
Tractor speed 2.4 kph 2.4 kph 2.4 kph 2.4 kph
Water pH 7.9 7.8 7.3 7.6









Florida Entomologist 77(3) September, 1994


Mite Sampling

Twenty fruit were examined at random around the canopy perimeter of each sam
ple tree at recorded time intervals before and after spraying of the treatments. Live
motile citrus rust mites were counted using a 10x hand lens equipped with a 1 cm2
grid subdivided into 4 mm2 subunits (Childers & Selhime 1983). Two places on oppo
site sides of each fruit outside of the exposed sunspot were examined. Population den
sity estimates of citrus rust mite were obtained by counting the number of living
motile mites present within the combined 2 cm2 area per fruit and recorded as one ob
servation. In test 2, 10 fruit were examined at random around the canopy perimeter
from each of 2 sample trees with a total of 20 fruit per plot.
Twenty leaves from the spring flush were picked at random around the canopy pe
rimeter of each sample tree at designated time intervals before and after spraying.
Each leaf sample was placed in separately labeled paper bags and returned to the lab
oratory. A stereomicroscope was used to examine a 1 cm2 area on both the upper and
lower leaf surfaces in test 1, or a 2 cm2 area on both leaf surfaces in tests 4 and 5. Live
motile citrus rust mites were counted within each area on both the upper and lower
leaf surfaces. The 2 counts were combined as 1 observation and equalled either 2 or 4
cm2 area of leaf surface. In test 3, 10 leaves were collected from each of 2 trees in the
center of each plot.
Cumulative citrus rust mite-days were determined for each treatment within a
grove each year by calculating the area under the population growth curve over time
(Allen 1976). Mite-days were calculated using the formula presented by Childers et al.
(1987).

Fruit Damage Ratings

One hundred fruit were examined randomly around the perimeter of each sample
tree following completion of a field experiment. The cumulative percentage of surface
area with russeting from citrus rust mite feeding injury was recorded for each fruit.
This was determined by estimating the percentage area of russeting on one side of
each fruit and then turning the fruit 180 and repeating the estimate of russeted area
again. The combined estimated russeted area of sides one and two for each fruit was
then divided by two. A series of plates with line drawings of mature sized fruit with
calculated proportional areas of surface injury were prepared to provide a consistent
reference for the person determining rind blemish injury. A fruit was classified as re
jected for the fresh market when rind blemish from citrus rust mite feeding exceeded
5% of the total surface area.

Statistical Analysis

In all experiments, data were subjected to analysis of variance; Duncan's (1955)
multiple range test was used to separate treatment means when the ANOVA provided
a significant F value (P < 0.05) (SAS Institute 1991). Motile citrus rust mite counts
were subjected to Log10 (X + 1) transformations for statistical analysis. Untrans
formed means are shown in all tables.

RESULTS

Site 1, 'Valencia' Orange

Sustained moderate numbers of citrus rust mites were present in the untreated
check trees through August 2 following treatment applications on May 26 and 27. Cit









Childers: Copper Formulations and Rust Mite Control 353

rus rust mite densities on treated trees were significantly higher between August 2
and 25 in the Vendex + copper sulfate or hydroxide combinations compared with the
Vendex only treatments at the same rate (Table 2). Cumulative mite-days were sig
nificantly higher with the 2.10 kg rate of Vendex + copper hydroxide and the 1.40 kg
rate of Vendex + copper sulfate compared with the same rates of Vendex applied alone.
The other combinations of copper + Vendex did not result in significantly higher cu
mulative mite-days than the corresponding rates of Vendex alone. Damage rating dif
ferences were not significantly different between the miticide treatments in this
experiment (Table 2). Based on these data, all Vendex treatments performed well.


Site 2, 'Hamlin' Orange

Moderate to high densities of citrus rust mites were present on the untreated trees
between July 27 and October 12 following treatment applications (Table 3). Vendex at
1.68 kg + copper hydroxide had a significantly higher citrus rust mite density on the
fruit by September 20 compared with Vendex at 1.68 kg applied alone. Vendex at 1.68
kg + copper sulfate had a significantly higher citrus rust mite density by September
27 compared with Vendex at the same rate applied alone. These differences continued
through October 18. Vendex at 2.24 kg + copper hydroxide had significantly higher cit
rus rust mite densities by September 27 compared with Vendex at 2.24 kg applied
alone. Again, these differences continued through October 18. The Vendex at 2.24 kg
+ copper sulfate combination was not significantly different from Vendex at 2.24 kg
until October 3. These differences in citrus rust mite densities continued to be signif
icantly higher through October 12. Vendex at 2.80 kg combined with either copper hy
droxide or copper sulfate had significantly higher citrus rust mite densities on the
fruit by September 20 and 27 and October 18 compared with Vendex at 2.80 kg ap
plied alone. These high rust mite densities occurred while the population was increase
ing in the untreated check trees.
Consistently higher citrus rust mite densities ranging from 1 to 4 times higher
were present in both copper hydroxide and copper sulfate treated trees between Au
gust 17 and September 27 compared with the untreated trees.
Cumulative mite-day comparisons were significantly different between the Ven
dex at 1.68 kg + copper hydroxide treatment compared with Vendex at 1.68 kg applied
alone (Table 4). Cumulative mite-day differences between the remaining miticide
treatments were not significantly different although numerically higher densities of
citrus rust mites were consistently obtained in the Vendex + copper treatments com
pared with the Vendex only treatments.
Significantly higher percentages of russeted and rejected fruit were obtained in
the low rate of Vendex combined with either copper hydroxide or copper sulfate com
pared with the low rate of Vendex applied alone (Table 4). The highest percentages of
russeted and rejected fruit were obtained in the copper-only treatments compared
with all other treatments including the untreated check trees.


Site 3, 'Valencia' Orange

Low to moderate citrus rust mite densities were present on both fruit and leaves
between July 11 and August 19 on the untreated check trees (Tables 5, 6). Low but sig
nificantly higher citrus rust mite densities were present on the Vendex + copper DF
treatment by August 12 compared with Vendex applied alone. Significantly higher cit
rus rust mite densities on the fruit were consistently obtained on the Vendex + copper
DF treatment between August 12 and September 17 compared with Vendex applied












354 Florida Entomologist 77(3) September, 1994





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Florida Entomologist 77(3) September, 1994


TABLE 4. CUMULATIVE CITRUS RUST MITE-DAYS AND DAMAGE RATING COMPARISONS
OF FRUIT INJURY ON 'HAMLIN' ORANGE (SITE 2) IN LAKE COUNTY, FLORIDA
1989.

Cumulative
Rate per Mite-days % Russeted % Rejected
Treatment and Formulation Hectare Oct 181 Fruit' Fruit'

Vendex 4L 1.68 kg 729 d 12 ef 9 cd
Vendex 4L 2.24 kg 1271 cd 10 ef 6 cd
Vendex 4L 2.80 kg 710 d 5f 3d

Vendex + 4L 1.68 kg
Copper hydroxide 77 WP 7.85 kg 2561 c 52 bc 32 b

Vendex + 4L 2.24 kg
Copper hydroxide 77 WP 7.85 kg 2007 cd 31 cde 20 bc

Vendex + 4L 2.80 kg
Copper hydroxide 77 WP 7.85 kg 843 d 10 ef 5 cd

Vendex + 4L 1.68 kg
Copper sulfate 99 GC 7.85 kg 1753 cd 40 bcd 31 b

Vendex + 4L 2.24 kg
Copper sulfate 99 GC 7.85 kg 1261 cd 22 def 11 cd

Vendex + 4L 2.80 kg
Copper sulfate 99 GC 7.85 kg 932 d 17 ef 8 cd
Copper hydroxide 77 WP 7.85 kg 6431 a 86 a 70 a
Copper sulfate 99 GC 7.85 kg 6056 a 85 a 71 a
Untreated 3573 b 58 b 42 b

1Means within columns followed by the same letter are not significantly different (P>0.05; Duncan's [1955]
multiple range test).

alone. All 3 Vendex + copper treatments had significantly higher citrus rust mite den
sities on both fruit and leaves between August 19 and September 10 compared with
Vendex applied alone.
P oleivora densities on fruit in the 3 copper-only treatments were significantly
higher between July 11 and September 10 compared with the untreated check tree
counts. Populations of citrus rust mites ranged from 2 to 36 times higher on copper
treated fruit compared with the untreated check trees.
The 4 miticide treatments provided comparable control of the citrus rust mite on
leaves through July 18 (Table 6). Low, but significantly higher, citrus rust mite den
sities on leaves were present on the Vendex + copper DF and Vendex + copper hydrox
ide treatments by August 12 compared with Vendex applied alone. These differences
increased dramatically between August 26 and September 10. The Vendex + copper
sulfate treatment was less disruptive to rust mite control compared with the other 2
Vendex + copper treatments. However, leaf counts in this treatment were significantly
higher compared with those in the Vendex only treatment between August 19 and
September 17.













Childers: Copper Formulations and Rust Mite Control 357


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Childers: Copper Formulations and Rust Mite Control 359

Cumulative mite-day comparisons on fruit and leaves showed that the Vendex +
copper DF treatment was significantly higher than Vendex alone or Vendex combined
with copper sulfate. Cumulative mite-days on fruit were not significant between Ven
dex combined with either copper hydroxide or copper sulfate while the cumulative
mite-day differences were significantly higher on leaves between the 2 treatments.
Also, the Vendex + copper sulfate cumulative mite-day value on leaves was signifi
cantly higher compared with Vendex applied alone while those on the fruit were not
(Table 7).
Cumulative mite-day values on fruit and leaves for the Vendex + copper DF and
Vendex + copper hydroxide treatments were not significantly different. Correspond
ing percentages of russeted and rejected fruit were not significantly different between
the same 2 Vendex + copper treatments compared with the untreated check trees.
This demonstrates the negative effect of tank-mixing either copper hydroxide formu
nation at the rates tested with Vendex.
P oleivora densities on leaves in the 3 copper-only treatments were somewhat
variable until July 31. Between that date and September 10, all 3 copper-only treat
ments had consistent and significantly higher citrus rust mite densities on leaves that
ranged from 2 to 12 times higher than those on the untreated check trees. Both copper
DF and copper hydroxide tank-mixes with Vendex resulted in significantly higher
percentages of russeted fruit compared with Vendex applied alone. Only the Vendex
+ copper DF combination had a significantly greater percentage of rejected fruit com-
pared with Vendex applied alone (Table 7). Percentages of both russeted fruit and re
jected fruit were significantly higher in the 3 copper-only treatments compared with
the untreated check trees (Table 7).


TABLE 7. CUMULATIVE CITRUS RUST MITE-DAYS AND DAMAGE RATING COMPARISONS
OF FRUIT INJURY ON 'VALENCIA' ORANGE (SITE 3) IN THE LAKE PLACID VI
CINITY, HIGHLANDS COUNTY, FLORIDA, 1991.

Cumulative mite-days' % %
Rate per Russeted Rejected
Treatment and Formulation Hectare Leaves Fruit Fruit' Fruit'

Vendex 4L 2.24 kg 43 d 77 e 14 d 0.4 c

Vendex + 4L 2.24 kg
Copper hydroxide DF 8.97 kg 865 b 912 bc 55 b 18.8 b

Vendex + 4L 2.24 kg
Copper sulfate 99 GC 8.97 kg 231 c 271 de 25 cd 2.6 c

Vendex + 4L 2.24 kg
Copper hydroxide 77 WP 8.97 kg 501 b 594 cd 31 bc 6.6 bc
Copper hydroxide DF 8.97 kg 1080 a 3930 a 89 a 63.8 a
Copper sulfate 99 GC 8.97 kg 1410 a 4540 a 97 a 79.8 a
Copper hydroxide 77 WP 8.97 kg 1303 a 3930 a 93 a 66.6 a
Untreated 507 b 756 b 33 bc 6.4 bc

'Means within columns followed by the same letter are not significantly different (P > 0.05; Duncan's [1955}
multiple range test).









Florida Entomologist 77(3) September, 1994


Site 4, 'Hamlin' Orange

Low to moderate citrus rust mite densities on fruit in the untreated check trees
were recorded between July 15 and September 10 following treatment applications on
July 8, 9 and 10. Population development of P oleivora was evident on the back sides
of fruit by August 8 with no citrus rust mites present on the front sides of the same
fruit in the Vendex treatments. Poor spray coverage resulted from inadequate pene
tration of pesticide spray through the canopy since only 720 liters of finished spray
were applied per hectare.
All Vendex treatments provided comparable control of citrus rust mite on the fruit
through August 13 (Table 8). The Vendex + copper hydroxide and Vendex + copper DF
combinations generally had significantly higher citrus rust mite densities on the fruit
between August 21 and September 10 compared with Vendex applied alone and the
Vendex + copper sulfate treatments. As in the previous field experiment (site 3), the
amount of metallic copper applied per hectare was highest in the copper sulfate for
mulation followed by copper hydroxide WP and copper hydroxide DF The 3 copper
only treatments did not have higher rust mite densities on fruit during this experi
ment compared with the untreated check trees (Table 8).
All 4 Vendex treatments had significantly lower cumulative mite-day values on
fruit compared with the untreated check trees (Table 9). Only the Vendex + copper hy
droxide 77% WP treatment had significantly more russeted fruit compared with the
other Vendex treatments in this experiment. Numerically higher percentages of re
jected fruit were found in the Vendex + copper DF and Vendex + copper hydroxide
treatments. However, they were not significantly different compared with the other 2
Vendex treatments.
P oleivora densities on leaves were low in all treatments throughout this experi
ment (Table 10). However, significantly higher citrus rust mite densities were re
corded on the Vendex + copper DF and Vendex + copper hydroxide treatments on
August 27 and September 5 compared with Vendex applied alone or Vendex + copper
sulfate. The 3 copper-only treatments did not have consistently higher citrus rust
mite densities on the leaves compared with the untreated check trees. Cumulative
mite-day differences between the copper-only and untreated check treatments were
not significantly different in this experiment (Table 9).


DISCUSSION

Results from this study show that the different copper formulations tested are in
compatible with Vendex when applied at the rates tested. The copper hydroxide DF
formulation demonstrated the greatest degree of incompatibility followed by copper
hydroxide WP The copper sulfate formulation in combination with Vendex showed
the least disruption in residual rust mite control of the 3 formulations tested. The
amount of available metallic copper applied per hectare in tests 3 and 4 differed be
tween the 3 formulations as follows: copper DF = 1.45 kg, copper hydroxide = 1.81 kg
and copper sulfate = 1.93 kg. This would suggest that one or more other components
of the copper DF formulation were involved in creating increased antagonism in com
bination with the organo-tin miticide compared with the metallic copper content
alone. This phenomenon is not unique and has been reported with other tank mix
tures. For example, Venkata Ram (1963) found that nickel chloride hexahydrate con
trolled the fungal pathogen, Exobasidium vexans, on tea as effectively as copper
oxychloride without increasing purple mite, Calacarus carinatus (Green) (Acari: Erio
phyidae).











Childers: Copper Formulations and Rust Mite Control 361



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Florida Entomologist 77(3) September, 1994


TABLE 9. CUMULATIVE CITRUS RUST MITE-DAYS AND DAMAGE RATING COMPARISONS
OF FRUIT INJURY ON 'HAMLIN' ORANGE (SITE 4) IN THE FT. MEADE VICINITY,
POLK COUNTY, FLORIDA, 1991.

Cumulative Mite-days' % %
Rate per Russeted Rejected
Treatment and Formulation Hectare Leaves Fruit Fruit' Fruit'

Vendex 4L 2.24 kg 100 c 157 b 16 e 9d

Vendex + 4L 2.24 kg
Copper hydroxide DF 8.97 kg 203 be 361 b 25 de 13 cd

Vendex + 4L 2.24 kg
Copper sulfate 99 GC 8.97 kg 76 c 117 b 14 e 7 d

Vendex + 4L 2.24 kg
Copper hydroxide 77 WP 8.97 kg 224 be 379 b 32 cd 18 cd
Copper hydroxide DF 8.97 kg 399 a 1367 a 55 ab 38 ab
Copper sulfate 99 GC 8.97 kg 408 ab 1057 a 56 ab 38 ab
Copper hydroxide 77 WP 8.97 kg 418 a 1136 a 43 be 29 be
Untreated 449 a 1558 a 68 a 53 a

'Means within columns followed by the same letter are not significantly different (P > 0.05; Duncan's [1955}
multiple range test).

Citrus rust mite populations increased in 2 of the 3 field experiments in which
copper-only treatments were included. In both instances, population increases in the
copper-only treatments occurred at the same time as those occurring in the untreated
checks. However, the mite densities were significantly higher suggesting a
stimulatory effect by the copper compounds. Johnson (1960b) found that copper
compounds had no effect on the percentage of fruit infested with P oleivora when the
population was increasing on unsprayed trees. However, copper sprays prolonged
high citrus rust mite populations, or caused an increase, if applied when unsprayed
populations were declining. This situation did not develop in 2 of the 3 field
experiments in this study. Adverse effects caused by copper compounds on the
pathogenic fungus, Hirsutella thompsonii Fisher, have been proposed (Eger et al.
1985, McCoy 1979, Griffith & Fisher 1949, Spencer 1939, Thompson 1939).
In many instances, the use of copper on Florida citrus is essential for effective dis
ease control. Great care must be exercised by citrus growers when selecting pesticides
for control of pest complexes. Oftentimes, tank-mixes are designed to minimize appli
cation costs and to optimize efficiency. The use of such tank-mixes requires sound as
surance of both physical and chemical compatibility. Potential risks of pesticide
failure or acceleration of pesticide resistance dictate the need for such information.
Vendex is a highly effective miticide with broad spectrum activity against various
phytophagous mite pests in the families Eriophyidae, Tetranychidae, Tenuipalpidae,
and Tarsonemidae. Studies presented here demonstrate that copper DF and copper
hydroxide formulations result in reduced residual effectiveness of Vendex, especially
at lower rates, i.e., 2.24 kg of 50 WP or 2.34 liters of 4 L per hectare (University of
Florida 1993 Spray Guide recommendations). In a single field experiment by Lye et
al. (1990), Vendex 4 L was applied at the rate of 3.34 liters per hectare in combination













Childers: Copper Formulations and Rust Mite Control 363


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Florida Entomologist 77(3) September, 1994


with copper hydroxide. No reduction in miticidal activity was recorded. However, this
rate exceeds that which is recommended in the Florida Citrus Spray Guide.
Additional studies are needed to identify those miticides currently recommended
for use during the postbloom and summer sprays that have minimal compatibility
problems with specific available copper formulations.


ENDNOTE

The technical assistance of Paul M. Keen, Jr., Michael G. Warmington, and
Deanna K. Threlkeld is gratefully acknowledged. Florida Agricultural Experiment
Station Journal No. R-03528.


REFERENCES CITED

ALLEN, J. C. 1976. A model for predicting citrus rust mite damage on Valencia orange
fruit. Environ. Entomol. 5:1083-1088.
BRUSSEL, E. W. VAN. 1975. Interrelations between citrus rust mite, Hirsutella thomp
sonii and greasy spot on citrus in Surinam. Agric. Exp. Sta. Surinam (Parama
ribo). Bull. 98.
BULLOCK, R. C., C. C. CHILDERS, J. L. KNAPP, C. W. McCOY, AND P. A. STANSLY. 1993.
Citrus rust mite and spider mites, in J. L. Knapp [ed.], Florida Citrus Spray
Guide. Florida Coop. Ext. Serv, IFAS, Univ. Florida, Gainesville, SP-43.
CHILDERS, C. C. 1987. Chemical control of phytophagous mite pests on Florida citrus.
Proc. Int. Conf. on Pests in Agric. Paris III, pp. 119-126.
CHILDERS, C. C., L. W. DUNCAN, T. A. WHEATON, AND L. W. TIMMER. 1987. Arthropod
and nematode control with aldicarb on Florida citrus. J. Econ. Entomol.
80:1064-1071.
CHILDERS, C. C., AND A. G. SELHIME. 1983. Reduced efficacy of fenbutatin-oxide in
combination with petroleum oil in controlling the citrus rust mite Phyllocop
truta oleivora. Florida Entomol. 66:310-319.
DEAN, H. A. 1979. Citrus rust mite control affected by certain pesticides. J. Rio
Grande Valley Hort. Soc. 33:55-61.
DUNCAN, D. B. 1955. Multiple range and multiple F tests. Biometrics 11:1 42.
EGER, J. E., JR., V. M. FERGUSON, AND K. G. TOWNSEND. 1985. Efficacy of selected
miticides and spray tank mixtures used to control rust mite in Florida citrus.
Proc. Florida State Hort. Soc. 98:11-14.
GRIFFITH, J. T., JR., AND F. E. FISHER. 1949. Residues on Citrus trees in Florida. J.
Econ. Entomol. 42:829-833.
HOLLOWAY, J. K., C. F. HENDERSON, AND H. V. McBURNIE. 1942. Population increases
of citrus red mite associated with the use of sprays containing inert granular
residues. J. Econ. Entomol. 35:348-350.
JOHNSON, R. B. 1960a. The effect of copper compounds on control of citrus rust mite
with zineb. J. Econ. Entomol. 53:395-397.
JOHNSON, R. B. 1960b. The effect of copper on rust mite control with four rust mite
miticides. Florida State Hort. Soc. 73:84-89.
JOHNSON, R. B., J. R. KING, AND J. J. MCBRIDE, JR. 1957. Zineb controls citrus rust
mite. Proc. Florida State Hort. Soc. 70:38-48.
LYE, B. H., C. W. McCOY, AND J. FOJTIK. 1990. Effect of copper on residual efficacy of
acaricides and population dynamics of citrus rust mite (Acari: Eriophyidae).
Florida Entomol. 73:230-237.
McCOY, C. W. 1977. Horticultural practices affecting phytophagous mite populations
on citrus. Proc. Int. Soc. Citriculture 2:459-462.
OOMEN, P. A. 1982. Population dynamics of the scarlet mite, Brevipalpus phoenicis, a
pest of tea in Indonesia. Meded. Landbouwhogeschool Wageningen. 82-1, 88 p.









Childers: Copper Formulations and Rust Mite Control 365

SAS INSTITUTE. 1991. SAS language and procedures: Usage 2, version 6, First edition.
SAS Institute. Cary, North Carolina.
SEAMAN, A. J., AND H. RIEDL. 1986. Preventing decomposition of agricultural chemi
cals by alkaline hydrolysis in the spray tank. Cornell Univ., New York's Food
and Life Science Bull. 118. 7 p.
SPENCER, H. 1939. Increases in citrus scale insect infestations from heavy residue and
from copper spray mixtures. J. Econ. Entomol. 32:686-688.
THOMPSON, W. L. 1939. Cultural practices and their influence upon citrus pests. J.
Econ. Entomol. 32:782-789.
TIMMER, L. W., AND R. McGOVERN. 1993. Fungus diseases, fruit and/or foliage, pp 11
13 in J. L. Knapp [ed.], Florida Citrus Spray Guide. Florida Coop. Ext. Serv.,
IFAS, Univ. Florida, Gainesville. SP-43.
VENKATA RAM, C. S. 1963. Application of copper and nickel fungicides with motorized
mist blowers for blister blight control. 3. Effect of yield and buildup of purple
mites. Rep. Unit Plant Assoc. South India Sci. Dept. (Tea Sect.) 1962-63
(Coonoor) pp. XVII-XXIV.
WHITESIDE, J. 0. 1988. Greasy spot, melanose and scab diseases, pp. 15-17, 20-21, 26
27 in J. O. Whiteside, S. M. Garnsey, and L. W. Timmer [eds.], Compendium of
citrus diseases. APS Press, St. Paul, Minnesota.


++++++++++++++++++++++++++++++++++++++++++++++++++++





































INCISITERMES FURVUS, A NEW DRYWOOD TERMITE
(ISOPTERA: KALOTERMITIDAE) FROM PUERTO RICO

RUDOLF H. SCHEFFRAHN
Ft. Lauderdale Research and Education Center
University of Florida, Institute of Food & Agric. Sciences
3205 College Avenue, Ft. Lauderdale, FL 33314

ABSTRACT

The imago and soldier of Incisitermes furvus n. sp. from western Puerto Rico are
described for the first time. This is the seventh species of Incisitermes reported from
the West Indies.

RESUME

Se described por vez primera el imago y el soldado de Incisitermes furvus sp. n. de
Puerto Rico. Esta es la septima especie de Incisitermes registrada de las Indias Occi
dentales.

Key Words: New species description, soldier, winged imago, West Indies.





The genus Incisitermes was established by Krishna (1961) to accommodate the re
classification of a large subgroup of kalotermitid species previously placed together in
the genus Kalotermes Hagen. The known distribution of Incisitermes is limited to both


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.









Florida Entomologist 77(3) September, 1994


the Nearctic and Neotropical Americas (Snyder 1949, Araujo 1977), Australia (Gay
1976), India (Roonwal & Verma 1973), and islands of the Pacific Ocean (Krishna
1961).
A new dark and rather small species of Incisitermes was collected during a recent
survey of the termites of Puerto Rico. The imago and soldier castes of Incisitermes fur
vus n. sp. are described herein.


MATERIALS AND METHODS

Fourteen colonies of I furvus n. sp. were sampled in four locations in Puerto Rico
including three colonies from Reserva Forestal Maricao (1809'N, 66'59'W) on
19-V 1992 by S.C. Jones and one on 2-VI-1993 by J. R. Mangold; eight from Bosque
Estatal de Guajataca (18'25'N, 66'58'W) on 3-VI-1993; one from Area Recreativa Rio
Abajo (18'20'N, 66'43'W) on 4-VI-1993; and one sample from Bosque Estatal de Cam
balache (18'26'N, 66'36'W) on 4-VI-1993; the latter ten collected by J. A. Chase, J. de
la Rosa Guzman, J. R. Mangold and R. H. Scheffrahn. Colonies were dwelling inside
standing and fallen native dead tree trunks and branches of sound condition. The ter
mites were collected by aspirator and field-preserved in 85% ethanol.
Measurements of specimens, made with a calibrated ocular micrometer to a max
imum accuracy of + 0.01 mm, follow the conventions of Krishna & Emerson (1962).
Thirty-four images (14 from Maricao, 18 from Guajataca and one each from Rio Abajo
and Cambalache) and 52 soldiers (14 from Maricao, 31 from Guajataca, four from Rio
Abajo, and three from Cambalache) were used for measurements. Scanning electron
micrographs were made with a Hitachi S-4000 field emission microscope (10kV) of
specimens dehydrated in absolute ethanol and 1,1,1,3,3,3-hexamethyldisilazane (Na
tion 1983) and then sputter-coated with gold.
The holotype imago from Guajataca and morphotype soldier from the Maricao site
(2 VI 1993) will be deposited in the collection of the National Museum of Natural His
tory, Washington, D.C. Paratype soldiers and alates will be deposited in the American
Museum of Natural History, New York, New York, in the Florida State Collection of
Arthropods, Florida Department of Agriculture and Consumer Services., Division of
Plant Industry, Gainesville, Florida, and in the author's collection.

Incisitermes furvus, new species

IMAGO (Figs. 1-5, Table 1). Head capsule, antennae, and notum very dark brown;
wing scales and membranes blackish, costal veins including costal margin, radius,
and radial sector very dark brown, in all, giving the winged imago a strikingly uni
form black appearance when viewed dorsally Clypeus concolorous at base with head
grading to pale yellow anteriorly; labrum light brown. Head with about 16 fine setae
projecting from vertex and frons in lateral profile. Compound eye subtriangular, com
posed of very small facets; ocellus small, oblong, and hyaline, margin sharply defined
surrounding head pigmentation, less than one half diam above eye. Antennae
composed of 15-17 segments, usually 16; third or fourth segment shortest. Mandible
dentition as in Fig. 4 and as described for Incisitermes by Krishna (1961); Pronotum
wider than long, about as wide as head; anterior margin squarely incised to form
about a 160 angle, anterior corners rounded; posterior margin shallowly concave,
posterior corners reduced to two rounded, about 45 curves; all margins with scat
tered long and short setae. Wing membrane tuberculate; median and cubitus veins
and their branches delineated by a single row of papillae smaller than those on mem
brane, costal veins densly covered with even smaller papillae. In forewing, radial sec









Scheffrahn: New Puerto Rican Incisitermes


Fig. 1 5. Scanning electron micrographs of Incisitermes furvus winged imago. Dor
sal views of head (1), thorax, and wing scales (2); lateral view of head and pronotum
(3); and dorsal views of mandibles (4) and right forewing (5).


tor with about eight anterior branches; unsclerotized media unbroken to apex;
cubitus with 11-13 posterior branches. Arolia present.
Comparisons. The imago of I. furvus differs from images of most Incisitermes
species by its very dark body and wings although three other small Neotropical In
cisitermes are rather dark and small and might be confused with I. furvus. These in
clude I. miller (Emerson), known from the Florida Keys and Jamaica (Emerson 1943)
and the Dominican Republic (R.H.S., unpublished), I. nigritus (Snyder) from Guate
mala (Snyder 1946) and Mexico (Emerson 1969), and I. bequaerti (Snyder) from Cuba
(Snyder 1929), the Bahamas and U.S. Virgin Islands (Snyder 1956), the Turks and









Florida Entomologist 77(3) September, 1994


TABLE 1. MEASUREMENTS OF INCISITERMES FURVUS IMAGO

Measurement in mm (n=34) Range Mean + SD Holotype

Number antennal segments 15-17 15.72+ 0.53 16
Head length to labrum tip 1.25-1.48 1.37+ 0.045 1.44
Head length to lateral base of mandibles 0.96-1.10 1.02+ 0.030 1.10
Head maximum width at eyes 1.04-1.16 1.09+ 0.029 1.12
Eye maximum diameter 0.26-0.31 0.29+ 0.015 0.30
Ocellus maximum diameter 0.10-0.14 0.11+ 0.009 0.11
Distance from eye to lower margin of head 0.14-0.20 0.17 0.016 0.16
Pronotum median length 0.58-0.72 0.65 + 0.029 0.64
Pronotum maximum length 0.68-0.84 0.76 + 0.036 0.76
Pronotum maximum width 0.98-1.16 1.09+ 0.037 1.08
Hind tibia length 0.83-0.93 0.87 + 0.030 0.93
Forewing length at suture 5.12-6.42 5.83+ 0.36 5.83
Forewing maximum width 1.46-1.88 1.68+ 0.12 1.69
Body length without wings 4.73-6.09 5.45 + 0.33 4.92
Total length with wings 7.45-8.94 8.23 + 0.49 8.23


Caicos (Scheffrahn et al. 1990), the Dominican Republic (R.H.S., unpublished), and
Puerto Rico [S.C. Jones (USDA-FS, Tucson, AZ) & R.H. S. unpublished]. Table 2 gives
relevant characters to separate these species from I. furvus and each other. The ima
gos of I. miller are smaller and have 1-4 fewer antenna segments than I. furvus.
Wings of I. bequaerti are hyaline and larger and wider than I. furvus. Incisitermes nig
ritus images are closest to I. furvus but can be separated by the larger and more ro
bust bodies, larger eyes and ocelli, and more tuberiferous wing membranes of I.
furvus.
SOLDIER (Figs. 6-9, Table 3). Head capsule, when viewed dorsally, red-brown in
front grading to pale yellow posteriorly; from above, sides parallel, anterior corners
nearly square at mandibular articulations, posterior margin evenly rounded; ventral
surface pale yellow except for reddish anterior margin. Head capsule, when viewed
laterally, with 40-50 dorsal setae of variable length to 0.13 mm; setae shorter but of
same density on ventral surface. Frons slopes about 30 from vertex, weakly and
broadly concave, surface very weakly rugose. Anteclypeus a narrow, white trapezoid.
Labrum light yellow-brown, spatulate. In most specimens, eyespot composed of a
poorly defined hyaline ellipse having diffuse border with surrounding pigmentation;
eyespot regions about same diam and standing about one-half diam behind, and at
same height, as antennal fossae. In a few specimens, a small more well-defined hya
line or gray eye rudiment visible within center of unpigmented region. Mandibles
rather slender with outer margins smooth, angled dorsally about 30 from plane of
head, dentition as in Fig. 9; left mandible with long, narrow distal blade and three
marginal teeth, first two pointed forward, prominently arising in distal third of man
dible, the third tooth less prominent, arising from a shelf on the basal half; right man
dible with serrate blade in distal half and two triangulate teeth in basal half.
Antennae with 10-14 segments, usually 12, rarely 10 as in Fig. 6; third longest, nar
rower than first; third as long as fourth and fifth together; in some specimens, fourth
fused to third. Postmentum as in Fig. 8. Pronotum with median incision in anterior
margin forming square angle of about 135; anterior corners sharply rounded; poste
rior margin with a slight or no concavity; posterior corners broadly rounded; setae on
margins of same length and density as head. Femora weakly inflated.









Scheffrahn: New Puerto Rican Incisitermes


TABLE 2. COMPARISON OF CHARACTERS OF NEOTROPICAL INCISITERMES IMAGOS
(RANGE OF MEASUREMENTS IN MM).

Incisitermes Species'

Character miller furvus nigritus bequaerti

No. of antenna
segments 13-14 15 17 14-16 15-17
Body color very dark very dark very dark castaneous to
brown brown brown2 very dark brown
Wing color hyaline
to dark3 very dark dark hyaline
Papillae on
wing membrane faint prominent faint absent
Maximum diam.
eye 0.20-0.27 0.26-0.31 0.19-0.25 0.26-0.34
Maximum diam.
ocellus 0.09-0.11 0.10-0.14 0.09 0.14-0.16
Hind tibia length 0.68-0.76 0.83-0.93 0.69-0.75 0.70-0.85
Length forewing
to suture 4.60-5.37 5.12-6.42 5.00-5.50 7.58-8.23
Width forewing 1.29-1.50 1.46-1.88 1.39-1.50 2.07-2.43
Body length 4.28-5.12 4.73-6.09 5.00-5.25 5.125.83 (6.75)
Total length
with wings 6.40-7.76 7.45-8.94 7.50-8.00 9.40-10.50
Body length -
total length 0.62-0.69 0.63-0.68 0.65-0.67 0.50-0.60 (0.64)

1Characters and measurements obtained from the following: I miller, Emerson 1943 and four specimens from
four locations in Barahona and Pedernales Provinces, Dominican Republic. L furvus: this study I nigritus: Snyder
1946 and paratypes from U. S. National Museum. L bequaerti: Snyder 1929 (his maximum measurements in pa
rentheses); five specimens from one site each in Barahona, San Pedro de Macoris, and Peravia, and two sites in
Pedernales Province Dominican Republic; Bosque Estatal de Guanica, Puerto Rico; and Providenciales, Turks and
Caicos B.WI.
2Pigmentation faded in paratypes examined. Coloration based on Snyder 1946.
Wing membranes of L miller from Jamaica and Florida hyaline (Emerson 1943) and dark from the Dominican
Republic (Scheffrahn, unpubl.).

Comparisons. Compared with Incisitermes nigritus, soldiers of I furvus have
longer, apically more slender mandibles with all marginal teeth more prominent,
larger and more diffuse eye spot, a longer third antennal segment, and a deeper
pronotal incision. The headcapsule of L bequaerti flatter than that of L furvus, and the
soldier of L miller is smaller than that of L furvus.
Etymology. This species name is taken from the Latin term "furvus" which de
scribes the dark, swarthy body color and dark wing tint of the imago.


DISCUSSION

Soldiers of I furvus so far collected neither vary as greatly in size nor possess very
distinct long and short-headed forms as is common among Incisitermes species. Both
fully pigmented and lighter freshly-molted alates were collected in colonies in May
and June suggesting an early to mid-summer flight season. Alates flew readily upon
extraction from wood during collection. This habit, along with the dark pigmentation









Florida Entomologist 77(3)


Figs. 6-9. Scanning electronic micrographs of Incisitermes furvus soldier. Dorsal
and lateral views of head and pronotum (6,7); ventral view of head (8); and dorsal view
of mandibles (9).

of I. furvus alates indicate that this species is a daytime flier. Incisitermes furvus ap
pears to be a relatively common species within the mountainous forests of its limited
range in western Puerto Rico.

Incisitermes furvus is the thirteenth Neotropical Incisitermes to be described
(Araujo 1977). Of these, seven species are reported from the West Indies including I.
bequaerti, I. incisus (Silvestri), I. miller, I. schwarzi (Banks), I. snyderi (Light), and
I. tabogae (Snyder) (Darlington 1992). Recent collections of Incisitermes in the West
Indies by the author and others hint that one or more new species resembling I. sny
deri may be identified. Our understanding of the West Indian Incisitermes is ham
pered by taxonomic difficulties common to other termite genera from the whole region
(Collins 1988). Further descriptions, redescriptions, and revisions of this region's ter
mites await completion.

ACKNOWLEDGMENT

I am indebted to S. C. Jones (USDA Forest Service, Tucson, AZ) for collecting and
sharing with me the first specimens of furvus;J.A. Chase and J. R. Mangold (Ter
minix International, L. P), and J. de la Rosa Guzman (Carib-Consult, Santo Domingo,
D. R.) for collecting specimens during a termite survey of Puerto Rico; D. S. Williams
of the ICBR Electron Microscope Core Facility at the University of Florida, Gaines
ville, for assisting with the S.E.M. photography; M.S. Collins (Natl. Mus. Nat. Hist.,
Washington, D.C.) for loan of nigritus specimens; and R.M. Giblin-Davis and N.-Y.
Su (University of Florida), J. Krecek (Acad. Sci. Czech Republic), and M. S. Collins for


September, 1994









Scheffrahn: New Puerto Rican Incisitermes


TABLE 3. MEASUREMENTS OF INCISITERMES FURVUS SOLDIERS.

Measurement in mm (n=52) Range Mean + SD Morphotype

Number antennal segments 11-14 12.4 + 0.78 13
Third antennal seg. length 0.15-0.24 0.19 +0.021 0.19
Third antennal segment
maximum width 0.088-0.11 0.10 + 0.006 0.10
Third antennal segment
minimum width 0.050-0.069 0.060 + 0.005 0.056
Head length with mandibles 2.53-3.33 2.90 + 0.20 2.86
Head length to lateral base of
mandibles 1.50-2.20 1.86 +0.16 1.87
Left mandible length to
dorsal base 1.13-1.40 1.26 +0.068 1.33
Pronotum median length 0.53-0.80 0.66 + 0.057 0.67
Pronotum maximum length 0.68-1.00 0.83 + 0.072 0.80
Head maximum width 1.03-1.33 1.20 + 0.064 1.21
Pronotum maximum width 1.00-1.35 1.20 + 0.071 1.20
Head maximum height 0.85-1.07 0.96 + 0055 0.95
Postmentum maximum width 0.44-0.58 0.51 + 0.030 0.51
Postmentum minimum width 0.16-0.27 0.21 + 0.021 0.22
Postmentum length 1.12-1.78 1.44 + 0.15 1.44
Hind tibia length 0.70-0.96 0.85 +0.063 0.88


critically reviewing and improving this contribution no. R-03458 of the University of
Florida Experiment Stations Series.


REFERENCES CITED

ARAUJO, R. L. 1977. Catalogo dos Isoptera do novo mundo. Acad. Brasileira de Cien-
cias, Rio de Janeiro, RJ. 92 pp.
COLLINS, M. S. 1988. Taxonomic problems with termites of North America, Canada
through Panama. Sociobiology 14: 207-210.
DARLINGTON, J. P. E. C. 1992. Survey of termites in Guadeloupe, Lesser Antilles
(Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae). Florida Entomol. 75:
104-109.
EMERSON, A. E. 1943. Kalotermes miller, a new species of termite from the Florida
Keys and Jamaica (Isoptera, Kalotermitidae). Psyche 50: 18-22.
EMERSON, A. E. 1969. A revision of the Tertiary fossil species of the Kalotermitidae
(Isoptera). American Mus. Nov. 2359: 157.
GAY, F. J. 1976. An Australian species of Incisitermes Krishna (Isoptera: Kalotermiti
dae). J. Australian Entomol. Soc. 14: 395-398.
KRISHNA, K. 1961. Generic revision and phylogenetic study of the family Kalotermiti
dae (Isoptera). Bull. American Mus. Nat. Hist. 122: 303-408
KRISHNA, K., AND A. E. EMERSON. 1962. New species of the Genus Glyptotermes Frog
gatt from the Papuan, Oriental, Ethiopian, and Neotropical Regions (Isoptera,
Kalotermitidae). American Mus. Novitates No. 2089: 165.
NATION, J. A. 1983. A new method using hexamethyldisilazane for the preparation of
soft insect tissue for scanning electron microscopy. Stain Technol. 55: 347-352.
ROONWAL, M. L., AND S. C. VERMA. 1973. First record of termite genus Incisitermes
Krishna (Kalotermitidae) from Indian Region, with description of a new species
from Rajasthan. Zool. Anzeiger 191: 390-397.









372 Florida Entomologist 77(3) September, 1994

SCHEFFRAHN, R. H., N.-Y. SU, AND B. DIEHL. 1990. Native, introduced, and
structure-infesting termites of the Turks and Caicos Islands, B.W.I. (Isoptera:
Kalotermitidae, Rhinotermitidae, Termitidae). Florida Entomol. 73: 622-627.
SNYDER, T. E. 1929. New termites from the Antilles and Middle America. Proc. Ento
mol. Soc. Washington 31: 79-87.
SNYDER, T. E. 1946. A small, dark-colored new Kalotermes from Guatemala. Proc. En
tomol. Soc. Washington 48: 158-160.
SNYDER, T. E. 1949. Catalog of the termites (Isoptera) of the world. Smithsonian Misc.
Coll. 122: 1490.
SNYDER, T. E. 1956. Termites of the West Indies, the Bahamas, and Bermuda. J. Ag
ric. Univ. Puerto Rico 40: 189-202.









Scientific Notes


FECUNDITY AND FERTILITY OF RHYNCHOPHORUS
CRUENTATUS
(COLEOPTERA: CURCULIONIDAE)

THOMAS J. WEISSLING' AND ROBIN M. GIBLIN-DAVIS
University of Florida, Institute of Food and
Agricultural Sciences, Ft. Lauderdale Research and Education
Center, 3205 College Ave., Ft. Lauderdale, FL 33314

The palmetto weevil (R. cruentatus F) breeds in a variety of stressed or dying
palms (Giblin-Davis & Howard 1988, 1989). These large (1.9 3.0 cm long) weevils are
associated with the native cabbage palmetto, Sabal palmetto (Walter), in Florida
(Woodruff 1967). Semiochemicals emanating from stressed or dying palms and male
conspecifics (Weissling et al. 1992, 1993, 1994, Giblin-Davis et al. 1994) are attractive
to R. cruentatus adults. Although not fully understood, mating apparently takes place
on dying palms and females lay their eggs in the leaf bases or directly into the wounds
of the host. Larvae develop primarily in the crown region but can occasionally be
found in the stem tissue. Last instar larvae migrate to the fibrous stem periphery or
petiolar bases and construct cocoons from fiber (Giblin-Davis & Howard 1988).
Research on an improved method to culture R. cruentatus has required the collec
tion of a large number of eggs to produce neonate larvae for evaluation of diets. Using
pineapple, Anana comosus (L.), as an ovipositional substrate, Giblin-Davis et al.
(1989) reported the mean lifetime fecundity of field-collected females as 26 + 15 eggs
per female. However, pineapple proved to be a difficult media to dissect for removal of
eggs. More suitable ovipositional substrates were investigated and we found that ap
ple (Pyrus malus L.) slices were easily dissected and were readily accepted by R.
cruentatus females. Using apple slices, we reinvestigated the fecundity of R. cruenta
tus females and determined fertility.
Cocoons were harvested in the field from infested palms or in the laboratory from
sugarcane (Saccharum officinarum L.) stem (Giblin-Davis et al. 1989). Cocoons were
placed individually in covered 100-ml plastic cups with moistened tissue paper
(Giblin-Davis et al. 1989) and were stored at 29 C until adult emergence. One male
and one female were placed in a 500-ml covered plastic container with moistened tis
sue. After 48-72 h, males were removed and a slice of apple ('Red Delicious') was
added. Slices were thin (5-15 mm; 10 18 g wet weight), convex segments covered by
peel. Females oviposit through the apple pulp and most eggs are found along the peel.
All containers were placed in an environmental chamber at 29C with a 13:11 (L:D)
photoperiod. Apple slices were usually replaced every 1-3 days until female death.
Slices removed from containers were carefully dissected and eggs were removed. This
test was repeated five times with four or five females per test (22 females total). Data
were converted to the number of eggs laid per female per week. In addition, the total
number of eggs laid per female was determined. During two of the tests, eggs removed
from apple slices were placed in 15 x 100 mm plastic petri dishes lined with water
moistened filter paper, sealed with parafilm, and placed in the environmental cham-
ber. Neonate larvae were removed from each dish at daily intervals and the dish re
sealed until all eggs hatched or decomposed. For the first test, fertility was
determined every one to three days for 45 d. During the second test, eggs were col


'Current address: Yakima Agricultural Research Laboratory, USDA-ARS, 3706 W Nob Hill Blvd., Yakima, WA
98902


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FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
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Florida Entomologist 77(3) September, 1994


elected every one to three weeks for 14 weeks. Data on overlapping dates were com-
bined and all data were converted to percent eclosion.
R. cruentatus produced an average of 207 + 19 (range 12-433) eggs per female. This
number is considerably higher than the previous estimate of 26 15 eggs per female
(Giblin-Davis et al. 1989) and is more consistent with fecundity reported for other
Rhynchophorus species (Wattanapongsiri 1966). A reduced estimate of fecundity by
Giblin-Davis et al. (1989) may have been caused by suboptimal conditions on pineap
ple slices, decreasing egg and larval survival. In addition, pineapple slices were diffi
cult to dissect and eggs or larvae may have been overlooked (R.M.G.D. pers. observe .
In this study, males were removed from containers after 48 -72 h while Giblin-Davis
et al. (1989) confined females with males throughout the course of the study. The pres
ence of males in a small container may have interfered with oviposition or increased
damage to larvae and eggs (Giblin-Davis et al. 1989). Rananavare et al. (1975) deter
mined that R. ferrugineus Oliver females laid less eggs when confined with males
than without. The mean number of eggs laid per R. cruentatus female per day de
clined until almost no eggs were laid 14 weeks after mating (Fig. 1). However, egg lay
ing by surviving females increased over the subsequent 6 weeks. Reasons for this
increase are unclear.
Fertility of eggs laid by R. cruentatus varied through time, however, eggs collected
eight weeks after female mating did not eclose (Fig. 2). These results suggest that fe
males had utilized all sperm indicating the need for multiple matings. Rananavare et
al. (1975) reported the need for multiple matings in R. ferrugineus to maintain fertil
ity.


8

0



E
LLJ
4-

w


z 2

r-

0 .......... .. .....

0 2 4 6 8 10 12 14 16 18 20 22 24

Weeks After Mating
Fig. 1. Mean ( SEM) weekly egg production by newly-emerged R. cruentatus fe
males (n = 22) confined individually on an apple slice at 29 C after 48 -72h confine
ment with males.









Scientific Notes


100



80


o 60



0
c 40


a_ 20



0 -. . ....,,,, \ ,,,,,, ,, ,,,, ,,,,,,,,,, ,,,,,
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14

Weeks After Mating
Fig. 2. Percent eclosion of eggs produced by newly-emerged R. cruentatus females
(n 9).

We thank J. Cangiamila for technical assistance, and R.H. Scheffrahn and F.W.
Howard for their suggestions. This research was supported by a USDA Special Grant
in Tropical and Subtropical Agriculture CRSR-90-34135-5233. This manuscript is
Florida Agricultural Research Stations Journal Series R-03674.

Summary

The mean fecundity of R. cruentatus when provided apple slices as an oviposition
media was almost eight times higher (207 + 19 eggs per female) than previously esti
mated (26 + 15 eggs per female). The rate of egg-laying decreased through time until
14 weeks after mating when there was a temporary increase. Fertility of R. cruenta
tus eggs remained between 40 and 100 percent eclosion until seven weeks after fe
males were mated but dropped to zero by nine weeks.

REFERENCES CITED

GIBLIN-DAVIS, AND F. W. HOWARD. 1988. Notes on the palmetto weevil, Rhynchopho
rus cruentatus (Coleoptera: Curculionidae). Proc. Florida State Hort. Soc. 101:
101-107.
GIBLIN DAVIS, R. M., AND F. W. HOWARD. 1989. Vulnerability of stressed palms to at
tack by Rhynchophorus cruentatus (Coleoptera: Curculionidae) and insecti
cidal control of the pest. J. Econ. Entomol. 82: 1185-1190.
GIBLIN DAVIS, R. M., K. GERBER, AND R. GRIFFITH. 1989. Laboratory rearing of Rhyn
chophorus cruentatus and R. palmarum (Coleoptera: Curculionidae). Florida
Entomol. 72: 480-488.









376 Florida Entomologist 77(3) September, 1994

GIBLIN-DAVIS, R. M., T. J. WEISSLING, A. C. OEHLSCHLAGER, AND L. M. GONZALEZ.
1994. Field response of Rhynchophorus cruentatus (F) (Coleoptera: Curculion
idae) to its aggregation pheromone and fermenting plant volatiles. Florida En
tomol. 77: 164-177.
RANANAVARE, H. D., K. SHANTARUM, M. R. HARWALKAR, AND G. W. RAHLKAR 1975.
Method of laboratory rearing of red palm weevil, Rhynchophorus ferrugineus
Oliv. J. Plantation Crops 3: 65-67.
WATTANAPONGSIRI, A. 1966. A revision of the genera Rhynchophorus and Dynamis
(Coleoptera: Curculionidae). Dept. Agric. Science Bulletin, Bangkok 1: 1328.
WEISSLING, T. J., R. M. GIBLIN-DAVIS, R. H. SCHEFFRAHN, AND N. M. MENDOZA. 1992.
Trap for capturing and retaining Rhynchophorus cruentatus (Coleoptera: Cur
culionidae) adults using Sabalpalmetto as bait. Florida Entomol. 75: 212-221.
WEISSLING, T. J., R. M. GIBLIN-DAVIS, AND R. H. SCHEFFRAHN. 1993. Laboratory and
field evidence for male-produced aggregation pheromone in Rhynchophorus
cruentatus (F.) (Coleoptera: Curculionidae). J. Chem. Ecol. 19: 1195-1203.
WEISSLING, T. J., R. M. GIBLIN-DAVIS, G. GRIES, R. GRIES, A. L. PEREZ, H. D. PIERCE,
JR., AND A. C. OEHLSCHLAGER. 1994. Aggregation pheromone of palmetto wee
vil Rhynchophorus cruentatus (Coleoptera: Curculionidae). J. Chem. Ecol. 20:
505-515.
WOODRUFF, R. E. 1967. A giant palm weevil, Rhynchophorus cruentatus (Fab.), in
Florida (Coleoptera: Curculionidae). Florida Dept. Agr. Div. Plant Industry. En
tomology Circ. No. 63.


++++++++++++++++++++++++++++++++++++++++++++++++++++









Scientific Notes


NEW SYNONYMY IN THE GENUS POLYTES STAL
(HETEROPTERA: SCUTELLERIDAE)

J. E. EGER, JR.
DowElanco
Ste. 780, One MetroCenter
4010 Boy Scout Boulevard
Tampa, FL 33607
and
Florida State Collection of Arthropods
Florida Department of Agriculture and Consumer Products
Gainesville, FL 32602

In a revision of the genus Polytes Stal (Eger 1990), I overlooked two previously de
scribed species. One of these species was originally described in the genus Pachycoris
Burmeister, the type of which I examined recently while researching the latter genus.
The other was an oversight of a species described in Polytes. The purpose of this paper
is to rectify those errors. Measurements are given in mm.

Polytes discrepans (Uhler 1875) New Combination

Pachycoris discrepans Uhler 1875:282-283.

DIAGNOSIS. Dorsum black with red markings as follows: 4 evenly spaced mac
ules on pronotum (lateral 2 oval, mesial 2 elongate, extending from near posterior


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Scientific Notes


pronotal margin to just anterad of cicatrices); cicatrices; two oval macules near lateral
margins on anterior half of scutellum. Dorsal punctation concolorous, dense, most
dense laterally. Venter entirely black, coxae and rostrum slightly lighter in color than
remainder of venter. Posterior margin of last abdominal sternite lacking colored line
or macule. Length 8.9.
Head very broadly rounded anteriorly; 2.1 long, 2.5 wide across eyes. Length of an
tennal segments 1-4 (5 missing): 0.6; 0.4; 0.5; 1.0, respectively.
Pronotum 2.7 long mesially, 5.4 wide across humeri. Anterolateral margins
slightly convex. Scutellum 5.4 long, 5.0 broad at widest point. Pleura unmarked,
densely punctate. Venter of abdomen densely punctate, less so mesially and on stri
ated areas.
Posterior margin of genital cup shallowly concave mesially, convex laterally. Ven
trally exposed portion of cup densely punctate.
TYPE MATERIAL. Holotype: male, labeled: (a) Peru, Orton. (b) PR Uhler Collec
tion. (c) Pachycoris discrepans, Peru, Uhler. The type is deposited in the National Mu
seum of Natural History, Smithsonian Institution, Washington, D. C.
Polytes discrepans resembles a number of other species in the P fenestra Breddin
species group (P bullock Eger, P bicolor Distant, P lattini Eger, P leopardinus Dis
tant, P speculigerBreddin, and P ruth Breddin) in having a broadly rounded head. It
differs from all of these except P ruth in lacking a large bilobed mesial macule on the
posterior margin of the last abdominal sternite. There is a thin pale line on the pos
terior margin of this sternite in P ruth, but P discrepans has no pale markings on this
sternite. Polytes ruth is also larger and is very differently colored than P discrepans.
The dorsal coloration of P discrepans is most similar to that of P lattini, but the head
of P discrepans is distinctly more broadly rounded than that of P lattini, and the
pronotal vittae are longer than those of P lattini.
Because P discrepans appears to be distinct from all known congeners, I opted not
to dissect the unique male holotype.
Polytes discrepans will key to couplet 12 in the key by Eger (1990), but does not fit
either option given for this couplet. The following changes to the key will allow recog
nition of P discrepans:


Polytes speculiger Breddin 1914


12(11). H ead broadly rounded anteriorly............................................................ 12'
H ead narrow ly rounded anteriorly ........................................................... 13
12'(12). Pronotum with two large yellow to red macules, each with distinct brown
to black lanceolate intrusion from posterior margin; larger, 9.5-10.9 long
.................................................................................... . .. ruth (B reddin)
Pronotum with four evenly spaced macules; smaller, 8.9 long....................
................................................................................ discrepans (U hler)


Polytes speculiger Breddin 1914:53-54.
Polytes quechuus Eger 1990:137-139, figs. 104-109. New Synonymy.

I overlooked Breddin's paper when researching the Polytes revision. His descrip
tion clearly indicates the presence of an impunctate area on the pronotum, a diagnos
tic character for this species, and leaves little doubt that my species is a junior
synonym of his. I have not been able to locate Breddin's types although one of the
paratypes of P quechuus was a syntype of P fenestra Breddin, 1903, and was cer









Florida Entomologist 77(3) September, 1994


tainly seen by him. It bore no labeling to indicate that it was a type of P speculiger,
though. Breddin also described two varieties (as 'var. a' and 'var. b) and suggested
that these may be synonymous with P bicolor and P leopardinus. The color forms he
described were also seen by me and are apparently typical for P speculiger.
I thank Drs. R. C. Froeschner (Smithsonian Institution) and T. J. Henry (USDA
ARS) for the loan of the type of Pachycoris discrepans and Drs. L. H. Rolston (Louisi
ana State University) and D. A. Rider (North Dakota State University) for assistance
in locating literature for this study.

SUMMARY

Corrections to a revision of the genus Polytes are presented. Pachycoris discrepans
Uhler is transferred to Polytes with a diagnosis and revised key to allow recognition
of this species. Polytes quechuus Eger is placed in the synonymy of P speculiger Bred
din.


LITERATURE CITED

BREDDIN, G. 1903. Neue Rhynchoten aus den Anden. Societas Entomologica 18:122
124.
BREDDIN, G. 1914. Neue odor wenig gekannte neotropische Hemiptera. Abh. Senck
enb. Naturforsch. Gesellschaft 36:53-59.
EGER, J. E., JR. 1990. Revision of the genus PolytesStal (Heteroptera: Scutelleridae).
Ann. Entomol. Soc. America 83(2):115-141.
UHLER, P. R. 1874 [1875]. List of the species of Hemiptera and Neuroptera obtained
by Prof. James Orton, in Northern Peru. Proc. Boston Soc. Nat. Hist. 17:282
286.


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Florida Entomologist 77(3) September, 1994


FIRST RECORDS OF ENTOMOPATHOGENIC DISEASES IN
THE PARAGUAY TEA AGROECOSYSTEM IN ARGENTINA

DANIEL R. SOSA-GOMEZ', ELLIOT W. KITAJIMA2 AND MARCELO E. ROLON3
'Centro Nacional de Pesquisa da Soja, EMBRAPA Caixa Postal 1061, 86001-970,
Londrina, PR, Brasil

2 Dept. Biol. Cel., Univ. Brasilia, 70919-970, Brasilia, DF, Brasil

3Establecimiento "Las Marias", 3342 Gobernador Virasoro, Corrientes, Argentina.


The northeastern region of Argentina leads that country in the production of Par
aguay tea (Ilexparaguariensis Saint Hilarie) with 170 metric tons per year. The most
important pests of tea in this region are: a psyllid Metaphalara spegazziniana
(Lizer) (Homoptera: Psyllidae), the "paraguay tea hornworm" Pengonia lusca ilus Bsd.
(Lepidoptera: Sphingidae), and the gall bud mite Dichopelnus notus Keifer (Acarina:
Eriophyid)]. M. spegazziniana is usually controlled by foliar sprays with dimethoate


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Scientific Notes


directed against adults to prevent damage by feeding and oviposition. P lusca ilus is
controlled mainly by pyrethroids.
The use of microbial control is a potentially valuable alternative to the high costs,
possible pest resurgence, development of resistance, and environmental contamina
tion associated with chemical insecticides. Thus, as a first step toward the develop
ment of a biocontrol program, an investigation was begun to find natural enemies of
these pests in field-collected individuals. This note reports the finding of a fungus in
the psyllid and a baculovirus in the "paraguay tea hornworm". These species are
known by the common names "psilido de la yerba mate" and "marandova de la yerba
mate", respectively.
Sampling was done by randomly collecting the terminal shoot of paraguay tea
plants from groves in Gobernador Virasoro, Corrientes, Argentina. Cadavers of the
psyllid were placed in humid chambers to induce the fungus to sporulate. After this
they were mounted in Hoyer's medium and observed under a stereomicroscope at 400
magnification. Percent disease estimates were made by counting cadavers with myce
lium and sporulating structures. The conidial stage of the fungus was observed dur
ing March, April and early May of 1991 and 1992. A high prevalence (82% mortality,
n=28) of the disease caused by the fungus was observed in the first week of May,1991.
In 1992, the infection rate on April 24, May 11 and July 10 was, respectively, 30%
(n 10), 93% (n 30) and 2% (n 88). We identified the fungus as Zoophthora radicans
(Brefeld) Batko based on the descriptions of Ben-Ze6v & Kenneth (1981), Balazy
(1986) and by careful comparisons of morphology and cultural characters with a spec
imen from a culture collection (ARSEF 2282). This fungus has potential value as a bi
logical control agent, and basic knowledge about it's production process is available
(McCabe & Soper, 1985).
In the summer of 1988, some larvae (1% mortality, n=102) of P lusca ilus displayed
symptoms typical of viral infection -the larvae changed color from typical green to
pale green, they fed less, and after death they were found hanging from the branches
of paraguay tea plants, attached by their anus and legs (Fig. 1). To confirm viral eti
ology, we examined hemolymph under the light microscope. This revealed large num
bers of polyhedra-like particles. The suspected viral polyhedra were concentrated by
homogenization of larval tissue, filtered through cheese cloth, and centrifuged at low
(120 g for 2 min) and high speeds (6,000 g for 15 min). The pellet resulting from high
speed centrifugation was fixed in a modified Karnovsky fixative (2% glutaraldehyde,
2% paraformaldehyde in 0.05 M cacodylate buffer, pH 7.2), postfixed in 1% osmium
tetroxide, dehydrated in acetone, and embedded in Spurr low viscosity medium.
Blocks were sectioned in a LKB Ultratome III microtome equipped with a diamond
knife, and the sections were stained with uranyl acetate and lead citrate before being
examined in a JEOL JEM 100C electron microscope. Also, the suspension of sus
pected viral polyhedra was applied directly onto a specimen holder, air-dried, sputter
coated with gold in a Balzer's sputter coater, and examined in a JEOL 840A scanning
electron microscope.
Electron microscopic examination demonstrated that the particles were typical
baculovirus polyhedra containing large numbers of singly embedded rod-shaped viri
ons (Fig. 2). Scanning electron microscopy revealed large numbers of polyhedral
structures as the sole component of the nuclear polyhedra suspension (Fig. 3). These
polyhedra measured 1 3 micrometers in diam, matching in size the polyhedra seen by
transmission electron microscopy.
A crude preparation of this virus has been successfully used by farmers to control
P lusca ilus larvae, thus avoiding defoliation. This was achieved in a preliminary
study conducted by farmers over approximately 900 ha in Gobernador Virasoro









Florida Entomologist 77(3) September, 1994


Fig. 1. Diseased larvae of Pengonia lusca ilus with symptoms typical of viral infec
tion.

county, in the province of Corrientes, Argentina in February of 1992. A large amount
of the virus was obtained by collecting larvae from artificially-infested fields and
stored frozen for use in the subsequent season. The virus was applied by airplane us
ing the hemolymph from 15 infected last instar larvae per ha although, at that time,
the etiology of the disease was not well understood. Cadavers were collected for use
the next year. In January of 1993, the treated area reached 2,362 ha. This baculovirus
is currently used empirically without the benefit of prior research on dosage, timing,
and population levels.
We thank Dr. James R. Fuxa of Louisiana State University for the critical manu
script review.

SUMMARY

We report for the first time natural and artificial epizootics of pest populations on
Paraguay tea in Corrientes Province, Argentina. The natural epizootic was caused by
Zoophthora radicans on Metaphalara spegazziniana and the artificial epizootic was
caused by a nuclear polyhedrosis virus on Pengonia lusca ilus.









Scientific Notes


Fig. 2. Transmission electron micrographs of thin sections of polyhedral inclusion
bodies of the virus. Note that the m nucleocapsids are wrapped individually by the
membrane characterizing a SNPV (x 40,000)









Florida Entomologist 77(3)


Fig. 3. Scanning electron micrographs of purified polyhedral inclusion bodies of
the virus.


REFERENCES CITED

BALAZY, S. 1986. Taxonomic criteria for inter and intraspecific differentiation in the
entomophthoraceae, exemplified by the subgenus Zoophthora, pp. 201-205 in
R.A. Samson, J.M. Vlak and D. Peters [eds.] Fundamental and Applied aspects
of Invertebrate Pathology. Foundation 4th International Colloquium.
BEN-ZEEV, I., AND R. G. KENNETH. 1981. Zoophthora radicans and Zoophthora petchi
sp. nov. [Zygomycetes: Entomophthorales], two species of "Sphaerosperma
group" attacking leaf hoppers and frog-hoppers [Hom.]. Entomophaga. 26: 131
142.
MCCABE, D., AND R. S. SOPER. 1985. US Patent 4530834 (July,23, 1985).


4444444444444444444444444444444444444444444444444444


September, 1994









Florida Entomologist 77(3) September, 1994


BEE FLIES OF THE BRITISH VIRGIN ISLANDS
(DIPTERA: BOMBYLIIDAE)

NEAL L. EVENHUIS AND SCOTT E. MILLER
Bishop Museum, Box 19000-A, Honolulu, HI 96817-0916

Heretofore no bee flies have been recorded from the British Virgin Islands (BVI)
(Evenhuis, 1983, 1992). Survey work for other insects in the BVI by The Conservation
Agency from 1984 to 1993 has recorded 6 species of bee flies from a total of 9 islands.
The intensity of survey has varied considerably from island to island. Guana Island



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Scientific Notes


has been extensively surveyed, while some islands were visited only briefly (e.g., Eus
tacia, Necker, Scrub).
Fieldwork was based on Guana Island and many specimens were collected in Mal
aise traps at North Bay. Guana is a small island on the north side of Tortola in the
British Virgin Islands (18 28'N, 64 35'W). While it is small, only 297 ha, and the max
imum elevation is 266 m, it bears a relatively rich biota and has sustained less dam
age by feral animals and man than have many adjacent islands (Becker & Miller,
1992). Scott E. Miller and collaborators were in the field VII-1984, VII-1985, VII-1986,
VII-1987, VII-1988, and X-XI-1990. Roy R. Snelling was in the field X-1991, X-1992,
IV 1993, and VI-VII-1993. Material is split between the Bishop Museum and Smith
sonian Institution, with synoptic vouchers in the Natural History Museum of Los An
geles County and The Natural History Museum (London). A few additional records
were provided from the private collections of Michael Ivie (Norman Island) and Rich
ard Miller.
The material available is summarized in Table 1. All the BVI species are known
from Puerto Rico, and all but Exoprosopa cubana are known from the U.S. Virgin Is
lands (Curran, 1928, 1931). Most of these are also more widespread in the West In
dies. Despite recent collecting in the BVI and U.S. Virgin Islands, Chrysanthaxnero
Fabricius remains known only from the type described from "Americae meridionalis
insulis" (either St. Thomas or St. Croix). Species known from Puerto Rico and the
South American continent, but not yet recorded from the BVI include: Anthrax insu
plans Marston, Poecilanthrax lucifer Fabricius, and Ligyra cerberus Fabricius. We be
lieve that with further collecting in the BVI, these species may also be found.
Field work by Miller and Snelling was supported by The Conservation Agency
(Rhode Island), through a grant from the Falconwood Corporation (New York). We
thank Michael Ivie, Tina M. Kuklenski, James D. Lazell, Richard Miller, and Roy R.
Snelling for assistance in obtaining specimens.


SUMMARY

Six species of bee flies are known from the British Virgin Islands, including records
from 9 islands. The BVI fauna is a subset of that of the Puerto Rican Bank.

TABLE 1. DISTRIBUTION OF BOMBYLIDAE IN THE BRITISH VIRGIN ISLANDS.

a-
0






Heterostylum ferrugineum (Fabr., 1805) x
Anthrax oedipus Fabr., 1805 x x
Chrysanthraxgorgon (Fabr., 1805) x x x x x
Neodiplocampta roederi (Curran, 1931) x x x
Villa lateralis (Say, 1825) x x x x x x x x
Exoprosopa cubana Loew, 1869 x









384 Florida Entomologist 77(3) September, 1994

REFERENCES CITED

BECKER, V. O., AND S. E. MILLER. 1992. The butterflies of Guana Island, British Vir
gin Islands. Bull. Allyn Mus. 136: 19.
CURRAN, C. H. 1928. Diptera. Scientific Survey of Porto Rico and the Virgin Islands
11: 1118. New York Academy of Sciences.
CURRAN, C. H. 1931. First supplement to the 'Diptera of Porto Rico and the Virgin Is
lands'. American Mus. Novitates 456: 123.
EVENHUIS, N. L. 1983. An indexed bibliography of Bombyliidae (Insecta, Diptera).
Theses Zoologicae 4: 1493.
EVENHUIS, N. L. 1992. An indexed bibliography of Bombyliidae (Insecta, Diptera):
Supplement I. Bishop Mus. Tech. Rep. 2: 1136.









Book Review


BOOK REVIEW

BEDDING, R. A., R. J. AKHURST, AND H. K. KAYA. (eds.). 1993. Nematodes and the
biological control of insects. CSIRO; East Melbourne. vi + 178 p. Hardback. ISBN 0
643-05479-0. (Available from ISBS, 5602 NE Hassalo St., Portland, OR 97213-3640 at
US $60.00 plus shipping).

Following closely on the heels of the successful (and out-of-print) CRC Press text
on entomopathogenic nematodes (Gaugler, R. and H. K. Kaya. 1990. Entomopatho
genic Nematodes in Biological Control. CRC Press, Boca Raton, Florida) is this con
tribution from Australia's Commonwealth Scientific and Industrial Research
Organization (CSIRO). Unlike its predecessor, which dealt exclusively with ento
mopathogenic nematodes in the families Steinernematidae and Heterorhabditidae,
this text also contains information on species in the families Phaenopsitylenchidae,
Aphelenchoididae, and Mermithidae which are considered insect-parasitic. Some
members of these families eventually kill their host and they have been used prima
rily as classical biological control agents. In contrast, entomopathogenic nematodes
kill their hosts rapidly and are used as biological insecticides.
The book's 25 contributors hail from seven nations and they offer a diversity of
subject matter. Seven chapters provide case studies of pest insect control with nema
todes in Australia, Korea, the United States and China. One chapter each is devoted
to Phaenopsitylenchidae, Aphelenchoididae and Mermithidae and the remaining four
deal with steinernematids and heterorhabditids. The account of the phaenopsity
lenchid Daledenus siricidicola and its use as a control agent for woodwasps makes for
the most interesting reading of the entire book. This parasitic nematode was the first
employed successfully against an insect pest. Its history as detailed here provides in
sights into the preparation for, and implementation of, a successful biological control
project including the selection of correct strains, continued monitoring after release to
assess effectiveness, and maintenance of lab colonies to ensure viable control agent
stock. Other chapters of note describe the control efforts for the banana weevil borer
and for scarab grubs in turf (the latter providing an excellent literature review). Un
fortunately, some chapters are superficial and provide very little more information
than in a typical scientific journal paper.
The remaining ten chapters pertain to the study and use of steinernematids and
heterorhabditids, and cover subjects of a more general nature including post-applica
tion biology, ecological genetics, integrated control, and bacterial symbionts. Three
chapters are devoted to bacterial symbionts (the "power behind the throne" as one au
thor describes them) which provide the toxin for target hosts and essential nutrients
for the nematodes. As with those of case studies, the form and information content of
these chapters vary somewhat. Most are engaging and informative, but a few are
quite dry, narrow in scope, and almost painful to read.
Due primarily to their increasing importance as commercially-available biological
control agents and their relative ease of rearing, research on steinernematids and
heterorhabditids has proliferated in the past few years, which explains why the great
majority of the book is devoted to them. Much of what is presented was not available
or known at the writing of the Gaugler and Kaya text.
The layout of the book is attractive and functional. Each chapter begins with an
abstract (summary) and the top of each right-hand page contains the chapter title.
The paper is of medium-heavy weight and is semi-glossy The contributing authors'
affiliations are listed as an addendum glued to the last page.



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386 Florida Entomologist 77(3) September, 1994

Moderately priced, this text would be a valuable (and affordable) addition to the li
brary of anyone studying insect-parasitic or entomopathogenic nematodes or those
wishing to stay abreast of this increasingly-important group of biological control
agents.

Patrick Parkman
Entomology & Nematology Dept.
University of Florida
Gainesville, FL 32611-0620









IN MEMORIAL


IN MEMORIAL


HARRY R. GROSS, JR.

(1939-1994)

Harry R. Gross, Jr., was born in New Orleans, LA, on March 16, 1939, and passed
away on May 3, 1994 in Wesley Woods Hospital in Decatur, GA. He is survived by his
wife, Marlyne Adam Gross of Tifton, and a daughter, Lisa Gross of Atlanta, GA. Dr.
Gross was a Supervisory Research Entomologist and Research Leader of the Insect
Biology/Management Systems Research Unit, USDA-ARS, Insect Biology and Popu
lation Management Research Laboratory at Tifton, GA.
He graduated from Louisiana State University, obtaining his B.S. (1960), M.S.
(1964), and Ph.D. (1967) degrees in Entomology. Throughout his career, Dr. Gross was
employed by the U.S. Department of Agriculture, Agricultural Research Service. He
began his research career in 1967 at Gulfport, MS, working on the white-fringed bee
tle before transferring in 1971 to the Insect Biology and Population Management Re
search Laboratory (formerly Southern Grain Insects Research Laboratory), Tifton,
GA. Dr. Gross was a member of Sigma Xi, the Entomological Society of America, the
Southeastern Branch of ESA, the Southeastern Biological Control Working Group
(SBCWG), and the Georgia and Florida Entomological Societies. Dr. Gross was Pres
ident of the SBCWG in 1990. He was awarded the USDA Certificate of Merit in 1983
for exemplary leadership and managerial accomplishments as Director and Research
Leader for the Insect Suppression Research Unit. He served as a member of an advi
sory panel to provide expertise in the development of a National Biological Control
Program for ARS. Dr. Gross published 74 articles and made over 70 scientific present
stations at professional meetings (10 of which were invitational) during his career.


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Florida Entomologist 77(3) September, 1994


Harry served on numerous committees within the Sigma Xi organization (Program
Chairman), the Georgia Entomological Society (Public Relations, Resolutions, and 0.
I. Snapp Awards committees), the Entomological Society of America (Fellows Commit
tee), and Southeastern Branch ESA (Committee on Constitutional Amendments, Pro
gram Chairman, Agricultural Awards, and Auditing Committees). Dr. Gross also
served as Chairman of the Southern Regional Project S-59, Fall Armyworm Confer
ence and the Heliothis Conference.
Dr. Gross was a dedicated scientist who spent many long and hard hours at his
work. His invention (08/062,516) on "Beehive-mounted device for utilizing honeybees
(Hymenoptera: Apidae) in the dissemination of biocontrol agents" was allowed in May
1994. Dr. Gross was an expert on integrated biocontrol strategies for key insect pests
and highly knowledgeable of their behavior and population dynamics. He unraveled
several mysteries of how parasites and predators use an array of chemical signals to
identify and locate pest insects, and how man might manipulate parasites and pred
ators to provide better control of insect pests. He also helped build a foundation for ba
sic knowledge that is, and will continue to be, used by entomologists throughout the
world. Dr. Gross was the first to demonstrate the potential for augmentative releases
of Archytas marmoratus, a primary parasitoid of the corn earworm and fall army
worm, and developed a system for its large scale rearing. He demonstrated that corn
earworm male moths use visual cues to locate mates and showed that these visual
cues could improve the efficiency of insect pheromone traps. Dr. Gross also demon
strated the presence of previously unknown indigenous pupal parasitoids for the fall
armyworm and the tobacco budworm. The frequent citations of his research in inter
national scientific journals offer but a glimpse of the significance with which Dr.
Gross's contributions are viewed by the world scientific community. Harry will long be
remembered by his friends, colleagues, coworkers, and especially by the technical
support personnel, because of his friendliness, fairness and leadership at the IBP
MRL. At the request of Mrs. Gross, a "Harry R. Gross, Jr. Memorial Fund" has been
established with the Tifton Sigma Xi Club. Contributions may be made in Harry's
name and sent to the Sigma Xi Club, c/o Dr. Larry Newton, CPES, P.O. Box 748, Tif
ton, GA. 31793-0748.

The Insect Biology and Population Management Research Laboratory:
J. E. Carpenter, L. D. Chandler, J. J. Hamm, W. J. Lewis, R. E.
Lynch, O. G. Marti, S. D. Pair, W. D. Perkins, C. E. Rogers, A. N.
Sparks, H. R. Sumner, N. W. Widstrom, and B. R. Wiseman.
USDA, ARS, IBPMRL, Tifton, GA 31793.









IN MEMORIAL


IN MEMORIAL


DR. YIN-CHI Hsu

The standard reference on mayfly biology remains the 1935 classic Biology ofMay
flies by J. G. Needham, J. R Traver, and Yin-Chi Hsu, originally published by Com-
stock Publishing of Ithaca, New York. Needham and Traver are familiar names, but
the third author is somewhat of an enigma to North American entomologists. Many
are surprised to learn that Hsu continued his research for more than 50 years and, in
addition to his role in North American entomology, Hsu was the founder of
Ephemeroptera study in China, as well as being that country's leading human para
sitologist. I was fortunate to have met him when he visited Florida A&M University
in 1983, and I am happy to have the opportunity to submit this tribute, prepared by
colleagues in Nanjing, to one historic American mayfly scientist.

William L. Peters, Florida A&M University, Tallahassee

OUR TUTOR DR. XU YINQI (HSU YIN-CHI)
(SEPT. 9, 1905-JAN. 29, 1986)

Prof. Xu Yinqi commanded the highest respect in China as a prominent entomolo
gist, parasitologist, and educator. A member of an educated family, he was a highly in
telligent and dedicated student, and earned a BS in Biology from Soochow University
in 1926, with honorary titles of Beta Beta Beta and Phi Tau Phi. In 1927, he entered
Yenching University, Peking, and graduated with an MS in Zoology in 1929; his MS
dissertation was entitled "Gryllidae in China." Because of his outstanding work at
Yenching, he was awarded a Rockefeller Foundation Fellowship to Cornell University,
completing there his PhD degree, with honors, in 1932. He was also offered the Prize


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Florida Entomologist 77(3) September, 1994


of Scientific Research under auspices of China's Cultural Fund and later received the
Sino-American Service Professor Fellowship at the University of Minnesota. In 1932,
he was elected a member of Sigma Xi; also in 1932, he attended the Vth International
Congress of Entomology in Paris, France.
Early in his professional career, Prof. Xu Yinqi was devotedly engaged in research
on Ephemeroptera taxonomy and morphology. After coauthoring (with J. G. Needham
and J. R. Traver) the Biology of Mayflies, published in 1935, a universally acknowl
edged classic in Ephemeroptera research, he (1936-1938) published a monograph on
The Mayflies of China. This work presented a comprehensive survey and summary of
all previous Ephemeroptera in China and added many descriptions of new species of
China. Prof. Xu Yinqi, considered the founder of Ephemeroptera research in China,
enjoyed international renown.
In the early 1950s, he turned his studies to Parasitology and Acarology, investigate
ing serious problems of mite typhus in China. He and his assistants began a system
atic probe into the classification, ecotype, and fauna of chigger mites and their role in
transmission of communicable diseases. His forty-odd pieces of published academic
treatises on the subject have been frequently quoted, and provide a foundation for this
field of medical acarology
In the latter part of the 1970's, he and his research team studied interrelation
ships between Demodicidae and demodicidosis-dermatopathy He conducted clinical
case analyses and pathological studies on the morphological structure, classification,
and distribution of Demodicidae on folliculitus, rosacea, etc. as well as treatments for
these conditions, publishing extensively on the subject. Prof. Xu Yinqi constantly pro
moted Chinese science and published numerous treatises, monographs and text
books.
Prof. Xu Yinqi dedicated his life to teaching in the biological sciences. He served as
a professor in, sequentially, Soochow University, Yenching University, St. John's Uni
versity, Shanghai First Medical College (with a long term as Dean of the Biology De
apartment Soochow University (1938 1952), and Professor-in-Charge of the Board of
Studies of Parasitology, Shanghai First Medical College (1952-1984). He was on the
Committee of Experts, Shanghai Medical University from 1984 until 1986. For 59
years, he taught courses on Invertebrate Zoology, Comparative Anatomy, Human Par
asitology, Entomology, Medical Entomology, Entomological Taxonomy, Medical Ac
arology, Histology, Embryology, etc. He was an erudite scholar, an animated lecturer,
and an indefatigable teacher who was loved and esteemed by his students. His public
cation, How to Deliver Successfully a Lecture in Class, elevated and elucidated the im
portance of instruction for younger faculty members. Numbered among his students
are academicians of the Academy of Sciences of China, presidents of institutions of
higher learning, professors, and doctors, all playing important roles in biological,
medical, and educational circles in China.
Prof. Xu Yinqi, one of the sponsors of the Zoology Society of China and Entomology
Society of China, served long terms on the Boards of Directors of both societies, as well
as acting as Deputy Director-General of the Shanghai Parasitology Society and as ac
ademic advisor to the Shanghai Museum of Natural Science. Concurrently, he was
Deputy Editor-in-Chief of the Journal of Parasitology and Parasitic Disease and a
member of the Editorial Board of Acta Entomologica Sinica, Acta Zootaxonomica Sin-
ica, Acta Academiae Medicinae Primae Shanghai, Lexicon (section on medicine), Eng
lish-Chinese Dictionary of Agricultural Entomology, Glossary and Nomenclature of
Acarology and others.
Professor Xu Yinqi adopted a vigorous scientific approach in his academic work
and was always well respected by his colleagues. As a professor, he was direct, decent,









IN MEMORIAL


enthusiastic and encouraging. He will be long remembered for his outstanding contri
butions to Chinese education and scientific research.


Publications on Ephemeroptera by Xu Yinqi [Hsu Yin-Chi]

1. Hsu, Yin-Chi. 1931. Two new species of mayflies from China. Peking Nat. Hist.
Bull. 6(2):39-41.
2. 1932. Morphology, anatomy, and ecology of genus Heptagenia. Ph.D. Diss.,
Cornell Univ. Graduate School, Dep. Entomol.
3. 1933. Some new morphological findings in Ephemeroptera. 5th Int. Congr.
Entomol., Paris, 1932, 2:361-368. 2 p1s.
4. 1935. New Chinese mayflies from Kiangsi Province. Peking Nat. Hist. Bull.
10(40):319-326.
5. Needham, J. G., J. R. Traver and Y.-C. Hsu. 1935. The Biology ofMayflies with
a Systematic Account of North American Species. Comstock Publ., Ithaca, New
York. xi + 759 pp., 168 figs., plates I-XL.
6. Hsu, Y.-C. 1936. Mayflies of Hong Kong with description of two new species
(Ephemeroptera). Hong Kong Nat. 7:233-238.
7. 1936-1937. The Mayflies of China I. Peking Nat. Hist. Bull. 11:129-148.
8. 1936-1937. The Mayflies of China II. Peking Nat. Hist. Bull. 11:287-296.
9. 1936-1937. The Mayflies of China III. Peking Nat. Hist. Bull. 11:433-440.
10. 1937-1938. The Mayflies of China IV. Peking Nat. Hist. Bull. 12:53-56.
11. 1937-1938. The Mayflies of China V Peking Nat. Hist. Bull. 12:123-126.
12. 1937-1938. The Mayflies of China VI. Peking Nat. Hist. Bull. 12:221-224.
13. You Da-Shou, Wu Tian, Gui Hong and Hsu Yin-Chi. 1980. Anew species of the
genus Choroterpes from Nanjing (Ephemeroptera: Leptophlebiidae). Acta Zoot
axonomica Sinica 5(4):388-391.
14. Xu Jia-zhu, You Da-shou, Su Cui-rong and Xu Yin-qi. 1980. Two new species of
genus Ephemerella (Ephemeroptera: Ephemerellidae). J. Nanjing Teacher's
College 1980(2):60-63.
15. You Da-shou, Wu Tian, Gui Hong and Hsu Yin-Chi. 1981. Two new species and
diagnostic characters of genus Cinygmina (Ephemeroptera; Ecdyoneuridae). J.
Nanjing Teacher's College 1981(3):26-32.
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Gui Hong
Zhang Jun
Department of Biology
Nanjing Normal
University









Florida Entomologist 77(3)


IN MEMORIAL

HARRY KYDD GOUCK

Harry Kydd Gouck, a retired research entomologist from the USDA-ARS Medical
and Veterinary Entomology Research Laboratory, Gainesville, FL died 9 May, 1994 af
ter a long illness.
Harry Gouck was born and raised in Andover, MA. He attended the University of
New Hampshire where he earned a B.S. degree in entomology in 1935 and played var
sity football. He continued his education at the University of Massachusetts where he
received his M.S. degree in entomology in 1936. While a student, and after receiving
his M.S. degree, he worked for the USDA under the direction of Dr. C. N. Smith on the
island of Martha's Vineyard, MA. Together, they studied the biology and ecology of the
American dog tick. When this laboratory was closed in 1942, the staff, including
Harry, moved with their families to Savannah, GA, to continue research on ticks.
The laboratory in Savannah was split in 1947, and some of the staff, including
Harry, joined other scientists at a newly organized facility expressly devoted to the
control of medically-important insects. Established at Orlando, FL in 1942, this labo
ratory was the Insects Affecting Man and Animals Research Laboratory which had
the primary task of solving insect-borne disease problems facing U.S. combat troops
during World War II. Ulii 11, I, this laboratory and its scientists received global rec
ognition for their contributions to the field of medical entomology.
In 1950, Harry was transferred to the USDA Corn Research laboratory at Cham-
paign-Urbana, IL, where he worked on the control of the corn earworm for two years.
In 1952, Harry and his family returned to the Orlando laboratory to find that his
colleague, Dr. C. N. Smith, had become laboratory director. Harry's new job was to
evaluate and develop insect repellents. He tested numerous chemicals as repellents


This article is from Florida Entomologist Online, Vol. 77, No. 3 (1994).
FEO is available from the Florida Center for Library Automation gopher (sally.fcla.ufl.edu)
and is identical to Florida Entomologist (An International Journal for the Americas).
FEO is prepared by E. O. Painter Printing Co., P.O. Box 877, DeLeon Springs, FL. 32130.


September, 1994









IN MEMORIAL 393

against mosquitoes, biting flies, ticks, chiggers, and leaches. This research took him
to field sites in Asia, Africa, Central America and the Bahama Islands. He collabo
rated in the discovery and development of the repellent deet which, to this day, is the
most widely used insect repellent in the world.
When the laboratory in Orlando was moved to the campus of the University of
Florida in 1963, Harry was named head of the mosquito repellents and attractants re
search project. In 1968, together with laboratory chemists, he participated in research
that led to the identification of lactic acid from human skin as a mosquito attractant.
During his career Harry wrote and/or co-authored more than 60 scientific public
tions and reported this research at major scientific meetings. He retired from the
USDAin 1973 and, together with his wife, started a second career as a flea marketeer
and antique dealer. He continued this work until his health declined three years ago.
Harry is survived by his daughter, Polly Hall of Atlanta, GA, son Peter E. Gouck
of North Attleboro, MA, and three grandsons.

Richard S. Patterson
Carl E. Schreck
Medical and Veterinary Entomology Research Laboratory
USDA-ARS, SAA MAVERL
1600 SW 23rd Drive
Gainesville, FL 32604




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