Title: Florida Entomologist
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Permanent Link: http://ufdc.ufl.edu/UF00098813/00085
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1986
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
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Bibliographic ID: UF00098813
Volume ID: VID00085
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text

(ISSN 0015-4040)


(An International Journal for the Americas)

Volume 69, No. 2 June, 1986


CHASE, A. R., L. S. OSBORNE, AND V. M. FERGUSON-Selective Isolation of
the Entomopathogenic Fungi Beauveria bassiana and Metarhizium anisop
liae From an Artificial Potting Medium ......................................... 285
FOSTER, R. E.-Monitoring Populations of Lyriomyza trifolii (Diptera: Ag-
romyzidae) in Celery with Pupal Counts .......................................... 292
WILLIAMS, R. N., AND L. A. B. DE SALLES--Nitidulidae Associated with Fruit
Crops in Rio Grande do Sul, Brazil ............................................... 298
ATKINSON, T. H., AND A. EQUIHUA-Biology of the Scolytidae and Platypodidae
(Coleoptera) in a Tropical Deciduous Forest at Chamela, Jalisco, Mexico .. 303
DE RODRIGUEZ, M. C., AND W. W. WIRTH-A New Species of Man-Biting
Culicoides from the High Andes of Columbia (Diptera: Ceratopogonidae) 311
GORDON, A. E., S. C. HARRIS, AND P. K. LAGO-Description of new species of
Cheumatopsyche (Trichoptera: Hydropsychidae) and the presumed female
of C. helma Ross from Alabama ..................................................... 314
EPLER, J. H.-A Novel New Neotropical Nanocladius (Diptera: (C I,-. r..,... I,
Symphoretic on Traverella (Ephemeroptera: Leptophlebiidae) .............. 319
DEYRUP, M., AND D. MANLEY-Sex-Biased Size Variation in Velvet Ants
(Hymenoptera: Mutillidae) ..................................... .................... 327
TRYON, E. H., JR.-The Striped Earwig, and Ant Predators of Sugar Cane
Rootstock Borer, in Florida Citrus ................................................. 336
PENA, J. E., R. M. BARANOWSKI, AND R. E. LITZ-Oviposition of the Papaya
Fruit Fly Toxotrypana curvicauda as Affected by Fruit Maturity .......... 344
SAKIMURA, K.-Thrips in and Around the Coconut Plantations in Jamaica, with
a Few Taxonomical Notes (Thysanoptera) ....................................... 348
FRANK, J. H.-A Preliminary (C... '.l", of the Staphylinidae (Coleoptera) of
Florida ................................................................... ................. 363
ING-Use of Elcar to Reduce the Number of Corn Earworms (Lepidoptera:
Noctuidae) Developing in Field Corn .............................................. 383
RILEY, T. J.-Greenhouse and Field Evaluation of Granular Soil Insecticidesfor
Control of Sugarcane Beetle, Eutheola rugiceps (Coleoptera: Scarabaeidae)
in F ield C orn ..................................................... ..................... 390
WOJCIK, D. P.-Bibliography of Imported Fire Ants and Their Control: Second
Supplement ..................................... 394

Continued on Back Cover

Published by The Florida Ent,,,lnld,,r;irl Society


President ..................................... .............. D. H. Habeck
President-Elect ................................. ....................... D. J. Schuster
Vice-President ............................................. ................ J. L. Taylor
Secretary ........................................... ............................... E R M itchell
Treasurer ............................................ ............... A. C. Knapp

M. L. Wright, Jr.
J. E. Eger, Jr.
R. C. Bullock
Other Members of the Executive Committee ....... Mtullin
C.G. Mathurin
C, G. Witherington
J. R. McLaughlin


Editor .......................................... .................. J. R. McLaughlin
Associate Editors .................... ........... ..... .................... A. Ali
C. S. Barfield
J. B. Heppner
M. D. Hubbard
O. Sosa, Jr.
H. V. Weems, Jr.
W. W. Wirth
Business Manager ....................................... ..... .................. A. C. Knapp

FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30.00 per year in advance, $7.50 per
copy. Membership in the Florida Entomological Society, including subscription to Flor-
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Inquires regarding membership, subscriptions, and page charges should be addressed
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Authors should consult "Instructions to Authors" on the inside cover of all recent
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Manuscripts and other editorial matter should be sent to the Editor, JOHN R.
MCLAUGHLIN, 4628 NW 40th Street, Gainesville, FL 32606.

This issue mailed August 8, 1986


The Florida Entomological Society will hold its 69th Annual Meeting on
6-8 August 1986 at the Sheraton Sand Key Hotel, Clearwater Beach, Florida. The
location is 1160 Gulf Boulevard, Clearwater Beach, Florida 33515; telephone-1-813-595-
1611. Room rates will be $66.00 either single or double. Pre-registration and registration
fees will be released in the June, 1986 Florida Entomologist and the April Newslbtt,1r.

Since many will present papers please copy the sheet and submit before 1 .June 1: 9

Program Committee, FES
P. O. Box 1893
Sanford, Florida 32771
Phone: 1-305-322-5716

Eight minutes will be allotted for presentation of oral papers, with 2 minutes for
discussion. In addition, there will be a separate session for members who may elect to
present a Project (or Poster) Exhibit. The three oral student papers judged to be the
best on content and delivery will be awarded monetary prizes during the n,..-tl,
Student authors must be Florida Entomological Society Members and must be regis-
tered for the meeting.


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Chase et al.: Isolation of Fungi


Agricultural Research and Education Center
2807 Binion Road
Apopka, FL 32703

A basal medium consisting of oatmeal agar was superior to seven other media for
isolation of Metarhizium anisopliae and Beauveria bassiana from an artificial potting
medium. A rate of 0.55 g/1 of dodine and 5 mg/1 of chlortetracycline allowed optimum
recovery of B. bassiana alone. Decreasing the amount of dodine to 0.46 g/1 and adding
0.38 g benomyl allowed recovery of both entomopathogens at high frequencies in the
abscence of most other fungi. Addition of 10 mg/1 crystal violet aided in visibility of the
target fungi without affecting recovery frequencies. Exposure of culture plates to as
little as 2 pmol m 2s-1 light delayed observation of colonies of both entomopathogens
up to 7 days compared to cultures incubated in the dark.


Un medio basico consistent de agar de avena fue superior a otros 7 medios para
aislar a Metarhizium anisoplia y Beauveria bassiana de un medio de siembra artificial.
Una raz6n de 0.55 g/1 de dodine y 5 mg/1 de clorotetraciclina permiti6 una recuperaci6n
de B. bassiana solamente. Disminuyendo la cantidad de dodine a 0.46 g/1 y afiadiendo
0.38 g de benomil permiti6 la recuperaci6n de ambos entomopat6genos en frecuencias
altas en la ausencia de la majoria de otros hongos. Afadiendo 10 mg/1 de cristales de
violeta ayud6 la visibilidad del hongo sin afectar las frecuencias de su recobro. Ex-
poniendo plates de cultural a niveles tan bajo de luz como 2 umol m- s-', demor6 la
observaci6n de colonies de ambos entomopat6genos hasta 7 dias en comparaci6n con
cultures incubadas en la oscuridad.

With increased concern over environmental impacts of chemical pesticides, consider-
able research has focused on biological pesticides. Entomopathogenic fungi such as
Beauveria bassiana (Bals.) Vuill. and Metarhizium anisopliae (Metschn.) Sorokin have
been investigated extensively over the past 10 to 15 years (Ferron 1981, Roberts and
Campbell 1977). Interest in commercial application of these biological pesticides has led
to development of information regarding their biology (Barnes et al. 1975, Campbell et
al. 1983), ecology (Doberski and Tribe 1980), pathology (Doberski 1981, Gardner and
Noblet 1978), and effects of chemical pesticides (Olmert and Kenneth 1974) under a wide
range of conditions. Both bioassay with a susceptible insect host and direct isolation
from soil, insect, or plant tissue have been utilized to evaluate reactions of these fungi
to various conditions. Direct isolation of conidia of B. bassiana and M. anisopliae have
primarily involved use of semi-selective culture media. One of the earliest media was
based on one used for general isolation of soil fungi and contained glucose, oxgall, and
peptone with rose bengal, chloramphenicol, and cycloheximide as antibiotics (Veen and
Ferron 1966). Doberski and Tribe (1980) developed a medium based on similar ingre-
dients with crystal violet substituted for rose bengal. This medium was used to isolate


286 Florida Entomologist 69(2) June, 1986

B. bassiana and M. anisopliae from elm bark and soil. A medium with V-8 juice,
glucose, yeast extract, and oxgall as the nutrient base amended with cycloheximide,
streptomycin sulfate, and tetracycline was developed for general isolation of B. tenella
from soils (Joussier and Catroux 1976). Sabouraud-dextrose medium and oxgall was
used as another basal medium amended with penicillin G, streptomycin sulfate, oxytet-
racycline, cycloheximide, and binapacryl (Morocide, a fungicide). This medium was
developed for use in isolations from field soil maintained under laboratory conditions
(Lingg and Donaldson 1981). More recently, Beilharz et al. (1982) discovered that dodine
(n-dodecylguanidine acetate) selectively inhibited some soil fungi other than B. bassiana
and M.anisopliae when added at the rate of 1.0 g formulated (Cyprex 65WP, American
Cyanamid) product per liter. Each of these media was developed in response to a need
by researchers to evaluate effects of various treatments on survival of these en-
tomopathogens. The present paper describes a modification of one of these media
(Beilharz et al. 1982) to maximize recovery of either B. bassiana or M. anisopliae from
an artificial potting medium.



The basal medium used for most trials was oatmeal agar prepared from 20 g Gerber
Oatmeal Cereal For Baby (Gerber Products Co., MFR, Fremont, MI 49412) autoclaved
for 20 min with 1 1 of deionized water. The suspension was immediately filtered twice
through four layers of cheesecloth, and the volume of filtrate adjusted to 1 1 with
deionized water. Agar (20 g/1) was ground in a mortar and pestle with various amounts
of dodine and added to the oatmeal filtrate while stirring. The final medium was autoc-
laved for 20 min and cooled to 480C prior to adding chlortetracycline (5 mg/1). One 1 of
medium was used to pour 50 plates, (90 x 15 mm).


An experimental wettable powder formulation of Beauveria bassiana (ABG 6112,
Abbott Laboratories, North Chicago, IL) and a formulation of Metarhizium anisopliae
(70-25 Mycogen Corporation, San Diego, CA) were used in all trials. A potting medium
consisting of 3 parts Canadian peat and one part builder's sand was amended with 4 kg
ground dolomitic limestone and 1 kg Micromax (micronutrient source from Sierra Chem-
ical Co., Milpitas, CA) per m3 and steam-treated at 950C for ca. 1 hr. Beauveria bassiana
or M. anisopliae or both were added to 500 g of the potting medium at the rate of 0.5
g formulation and mixed thoroughly by hand. The medium was used the same day it
was prepared.


Ten g samples of the potting medium-conidial preparation were added to 100 ml of
sterilized deionized water (SDW) and placed on a rotary action shaker for 15 min. Serial
dilutions were made with SDW, and 0.5 ml aliquots of the 10-3 and 10-4 dilutions were
spread on the surface of five plates of each medium. Plates for all trials except those
concerning light levels were incubated at 25 to 27C with 10 pImol m-2s-1 light (Sylvania,
Gro-lux, fluorescent bulbs) for 16 hr daily. Colony counts were determined at various
times depending upon the trial. Specific trials with media and incubation conditions are
described below.

Chase et al.: Isolation of Fungi 287


The oatmeal-dodine medium was compared to several media developed by others as
well as basal media amended with dodine (variable rates) and chlortetracycline (5 mg/1).
The following media were tested at least twice: 1) Lingg and Donaldson basal medium
(1981) with dodine (0.62 g/l) and chlortetracycline amendments: 2) Doberski and Tribe
(1980); 3) Martin's RB-M2 medium (Tuite, 1969); 4) Sabouraud's Dextrose agar (Difco)
amended with 5 g per liter of yeast extract, dodine (0.62 g/l) and chlortetracycline; 5)
Czapek's-Dox medium (Difco) amended with dodine (0.58 or 0.62 g/l) and chlortetracyc-
line; 6) Pasteur's-MC sporulation medium (0.36 g KH2PO4, 1.05 g NaHPO4 7H20, 0.6
g MgSO4, 1.0 g KC1, 10.0 g dextrose, 0.7 g NH4NO3, 5.0 g yeast extract, and 20.0 g
agar per liter) amended with dodine (0.58 or 0.62 g/1) and chlortetracycline; and 7)
Potato Dextrose agar medium (Difco) amended with dodine (0.62 g/l) and chlortetracyc-


Dodine rates were varied from 0.16 to 0.62 g per liter using oatmeal agar as the
basal medium. The following rates were used in at least one of five tests: 1) 0.65, 0.49,
0.32, and 0.16 g1l; 2) 0.65, 0.58, 0.52, and 0.46 g1l; 3) 0.52, 0.49, 0.46, and 0.39 g1l; 4)
0.52, 0.49, 0.46, and 0.39 g1l; and 5) 0.65, 0.62, 0.58, 0.55, 0.52, 0.49, 0.46, and 0.39 g/1.


Benomyl (Benlate 50WP, E. I. Dupont Nemours, Inc., Wilmington, DE) is known
to inhibit in vitro growth of both B. bassiana and M. anisopliae (Olmert and Kenneth
1974, Tedders 1981) at rates used to treat plants. The effect of very low rates on
recovery of these two entomopathogens was tested. Benomyl (prepared as a stock
suspension in SDW) was added to the basal medium prior to autoclaving. The following
rates were used in three tests: 0, 0.25, 0.5, and 1.0 mg/1. Two additional tests were
performed using 0, 0.25, 0.38 and 0.5 mg benomyl/1.


The medium of Doberski and Tribe (1980) contains 10 ppm crystal violet. When this
rate of crystal violet was added to the oatmeal medium, M. anisopliae was not reco-
vered, although B. bassiana recovery was unaffected in a preliminary trial. Various
rates of crystal violet were therefore tested for a differential effect on recovery of the
two fungi. Rates from 0 to 20 ppm were included: (Test 1) 0, 2, 10 and 20 mg/1 (Test 2)
2.5, 5.0, 7.5 and 10 mg/l, and (Test 3) 0, 5, 10 and 15 mg/1.


Two types of tests were performed to evaluate the effect of light level on recovery
of B. bassiana and M. anisopliae on oatmeal medium amended with 0.46 g dodine/1. In
the first test, 25 plates were treated with 0.5 ml of a 104 soil dilution of either M.
anisopliae or B. bassiana. Plates were arranged in five stacks of five plates each with
the bottom plate in each stack wrapped in foil for each fungus. Stacks were incubated
with 10 pmol m-'s-1 fluorescent light and colony counts recorded as soon as possible.
The second type of test was performed with plates in a single layer under four light
levels (10 plates per treatment): 1) wrapped in foil; 2) 2 pimol m-2 s-1; 3) 5 xmol m-2 s-';
and 4) 10 gmol m2 s-'. The second type of test was performed twice.
All data were analyzed for variance using the F test for significance.

Florida Entomologist 69(2)

June, 1986



The mean number of colonies per plate was significantly affected by the medium
used for isolation. Beauveria bassiana was recovered in high frequencies on oatmeal
agar and Sabouraud medium amended with yeast extract, but not from other culture
media utilized. Metarhizium anisopliae was recovered only from oatmeal agar medium.
Mean recovery per plate for B. bassiana on oatmeal medium ranged from 29 to 92
colonies (10-4 dilution rate), while recovery for M. anisopliae ranged from 17 to 45 (10-4
dilution rate). These recovery frequencies relate to a percent recovery between 50 and
80 for the majority of trials.


An optimum rate of dodine was identified for isolation of each target fungus. The
rate of 0.62 g/1 dodine significantly reduced recovery of M. anisopliae (Table 1). De-
creasing the rate/l to 0.55 g allowed a high recovery of B. bassiana, but was still too
high for recovery of M. anisopliae which grew out only when the rate was decreased
to 0.46 g/1. Rates of dodine from 0.32 to 0.62 generally did not influence recovery of B.
bassiana (Table 1). Rates of dodine lower than 0.32 allowed good recoveries of the
target fungi, but proved to be too low to inhibit germination and growth of several
saprophytic soil fungi such as Trichoderma and Rhizopus spp. (data not included).


Mean recovery at 10-4 dilution
Beauveria bassiana (Tests) Metarhizium anisopliae (Tests)
Dodine g/1 1 2 3 4 5 1 2 3 4 5

0.32 19.2a NTb NT NT NT 193.0 NT NT NT NT
0.39 NT NT 10.2 17.4 21.0 NT NT 20.2 8.2 19.6
0.46 NT 22.6 8.2 22.2 16.6 NT 67.8 25.2 10.8 21.0
0.49 25.0 NT 10.2 19.6 23.8 23.6 NT 0.8 0 16.0
0.52 NT 20.2 9.6 11.0 15.8 NT 4.0 7.6 0 5.8
0.55 NT NT NT NT 19.2 NT NT NT NT 5.2
0.58 NT 18.6 NT NT 13.8 NT 0.8 NT NT 4.0
0.62 NT NT NT NT 13.4 NT NT NT NT 2.6
0.65 16.4 0.4 NT NT 0 0.4 0 NT NT 0

Significance ** ** ns ns ** ** ** ** ** **

aMean number of colonies for five replicate plates for each test.
bNT = Not tested.
'Significance level of the F test denoted as follows: ns = not significant, and **

Chase et al.: Isolation of Fungi


A rate of 0.25 mg/1 of benomyl did not affect recovery of either fungus significantly
while a rate of 0.38 mg/1 reduced recovery of B. bassiana to undetectable levels, but
did not affect recovery of M. anisopliae. Rates of 0.5 to 1.0 mg/1 significantly decreased
recovery frequencies of both (Table 2). Tests were performed using the oatmeal-dodine
medium with dodine used at 0.46 g/l to allow recovery of M. anisopliae. Since this rate
does allow occasional colonies of Trichoderma spp. to develop, the effect of low rates
of benomyl proved beneficial in eliminating these contaminants. A rate of 0.38 mg/1 of
benomyl was chosen for addition to oatmeal-dodine medium when the 0.46 g/1 rate of
dodine was employed to eliminate saprophytes and select M. anisopliae over B. bas-


Although a preliminary test showed differential effects of crystal violet concentration
on recovery of the entomopathogens, the results were not repeated in any subsequent
test. Rates up to 10 mg/1 did not affect recovery of either fungus, while those above 10
mg/1 reduced recovery of the two entomopathogens similarly. Because oatmeal agar is
an opaque, white medium and both organisms initially produce white to creamy colored
colonies, the addition of 10 mg/1 of crystal violet greatly improved colony visibility.


When plates of either B. bassiana or M. anisopliae were incubated in stacks under
10 p.mol m-2 s-' light, the recovery frequency and the number of days required for
incubation were affected by their position in the stack (data not included in a table).
Plates on the bottom of the stacks, which were wrapped in foil, could be counted two


Mean recovery at 10-4 dilution
Beauveria bassiana (Tests) Metarhizium anisopliae (Tests)

Benomyl (mg/1) 1 2 3 4 1 2 3 4

0 12.2a 13.5 25.8 27.0 7.6 3.4 30.4 63.6
0.25 13.4 14.2 21.8 22.4 9.0 4.8 6.0 27.8
0.38 NTb NT 0 0 NT NT 9.7 20.2
0.50 0 0 0 0 10.1 0 0 0
1.00 0 0 NT NT 0 0 NT NT

Significance' ** ** ** ** ** ** ** **

aMean number of colonies for five replicate plates for each test.
bNT = Not tested.
CSignificante level of the F test denoted as follows: ** P<0.01.

290 Florida Entomologist 69(2) June, 1986

to five days earlier than those on the top of the stack. In addition, recovery frequencies
of both target fungi were reduced up to 60% when incubated on the top of the stack
compared to the foil wrapped plates. B. bassiana plates at the top of the stack yielded
a mean of nine colonies/plate while those at the bottom yielded a mean of 28. Similarly,
the number of M. anisopliae colonies was only 48 on the top of the stack compared to
a mean of 115 at the bottom of the stack. Exposure of plates to various levels of light
during incubation demonstrated that even 2 |.mol m-2 s-' light reduced recovery fre-
quencies of the target fungi in some tests (Table 3). As in the previous test, colony
counts for plates wrapped in foil were made approximately two to seven days earlier
than plates in 10 Rmol m-2 s-1 for B. bassiana and M. anisopliae respectively.
From these studies it can be concluded that a basal medium of oatmeal agar supports
the highest recovery of both B. bassiana and M. anisopliae from artificial potting
medium. Comparisons to most of the media developed by other researchers suggest
that a single medium may not be effective under all the conditions encountered in
research on these entomopathogens. It should be remembered, however, that there is
a certain amount of natural variability of isolates of fungi and that our media may not
be adequate for recovery of naturally occurring B. bassiana or M. anisopliae.
The effects of both dodine and benomyl on the recovery of these entomopathogens
have led to the development of two selective media. Selective isolation of B. bassiana
can occur if dodine is incorporated in the oatmeal medium at the rate of 0.55 g/1. Higher
rates of dodine generally do not decrease recovery of the B. bassiana, but the previously
used rate of 0.65 g1l significantly extends the period of time needed to incubate the
plates. A much lower rate of dodine (0.46 g/l) must be used to allow high recoveries of
M. anisopliae. Since this rate of dodine also allows good recovery of B. bassiana and
certain soil saprophytes, the addition of benomyl to the medium at the rate of 0.38 g/1
inhibits germination of these organisms and allows selective isolation of M. anisopliae.
Decreasing the benomyl rate to 0.25 ppm will allow high recovery frequencies of both
target fungi while keeping recovery of other soil fungi to a minimum.
A natural tendency of many researchers is to incubate culture plates under some
level of artificial or natural light. Some effects of light on B. bassiana and M. anisopliae
have been studied (Alves et al. 1979 1980, Zimmerman 1982). These studies do not


Mean recovery at 10-4 dilution
Beauveria bassiana (Tests) Metarhizium anisopliae (Tests)
Light level
pmol m 2 s-' 1 2 3 1 2 3

0 (in foil) 16.5a 34.4 41.4 30.2 10.4 23.4
2 11.6 33.0 24.0 30.0 8.6 22.3
5 9.8 30.0 25.6 22.2 13.0 20.9
10 8.4 25.4 32.8 17.0 10.0 18.2

Significance ns ns ** ** ns **

"Mean number of colonies for five replicate plates for each test.
"Significance of the F test denoted as follows: ns = not significant, and ** P>0.01.

Chase et al.: Isolation of Fungi 291

report the effects of relatively low light levels (10 pLmol m-2 s-') during incubation of
isolation plates. In the case of B. bassiana, light increases the time needed for recovery
plates by 2 to 5 days. Incubation of M. anisopliae in as little as 10 gmol m-2 s-1 not
only increases the incubation time, but also results in a significant decrease in recovery
frequencies compared to plates incubated in the dark.


Appreciation is extended to J. M. F. Yuen and B. Hewitt for their assistance in
conducting this research and to D. Kennedy for preparation of this manuscript. Florida
Agricultural Experiment Stations Journal Series No. 6380. Mention of a commercial or
proprietary product or of a pesticide in this paper does not constitute a recommendation
by the authors, nor does it imply registration under FIFRA as amended.


ALVES, S. B., L. C. FORTI, AND F. J. CIVIDANES. 1980. Influence of light color on
some biological activities of Metarhizium anisopliae (Metsch) Sorok. an en-
tomopathogenic fungus. Rev. Bras. Ent. 24: 123-125.
ALVES, S. B., J. M. MILANEZ, AND P. KASTEN, JR. 1979. Influencia do fotoperiodo
no crescimento e esporu lacno do fungo Beauverai bassiana (Bals.) Vuill. Anais
da S. E. B. 8: 203-206.
GENTRY. 1975. Growth and sporulation of Metarrhizium anisopliae and
Beauveria bassiana on media containing various peptone sources. J. Invert.
Pathol. 25: 301-305.
BEILHARZ, V. C., D. G. PARBERRY, AND H. J. SWART. 1982. Dodine: A selective
agent for certain soil fungi. Trans. British Mycol. Soc. 79: 507-511.
1983. Growth and sporulation of Beauveria bassiana and Metarhizium anisopliae
in a basal medium containing various carbohydrate sources. J. Invert. Pathol.
41: 117-121.
DOBERSKI, J. W. 1981. Comparative laboratory studies on three fungal pathogens of
the elm bark beetle Scolytus scolytus: Effect of temperature and humidity on
infection by Beauveria bassiana, Metarhizium anisopliae, and Paecilomyces
farinosus. J. Invert. Pathol. 37: 195-200.
DOBERSKI, J. W., AND H. T. TRIBE. 1980. Isolation of entomogenous fungi from elm
bark and soil with reference to ecology of Beauveria bassiana and Metarhizium
anisopliae. Trans. British Mycol. Soc. 74: 95-100.
FERRON, P. 1981. Pest control by the fungi Beauveria and Metarhizium. In: H. D.
Burgers (ed.). Microbial control of Pests and Plant Diseases 1970-1980. Academic
Press, London, 949 pp.
GARDNER, W. A., AND R. NOBLET. 1978. Effects of host age, route of infection, and
quantity of inoculum on the susceptibility of Heliothis virescens, Spodoptera
eridania, S. frugiperda to Beauveria bassiana. J. Georgia Entomol. Soc. 13:
JOUSSIER, D., AND G. CATROUX. 1976. Mise au point d'un millieu de culture pour le
denombrement de Beauveria tenella dans les sols. Entomophaga 21: 223-225.
LINGG, A. J., AND M. D. DONALDSON. 1981. Biotic and abiotic factors affecting stabil-
ity of Beauveria bassiana conidia in soil. J. Invert. Pathol. 38: 191-200.
LORIA, R., S. GALAINI, AND D. W. ROBERTS. 1983. Survival of inoculum of the
entomopathogenic fungus Beauveria bassiana as influenced by fungicides. Envi-
ron. Entomol. 12: 1724-1726.

Florida Entomologist 69(2)

June, 1986

OLMERT, I., AND G. KENNETH. 1974. Sensitivity of the entomopathogenic fungi,
Beauveria bassiana, Verticillium lencanii, and Verticillium sp. to fungicides
and insecticides. Environ. Entomol. 3: 33-38.
ROBERTS, D. W., AND A. S. CAMPBELL. 1977. Stability of entomopathogenic fungi.
Misc. Publ. Entomol. Soc. America 10: 19-76.
TEDDERS, W. L. 1981. In vitro inhibition of the entomopathogenic fungi Beauveria
bassiana and Metarhizium anisopliae by six fungicides used in pecan culture.
Environ. Entomol. 10: 346-349.
TUITE, J. 1969. Plant pathological methods. Burgess Publishing Co. Minneapolis, MN
239 pp.
VEEN, K. H., AND P. FERRON. 1966. A selective medium for the isolation of Beauveria
tenella and Metarrihizium anisopliae. J. Invert. Pathol. 8: 268-269.

l- ^ ^- -- ^ --^- ---^-


University of Florida
Institute of Food and Agricultural Sciences
Everglades Research and Education Center
P.O. Drawer A
Belle Glade, FL 33430


Populations of Liriomyza trifolii (Burgess) were monitored in 16 commercial fields
of celery during 1985 by taking 10 sets of 10 leaflet samples per field and counting
puparia after 3, 7, and 14 days. Three indices of dispersion showed that the population
had an aggregated distribution. Sample size for 3-day pupae counts were calculated to
be 7 sets of 10 leaflets for 25% precision at densities of 5 or more puparia per 10 leaflets.
The proposed sampling scheme requires between 30 and 45 minutes total sampling time
per field and will allow the grower to monitor L. trifolii population trends and evaluate
previous control actions. Sequential sampling plans may further reduce sampling time.


Las poblaciones de Liriomyza trifolii (Burgess) se esudiaron en 16 parcelas comer-
ciales de apio durante 1985. Diez grupos de muestras de diez hojas se obtuvieron de
cada parcela, contandose las crisalidas despu6s de 3, 7 y 14 dias. Tres indices de disper-
si6n montraron que la poblacion tenia una distribution aglomerada. Se determine que
el tamafo de la muestra para el conteo de crisalidas despues de 3 dias debe ser 7 grupos
de 10 hojas para obtener una precision de 25% en densidades de 5 o mas crisalidas por
cada 10 hojas. El sistema de muestreo propuesto require entire 30 y 45 minutes de
tiempo para el muestreo de cada parcela y permitirA al productor el estudiar las tenden-
cias de las poblaciones de L. trifolii y evaluar acciones de control previas. Planes de
muestreo de secuencia pueden reducir atin mAs el tiempo de muestreo.

Foster: Monitoring Leafminers 293

Liriomyza trifolii (Burgess), a serpentine leafminer, is a serious pest of celery,
lettuce, tomatoes, and other vegetable and ornamental crops in Florida. This insect
damages celery in at least 3 ways; by removing photosynthetic material from the leaves,
by making avenues of entry for plant pathogens, and by causing cosmetic damage to
the marketable portion of the crop.
There are a number of methods that have been considered as possible sampling
methods for leafminers. Musgrave et al. (1979) calculated sample sizes for sweep net
estimations of adult densities and for counts of live larvae of L. sativae in the foliage.
A problem with sweep net samples is that on windy days the adult leafminers tend to
stay low in the foliage and are not caught by the net. Also, if the foliage is wet from
rain or dew, a sweep net is not useful.
Growers in south Florida tend to avoid counting live leafminer larvae in the leaves
because it is tedious work and because it is easy to miss small larvae. Michelbacher et
al. (1953) counted the number of mines in tomato leaves but, as Johnson et al. (1980)
point out, this sampling method does not accurately estimate the current density be-
cause the mines remain long after the leafminers are gone. Parella and Jones (1985)
developed sampling plans for yellow cards covered with sticky material to monitor adult
leafminer populations in chrysanthemum greenhouses. Zehnder and Trumble (1985)
developed sequential sampling plans for sticky cards in tomatoes as well as for the pupal
tray method to be discussed later. At least one major celery grower in Florida has used
sticky card to monitor leafminer adults. However, sticky cards are inconvenient to work
with and often are fouled by blowing soil.
Johnson et al. (1980) recommended placing styrofoam trays under tomato foliage to
catch the larvae as they emerge from the leaves and drop to the ground to pupate. This
method is not practical in Florida because frequent strong winds blow the pupae from
the trays or blow the trays away. Also, there are usually no rows in which to place the
trays that will not be run over by farm equipment (J. T. Shaw, pers. comm.). A modifi-
cation of this method that is used by Florida growers is to hold leaflet samples in a
container for several days and then count the number of puparia that have emerged
from the leaves. A drawback to this sampling method is the time lag between when
samples are taken and when the information becomes available. The pupal rearing
method provides information about leafminer ovipositional activity during some previ-
ous time interval. The current uses for this sampling method are to monitor population
trends and to evaluate the effectiveness of previous control measures. An advantage of
pupal rearing over the pupal trays (Johnson et al. 1980) is that it can also be used to
estimate parasitism.
The objectives of this study were to determine the appropriate time interval to hold
celery leaflet samples before counting the puparia, to determine the appropriate sample
sizes for estimating L. trifolii densities, and to develop constant-precision-level sequen-
tial sampling plans at three levels of precision.


Sixteen celery fields near Belle Glade were sampled between February 5 and April
8, 1985. The approximately 11-ha fields were planted sequentially in time to provide an
even supply of celery for market. Each field was sampled between 1 and 6 times,
depending on the level of maturity of the crop at the initiation of the study. A total of
51 sets of sampling data was collected.
A field sample consisted of 10 sets of 10 terminal leaflets per set. The leaflets selected
were from petioles that were mature, but had not yet begun to senesce. The ten leaflets
collected at a site were selected randomly from 10 different plants and the 10 sites were

294 Florida Entomologist 69(2) June, 1986

arranged systematically throughout the field to insure that all areas of the field were
sampled. No sample sites were located within 15 m of the sides or ends of the fields to
remove possible edge effects. Each field was treated uniformly with insecticides when
determined to be necessary by the grower.
After collection in the field, the leaflets were placed in plastic bags and returned to
a laboratory. Leaflets then were placed in 0.47-liter (1-pint) styrofoam cups and held in
a laboratory at room temperature (23.0 0.56C). Three days after field collection, the
leaves in each carton were removed and the leafminer puparia were counted. The leaves
were then returned to the carton and again held at room temperature. The counting
procedure was repeated 7 and 14 days after field collection, at which time the leaves
were discarded because 14 days is an adequate length of time to insure that all eggs
and larvae present had pupated (Leibee 1984).
There are a number of indices of dispersion that have been used to analyze the
distribution of insect populations. The simplest of these is the variance to mean ratio.
If s2 = m, the population is assumed to be randomly distributed. s2 > m indicates
aggregation and s2 < m indicates uniformity. Iwao (1968) introduced a regression
method for measuring spatial dispersion. He used a parameter, mean crowding (me),
proposed by Lloyd (1967): m, = m + ((s2/m)-1). Mean crowding is defined as the mean
number of other individuals per individual. Iwao found that when mean crowding was
regressed on mean density, there was a direct linear relationship. Regression parame-
ters a, the intercept, and b, the slope, are both indices of dispersion. The a-value is an
index of basic contagion, and the b-value represents the density-contagiousness coeffi-
cient. Random distribution is suggested by a = 0 and b = 1, and aggregated distribution
is indicated by a > 0 and b > 1.
According to Taylor (1961) the sample variance can be related to the sample mean
by the power law:
s2 = ab (1)
The mean (R) and the variance (s2) were calculated for each data set in this study and
a linear regression of log s2 on log R was computed. The a and b values determined from
the regressions were used to calculate necessary sample sizes for various desired levels
of precision using the equation published by Ruesink (1980):
n = ab-2 (2)
where n = the number of samples and c = precision expressed as a fraction of the mean.
The a- and b-values from Taylor's power law also were used to develop constant-pre-
cision-level sequential sampling plans for three day pupal counts. The stop points were
calculated by the formula (Green 1970):
log T = log (c/a) + b log n (3)
b-2 b-2
where Tn = cumulative number of pupae. Southwood (1978) considered 25% precision
to be adequate for use in pest management programs.


Holding leaflets for 7 days resulted in approximately twice as many pupae as holding
them for 3 days (Table 1). Holding the leaflets for 14 days produced a mean of 71.4
puparia per 10 leaflets, which is only 7.4% more than holding them for 7 days. Therefore,
holding the leaves for longer than 7 days is not worthwhile. There was a wide range of
mean densities, which is considered by Ruesink (1980) to be one of the requirements
for using Taylor's power law.

Foster: Monitoring Leafminers


No. of days held Mean Range

3 32.1 1-224
7 66.5 3-258
14 71.4 3-270


Taylor's Power Law Iwao's Regression
No. of Days held a b Variance/Mean a b

3 1.429 1.297 4.39 0.408 1.189
7 1.432 1.352 4.01 1.167 1.140
14 1.346 1.378 4.42 1.087 1.150

All 3 indices of dispersion calculated showed that leafminers have an aggregated
distribution for each of the 3 durations for which leaflets were held (Table 2). These
results agree with those of Beck et al. (1981), who found that active mines on whole
plants were aggregated in their distribution in the field (Taylor's b-value = 1.51).
Figure 1 illustrates the distribution of data points around the regression line generated
by Taylor's power law for pupal counts.
The a- and b-values from Taylor's power law (Table 2) can be used to calculate
sample sizes necessary to estimate leafminer densities (Table 3). Although there are no
economic thresholds in use for leafminers in celery, means of 5 or fewer pupae reared
from 10 leaflet samples held for 3 or 7 days would pose little threat of causing economic
damage. Therefore, it appears from these data that the sample size of 10 sites used in
this study is adequate for estimating leafminer densities of i > 5 per 10 leaflets with
an acceptable level of precision (c= 0.25). In most situations, it would be preferable to
hold the samples for only 3 days before counting to reduce the lag time. If the leaves
were returned to the cup after counting and held for another 4 days, information about
oviposition nearer to the time of sampling could be obtained.
It takes approximately 20 to 30 min to take 10 sets of 10 leaflet samples from a field
of about 11 ha. If the sets of 10 leaves are held in individual cups, it takes approximately
15 min to count the puparia at a density of 10 pupae per 10 leaflets. If the leaflets are
small enough to be composite into a single or a couple of containers, the time required
is reduced to about 8 minutes. Therefore, the total time required to sample a field is
about 30 to 45 minutes. Because most of the field time is expended while walking, not
much time is saved by taking fewer than 10 samples per field. Sampling time could be
reduced further using sequential sampling (Table 4). This would not reduce the amount
of time spent in the field, but could reduce the counting time considerably.
Pupal counts, then, are a relatively efficient method for monitoring leafminer popu-
lation activity in celery. Precise measurements of leafminer densities can by achieved
with few samples in a short period of time, although there is a 3- or 7-day lag time
before the information is available. This sampling method is useful for monitoring pop-
ulation trends or evaluating the success of previous control measures. If economic






Fig. 1. Regression line describing relationship
day pupal counts of Liriomyza trifolii.

between log s2 and log i for three


No. of samples
3-Day Drops 7-Day Drops
x c = 0.2 c = 0.25 c = 0.3 c = 0.2 c = 0.25 c = 0.3

0.5 58 37 26 56 36 25
1 36 23 16 36 23 16
3 17 11 7 18 11 8
5 12 7 5 13 8 6
10 7 5 3 8 5 4
20 4 3 2 5 3 2
50 2 1 1 3 2 1





Florida Entomologist 69(2) June, 1986

LOG S 1 .lSS 1.297 LOG X
tRt "|.91)



*, *

I I I ___1








Foster: Monitoring Leafminers


Cumulative pupae counted
No. samples c = 0.20 c = 0.25 c = 0.30

1 162 86 51
2 121 64 38
3 102 54 32
4 90 48 28
5 82 43 26
6 76 40 24
7 71 38 22
8 67 36 21
9 64 34 20
10 61 32 19

thresholds are developed for leafminers on celery, a sampling method that will make
estimates of the actual leafminer density may be required. But until that time, the pupal
rearing method should be useful to celery growers in southern Florida to monitor L.
trifolii populations and to evaluate the effectiveness of control practices.


I wish to thank J. Shaw, D. Remick, D. Malloy, P. Yance, R. Wojciak, and G.
Mitchell of A. Duda & Sons, Inc. for their willing cooperation in this study. Florida
Agricultural Experiment Station Journal Series No. 6657.


Spatial dispersion patterns of Liriomyza spp. on celery. Proc. 1st Annual Indus-
try Conf. Leafminer: 129-140.
GREEN, R. H. 1970. On fixed precision level sequential sampling. Res. Pop. Ecol. 12:
IWAO, S. 1958. A new regression method for analyzing the aggregation pattern of
animal populations. Res. Popul. Ecol. 10: 1-20.
A technique for monitoring Liriomyza sativae in fresh market tomatoes. J. Econ.
Entomol. 73: 552-555.
LEIBEE, G. L. 1984. Influence of temperature on development and fecundity of
Liriomyza trifolii (Burgess) (Diptera: Agromyzidae) on celery. Environ. En-
tomol. 13: 497-501.
LLOYD, M. 1967. 'Mean crowding.' J. Anim. Ecol. 36: 1-30.
Tomato insect investigations in northern California in 1951. J. Econ. Entomol.
46: 73-6.
M. WHITE, W. G. GENUNG, AND V. L. GUZMAN. 1979. Dispersion analysis and
sampling plans for insect pests in Florida celery. Proc. Florida State Hort. Soc.
92: 106-108.


Florida Entomologist 69(2)

June, 1986

PARELLA, M. P., AND V. P. JONES. 1985. Yellow-traps as monitoring tools for
Liriomyza trifolii (Diptera: Agromyzidae) in chrysanthemum greenhouses. J.
Econ. Entomol. 78: 53-56.
RUESINK, W. G. 1980. Introduction to sampling theory. Pages 61-78 in M. Kogan and
D. C. Herzog, eds. Sampling methods in soybean entomology. Springer-Verlag,
New York. 587 pp.
SOUTHWOOD, T. R. E. 1978. Ecological methods, with particular reference to the study
of insect populations. Chapman and Hall, London.
TAYLOR, L. R. 1961. Aggregation, variance and the mean. Nature 189: 732-735.
ZEHNDER, G. W. AND J. T. TRUMBLE. 1985. Sequential sampling plans with fixed
levels of precision for Liriomyza species (Diptera: Agromyzidae) in fresh market
tomatoes. J. Econ. Entomol. 78: 138-142.


The Ohio State University
Ohio Agricultural Research and Development Center
Wooster, Ohio 44691
Centro Nacional de Pesquisa de Fruteiras
de Clima Temperado
Pelotas, RS, Brazil


This is the first of a series of studies designed to compare sap beetles (Coleoptera:
Nitidulidae) present at various locations in the state of Rio Grande do Sul in southern
Brazil. Collections from strawberries, apple, peach, plum, and quince drops yielded 16
species in 7 genera. Almost twice as many species were found in the Pelotas area as
near Vacaria. The most abundant species was Carpophilus fumatus Boheman. This is
the first report of C. fumatus and of Haptoncus sobrinus Grouvelle from South


Este articulo es el primero de una series de studios para comparar los nitiddlidos
(Coleoptera: Nitidulidae) que se encuentran en el Estado de Rio Grande do Sul, en el
sur del Brasil. El analisis de los frutos de fresa, manzana, durazno, ciruela, y membrillo,
indic6 que 16 species pertenecientes a 7 g6neros estaban presents. Casi el double
ndmero de species se encontraron en el area de Pelotas cerca de Vacaria. La especie
mas abundante fue Carpophilus fumatus Boheman. Esta 6s la primera vez que se
report a C. fumatus y a Haptoncus sobrinus Grouvelle en Sudambrica.


Williams & Salles: Nitidulids in Fruit Crops 299

This is the first of a series of studies on the sap beetles (Coleoptera: Nitidulidae)
collected from fruit in Rio Grande do Sul. This, the southernmost State in Brazil is
noted for its production of grains and fruit.
Literature documenting the occurrence of sap beetles on fruit and vegetable prod-
ucts in Brazil is scarce. References containing information about sap beetles in Brazil
can be found in Lima (1953), Blackwelder (1945), and Silva et al. (1968). Salles and
Williams (1983) were first to publish information on the pest status of Lobiopa insularis
(Castelnau), the most serious strawberry pest in southern Brazil. Loss estimates range
between 10-20% of total strawberry production, which during the 1983 season alone,
amounted to 450-600 metric tons of strawberries.


We have compared nitidulid collections from the two main fruit growing areas of Rio
Grande do Sul near the cities of Pelotas (Lat: 31S; Long: 53W) and Vacaria (Lat: 29S;
Long: 51W).
Pelotas is the most important peach and strawberry producing area in Brazil; Vac-
aria is the most important apple producing region in R. G. do Sul. Pelotas is in rolling
country a few meters above sea level, and the elevation of Vacaria is ca. 760 meters
above sea level. These two fruit-growing regions are 600 km apart (see map Fig. 1).
During the 1982 and 1983 growing seasons, fruits were collected at intervals from
November to February. Samples (ca. 1 kilo/collection) of various kinds of ripe and
overripe fruit from the ground surface were taken to the laboratory in plastic bags. Sap
beetles were separated from the fruit and placed in small vials containing 70% alcohol.
Collections were labeled according to location, date and fruit type before forwarding
to the following specialists for identification: Colopterus-Dr. Larry E. Watrous, Field
Museum of Natural History, Chicago; Haptoncus-Lorin R. Gillogly, USDA APHIS,
San Pedro, CA; all others-Dr. W. A. Connell, University of Delaware, Newark. Num-
bers of each species were recorded from each collection.


There were twice the species at Pelotas (15) a warmer, more humid area as at
Vacaria (8). Vacaria had only one species that was not reported from Pelotas (Table 1).
The most prevalent sap beetle species in these collections was Carpophilusfumatus
Boheman. It was found at both locations in all kinds of fruit examined. This beetle is
widely distributed in the tropics, but has not previously been reported from South
America (W. A. Connell, Pers. Comm. 1983). Over half of the C. fumatus collections
were from apple; quince was the second most common host. Apple and quince are
closely related pome fruits. Even though Carpophilus is the most important genus of
nitidulids causing injury to agricultural commodities world wide (Williams et al. 1983),
with the exception of a single specimen of C. humeralis (Fab.), the only member of this
genus encountered was C. fumatus.
The genus Colopterus was second in abundance with 108 individuals representing at
least 4 species. Colopterus spp. are common in ripe and overripe fruits (L. E. Watrous,
Pers. Com. 1984).
Two species of Haptoncus are present in Rio Grande do Sul; however, H. sobrinus
Grouvelle was found only at Pelotas. This is a new locality record for H. sobrinus (L.
R. Gillogly, Pers. Com. 1983). Previous distribution for this species was Bourbon,
Seychelles, Madagascar, and the Caroline Islands (Gillogly 1962). Gillogly believes that

Florida Entomologist 69(2)

Fig. 1. Map of Brazil denoting sap beetle sampling places in the state of Rio Grande
do Sul. (Pelotas and Vacaria). 1982/83.

unusual discontinuous distributions of nitidulids about the world may be explained by
movement of "copra ships" in international commerce.
The "tropical strawberry sap beetle," Lobiopa insularis, is one of the most destruc-
tive nitidulids in both direct injury, caused to strawberry fruit by larvae and adults,
and severe impact on the canning industry when larvae are found in cans of strawberries
(Salles and Williams 1983). Collections of L. insularis adults presented in Table 1 are


June, 1986

Williams & Salles: Nitidulids in Fruit Crops

Cl. C~O -~lC~h Cl

1-4 00 0 -I 1

10 -


I:- K- i--
0- a

3 ';G %

C C3CC C- 6 CZ c3^

i 3 .-o .
~ 00 0 0 0 0 0~ ~


I10 c


e -



-. w

a3 .

ii o
II 0

I ^

,~ ~~~~~~~~\~\\~\

Florida Entomologist 69(2)

June, 1986

not representatives of peak populations on strawberries since they were made after the
usual harvest period. L. insularis was encountered on apple, peach, plum, and straw-
berry, but not on quince.
Diversity in Stelidota was equal to that in Colopterus with four species represented
in each. There were not large numbers of any of the Stelidota species. However, all
four species were found around Pelotas and only one near Vacaria.
No attempt was made to establish level of damage or importance for the species of
nitidulids encountered. Rather in this study we compared the nitidulid complex on fallen
fruit in two distinct ecological regions in Rio Grande do Sul.


We express our appreciation to the nitidulid specialist who identified the sap beetles
in this study: Walter A. Connell, Larry E. Watrous, and Lorin R. Gillogly. Without
their assistance this study would have been impossible.


BLACKWELDER, R. W. 1945. Checklist of the coleopterous insects of Mexico, Central
America, the West Indies, and South America. Smithsonian Institution, USNM,
Bull. 185(3): 1-550.
GILLOGLY, L. R. 1962. Insects of Micronesia. Coleoptera: Nitidulidae. B. P. Bishop
Museum, Honolulu. 16(4): 133-188.
LIMA, A. M. DA COSTA. 1953. Insetos do Brasil. 8 Tomo: Coleopteros 2a Parte. Esc.
Nac. Agron., Tip. Jornal do Commercio, Rio de Janeiro. 323 pp.
SALLES, L. A. B. DE, AND R. N. WILLIAMS. 1983. Broca do morango (Lobiopa in-
sularis). EMBRAPA/CASCATA, Documentos No. 17, 10 pp.
M. N. SILVA, AND L. SIMONI. 1968. Quarta catAlogo dos insects que vivem nas
plants do Brasil seus parasites e predadores. Parte II, Tomo I. Insectos, hos-
pediros e inimigos naturals. Min. Agr. Depto. Def. e Inspecao Agropecuaria, Rio
de Janeiro. 622 pp.
1983. Bibliography of the genus Carpophilus Stephens (Coleoptera: Nitidulidae).
Ohio State University/Ohio Agricultural Research and Development Center,
Res. Circ. 278: 1-95.


Atkinson & Equihua: Biology of Wood Borers



Institute de Biologia
Universidad Nacional Aut6noma de M6xico
Apdo. 21 San Patricio 48980,
Jalisco, Mexico
Centro de Entomologfa y Acarologia
Colegio de Postgraduados
56230 Chapingo, Mexico, Mexico


The feeding habits, degree of host specificity, and mating systems were examined
for 96 species of Scolytidae and Platypodidae in a tropical deciduous forest at Chamela,
Mexico. The dominant feeding habit was phloeophagy (59.4%), followed by
xylomycetophagy (13.5%), myelophagy (13.5%), and xylophagy (12.5%). Most
phloeophagous species were monophagous (limited to one plant genus); the other three
groups were largely polyphagous. The mating systems found in order of importance
were monogyny (40%), harem (heterosanguineous) polygyny (20.2%), inbred (consan-
guineous) polygyny (19.1%), and bigyny (18.1%). Monogynous species were mostly
phloeophagous, to a lesser extent xylomycetophagous. Phloeophagy and xylophagy
were equally represented among bigynous species. Harem polygynous species were
almost exclusively phloeophagous while inbred polygynous species were mostly
myelophagous or xylomycetophagous. All inbred polygynous species were polyphagous.
The overall biological patterns of these beetles differed from those reported for
Scolytidae and Platypodidae of temperate or tropical regions.


Los habitos alimenticios, grado de especificidad respeto al hospedero y sistemas de
apareamiento se examinaron en 96 species de Scolytidae y Platypodidae en un bosque
tropical caducifolia en Chamela, M6xico. El hAbito alimenticio dominant era fleofagia
(59.4%) seguido por xilomicetofagia (13.5%), mielofagia (13.5%) y xilofagia (12.5%). La
mayoria de las species fle6fagas eran mon6fagas (limitadas a un g6nero de plantss; los
otros 3 grupos eran polifagos. Los sistemas de apareamiento encontrados en orden de
importancia eran monoginia (40%), poliginia de harem (heterosanguinea) (20.2%),
poliginia end6gama (consanguinea) (19.1%), y biginia (18.1%). Especies monoginas eran
principalmente fle6fagas, o xilomicet6fagas en menor grado. Floeofagia y xilofagia esta-
ban igualmente representadas entire species biginas. Especies poliginas de harem eran
casi exclusivamente fle6fagas; species poliginas end6gamas eran principalmente
miel6fagas o xilomicet6fagas. Todas las species poliginas end6gamas eran polifagas.
Los patrons globales de la biologia de estos cole6pteros diferieron de los reportados
para Scolytidae y Platypodidae de otras regions templadas o tropicales.

Florida Entomologist 69(2)

June, 1986

The Scolytidae and Platypodidae form a compact group, both taxonomically and
ecologically. Most are borers of the woody tissues of shrubs, vines, and trees. The
common names "bark beetles" and "ambrosia beetles" refer to the most common feeding
habits found in the group; i.e. consuming phloem (inner bark) (phloeophagy) and ec-
tosymbiotic fungi that grow in their galleries (xylomycetophagy), respectively. Other
habits include the consumption of wood (xylophagy), pith in twigs and branches
(myelophagy), herbaceous plants (herbiphagy) and fruits or seeds (spermatophagy)
(Wood, 1982). Generally, feeding habit is characteristic of a given species although there
is some overlap of feeding guilds.
Reproductive biology in the Scolytidae and Platypodidae has recently been reviewed
by Kirkendall (1983). Mating systems of outbreeding species are monogyny or harem
polygyny (heterosanguineous polygamy of Wood (1982)). Depending on the species
either males or females initiate gallery construction in monogynous species, while males
initiate attack in polygynous species. Bigyny is a special case of polygyny, in which a
male initiates attacks and is associated with only two females. Another important
mating system is inbred polygyny (spanandry of Beaver (1977, 1979); consanguineous
polygamy of Wood (1982)) in which females mate with their siblings prior to emergence
and then construct new galleries alone. In this last case, males are few in number,
reduced in size and are flightless.
In a comparison of 2 tropical and 2 temperate areas Beaver (1979) reported that the
ambrosial habit was more common in the humid tropics and that the overall degree of
host specificity was lower (i.e., more polyphagous) there. However, 3 of the 4 areas
compared were large and ecologically heterogeneous (France, California, West
Malaysia). The practice of inbred polygyny is more prevalent in tropical areas and is
related to low host specificity (Browne 1961, Beaver 1977, Wood 1982).
We present here an analysis of the host tissues consumed, degree of host specificity,
and mating systems for the Scolytidae and Platypodidae in tropical deciduous forest on
the Pacific Coast of Mexico. Preliminary results were presented by Equihua et al.
(1984). Our current analysis is based on our recent, more complete annotated checklist
of the 96 species of Scolytidae and Platypodidae in the region (Equihua and Atkinson,
in press).


The Estaci6n de Biologia Chamela is a biological reserve and field station of the
National University of Mexico on the coast of Jalisco, Mexico (1930' N, 105'03' W).
The station covers 1600 ha, mostly below 150 m above sea level and is 2 km from the
coast at its closest point. The mean annual temperature is 24.90C and the 10 year
average rainfall is 748 mm, most of which (80%) falls within the 4-month period from
July through October (S. H. Bullock, Estaci6n de Biologia Chamela, personal communi-
cation). The vegetation of the station is mostly tropical deciduous forest with some
tropical subdeciduous forest along the courses of the larger drainages (forest types from
Rzedowski, 1978). Estuarine and riparian communities are found nearby. Despite the
low rainfall and its strong seasonality a diverse flora occurs in this forest type, both in
terms of numbers of species and variety of different life forms (Rzedowski 1978, Lott
1985). More than 750 species of vascular plants are known from the station (Lott 1985),
of which approximately 60% are woody (Lott, personal communication).
We observed feeding habits (type of tissue consumed), mating systems, and degree
of host specificity for most of the Scolytidae and Platypodidae at the station and nearby
areas (within 5 km) from early 1982 until mid 1985. Feeding habits were inferred from
observations on the location of adult and larval galleries. Degree of host specificity was


Atkinson & Equihua: Biology of Wood Borers


assigned based on our observations in the area, subjective judgment, and a critical
review of the literature; doubtful cases were not included. We consider species that
utilize host plants of the same genus as monophagous; those which utilize hosts within
the same family or occasionally from related families as oligophagous; those that utilize
several to many unrelated hosts are classified as polyphagous. Mating systems were
inferred from the number and sex of adults constructing galleries and gallery architec-


Phloeophagy is the dominant feeding habit; xylomycetophagy, xylophagy, and
myelophagy are of approximately equal importance (Table 1). Only one spermatophag-
ous species and no herbiphagous species were found. In the 2 temperate areas compared
by Beaver (1979) phloeophagy was the dominant habit (more than 80%), followed by
xylomycetophagy; in his two humid tropical areas xylomycetophagy predominated (57%
in Fiji, 76% in West Malaysia) while phloeophagy was a distant second (30.3% and
11.6%, respectively). In the same study none of the other four feeding guilds included
more than 10% of the total species in any locality. The relatively high importance of
xylophagy (12.5%) and myelophagy (13.5%) in the Chamela fauna is unprecedented. The
dominance of phloeophagy at Chamela is more similar to the pattern characteristic of
temperate areas than to that of humid tropical ones. The relatively low importance of
xylomycetophagous species at Chamela (13.5%) compared with that in humid tropical
areas (Beaver 1979) may be related to the long (8 months) dry season, during which
moisture may limit growth of ambrosial fungi in wood. In the Chamela area, ambrosia
beetles were primarily limited to the shadier, more humid watercourses, particularly
during the dry season, lending credence to the above hypothesis. In addition the rela-
tively large number of xylophagous species, mostly Micracini, may partly exclude am-
brosia beetles by competition for wood. Fungi are apparently present in the wood
surrounding gallery systems of most (all?) xylophagous Scolytidae and Platypodidae and
stain and/or alter the texture of the wood (Wood 1982, Atkinson, personal observations).
Since xylophagous Scolytidae and Platypodidae directly consume xylem along with any
included fungal hyphae they may be less limited by low atmospheric moisture than
xylomycetophagous species that require an abundant growth of ambrosial fungi on their
gallery walls.


Monophagous species (50%) were slightly more numerous than polyphagous species
(41%) in the Chamela fauna (Table 1). Eighteen species were excluded from the analysis
because their degree of host specificity was not known; most belonged to the genera
Pseudothysanoes, Araptus, and Pityophthorus. Based on our experience with other
species of these genera, most are expected to be monophagous. Oligophagy is by far
the least common of the 3 host-specificity categories, suggesting that it is the least
successful adaptive strategy in tropical deciduous forest. The relative importance of the
different host specificity categories at Chamela more nearly resembles temperate (57%
in France, 70% in California restricted to single genus or species of host) than humid
tropical (14% in Fiji, 4% in Malaysia) patterns (Beaver 1977, 1979).
Apparently the type and condition of host tissue consumed influences the degree of
host specificity (Table 1). Although 15 combinations were possible (3 degrees of host
specificity x 5 feeding habits), only 3 were well represented: phloeophagous

Florida Entomologist 69(2)

June, 1986


Degree of Host Specificitya (No. of spp.) Total spp.
Habit MonophagyOligophagy Polyphagy Unknown No. %

Phloeophagy 34 5 4 14 57 59.4
Xylomycetophagy 1 12 13 13.5
Myelophagy 2 11 13 13.5
Xylophagy 3 5 4 12 12.5
Spermatophagy 1 1 1.0
Total spp. No. 39 7 32 18 96
% 50.0 9.0 41.0 -c

aMonophagy = restriction to one plant genus; Oligophagy = restriction to one plant
family; Polyphagy = use of hosts from 2 or more unrelated plant families.
bValues based on the 78 species with known host specificity.
c18.8% of all 96 species.

monophages, xylomycetophagous polyphages, and myelophagous polyphages (35.4%,
12.5%, and 11.5% of all 96 species, respectively). Beaver (1977, 1979) reported that
ambrosia beetles are basically polyphagous in both temperate and humid tropical areas,
a conclusion supported here. He attributed decreased host specificity in
xylomycetophagous species to several factors, of which the most important was that
these feed on fungi rather than directly on host tissues. Perhaps more important is the
association between utilization of dead tissues and polyphagy. Most ambrosia beetles
breed in heartwood which consists of dead cells. A relatively high percentage of the
myelophages (Hypothenemus, Cryptocarenus) and xylophages (Hylocurus, Thysanoes)
at Chamela are also polyphagous. The species of these genera at Chamela breed in
obviously dead hosts. Three of the 4 polyphagous phloeophages (Hypothenemus) are
also found in dead, dry material.
The high degree of host specificity among the phloeophages probably reflects the
fact that they usually breed in living or recently killed hosts which produce resin or
latex (Table 2). The strong association of host-specific phloeophages with this type of
defense system suggests that latex/resin defenses are effective against generalist
phloeophages and have been a selective factor leading to host specialization (see Cates
and Alexander 1982). The relative scarcity of host-specific Scolytidae in plants which
do not produce latex or resin may indicate that they are at a competitive disadvantage
with respect to generalist species or to other organisms (Cerambycidae, Buprestidae,
Bostrichidae) in those hosts.
Although relatively few plants at Chamela are associated with host-specific
Scolytidae and Platypodidae, all woody plants which we observed were subject to attack
by one or more polyphagous species belonging to different feeding guilds. Apparently
the presence or absence of a given species or guild is determined mostly by physical
characteristics of the host plant and microhabitat.


Monogyny is the most prevalent mating system in the Chamela fauna; bigyny, harem
polygyny, and inbred polygyny are of similar rank (Table 3). The 2 species with unknown


Atkinson & Equihua: Biology of Wood Borers



Plant Family Plant Species Insect Species











Astronium graveolens (RbNC)
Mangifera indica (R, N C)
Spondias purpurea (R,NC)

Plumeria rubra (L,NC)

Thevetia ovata (L,NC)

Sarcostemma clausum (L,NC)
several genera, fruits (1)
Bursera arborea (R)
Bursera instabilis (R)

Stenocereus chrysocarpus (R)
Stenocereus standleyi (R)
Acanthocereus occidentalis
Conocarpus erecta (NC)
Laguncularia racemosa (NC)
Ipomoea wolcottiana (L)
Celaenodendron mexicanum
Croton spp. (L)
Euphorbia colletioides (L)
Hippomane mancinella (L,NC)
Hura polyandra (L,NC)
Amphipterygium adstringens
Acacia spp. (R)
Cynometra oaxacana (NC)
Lonchocarpus spp. (R)
several genera

Heteropterys laurifolia
Brosimum alicastrum (L,NC)

Chlorophora tinctoria (L,NC)
Ficus spp. (L)

(Cont. next page)

Pityophthorus indefessus (ph)
Hypocryphalus mangiferae (ph)
Pityophthorus nanus (ph)
Dendroterus luteolus (ph)
Scolytodes plumeriacolens (ph)
Scolytodes plumeriae (ph)
Pityophthorus costabilis (ph)
Pityophthorus costatulus (ph)
Araptus delicatus (m)
Araptusfossifrons (s)
Dendroterus sallaei (ph)
Phloeoterus burserae (ph)
Dendoterus luteolus (ph)
Cactopinus atkinsoni (ph)
Cactopinus setosus (ph)

Cactopinus setosus (ph)
Scolytopsis puncticollis (ph)
Scolytopsis puncticollis (ph)
Scolytogenes rusticus (ph)

Pseudothysanoes thomasi (ph)
Chramesus exul (ph)
Araptus sp.4 (m)
Cnemonyx splendens (ph)
Cnemonyx equihuai (ph)

Pityophthorus ingens (ph)
Hylocurus scitulus (x)
Pseudochramesus sp. (ph)
Chramesus vitiosus (ph)
Araptus sp.1 (ph)
Pseudothysanoes squameus (ph)
Chaetophioeus minimum (ph)
Chramesus securus (x)
Platypus excisus (sm)
Scolytus propinquus (ph)
Scolytus cristatus (ph)
Phloeotribus setulosus (ph)
Pycnarthrum amersum (ph)
Pseudothysanoes spinatus
Araptus consobrinus (ph)
Gymnochilus reitteri (ph)
Pycnarthrumfurnerium (ph)
P. hispidum (ph)
P. reticulatum (ph)
Scolytodes amoenus (ph)
S. tennis (ph)

Florida Entomologist 69(2)

June, 1986


Plant Family Plant Species Insect Species

Nyctaginaceae Guapira sp. (NC) Dendrosinus mexicanus (x)
Ulmaceae Celtis iguaneus Chramesus subopacus (ph)
Phloeotribus opimus (ph)
Phyllostylon brasilense (NC) Phloeotribus (sp. (ph)

aBased on Lott (1985, personal communication).
bPlant and host-specific insect introduced.
eGenera not the same for different beetles.

mating habits are species of Pseudothysanoes. All species of this genus with known
mating habits are bigynous (Wood 1982, Atkinson and Equihua 1985a,b, Equihua and
Atkinson, in press). Little comparative information is available on the relative impor-
tance of the various mating systems in different parts of the world except that inbred
polygyny is the dominant habit in the humid tropics. (Browne 1961, Beaver 1977, 1979,
Wood 1982). In this context, the relatively low ranking of inbred polygyny in the
Chamela fauna is noteworthy. The high incidence of bigyny (18.1%) is unusual. If bigyny
were considered a special case of harem polygyny then combining these 2 categories
would make harem polygyny about equal in rank to monogyny in the Chamela fauna.
Mating systems were related to feeding habits and degree of host specificity.
Monogyny was the most frequent mating system exhibited by phloeophagous insects,
followed by harem polygyny and bigyny (Table 3). This mating system was also impor-
tant among the xylomycetophagous species. Inbred polygyny was most common among
the myelophages and xylomycetophages. Most xylophages were bigynous. All species
exhibiting inbred polygyny were also polyphagous while species with harem polygyny
were monophagous (8 spp.) or oligophagous (1 sp.) (Table 4). The largest group of
bigynous species of known specificity were polyphagous. Monogynous species were
mostly monophagous although an appreciable number were polyphagous.
Harem polygynous species at Chamela were basically phloeophagous and monophag-
ous. This is consistent with the hypothesis of Kirkendall (1983) regarding the origin of
this mating system, which has evolved at least 7 times independently in the Scolytidae.
Kirkendall (1983) proposed that harem polygyny arose from monogyny in situations in
which resource quality varied widely. This made it "profitable" for males to find and
defend patches of high quality tissue and attract females instead of searching for females
directly. Of all the tissues consumed by Scolytidae in living woody plants, phloeom
(including the cambium), is initially the highest in nutrient value, and would change
most in quality after the death of the plant.
Bigynous species were either phloeophagous or xylophagous (Table 3). The former
were mostly monophagous while the latter were mostly polyphagous. In all but one case
(Scolytus propinguus), these insects belong to the tribe Micracini. Most Micracini with
known mating habits are bigynous except Micracisella spp. which are monogynous
(Wood 1982, Kirkendall 1983, Atkinson and Equihua 1985a, b).
Species exhibiting inbred polygyny were associated with several feeding habits
(Table 3) but all were polyphagous (Table 4). The Chamela species belong to two unre-

Atkinson & Equihua: Biology of Wood Borers



Mating System (No. of spp.) Total spp.
Feeding Monogyny Bygyny Polygyny Inbred Unknown No. %
Habit Polygyny

Phloeophagy 27 9 18 3 57 59.4
Xylomycetophagy 8 5 13 13.5
Myelophagy 3 10 13 13.5
Xylophagy 2 8 2 12 12.5
Spermatophagy 1 1 1.0
Total spp. No. 40 17 19 18 2 96
%a 42.6 18.1 20.2 19.1 -b

aValues based on the 94 species with known mating systems.
b2.2% of all 96 species.


Mating System Total spp.
Degree Monogyny Bygyny Polygyny Inbred Unknown No. %"
Specificity Polygyny

Monophagy 28 2 9 39 50.0
Oligophagy 3 2 2 7 9.0
Polyphagy 8 6 18 32 41.0
Unknown 1 7 8 2 18
Total spp. No. 40 17 19 18 2 96
%" 42.5 18.1 20.2 19.1

aValues based on the 78 (host specificity) or 94 (mating systems) species for which
information is available.

lated tribes, the Cryphalini (Hypothenemus, Cryptocarenus) and the Xyleborini (all
species). This strong correlation of inbred polygyny and polyphagy agrees strongly with
the results of Beaver (1977) although the species he examined were mostly
xylomycetophagous (Xyleborini) while most at Chamela were myelophagous (Crypha-
lini). No completely satisfactory explanation has been advanced for the origin of inbred
polygyny, although it has evolved at least 5 times in the Scolytidae. Comparative studies
of biology of species with comparable feeding habits but different mating systems would
allow a better understanding of the relationship between resources and reproductive
biology of the Scolytidae and Platypodidae.


Field work for this study was supported in part by the Centro de Entomologia y
Acarologia, Colegio de Postgraduados, Chapingo, Mexico. Emily Lott and Stephen Bul-
lock critically reviewed the manuscript. Special thanks are due R. A. Haack for his
editorial comments.

310 Florida Entomologist 69(2) June, 1986


ATKINSON, T. H., AND A. EQUIHUA M. 1985a. Listado comentado de los Scolytidae
y Platypodidae (Coleoptera) del Valle de Mexico. Folia Ent. Mex. 65: 63-108.
-- AND 1985b. Notes on Biology and Distribution of Mexican and Central
American Scolytidae. I. Hylesininae, Scolytinae except Cryphalini and Cor-
thylini. Coleopterists Bull. 39: 227-238.
BEAVER, R. A. 1977. Bark and ambrosia beetles in tropical forests. Pages 133-147 in
Proc. Symp. on Forest Pests & Diseases in Southeast Asia, Bogor, Indonesia,
April, 1976. BIOTROP. Spec. Publ. 2.
--. 1979. Host specificity of temperate and tropical animals. Nature 281: 139-141.
BROWNE, F. G. 1961. The biology of Malayan Scolytidae and Platypodidae. Malayan
For. Rec. 22: 1-225.
CATES, R. G., AND H. ALEXANDER. 1982. Host resistance and susceptibility. Pages
212-263 in J. B. Mitton & K. Sturgeon, eds., Bark Beetles of North American
Conifers. Univ. Texas Press, Austin, Texas.
EQUIHUA, M. A., T. H. ATKINSON, AND E. J. LOTT. 1984. Scolytidae y Platypodidae
(Coleoptera) de la Estaci6n de Biologia Chamela, Jalisco. Agroci6ncia 57: 179-193.
AND 1986. Annotated checklist of the bark and ambrosia beetles (Col-
eoptera: Scolytidae and Platypodidae) associated with tropical deciduous forest
in Chamela, Jalisco, Mexico. Florida Ent. (in press).
KIRKENDALL, L. R. 1983. The evolution of mating systems in bark and ambrosia
beetles (Coleoptera: Scolytidae and Platypodidae). Zool. J. Linnean Soc. 77: 293-
LOTT, E. J. 1985. Listado floristico de la Estaci6n de Biologia Chamela. Institute de
Biologia, Univ. Nacional Aut6noma de M6xico, Mexico City. (in press).
RZEDOWSKI, J. 1978. Vegetaci6n de M6xico. Editorial Limusa, Mexico City. 432 pp.
WOOD, S. L. 1982. The bark and ambrosia beetles (Coleoptera: Scolytidae) of North
and Central America, a taxonomic monograph. Great Basin Natur. Mem. 6, 1359

Rodriguez & Wirth: New Andean Midge


Institute Nacional de Salud
Ministerio de Salud
Apartados AMreos 80080
Bogota, Republica de Colombia
Cooperating Scientist
Systematic Entomology Laboratory, USDA,
and Research Associate
Florida State Collection of Arthropods
1304 NW 94th St.
Gainesville, FL 32606 U.S.A.


Culicoides suarezi Rodriguez and Wirth, NEW SPECIES, is described from 3230 m
elevation on Monserrate near Bogota, Colombia, where it was collected while biting
man. A key is presented to distinguish the 4 described species of the andicola group of
the subgenus Avaritia, to which the new species belongs. All are anthropophilic and
confined to the paramo regions of the Colombian Andes.


Se describe una nueva especie, Culicoides suarezi Rodriguez y Wirth, colectada
cuando picaba al hombre a una altura de 3230 m en Monserrate cerca de Bogota, Colom-
bia. Se present una clave para separar las 4 species descritas del grupo de andicola
del subg6nero Avaritia al cual pertenece la nueva especie. Todas son antropolificas y
limitadas a las regions de paramo de los Andes colombianos.

Very little has been published on the Culicoides biting midges of Colombia, although
extensive collections have been made by V. H. Lee and P. Barreto of the Universidad
de Valle on the Rio Raposo, Dpto. Valle; J. E. Browne and M. A. Tidwell of Tulane
University and the Universidad de Valle in the vicinity of Cali, Depto. Valle; and G.
R. Defoliart and associates of the University of Wisconsin on the Rio Anori near Pro-
videncia, Dpto. Antioquia. The principal publications were descriptions of 6 new species
from near Cali by Browne (1980), 8 new species from the Rio Raposo by Wirth and
Barreto (1978), and 1 new species from Florencia by Messersmith (1972). Wirth and
Blanton (1973) reported 18 species from the Colombian Amazon and Wirth (1974) listed
Colombian records of 17 additional species.
However, Wirth and Lee (1967) reported a very interesting Culicoides fauna from
cool, wet situations at high altitudes in the central and western ranges of the Andes,
describing 7 new species. Three of these species belong to a distinct group of the
subgenus Avaritia that are endemic to the Paramo de Purac6 between 3100 and 3320

312 Florida Entomologist 69(2) June, 1986

m elevation. In addition they mentioned a 4th undescribed species of this group from
similar elevations in Costa Rica. The purpose of the present contribution is to report
and describe a 5th species of the andicola group from a paramo situation on Monserrate,
3230 m in Dpto. Cundinamarca near Bogota. All the Colombian species of this group
are avid feeders on human blood and 1 species, A. puracensis Wirth and Lee, is so
abundant as to be an annoying pest on the Paramo de Purace. A good description of
the paramo habitats and an explanation of the morphological terminology used in our
description can be found in the paper by Wirth and Lee (1967).


1. Pale wing markings extensive, poststigmatic pale spot in cell r5 covering distal 1/
of 2nd radial cell, and broader than the dark area distal to it ..................... 2

t .. ....



Fig 1. Culicoides suarezi female: A, head; B, wing; C, abdomen showing sper-
mathecae; D, thorax including legs.


Rodriguez & Wirth: New Andean Midge


- Pale wing markings very restricted, poststigmatic pale spot in cell r5 covering less
than distal /4 of 2nd radial cell and narrower than the dark area distal to it .. 3
2. Third palpal segment slender, palpal ratio 3.4, sensory pit shallow; proboscis
longer, P/H ratio 1.17 ........................................................ suarezi n. sp.
- Third palpal segment swollen distally, palpal ratio 2.5-2.9, sensory pit depth; P/H
ratio 1.00 ........................................................... andicola W irth and Lee
3. Larger species, wing length 1.52 mm; antennal segments 3, 11-15 with sensilla
coeloconica; 3rd palpal segment with shallow sensory pit orjuelai Wirth and Lee
- Smaller species, wing length 1.33 mm; antennal segments 3,13-15 with sensilla
coeloconia; 3rd palpal segment with deep sensory pit puracensis Wirth and Lee

Culicoides (Avaritia) suarezi Rodriguez and Wirth, NEW SPECIES

FEMALE HOLOTYPE: Wing length 1.39 mm; breadth 0.63 mm.
Head (Fig. 1A): Brown including antennae and palpi. Eyes hairy, contiguous for a
distance equal to diameter of 2 facets. Antenna with lengths of flagellar segments in
proportion of 25-20-20-21-21-20-23-26-31-31-40-42-65, antennal ratio (11-15/3-10) 1.18;
sensilla coeloconica present on 3,11-15. Palpal segments with lengths in proportion of
10-45-45-21-30; 3rd segment slender with a shallow, round, distal sensory pit, palpal
ratio 3.4. Proboscis long, P/H ratio 1.17; mandible with 17 teeth.
Thorax (Fig. 1D): Uniformly brown including legs. Wing (Fig. 1B) with distinct
pattern of pale areas typical of the subgenus Avaritia; 2nd radial cell with distal /2
included in the poststigmatic pale spot; the pale markings intermediate in extent be-
tween those of C. andicola and those of C. puracensis. Costal ratio 0.61. Halter pale.
Abdomen (Fig. 1C): Brown. Spermathecae 2, oval with short slender necks; sub-
equal, each measuring 0.048 by 0.039 mm; rudimentary 3rd spermatheca and sclerotized
ring present.
HOLOTYPE: 9 Colombia, Dpto. Cundinamarca, Bogota D.E., Monserrate, 3230 m,
biting human, 7.ix.1978, H. Strumm (deposited in National Museum of Natural History,
Washington, D.C.).
DISCUSSION: This species is dedicated to Marco F. Suarez of the Servicio de Er-
radicaci6n de Malaria in BogotA in recognition of his long-standing interest in the
Culicoides fauna of Colombia. Sefor Suarez has worked since 1976 in assembling a
collection and check list of Colombian Culicoides and first submitted the holotype of the
new species to the junior author in 1978.
Culicoides suarezi can be distinguished from the 3 other described species of the
andicola group by the characters given in the key. It is most similar to C. andicola,
resembling it more than orjuelai and puracensis in its more prominent wing markings,
the 5th palpal segment noticeably longer than the 4th, and the spermathecae with
shorter, less tapering necks. C. andicola, however, has much more distinct wing mark-
ings than suarezi, and the 3rd palpal segment is stouter with deeper sensory pit.


This research was partly supported by NIAID Contract 5PolA120108-02.


BROWNE, J. E. 1980. New species of biting midges of the genus Culicoides from Colom-
bia and the first description of the male of C. florenciae (Diptera:
Ceratopogonidae). J. Med. Entomol. 17: 533-544.

314 Florida Entomologist 69(2) June, 1986

MESSERSMITH, D. H. 1972. A new species of Culicoides from Colombia (Diptera:
Ceratopogonidae). Proc. Entomol. Soc. Washington. 74: 165-169.
WIRTH, W. W. 1974. A catalogue of the Diptera of the Americas south of the United
States. 14. Ceratopogonidae. Mus. Zool. Univ. Sao Paulo 14: 1-89.
WIRTH, W. W., AND P. BARRETO. 1978. New species of Culicoides biting midges
(Diptera: Ceratopogonidae) from Colombia. J. Med. Entomol. 14: 553-564.
WIRTH, W. W., AND F. S. BLANTON. 1973. A review of the maruins or biting midges
of the genus Culicoides in the Amazon Basin. Amazoniana 4: 405-470.
WIRTH, W. W., AND V. H. LEE. 1967. New species of Culicoides from high altitudes
in the Colombian Andes (Diptera: Ceratopogonidae). Proc. United States Natl.
Mus. 124: 1-22.


Entomology Department
Florida A&M University
Tallahassee, FL 32303
Aquatic Biology Program
Biology Department
University of Alabama
Tuscaloosa, AL 35486
Biology Department
University of Mississippi
University, MS 38677


The presumed female of Cheumatopsyche helma Ross and 3 new species, C. bibben-
sis, C. cahaba and C. kinlockensis, are described and illustrated. Each is compared to
the most similar congeners.


Se described e ilustra la supuesta hembra de Cheumatopsyche helma Ross y 3
species nuevas, C. bibbensis, C. cahaba y C. kinlockensis. Cada una se compare con
las cong6neres mas similares.

A recent survey of the caddisfly fauna of northern Alabama revealed the presence
of 3 undescribed species of Cheumatopsyche. In collections from 2 localities in this

Gordon et al.: New Caddisflies

portion of the state, males of C. helma Ross were identified. Because the female of this
species is unknown, these 2 collections were resorted and the female Cheumatopsyche
identified. A single female could not be assigned to any species and is presumed to be
C. helma.
Terminology for genitalic structures generally follows that of Gordon (1974). In all
descriptions, total specimen length is from the front of the head to the tip of the folded
wings. Holotypes will be deposited in the United States National Museum of Natural
History. Paratypes will be deposited at the Florida State Collection of Arthropods and
the Illinois Natural History Survey.

Cheumatopsyche helma Ross
Fig. 1.
This female will key to couplet 34 in Gordon's (1974) key. This species is easily
distinguished from all other Cheumatopsyche females by the shape of the clasper recep-
tacles which have a short, broad chimney and an evenly bowed ventral margin and by
the unique membranous areas in the median plate.
FEMALE: Length-8 mm. Color of head and body brown, legs light brown proxi-
mally, brown distally; antennae light brown, without distinct markings; wings general.
Each clasper receptacle wide and near dorsum of tenth tergum; in lateral view, with
ventral margin strongly bowed, anterior and posterior corners level; chimney broad and
twisted mesad; in dorsal view, long, tunnel-like, slightly bulbous at tip, lateral edge
higher than mesal, inner opening directed caudad. Median plate with dorsal sclerite
large and tongue-like at apex, membranous at sclerite midpoint and posterior edge;
lateral extension cupped at tip and with membranous fold that loops verntrad; extensive
membranous fold attached ventrally to tongue of dorsal sclerite and anterior margin of
lateral extension; tip of dorsal sclerite bifid in dorsal view.
LOCALITY.--ALABAMA: Clay Co., Cheaha Creek, 100 m. upstream from Lake
Chinnabee, Cheaha State Park, 22-V-81, S. Harris and P. O'Neil, 1 female and 1 male.
REMARKS.-This species was previously known only from the type locality, Gatlin-
burg, Tennessee. In Alabama, specimens were collected on the Piedmont Plateau,

\ /


Fig. 1. C. helma female. Tenth and eleventh terga: A, lateral view; B, dorsal view.
Median plate: C, lateral view; D, dorsal view.

Florida Entomologist 69(2)

June, 1986

Cheaha Creek, and lower Appalachian Mountains, Little River, DeKalb County; both
fast-flowing, rocky streams.

Cheumatopsyche cahaba, NEW SPECIES
Fig. 2.
This species will key to couplet 13 in Gordon's (1974) key and can be distinguished
from all other Cheumatopsyche spp. males by the long narrow cercus adhering to the
anterior edge of the apical lobes of the tenth tergum and the flange on the dorsal edge
of the phallotheca.
MALE: Length-7.5mm. Color of head and body brown, venter light brown; anten-
nae yellow with distinct dorsal V-marks on the five basal segments; wings irrorate with
large, light area at anal angle. Ninth segment annular, bearing long setae on dorsal
lobes and latero-posterior margin, lateral apical lobe angular. Tenth tergum moderately
elongate with long, narrow setose cerci adhering to anterior edge of apical lobes; in
lateral view, lobes long and upright, projecting above dorsum and exposing mesal apex
of tenth tergum, space between lobes and mesal apex filled by large cercus; in caudal
view, lobes long and irregular, tapered at apices and widely separated. Clasper short,
apex of basal segment not clearing dorsum of tenth tergum; apical segment approxi-
mately one-third length of basal segment, evenly tapering to abruptly narrowed tip.
Phallotheca long with a dorsal flange near apex, base moderately bulbous.
HOLOTYPE MALE.-ALABAMA; Jefferson Co., Cahaba River at Interstate 59, 25-
VII-81, S. Harris and P. O'Neil.
ETYMOLOGY.--Named for the Cahaba River.
REMARKS.-C. cahaba appears to be restricted to the headwaters of the Cahaba
River. At the type locality, the river is narrow, rocky bottomed and fairly rapid. Exten-
sive collecting throughout the Cahaba basin in the lower Appalachian mountains by
Harris and colleagues (Harris et al. 1984) failed to yield additional specimens.

Cheumatopsyche bibbensis, NEW SPECIES
Fig. 3.
This species will key to couplet 30 in Gordon's (1974) key. It is separated from all
other Cheumatopsyche spp. males by the nipple-like apices of the apical lobes of the
tenth tergum.
MALE: Length-6 mm. Color of head and body brown, legs light brown; antennae
brown with no apparent markings; wings general. Ninth segment annular, bearing long
setae on dorsal lobes and latero-posterior margin, lateral apical lobe angular. Tenth
tergum long with large, setose cerci anteriad of apex, apically divided into long, dorsally
projecting lobes; in lateral view, lobes at oblique angle, not projecting above dorsum of
tenth tergum; in dorsal view, lobes widest at midpoint, constricting into nipple-like
apices. Claspers short and exceedingly thin, apex of basal segment not clearing dorsum
of tenth tergum; apical segment one-third length of basal segment, linear and narrowed
slightly at tip in caudal view. Phallotheca with base moderately enlarged.
FEMALE .-Unknown.
HOLOTYPE MALE.-ALABAMA: Bibb Co., Cahaba River at Co. Hwy. 27, 6-X-82,
S.C. Harris.
ETYMOLOGY.-Named for county of type locality.
REMARKS.-This species has been collected only from the type locality on the
Cahaba River in an area where the river flows across an extensive limestone outcrop-


Gordon et al.: New Caddisflies



\ ''

- /i/=
-z L
!^E^ ^ ^~/


Figs. 2-4. Male genitalia. A, lateral aspect; B, caudal view of apical lobes of tenth
tergum; C, caudal view of claspers. Fig. 2. C. cahaba n.sp.; Fig. 3. C. bibbensis, n.sp.;
Fig. 4. C. kinlockensis, n.sp.

318 Florida Entomologist 69(2) June, 1986

Cheumatopsyche kinlockensis, NEW SPECIES
Fig. 4.
This species will key to couple 38 in Gordon's (1974) key, but can be distinguished
from C. ela Denning and C. campyla Ross by its short, tapered lobes of the tenth
tergum. Overall, the genitalia of this species are most suggestive of C. (harwoodi)
harwoodi Denning, but the diagonal upward slant of the dorsal edge of the apical lobes
in lateral view and their tapered apices in caudal view render C. kinlockensis distinct.
MALE: Length-7.5 mm. Color of head and body brown, venter and antennae light
brown; wings general. Ninth segment annular, bearing long setae on dorsal lobes and
latero-posterior margin, lateral apical lobe broadly rounded. Tenth tergum long with
small nodiform setose cerci anteriad of apex, divided apically into moderately long,
dorsally projecting lobes; in lateral view, lobes at an oblique angle, extending slightly
above mesal apex of the tenth tergum, dorsal edge of lobes slanted upward from an-
teriad to posteriad; in caudal view, lobes widest at midpoint, tapered at apices and
widely separated. Claspers short, apex of basal segment slightly enlarged and not clear-
ing dorsum of tenth tergum; apical segment approximately one-third length of basal
segment, sinuate with base widened in caudal view. Phallotheca long, base moderately
HOLOTYPE MALE.-ALABAMA; Lawrence Co., Hubbard Creek below Kinlock
Falls, on Forest Rd. 210, Bankhead National Forest, 28-V-83, S. C. Harris.
PARATYPES.-ALABAMA: as above, 1 male; as above except 28-V-85, 2 males.
ETYMOLOGY. -Named for Kinlock Falls.
REMARKS. -This species is known only from the type locality, a fast-flowing, rocky
stream of the Cumberland Plateau.


This research was supported by a research program (Flax 79009) of the SEA/CSRS,
USDA. The Geological Survey of Alabama provided facilities and supplies to the junior
author (SCH) during the course of the study and they are gratefully acknowledged. The
assistance of Patrick O'Neil in the collection of specimens is also appreciated.


GORDON, A. E. 1974. A synopsis and phylogenetic outline of the Nearctic members of
Cheumatopsyche. Proc. Acad. Nat. Sci. Philadelphia 126: 117-160.
HARRIS, S. C., P. K. LAGO AND P. E. O'NEIL. 1984. Trichoptera of the Cahaba River
system in Alabama. Entomol. News. 95: 103-112.

Epler: Nanocladius Symphoretic on Traverella



Department of Entomology
Florida A&M University
Tallahassee, FL 32307


The pupa and larva of Nanocladius (Plecopteracoluthus) bubrachiatus sp. nov.
(Diptera: Chironomidae) are described from Honduras. This species lives in symphoretic
association with the leptophlebiid mayfly Traverella sp. This is the first record of a
Nanocladius living in symphoretic association with a mayfly. The larva is unusual in
that it possesses a premento-hypopharyngeal fulcrum with long dorsocaudally directed
arms. A key is included for the immature stages of Nanocladius (Plecopteracoluthus).


Se described la pupa y la larva de Nanocladius (Plecopteracoluthus) bubrachiatus
sp. nov. (Diptera: Chironomidae) de Honduras. Esta especie vive en asociaci6n sin-
for6tica con el leptoflebiido Traverella sp. Este es el primer record de un Nanocladius
viviendo en associaci6n sinfor6tica con una efimera. La larva es fuera de lo comdn ya
que posee un fulcro del complejo premento-hipofaringeal con brazos largos dirigidos
dorsocaudalmente. Se incluye una clave para los estados inmaduros de Nanocladius

Larvae of the orthoclad genus Nanocladius Kieffer are well known as symphoretic
associates of Plecoptera (Steffan 1965, Saether 1977) or Megaloptera (Hilsenhoff 1968,
Gotceitas & MacKay 1980). Nanocladius is composed of 2 subgenera, N. (Nanocladius)
and N. (Plecopteracoluthus). Saether (1977) considered Plecopteracoluthus Steffan to
be a subgenus of Nanocladius modified for symphoretic life. At least one species of
Nanocladius (Nanocladius) is also symphoretic on Megaloptera (Gotceitas and MacKay
Most records of chironomid-mayfly symbiosis deal with parasitic relationships usu-
ally involving the chironomid genus Symbiocladius Kieffer. Arvy & Peters (1973) pro-
vide a detailed account of the chironomids and mayflies involved in such relationships.
I recently examined some mayfly nymphs of Traverella Edmunds (Ephemeroptera:
Leptophlebiidae) from Honduras which had chironomid larvae attached to their bodies.
The mayfly nymphs are similar to Traverella sp. A of Allen (1973). This paper describes
the larva and pupa of a new species of Nanocladius (Plecopteracoluthus) from those
nymphs and presents the first record of a symphoretic association between N. (Plecop-
teracoluthus) and a mayfly.
Terminology and abbreviations follow Saether (1980). All measurements are in
micrometers unless otherwise stated, and consist of a range, mean, and the number of
specimens measured in parentheses if different from the number (n) stated at the begin-
ning of the description. Means are not given for samples of 3 or less. Postmentum length

Florida Entomologist 69(2)

June, 1986

is measured from the bottom of the division of the median teeth to the caudal margin
of the postmentum.


Nanocladius bubrachiatus sp. nov.

PUPA (n = 1).
Color. Light brown.
Total length 3.15 mm.
CEPHALOTHORAX. Thoracic horn elongate-ovate, with small spines (Fig. 1); 76 long,
22 wide at widest point; length/width ratio 3.45. Frontal setae 58 long, on small, low
tubercles. Thoracic setal lengths: LAps 35; MAps 78; Pc1 30; Pc2 130; Pc3 105; Dce 35;
Dc2 lost; Dc3 and Dc4 25; Dce 140 anterior to Dc3, Dc3 25 anterior to Dc4.
ABDOMEN (Fig. 2). PSA present on S IV-VII, strongest on IV, becoming progres-
sively weaker. PSB absent. Segment I with 1 L-seta (segment damaged), II and III
with 2 L-setae, IV-VI with 3 L-setae, VII with 4 filamentous L-setae, VIII with 4 or
5 filamentous L-setae (one side with 4, the other with 5). T II without prominent
caudomesal protuberance; with approximately 73 caudomesal hooklets, the longest 9
long. T III-VI with caudal rows of spines; T IV-VI also with weak median shagreen
area; median spine patches absent on all tergites. Lengths and numbers of tergal caudal
spines: T III with 150, longest 12; IV with 115, longest 8; T V with 71, longest 12; T
VI with 43, longest 14. T VII-VIII with median shagreen patches. Anal lobe with fine
anterodorsal shagreen; with 30-31 setae/side. Anal macrosetae weak, about 50 long, 2
FOURTH INSTAR LARVA (n = 2, unless otherwise stated).
Color. Head capsule light brown with darker posterior margin.
Total length 3.15 mm (1).
HEAD. Postmentum length 135-140. All labral setae simple; pecten epipharyngis of
3 simple, smooth, sharply-pointed spines. Antenna 4-segmented, with 3rd segment dis-
tally thinner (Fig. 3). Antennal segment lengths: 48-49; 15-16; 7-8; 4-5. AR 1.66-1.88.
Antennal segment I 14-15 wide; apical blade 20-24 long; ring organ 2-3 from base; seta
1 from base; apical setal mark 27-29 from base. Mandible (Fig. 4) 87-96, 91 (4) long.
Premandible (Fig. 5) 37-46 (3) long, with sharply bifid apex and 3 inner teeth. Mentum
(Fig. 6) with 2 pointed ventromental teeth flanked by 2 anterolateral ventromental
teeth, and 6 pairs of darker dorsomental teeth. Ventromental plates elongate and nar-
row (Fig. 6). Maxilla reduced (Fig. 7). Fulcrum of premento-hypopharyngeal complex
with 2 large dorsocaudally directed arms, each arm about 90 pm high (Figs. 10-12).
BODY. Distal claws of anterior parapods smooth (Fig. 8), mesal and proximal claws
strongly serrated (Fig. 9). Body setae simple. Procercus 7-11 wide, 8-10 high; with 2
small (14-15 long) setae, 2 medium (45-65 long) setae and 3 large (130 long) setae. Dorsal
anal tubules appear slightly larger than ventral anal tubules, both with an apparent
medial constriction (Fig. 13).
IMAGINES. Unknown.
DIAGNOSIS. The pupa is separated from N. (P.) branchicolus Saether and N. (P.)
downesi (Steffan) by the lack of median spine patches on T III-VII, and by the number
of anal lobe setae: 30-31 in N. (P.) bubrachiatus, 15-20 in N. (P.) downesi, 49 in N.
(P.) branchicolus. The larva is distinguished by the large dorsal arms of the fulcrum,
the shape and number of teeth of the mentum and the 4-segmented antennae. Both
larvae and pupae can also be distinguished by their phoretic association with Traverella

Epler: Nanocladius Symphoretic on Traverella 321

r-- -1
I/I l %V .

Figs. 1-3. Nanocladius (Plecopteracoluthus) bubrachiatus: 1) Pupal thoracic horn
and precorneal setae; 2) pupal abdomen, dorsal (segment I missing); 3) larval antenna.
Figs.~~ 1-3 Nocaius(Pecptracltu)bbahau:1 ua hrcchr

and ~ O prcoea seae 2) pupal abdme, dosa (egmen I isn))lavlatna

322 Florida Entomologist 69(2) June, 1986

4 6

5 7

8 9

Figs. 4-9. Nanocladius (Plecopteracoluthus) bubrachiatus, larval structures: 4)
mandible; 5) premandible; 6) mentum; 7) maxilla; 8) distal claws of anterior parapods;
9) proximal and mesal claws of anterior parapods.

ETYMOLOGY. From the Latin bu, meaning large, great; and the Latin brachiatus,
meaning with arms or branches. The name refers to the large dorsolateral arms of the
fulcrum of the larval premento-hypopharyngeal complex.
MATERIAL EXAMINED. Holotype: female pupa with larval exuviae, HONDURAS:
Comayagua; Rio Humuya, 12 km NW Comayagua (milky R., thorn scrub), 19-VII-1977,

Epler: Nanocladius Symphoretic on Traverella

12 13

Figs. 10-13. Nanocladius (Plecopteracoluthus) bubrachiatus, larval structures: 10)
premento-hypopharyngeal complex, frontal view; 11) same, dorsal view; 12) slightly
oblique lateral view of head capsule, showing position of dorsal arms of fulcrum; 13)
anal end (DA, dorsal arms of fulcrum; LP, labial palpi; SD, salivary ducts).

leg. L. B. & C. W. O'Brien, G. B. Marshall [on Traverella sp.]. Paratypes (4): 4 larvae
on Traverella sp., same data as holotype.
TYPE INFORMATION. The holotype and one paratype will be deposited in the Florida
State Collection of Arthropods, Tallahassee. Paratypes will be placed in my personal
collection and in the Zoologische Staatssammlung, Munich, West Germany. One
paratype remains attached to its mayfly host, preserved in alcohol; the other specimens
are mounted in Euparal on microscope slides.

Florida Entomologist 69(2)



Figs. 14-15. Traverella sp. : 14) nymph with N. (P.) bubrachiatus larva attached
above gills; 15) nymph with N. (P.) bubrachiatus larva attached on outside of wing pad;
note empty tube at midline of wing pads.

June, 1986

Epler: Nanocladius Symphoretic on Traverella



The larvae of N. (P.) bubrachiatus live in symphoretic association with Traverella
sp. Five Traverella nymphs were examined. Chironomid larvae and one pupa (with its
larval exuviae still attached) were found in silken tubes attached to both the outside
and inside edges of the nymphal wing pads, along the dorsolateral margin of the abdo-
men above the gills (Figs. 14-15), and between the hind coxae. Three of the nymphs
had one larval tube on each; 2 nymphs had 2. Larvae (and the empty larval tubes) were
located on both sides of the nymphs (3 on the right side, 3 on the left, one in the middle).
All larvae and the one pupa examined were positioned with their heads facing the caudal
end of the mayfly. None of the mayfly nymphs appeared to be damaged or malformed.
Traverella nymphs inhabit the rapids of medium-sized to large rivers, where they
live under rocks (Edmunds et al. 1976), a habitat not unlike that of the Plecoptera which
are utilized by other members of the subgenus Plecopteracoluthus. The "milky R.,
thorn scrub" listed in the collection site data refers to the milky color of the Rio Humuya
where it flows through an area of thorn scrub (C. W. O'Brien, personal communication).


The lack of both PSB and a caudomesal projection on T II of the pupa, and the
straight elongate caudal apices of the ventromental plates of the larva place N. (P.)
bubrachiatus in the subgenus Plecopteracoluthus as recognized by Saether (1977).
The most striking character distinguishing larval N. (P.) bubrachiatus from the
other species in the subgenus is the premento-hypopharyngeal complex (Figs. 10-12).
From each side of the fulcrum of the premento-hypopharyngeal complex (Saether
1980:25), a heavily sclerotized dorsal arm projects and bows gently caudad (Fig. 12).
The arms are apparently not fused with the head capsule, as are the hypopharyngeal
suspensorial bars of Anopheles quadrimaculatus Say illustrated by Menees (1962: Fig.
21). Hirvenoja (1973: Fig. 241) illustrated extended fulcrum arms in Protanypus spp.
They are, however, much smaller than those in N. (P.) bubrachiatus. It was not possible
to discern any muscle attachments to this structure in N. (P.) bubrachiatus.
The larvae of most Nanocladius species possess a ventromentum with 2 pointed
median teeth flanked by low, rounded humps or in a planar area which extends to the
anterolateral corners of the ventromentum (see Saether 1977: Figs. 10F, 16F, 19D).
The ventromentum of N. (P.) bubrachiatus has 2 pointed median teeth flanked by
another pair of pointed teeth at the anterolateral corners of the mentum (Fig. 6).
The larval antennae in N. (P.) bubrachiatus and N. (P.) downesi are apparently
4-segmented. In N. (P.) bubrachiatus the 3rd and 4th antennal segments appear to
have been fused, indicated by a change in the antennal wall; the distal 1/5 of the 3rd
segment is thinner than the basal portion (Fig. 3). Indications of a slight ring-like
thickening of the antennal wall at this point are evident. Steffan's (1965) figure does not
illustrate a similar construction for N. (P.) downesi.
Mature adults of N. (P.) bubrachiatus were not available. Saether (1977:6) stated:
"because there are very few characters to separate the adults, an identification without
associated pupae can only be regarded as tentative. The pupae, however, are easily
distinguishable .. ." The pupa (and larva) of N. (P.) bubrachiatus are easily distin-
guished from the other described species of Plecopteracoluthus (see Diagnosis).
The developing female imago within the pupal skin is not sufficiently mature to
examine thoroughly; developing ommatidia and seminal capsules are barely visible.
However, the tarsi are sufficiently developed to observe that pulvilli are present. The
2 other species of N. (Plecopteracoluthus) possess pulvilli, although Steffan (1965)

Florida Entomologist 69(2)

June, 1986

stated that pulvilli were absent in Plecopteracoluthus downesi. However, his illustra-
tion (Steffan 1965: Fig. 7) clearly shows a pulvillus. Saether (1977) described N. (P.)
branchicolus with pulvilli, and used the lack of pulvilli as a character to separate N.
(P.) downesi from N. (P.) branchicolus. Because both species possess pulvilli, this
character should be eliminated from Saether's adult keys; the other characters given in
the keys will separate the 2 species.
The following key to the immature stages of N. (Plecopteracoluthus) is based on the
descriptions of Steffan (1965), Saether (1977) and the present paper.

KEY TO IMMATURE STAGES OF Nanocladius (Plecopteracoluthus)

1. Pupae ...................................................... ................... ......... ....... 2
Larvae .............................................. .......................................... 4
2. Abdominal tergites V-VI without median spine patches; 30-31 setae on anal lobe;
symphoretic on Leptophlebiidae (Ephemeroptera)
............... ............................................... N (P.) bubrachiatus sp. nov.
Abdominal tergites V-VI with median spine patches; either 15-20 or about 50 setae
on anal lobe; symphoretic on Perlidae (Plecoptera) ................................... 3
3. Anal lobe with 15-20 setae; median spine patches well developed only on T V-VI
............................................................................. N (P .) dow nesi Steffan
Anal lobe with about 49 setae; median spine patches well developed on T IV-VII
............ ................ ........ ....... N. (P.) branchicolus Saether
4. Ventromentum with 4 pointed teeth; premento-hypopharyngeal complex with ex-
tremely large dorsocaudally directed fulcrum arms
......................................... ..... ........... N. (P.) bubrachiatus sp. nov.
Ventromentum with only 2 pointed median teeth; premento-hypopharyngeal com-
plex without extremely large fulcrum arms ......................................... 5
5. First 2 pairs of dorsomental teeth fused, producing mentum with 11 teeth; antennae
4-segmented, AR about 2.3; living in gelatinous cases N. (P.) downesi Steffan
Mentum with 13 teeth; antennae 5-segmented, AR about 1.5; living in silken tubes
.................. ......................................... N. (P.) branchicolus Saether


I would like to thank Dr. R. Wills Flowers for making the specimens available to
me. Dr. Annelle R. Soponis provided helpful comments on an earlier draft of this paper.
This research was partially supported by a CSRS/USDA grant (FLAX 79009) at Florida
A&M University.


ALLEN, R. K. 1973. Generic revisions of mayfly nymphs. 1. Traverella in North and
Central America (Leptophlebiidae). Ann. Ent. Soc. America 66: 1287-1295.
ARVY, L. AND W. L. PETERS. 1973. Phor6sies, biocoenoses et thanatocoenoses chez
les Eph6meropteres. pp. 254-312 in Peters, W. L. and J. L. Peters (eds.): Pro-
ceedings of the First International Conference on Ephemeroptera. E. J. Brill,
Leiden, Netherlands.
EDMUNDS, G. F., JR., S. L. JENSEN, AND L. BERNER. 1976. The Mayflies of North
and Central America. University of Minnesota Press, Minneapolis. 330 pp.
GOTCEITAS, V. AND R. J. MACKAY. 1980 The phoretic association of Nanocladius
(Nanocladius) rectinervis (Kieffer) (Diptera: Chironomidae) on Nigronia ser-
ricornis Say (Megaloptera: Corydalidae). Canadian Ent. 58: 2260-2263.


Deyrup & Manley: Mutillid Size Variation

HILSENHOFF, W. L. 1968. Phoresy by Plecopteracoluthus downesi on larvae of Nig-
ronia serricornis. Ann. Ent. Soc. America 61: 1622-1623.
HIRVENOJA, M. 1973. Revision der Gattung Cricotopus van der Wulp und ihrer Ver-
wandten (Diptera, Chironomidae). Ann. Zool. Fennici 10: 1-363.
MENEES, J. H. 1962. The skeletal elements of the gnathocephalon and its appendages
in the larvae of higher Diptera. Ann. Ent. Soc. America 55: 607-616.
SAETHER, O. A. 1977. Taxonomic studies on Chironomidae: Nanocladius,
Pseudochironomus, and the Harnischia complex. Bull. Fish. Res. Bd. Canada
196: 143 pp.
--. 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae).
Ent. Scand. Suppl. 14: 1-51.
STEFFAN, A. W. 1965. Plecopteracoluthus downesi gen. et sp. nov. (Diptera,
Chironomidae), a species whose larvae live phoretically on larvae of Plecoptera.
Canadian Ent. 97: 1323-1344.


Archbold Biological Station
P.O. Box 2057
Lake Placid, FL 33852
Clemson University
Pee Dee Research and Education Center
P.O. Box 5809
Florence, SC 29502


The relative sizes of male and female Mutillidae were studied at the Archbold Biolog-
ical Station in south-central Florida. Females are larger than males in Dasymutilla
pyrrhus (Fox), Pseudomethoca sanbornii aeetis (Fox), and P. oculata (Banks). Males
are larger than females in Timulla d. dubitata (Smith), T. floridensis (Blake), Ephuta
f. floridana Schuster, E. m. margueritae Schuster, E. s. slossonae (Fox), Dasymutilla
archboldi Schmidt and Mickel, and Photomorphus paulus (Bradley). There is no signif-
icant difference in Dasymutilla asopus bexar (Blake), D. nigripes (Fabricius), and D.
castor (Blake). Sex-biased size variation may be associated with courtship behavior or
host-seeking behavior.


Se estudi6 el tamafio relative de machos y hembras de Mutillidae en la Estaci6n
Biol6gica de Archbold en el centro-sur de la Florida. Las hembras son mas grande que
los machos en Dasymutilla pirrhus (Fox), Pseudomethoca sanborinii aeetis (Fox), y
P. oculata (Banks). Los machos son mas grande que las hembras en Timulla d. dubitata
(Smith), T. floridensis (Blake), Euphata f. floridana Schuster, E. m. margueritae
Schuster, E. s. slossonae (Fox), Dasymutilla archboldi Schmidt y Mickel, y Photomor-
phus paulus (Bradley). No hay diferencia significant en Dasymutilla asopus bexar

328 Florida Entomologist 69(2) June, 1986

(Blake), D. nigripis (Fabricius), y D. castor (Blake). Variaciones de tamafo parciales
al sexo pudieran estar asociadas con el comportamiento de cortejo o la buisqueda de

Among solitary parasitoid wasps, the size of the adult insect is limited by the size
of the host consumed by the larva. The sex of the wasp is normally determined by
whether the individual is haploid (male) or diploid (female) (Flanders 1946). A mated
female can potentially determine the sex of an offspring by releasing or withholding
stored sperm at the time of oviposition (Flanders 1946). Parasitoid wasps that attack
larvae of varying sizes have the opportunity, if the wasp is able to gauge the size of the
host, to allocate hosts to one sex or the other on the basis of size. Such sex-biased host
allocation occurs in the braconid Coeloides brunneri Viereck (Ryan 1961), in the
braconid Heterospilus prosopidis Viereck, and the pteromalid Lariophagus distinguen-
dus (Foerster) (Charnov 1982), all of which tend to lay male eggs on small hosts and
female eggs on large hosts.
Velvet ants (Mutillidae) are, in certain ways, particularly well suited for a survey
of sex-correlated size variation in natural populations. All species for which hosts are
known are solitary parasitoids that feed on fully grown host prepupae and pupae (Mickel
1928); continued host growth after the parasitoid has chosen the sex of her offspring
does not complicate analysis. We assume that the larvae completely consume their
hosts, as we know of no examples of solitary parasitoids that share a cell with a partially
consumed host. There is obvious size variation in some species of velvet ants. At our
study site, for example, female Dasymutilla pyrrhus (Fox) vary from 8.9 to 15.5 mm
in length, and male Timullafloridensis (Blake) vary from 6.0 to 13.6 mm. Mickel (1928)
showed that size variation in adult velvet ants is related to host size in the case of
Dasymutilla bioculata (Cresson), whose bimodal size distribution (in both sexes) is
correlated with the different sizes of its two host species. Preliminary surveys of collec-
tions of velvet ants from various sites indicated tendencies toward sex-biased size differ-
ences in some species, random size variation in other species, and little size variation
in other species. Of particular interest was the relatively large size of males of a number
of species. Velvet ants seem to offer an intriguing diversity in allocation of resources
between the sexes.
Although museum collections give an indication of patterns in size variation, these
collections do not make a good data base because many collectors preferentially capture
large "fine" specimens. Moreover, the specimens in a normal collection do not reflect
the consensus of size allocation by females of a population faced with the same spectrum
of hosts. The range of host sizes available to polyphagous mutillids is likely to vary from
place to place, and site data on museum specimens are seldom precise enough to ensure
that specimens are from the same populations. As an example of this site effect, we
have found that Dasymutilla pyrrhus on the coastal dunes of Sanibel Island (Florida)
average significantly smaller than the same species at the Archbold Biological Station,
presumably because the only abundant potential host at Sanibel seemed to be a rela-
tively small sphecid, Microbembex monodonta (Say). The range of host sizes is also
likely to vary from year to year and even season to season, depending on fluctuation in
populations of various hosts. Therefore, study of size variation in a population of velvet
ants requires specimens collected within the same time period and from a small area,
preferably one with a rich variety of hosts. This study controls these variables in a
survey of mutillid populations in one location over one season.

Deyrup & Manley: Mutillid Size Variation



The Archbold Biological Station (ABS) is an ecological reserve of about 4,300 acres
in Highlands County in south-central Florida. The ABS is on the Lake Wales Ridge,
whose well-drained sand soils are ideal for fossorial hosts of velvet ants; the ABS has
over 30 species of mutillids, some of which are very abundant. Collections were made
in an area of sand pine (Pinus clausa Chapman) scrub and adjacent firelanes. The study
site was a triangular area roughly 300 x 200 x 250 m. The high mobility of mutillids
observed within this area makes it unlikely that there are discrete subpopulations with
different host populations within the study site. All specimens were collected in summer
of 1984.
The hosts of many species of velvet ants are not known. Many hours of watching
velvet ants at the ABS produced only a few observations of burrow-entering. The size
allocations made by ovipositing velvet ants were therefore studied indirectly by captur-
ing, drying, and weighing the adults that result from these allocations. Specimens were
weighed rather than measured because the extreme sexual dimorphism in all species
prevents comparisonof linear dimensions. Males were collect in flight with an insect net
and in Malaise traps. Females were collected in #10 tin can pitfall traps and from the
surface of the ground. These methods may have biased our results if there is size-related
mortality in one sex that is not matched in the other sex. Dried specimens were weighed
on a Mettler H6 digital analytical balance (Mettler Instrument Co., Hightown, N.J.).
We were not able to separate weights of soft and hard tissue; the former may vary with
the age and nutritional state of the adult. We believe that in all specimens the age-re-
lated differences in dried soft tissues are insignificant compared with the weight of the
non-varying soft tissues and the extraordinarily massive exoskeleton. The results were
analyzed with the Mann-Whitney U-test (two-tailed) Siegel, 1956). Most of the speci-
mens are in the collection of the Archbold Biological Station, with voucher series in the
collection of Dr. Donald Manley (Florence, S.C.) and the Florida State Collection of
Arthropods, (Gainesville, FL).


The results are summarized in Table 1. The 13 species can be divided into 3 groups:
those in which the females tend to be heavier (3 species), those in which the males tend
to be heavier (7 species), and those in which there is no clear correlation between six
and weight (3 species).


The 3 species with large females follow the general rule among arthropods that
where size dimorphism occurs the female is usually the larger sex, as can be seen in
any collection of spiders or Orthoptera. The usual explanation for such dimorphism is
that egg production and storage is directly related to size, whereas male reproductive
success often depends more on agility and rapid development than on size (Darwin,
1881). Among provisioning and parasitoid Hymenoptera, which are able to allocate
resources on the basis of sex, the limitation of resources results in a tendency to allocate
more to females, for whom the reproductive penalty for small size is likely to be more
severe. Additionally, it may be more efficient for a female to invest in a large number
of small males rather than a few large males even when large males have higher repro-
ductive success (Alcock et al. 1977).

Florida Entomologist 69(2)

June, 1986

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Deyrup & Manley: Mutillid Size Variation

Dasymutilla pyrrhus (Fig. 1) was the only species of 5 studied in the genus in which
females are clearly heavier than males. The females also show great variation and a
high standard deviation (18.2) in weight. At the ABS, we have often observed this
species around nest aggregations of Bembix sayi Cresson and we have seen D. pyrrhus
entering the burrows of B. sayi and B. texana Cresson. A large number of D. pyrrhus
were found around the nesting aggregations of Microbembex monodonta on Sanibel
Island. Since this species tends to attack aggregating burrowers, the ability to develop
and store a number of eggs at a time may be an advantage of large females. On the
other hand, there is clearly no absolute size criterion for allocation of hosts to a particu-
lar sex in this species, and this flexibility may be important in a species that attacks
aggregating hosts, as a rigid adherence to host size as a determinant of sex could result
in highly skewed sex ratios where there are aggregations of a uniform host.
In Pseudomethoca sanbornii aeetis (Fox) (Fig. 2) and P. oculata (Banks) (Fig. 3)
there may be an additional factor favoring large size in females. The head of the female
of both species is greatly enlarged and might be used to block the host gallery if these
species tend to spend a period of days in the host burrow, as in the case of
Pseudomethoca frigida (Smith) (Brothers, 1972). The macrocephalic queens of eusocial
halictid bees described by Wilson (1971) show a remarkable morphological convergence
with female Pseudomethoca species. The known hosts of Pseudomethca are halictids
(Krombein, 1979), and it appears that the mutillids and the eusocial species of halictids
may have independently evolved the same features that permit domination of a halictid


In the genera Timulla (Figs. 4, 5) and Ephuta (Figs. 6-8), the larger size of males
is probably related to the male habit of carrying the female during courtship and copu-
lation. Phoretic copulation has been reported in Timulla oajaca (Blake) (Linsley, 1960)
and T. dubitata (Smith) (Sheldon, 1970). We have captured phoretic couples of T.
floridensis (Blake), Ephuta sabaliana Schuster and E. floridana Schuster. Phoretic
copulation with concomitant large males has been described in the families Bethylidae
and Tiphiidae as well as the Mutillidae (Evans, 1969). Large male size in certain
Hymenoptera was noted by Darwin (1881, p. 279) who mentions the apterogynous
tiphiid Methoca ichneumonides Latreille. "The explanation of this anomaly is that a
marriage flight is absolutely necessary with the species, and the male requires great
strength and size in order to carry the female through the air." In species with wingless
females (including all Mutillidae), phoretic copulation may have an important dispersal
function, as the female may thus cross barriers such as streams (Evans, 1969). Phoret-
ically copulating males may gain an advantage over potential rivals by obtaining sole
possession of the female during flight. It has recently been shown (O'Neill and Evans,
1983) that phoretic behavior is a highly effective strategy for larger individuals among
males of a species of sphecid; the female in this case is fully winged. Whatever the
selective pressures that led to phoretic behavior in Timulla and Ephuta, the antiquity
of phoresy in these genera is shown not only by the consistently large male size, but
also by tle modifications of the male clypeus and frons to accommodate the head of the
female (Sheldon, 1970).
While phoretic behavior may select for size dimorphism, another factor may also
select for small size in females of non-phoretic and phoretic species. If a species attacks
burrowing hosts of differing sizes, large females may have difficulty entering the bur-
rows of small host species. Where large size is not important for defense or for rapid
egg storage and production, one would expect selection for small size in females of
polyphagous velvet ants.

332 Florida Entomologist 69(2) June, 1986


Dasymutilla 5 Pseudomethoca
4 pyrrhus sanbornii


L a w
3 3
a a 3n

z 2 z

5 0 i 20 2 30 35 40 5 55 60 5 70 75 80
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Deyrup & Manley: Mutillid Size Variation


IllA Ill .
6 8 10 12 14 16 18 20 22 24 26 28

I asopus


14 16 18

20 22 24 26

28 30

2 4 6 8 10 12 14 16


2 4 6 8 10121416182022242628303234363840

Fig. 1-13: Weights of Mutillidae spp. Solid bars are males, open bars are females.

This latter explanation for small female size may apply to Dasymutilla archboldi
Schmidt and Mickel (Fig. 9), which shows a sexual size dimorphism as dramatic as those
of Ephuta and Timulla species. There are no reports of phoretic copulation in
Dasymutilla species, and male Dasymutilla show no modifications of the head or man-
dibles correlated with phoresy (Sheldon, 1970) or combat between males. Phoretic be-
havior in D. archboldi cannot be dismissed but seems highly unlikely. Dasymutilla
archboldi appears to be a Florida endemic confined to a few former islands in the central
part of the state, where the soil is very deep well-drained sand (Schmidt and Mickel,
1979). In the absence of some potential competitors, D. archboldi may have evolved an
expanded host range to efficiently use both large and small hosts. No other eastern



334 Florida Entomologist 69(2) June, 1986

Dasymutilla shows such striking and consistent size dimorphism. The most obvious
relevant characteristic of the habitat of D. archboldi is the uniform sandy soil occurring
over a large are; the consequence of this edaphic uniformity is that no burrowing bees
or wasps, with the exception of Bembix and Cerceris species, appear to form aggrega-
tions. When hosts are diverse and very widely dispersed, small size in the female velvet
ant is not limited by selection for rapid production and storage of many eggs, and
adherence to host size as the determinant of sex is not likely to lead to gross imbalance
in sex ratios.
Photomorphus paulus (Bradley) (Fig. 10) belongs to a genus whose biology is so
poorly known that we hesitate to speculate on the significance of the relatively small
size of the females. We suspect that phoresy does not occur in this species as we have
captured hundreds of males in Malaise flight traps, and not one female. Out of a much
smaller total number of Ephuta and Timulla, there were a number of females found in
the traps, sometimes still in the grasp of the male.
There is a certain amount of overlap in the weights of males and females in all these
species with large males. This overlap may be due to imprecise host measurement by
ovipositing females, to negligible effects of size on fitness in either sex within the
overlapping range, or to a tendency to balance sex ratios without ignoring any hosts.


Dasymutilla asopus bexar (Blake) (Fig. 11) and D. nigripes (Fabricius) (Fig. 12)
show no significant sex-bias in size variation, though both species vary considerably in
weight in both sexes. Females in these species may be indiscriminate in resource alloca-
tion, or there might be a pattern of resource allocation that is too subtle to be revealed
in the small number of individuals in these samples. Dasymutilla castor (Blake) (Fig.
13) seems to show a pattern in host size utilization even though the statistical analysis
shows no difference in sizes of males and females. The graph of frequency distribution
shows in the range of 4 to 14 mg a pair of more or less normal distribution curves, with
the females clearly heavier, much as in D. pyrrhus. Above 14 mg, however, all but one
of the 13 specimens were males. It is possible that in the lower size ranges the normal
advantages of large female size prevail, but above a certain threshold the problem of
gaining access to burrows may come into play as a selective pressure influencing the
behavioral repertoire of the female.


The explanations of the data presented in this study are highly speculative and
would not have been advanced if we had felt there was a practical method for eventually
obtaining direct observations on the oviposition behavior and direct evidence of the
effects of natural selection on various size classes of males and females. The principle
value of the study is that it shows diversity of male-female size ratios in Mutillidae and
focuses attention on the importance of resource allocation, which is only possible because
of the male haploidy of higher Hymenoptera. Male haploidy might arise and be main-
tained as a mechanism to increase the maternal genome at the expense of the paternal
genome (Bull, 1979); moreover male haploidy may be important in social Hymenoptera
because it makes sisters more closely related to each other than to their offspring
(Wilson, 1971). But the most direct benefit of male haploidy is that it permits optimal
reproductive success by differential allocation of resources between the sexes in re-
sponse to differences in the biological roles of the male and female.

_ __~__

Deyrup & Manley: Mutillid Size Variation 335


The work of Donald Manley at the Archbold Biological Station was supported in part
by a grant from Archbold Expeditions, Inc. Dr. John Alcock (Arizona State University)
and Dr. John MacDonald (Purdue University) provided valuable suggestions on the
manuscript. Nancy Deyrup and Rebecca Kimball weighed the velvet ants. The graphs
were prepared by Nancy Deyrup. Todd Coleman, Richard Shepard, and Edward Fabian
assisted in collection of female mutillids. Dr. Lawrence Battoe helped with computer
analysis of data. The manuscript was typed by Dorothy Carter.


ALCOCK, J., C. E. JONES, AND S. L. BUCHMANN. 1977. Male mating strategies in the
bee Centris pallida Fox. (Anthophoridae: Hymenoptera). Amer. Natur. 111:
BROTHERS, D. J. 1972. Biology and immature stages of Pseudomethocaf. frigida, with
notes on other species (Hymenoptera: Mutillidae). Univ. Kans. Sci. Bull. 50: 1-38.
BULL, J. J. 1979. An advantage for the evolution of male haploidy and systems with
similar genetic transmission. Heredity 43: 361-381.
CHARNOV, E. L. 1982. The theory of sex allocation. New Jersey. Princeton University
Press. x + 355 pp.
EVANS, H. E. 1969. Phoretic copulation in Hymenoptera. Entomol. News 80:113-124.
FLANDERS, S. E. 1946. Control of sex and sex-linked polymorphism in the Hymenopt-
era. Quart. Rev. Biol. 21: 135-143.
KROMBEIN, K. V. 1979. Superfamily Scolioidea. Pp. 1253-1321, in K. V. Krombein, P.
D. Hurd, Jr., D. R. Smith, and B. D. Burks (eds.), Catalog of Hymenoptera in
America North of Mexico. Vol. 2, pp. i-xvi, 1199-2209.
LINSLEY, E. G. 1960. A fragmentary observation on the mating behavior of Timulla.
Pan-Pac. Entomol. 36:36.
MICKEL, C. E. 1928. Biological and taxonomic investigations on the mutillid wasps.
Bull. U.S. Nat. Mus. 143: ix + 351 pp.
O'NEILL, K. M., AND H. E. EVANS. 1983. Alternative male mating tactics in Bem-
becinus quinquespinosus (Hymenoptera: Sphecidae): correlations with size and
color variation. Behav. Ecol. Sociobiol. 14: 39-46.
RYAN, R. B. 1961. A biological and developmental study of Coeloides brunneri Vier.,
a parasite of the Douglas-fir beetle, Dendroctonus pseudotsugae Hopk. Unpub-
lished Ph.D. dissertation, Oregon State University, Corvallis. 172 pp.
SCHMIDT, J. O., AND C. E. MICKEL. 1979. A new species of Dasymutilla from Florida
(Hymenoptera: Mutillidae). Proc. Entomol. Soc. Wash. 81: 576-579.
SHELDON, J. K. 1970. Sexual dimorphism in the head structure of Mutillidae
Hymenoptera: a possible behavioral explanation.Entomol. News 81: 67-61.
SIEGEL, S. 1956. Nonparametric statistics for the behavioral sciences. New York,
McGraw-Hill Co. 312 pp.
WILSON, E. 0. 1971. The insect societies. Cambridge, Mass. Harvard University Press.
x + 548 pp.

336 Florida Entomologist 69(2) June, 1986


Entomology Department
International Rice Research Institute
P.O. Box 933, Manila, Philippines


The relative abundance of ant and striped earwig (Labidura riparia) (Pallas) (Der-
maptera: Libiduridae) populations was assessed in 78 central Florida citrus groves.
Ants were predominant in 74 groves while the striped earwig was the dominant pred-
ator in 4 groves where ants were absent. Detailed studies were made in Winn and High
Acres citrus groves. Winn grove, with residues of cyclodine chlorinated hydrocarbon
insecticide in the soil, had a dense population of striped earwigs and no ants. High Acres
citrus grove, with no insecticide soil residues, had a dense population of ants (ca. 30
species) and few striped earwigs, typical of the other 74 groves sampled. Without a
natural population of ants, the striped earwig became a major nocturnal soil surface
predator in the Winn grove.
Striped earwig predation was measured on monitored populations of Diaprepes ab-
breviatus (L.) (Coleoptera: Curculionidae) first instar larvae placed on the soil surface
under the citrus tree canopy. These earwigs, quiescent during the day, had a peak
predation period of 3 to 4 hours after sunset. Thereafter predation decreased gradually
and ceased before sunrise. Seasonal predation activity of the striped earwig was great-
est between May and October.


La abundancia relative de poblaciones de hormigas y de la tijerata rayada (Labiduro
riparia) (Pallas) (Dermaptera: Libiduridae) fue evaluada en 78 arboledas de citricos del
centro de la Florida. Las hormigas predominaron en 74 arboledas mientras que la
tijereta rayada fue el depredador mis dominant en 4 arboledas donde las hormigas
estaban ausentes. Se hicieron studios detallados en arboledas en Winn y High Acres.
En la arboleda de Winn, con residues en la tierra del insecticide ciclodine hidrocarburo
clorinado, huvo una densa poblacion de tijeretas rayadas y no de hormigas. La arboleda
de citricos de High Acres, sin residue de insecticides en la tierra, tuvo una densa
poblaci6n de hormigas (aprox. 30 species) y pocas tijeretas rayadas, que fue lo tipico
de las muestras de las otras arboledas. Sin una poblaci6n natural de hormigas, la tijereta
rayada se convirti6 en un important depredador nocturno en la superficie de la tierra
en la arboleda de Winn.
Se midi6 la depredaci6n de la tijereta rayada en poblaciones chequeadas del primer
estadio larval de Diaprepes abbreviatus (L.) (Coleoptera: Curculionidae) puesto en la
superficie de la tierra debajo del dosel de los arboles citricos. Estas tijeretas, en reposo
durante el dia, tuvieron el period de depredaci6n mAs alto e 3 a 4 horas despues de la
puesta del sol. De aqui en adelante, depredaci6n disminuy6 gradualmente y ces6 antes
de la salida del sol. La actividad estacional depredadora de la tijereta rayada fue mayor
de Mayo a Octubre.

The sugarcane rootstalk borer weevil, Diaprepes abbreviatus (L.), is an important
curculionid pest of sugarcane and citrus in the West Indies. It was first reported attack-

Tryon: Predators of Diaprepes 337

ing citrus near Apopka, Florida, in 1964 (Woodruff 1964). The weevil egg masses are
deposited between leaves which are bound together with an adhesive secretion. Jones
and Schroeder (1983) reported the freshly hatched (144 to 224 h after oviposition) neo-
nate larvae drop immediately (235 to 248 h after oviposition) to the soil surface under
the tree canopy and remained on the soil surface no longer than 3 h, between 1100 and
2400 h before burrowing to feed on the roots. This larval root feeding eventually causes
severe host decline and decreases fruit production. An area of about 20,000 ha, including
Apopka, Florida is presently under quarantine.
Since the ban on the use of cyclodiene insecticides, effort has been directed toward
evaluating alternative control methods. Whitcomb et al. (1982) and Richman et al.
(1985) reported that D. abbreviatus neonate larvae are vulnerable to soil-surface ar-
thropod predators. They documented the primary soil-surface predators in a Florida
citrus grove to be a complex of ant species. Several of these ant species were effective
predators when D. abbreviatus neonate larvae were placed on the soil surface for obser-
The striped earwig, Labidura riparia (Pallas) (Dermaptera: Labiduridae), is also an
important predator of insect pests in cultivated fields in California (Schlinger et al. 1959)
and several southeastern states (Clements 1968, Whitcomb 1973, Neal 1974, Nguyen &
Workman 1975, Walker & Newman 1976, Buschman et al. 1977, Travis 1977, and
Reinert 1978). Clements (1968) and Travis (1977) reported that the striped earwig was
common in cultivated fields, including groves, in central Florida. Uncultivated fields,
pastures, and wooded areas had few striped earwigs (Walker & Newman 1976). Tawfik
et al. (1972) reported that "the striped earwig was strictly an animal tissue feeder; in
other words a strict predator." Its life history was described by Tawfik et al. (1972)
and Shepard et al. (1973).
To identify and evaluate possible biotic factors affecting first instars of sugarcane
rootstalk borer larvae, D. abbreviatus (Coleoptera: Curculionidae), a survey for soil
surface predators was initiated in 78 commercial citrus groves near Orlando, Florida.
Two groves were selected to evaluate predation rates on the soil surface.


Arthropods were collected and identified from 78 citrus groves in Lake, Orange, and
Polk counties in central Florida. Baited traps were placed monthly for a 72-h sampling
period in the groves during June and July 1979 and for 12 months in 1980. Moist cat
food was used as bait in open 7-dram snap vials. The vials with the bait were placed
upright in 50 ml of 70% ethyl alcohol at the bottom of widemouthed 946 ml Mason jars.
Alcohol was necessary to prevent the trapped insects from being eaten or decaying.
Ten baited Mason jars per grove were randomly buried with the lid at soil surface under
the citrus canopy for 72 h. The lids to the Mason jars had six holes (21.5 cm diameter)
to allow insect predators to enter but prevent mammals and reptiles from reaching the
bait or the trapped insects.
Detailed predation experiments were conducted in 2 commercial citrus groves, High
Acres and Winn, located within the D. abbreviatus quarantine area northwest of Or-
lando, Florida. These 2 groves characterized the 2 extremes in soil-surface predator
populations. Recommended pesticide practices (fungicides, scalicides, miticides, spray
oils, etc.) were used in both groves by the commercial operator. Unlike High Acres,
Winn had a history of chlorinated hydrocarbon soil insecticides used to control several
weevil species between 1950 and the early 1970s. Since 1968, dieldrin (2/16/73, date
sprayed), chlordane (9/15/71), 4/21/70), and heptachlor (5/13/69, 11/20/69) have been

Florida Entomologist 69(2)

June, 1986

Weed species complexes (measured using 10 l-m2 samples in each grove), relative
density of soil surface arthropod predators, and pesticide soil residues present were
used to characterize the two groves. Predators were collected and identified from baited
Sampling L. riparia is difficult because it is nocturnal. Maximum feeding activity of
the striped earwig occurred just after sunset and continued at decreasing levels until
all activity ceased before sunrise (Clements 1968, Shepard et al. 1973, Walker & New-
man 1976, and Buschman et al. 1977). Striped earwig predation rates were evaluated
by using first instars of D. abbreviatus larvae as prey in open 55 mm (depth) petri
dishes. The D. abbreviatus larvae were obtained from USDA laboratory in Apopka,
Florida. The open petri dishes were placed into the soil within the citrus tree drip line.
This approximated the area of larval drop after hatching from foliar egg masses. Ten
petri dishes with 20 weevil larvae each were placed beneath each of 4 randomly selected
citrus trees for 20 minutes. This was repeated hourly for 24 h each Monday during the
summer and fall of 1980. A camel hair brush was used to keep the crawling weevil larvae
inside the dishes. Because of the small size of the weevil larvae (avg 1.2 mm by 0.5 mm)
and soil surface debris, petri dishes were necessary for accurate nocturnal observations
of larval predation by the striped earwig and other arthropods. Nocturnal observations
were made with flashlights covered by transparent red cellophane. The number of
larvae missing from each dish after each trial was calculated as predation percentage.
The predation source was identified by observation.



Preliminary insect pitfall trap data collected during the second week of June 1979
demonstrated that the 2 groves selected for detailed experiments presented an extreme
contrast in the relative abundance of insect predators on the soil surface. The striped
earwig was the predominant predator (more than 97%) in Winn grove in 1980 (Tables
1 and 2). Few ants (0.2%) were observed or collected in this grove. The striped earwig
was the predominant arthropod (avg 98%) in only 4 of the 78 groves sampled for surface
predators. High Acres had an ant population, which was more typical of the 74 groves
with a large ant population and no L. riparia (Table 3). Whitcomb et al. (1982), who
surveyed High Acres, reported 30 ant species with 13 species predominating. High
Acres was typical of 95% (74 of 78) of the groves sampled. The total number of ants
ranged from 92.4% to 100% of the total predators (avg. 98.7%) in these groves. Whit-
comb et al. (1982) also reported that of 2,613 first-instar D. abbreviatus larvae collected
in High Acres, more than 75% were taken by 3 ant species Pheidole dentata Mayr
(31.1%), P. floridana Emery (24.0%), and Tetramorium simillimum Roger (20.8%).
The striped earwig comprised less than 3% (avg 0.9%) of the arthropod predators in
any one grove. Richman et al. (1986) also demonstrated that the ants collected in these
groves were the most important predator of the sugarcane rootstock borer neonate
Baited pitfall traps in Winn grove between 2100 and 2300 h (June 1980) attracted
significantly (P < 0.01) more striped earwigs than unbaited traps (49.7:18.4 striped
earwig/trap/2 h). The most effective baits were moist cat food with a strong odor (Fris-
kies' tuna and liver, etc.)


Tryon: Predators of Diaprepes



June 8-14, 1979 June 12-18, 1980
Insects Winn HA Winn HA

Labiduria riparia 370.0 0.0 264.8 0.0
Euborella annulipes
(Lucas) 13.8 6.8 9.1 6.4
Ants 1.4 81.2 0.0 74.7
Others (Hemiptera, Coleoptera) 0.7 13.7 0.0 9.8

RUS GROVE, 1980.

Predatory Insects (No./trap)
Labidura Euborellia Ants3
riparia annulipes
Sampling Dates (Pallas) (Lucas)

January 10-16 11.3 0.4 0.0
February 14-20 2.7 0.3 0.0
March 9-15 3.6 0.7 0.0
April 10-16 48.81 4.8 0.3
May 9-15 152.7 4.1 0.1
June 12-18 264.8 9.1 0.0
July 20-26 103.92 12.8 1.4
August 14-20 241.01 8.4 0.3
September 9-15 79.4 2.8 0.1
October 11-17 97.6 3.4 0.0
November 13-19 40.3 1.8 0.0
December 14-20 29./9 2.5 0.0

'A large percentage (50%) of the L. ripaira trapped were immature.
2Consist of adult females observed in nests and not scavenging for food.
3The 2.2 ants collected per trap consisted of 0.9 Pseudomyrmex elongata, 0.7 P.
brunnea, 0.4 Pheidole moerens, and 0.2 Camponotus impressus.


To help explain the two contrasting insect predator populations, weed species and
density in Winn and High Acres groves were assessed. Regular cultivation and an
annual use of herbicide kept Winn grove much freer of weeds than the seldom-cultivated
High Acres grove. In monocultures, population levels of native predators, e.g. ants and
earwigs, are directly influenced by weed complexes (Alteiri & Whitcomb 1979). Abun-
dant species exclusive to each grove were hairy signalgrass (Brachiaria piligera (F.
Meull.) and guineagrass (Panicum maxumum Jacq.) in the Winn grove, and crabgrass

Florida Entomologist 69(2)

June, 1986


Insects collected (No./trap)
9-15 12-18 20-26 14-20
Insect Species May June July August

Pheidole floridana
Emery 8.3 6.5 21.2 32.4
Pheidole dentata Mayr 14.9 13.7 28.4 15.1
Tetramorium similimum
Roger 15.8 17.6 13.7 19.6
Conomyra edeni Buren 9.2 4.9 2.8 4.0
Pheidole moerens Wheeler 1.6 3.7 8.7 3.7
Solenopsis invicta Buren 3.7 1.8 13.0 4.1
Paratrechina bourboniza
(Florel) 0.2 1.1 1.6 3.2
Others 0.1 5.9 2.3 7.0
Total 62.8 55.2 91.7 89.1
Labidura riparia (Pallas) 0.0 0.0 0.0 0.0
Euborellia annulipes (Lucas) 4.6 5.3 8.9 6.8
Total 4.6 6.3 8.9 6.8
Coleoptera 7.1 6.7 3.8 5.2
Hemiptera 0.9 2.4 4.9 3.3

Total (insects/trap) 75.4 69.6 109.3 104.4

'Identified with the help of
Gainesville, Florida.

Dr. Everett Nikerson, Division of Plant Industry,

(Digitaria ciliaris (Retz.) Koel.), southern sida (Sida acuta Burn. f.), two pigweeds
(Amaranthus viridus L. and A. hybridus), and Bidens alba (L.) in High Acres.
Amaranthus sp. has served as a reservoir for numerous phytophagous insects and
associated predators, including a ground beetle (Lebia analis Dej.) and several ant
species (Alteiri & Whitcomb 1979). Ants trapped on Amaranthus sp. in High Acres
included Conomyrma sp., Solenopsis invicta Buren, and several Pheidole sp., Bidens
alba (L.), also found in High Acres, always had mealybug populations, Planococcus
ficus (Signoret), at the soil root interface that were tended by the imported fire ant (S.
invita). Thus, Amaranthus sp. and Bidens alba, abundant only in High Acres, were
beneficial in maintaining ant populations.


Surface soil from Winn and High Acres groves was analyzed for the presence of all
soil insecticide residues by the Pesticide Research Laboratory, Institute of Food and
Resources, University of Florida. Trace amounts (0.003 ppm) of DDD and DDE were
found in soil from both High Acres and Winn groves. Soil samples from Winn grove
also had residues from dieldrin (avg. 0.0009), chlordane (avg 0.100), heptachlor (avg
0.093), endrin (avg 0.039) and two unidentifiable cyclodiene insecticides (ppm between
0.002 and 0.115).


Tryon: Predators of Diaprepes

The extensive use of the cyclodiene chlorinated hydrocarbon insecticides and the
continued existence of their residues help explain the eradication of soil-surface ant
populations and the resulting buildup of striped earwig populations at Winn grove.
Stoddard (1962), Workman (1963), and Gross & Spink (1969) reported that the striped
earwig may be resistant to the chlorinated hydrocarbons heptachlor and chlordane.
Clements (1968) found chlordane did not control the striped earwig. Striped earwig
populations were maintained at low density levels by ant predation (Stoddard 1962,
Gross & Spink 1969, and Price & Shepard 1977). Observations of earwig egg predation
by ants were supported by laboratory studies by Watts and Whitcomb (unpublished).
They found several ant species (Pheidole dentata Meyr, Solenopsis invicta Buren, and
Tetramorium bicarinatum (Nylander)) each removed 100% of eggs from L. riparia egg
masses (avg 36.4 eggs/batch) in less than 60 minutes. Dense striped-earwig populations
increased in lawns and fields only after ant populations had been eliminated by the use
of heptachlor (Stoddard 1962, Gross & Spink 1969).


Results from observations of predation by the striped earwig on first-instar D.
abbreviatus weevil larvae are presented in Figure 1. Earwigs fed on all or none of 20
weevil larvae per trial. One earwig would consume a dish of 20 weevil larvae in less
than one minute and only one adult earwig was found inside a petri dish at one time.
In an encounter between two earwigs, the smaller earwig usually left instantly or risked
being cannibalized. The earwigs remained motionless or hid when white lights were
used. This required the use of a flashlight with a dim red light to avoid this interruption
during night observations. Once feeding was initiated, however, light, regardless of
color, no longer inhibited their activity.

100 r ,

SS= Sunset
S 80 o SR= Sunrise

So 60

o 640 I

20 SR-


1800 h 2100 h 2400 h 300 h 600 h 900 h

Figure 1. Percentage of D. abbreviatus neonate larvae consumed by striped earwig
in open petri dishes in Winn citrus grove (12 observation days, 4 trial/h/day, 20 larvae/

342 Florida Entomologist 69(2) June, 1986

Observations of the striped earwig activity confirmed their nocturnal habits. Peak
foraging and feeding activity occurred during the first 3 hours after sunset and gradually
decreased throughout the night and ceased before sunrise. During the peak feeding
period, more than 75% of all weevil larvae placed under the citrus canopy during the
summer and fall of 1980 were consumed. The percentage eaten was calculated from the
number of weevil larvae taken from an open petri dish during 20-minute intervals (Fig.
1). The amount of the first-instar weevil larvae consumed per night during this 3 hour
time period ranged from 100% (June 16) to 64% (July 21). Feeding tests were conducted
only from May to November when the weevil neonate larvae were available. Only in
late November did earwig feeding activity decrease to less than 30%. This reflected the
decrease in L. riparia activity in late fall (Table 2).
During daylight hours, striped earwigs were quiescent and hid in shaded resting
sites protected by several layers of soil debris. Earwigs were rarely observed foraging
during the day (0700-1900 h). However, at dusk (2000-2100 h) feeding activity of the
striped earwig began. During the first few hours (2100-2400 h) they were most effective
at removing the introduced D. abbreviatus larvae from the petri dishes. This was
explained by the aggressive foraging and their concentration under the citrus tree
canopies in the early evening. Feeding continued during most of the dark hours (2100-
0700) but gradually became less effective after midnight as they dispersed and became
less active. Before sunrise, they disappeared into protected resting sites.
Periodic decreases in feeding activity by the striped earwig during summer (i.e.
64%, July 21) parallel a lack of feeding activity by several adult females observed in
shallow burrows near the feeding trials (Table 2). These females remained in their
burrows protecting their brood for several days despite feeding by other striped ear-
wigs. Even minor disturbances with various pointed devices failed to dislodge the
female. Caussanel (1968), Tawfik et al. (1972), and Shepard et al. (1973), reported that
females did not feed or leave their burrows while brooding eggs for up to 12 days.
Ants and the striped earwig are the predominant arthropod soil surface predators
in central Florida citrus groves. However, they may be mutually exclusive. Ants were
the predominant soil surface predator in the typical grove. Ant populations or the
striped earwig may play a major role in reducing larval populations. Chlorinated hydro-
carbon soil insecticide residues remain for years and effectively eliminate many soil-sur-
face predators including Formicidae species. Groves without ant populations were
characterized by nocturnal predation by dense populations of the striped earwigs. This
unique condition was observed in 4 of 78 surveyed citrus groves in central Florida.
Without the various ant species common to the citrus grove soil surface, large popula-
tions of striped earwigs become the dominant arthropod predator.


This is Florida Agricultural Experiments Station Journal Series No. 3402.


ALTEIRI, M. A., AND W. H. WHITCOMB. 1979. Manipulation of insect population
through seasonal disturbance of weed communities. Protection Ecology. 1: 185-
NGUYEN, N. C. LEPPLA, AND B. J. SMITTLE. 1977. Predators of velvetbean
caterpillar eggs in Florida soybeans. Environ. Entomol. 6: 403.

Tryon: Predators of Diaprepes 343

CAUSSANEL, C. 1968. Factors conditionant 7 maintien des soins aux feufs chez Labi-
dura riparia (Dermaptera: Labiduridae). Int. Cong. Entomol. Moscow. 13(1):
CLEMENTS, R. H. 1968. Important earwigs, Dermaptera, of central and south Florida
and the biology and control of the primary species, Labidura riparia (Pallas)
under laboratory conditions. MS thesis, Univ. of Florida. 87 pp.
GROSS, H. R., AND W. T. SPINK. 1969. Response of striped earwigs following applica-
tion of heptachlor and mirex, and predation-prey relationship between imported
fire ants and striped earwigs. J. Econ. Entomol. 62(3): 686-689.
JNOES, I. F., AND W. J. SCHROEDER. 1983. Study of first-instar Diaprepes ab-
breviatus (Coleoptera: Curculionidae) activity for control purposes. J. Econ. En-
tomol. 76(3): 567-569.
NEAL, T. M. 1974. Predaceous arthropods in the Florida soybean agroecosystem. MS
thesis, Univ. of Florida. 196 pp.
NGUYEN, RU, AND R. B. WORKMAN. 1975. Biotic agents limiting the cabbage looper
in northeast Florida. J. Georgia Entomol. Soc. 13(2): 152-155.
PRICE, J. F., AND MERLE SHEPARD. 1977. Striped earwig. Labidura riparia coloniza-
tion of soybean fields and response to insecticides. Environ. Entomol. 6(5): 679-
REINERT, J. A. 1978. Natural enemy complex of the southern chinch bug in Florida.
Ann. Entomol. Soc. of America 71(5): 728-731.
RICHMAN, D. B., W. H. WHITMAN, AND W. F. BUREN. 1985. Predation on neonate
larvae of Disprepes abbreviatus (Coleoptera: Curculionidae) in Florida and
Puerto Rico citrus groves. Florida Entomol. (in press).
SCHLINGER, E. I., R. VAN DEN BOSCH, AND E. J. DIETRICK. 1959. Biological notes
on the predaceous earwig Labiduria riparia (Pallas), a recent immigrant to
California (Dermaptera: Labiduridae), J. Econ. Entomol. 52(2): 247-294.
SHEPARD, M., V. WADDILL, AND W. KLOFT. 1973. Biology of the predaceous earwig,
Labidura riparia (Dermaptera: Labiduridae). Ann. Entomol. Soc. America 66(4):
STODDARD, H. J. SR. 1962. Bird casualties at a Leon County, Florida TV Tower,
1955-61. Bull. 1, Tall Timbers Research Station. Tallahassee, Fla. 36-41.
of Labiduria riparia (Pallas). Bull. Soc. Entomol. 56: 75-92.
TRAVIS, P. A. 1977. Population dynamics of Labidura riparia (Pallas) (Dermaptera:
Labiduridae). MS thesis, Univ. of Florida. 85 pp.
WALKER, J. T., AND G. G. NEWMAN. 1976. Seasonal abundance, diet periodicity and
habitat preference of the striped earwig Labidura riparia in the coastal plain of
South Carolina. Ann. Entomol. Soc. American 69(4): 571-574.
WHITCOMB, W. H. 1973. Natural populations of entomophagous arthropods and their
effect on the agrocosystem. Proc. Miss. Symp. Biol. Control. Univ. of Mississippi
pp. 150-169.
WHITCOMB, W. H., T. D. GOWAN, AND W. F. BUREN. 1982. Predators of Diaprepes
abbreviatus (Coleoptera: Curculionidae) larvae. Florida Entomol. 65: 150-158.
WOODRUFF, R. E. 1964. A Puerto Rican weevil new to the United States (Coleoptera:
Curculionidae). Florida Dept. of Agric., Div. Plant Ind., Ent. Cir. 30, 1-2.
WORKMAN, R. B. 1963. Laboratory test for control of earwigs. Florida Entomol. 46:

Florida Entomologist 69(2)

June, 1986


University of Florida, IFAS
Tropical Research and Education Center
18905 S.W. 280 St.
Homestead, FL 33031


A scale for assessing damage to papaya fruit by the papaya fruit fly Toxotrypana
curvicauda Gerstaecker was used in the field to help detect egg and larval infestation.
There were more eggs in green fruits than in ripe fruit, whereas more larvae were
found in ripe fruit than in green fruit. Under laboratory conditions, papaya fruit fly
preferred to oviposit in green papayas than in ripe fruit. In an experiment of the choice
type, papaya fruit flies showed strong preference for oviposition in green wax domes
than in yellow wax domes. The color preference of papaya fruit fly females was con-


Se utilize en el campo una escala para evaluar el dafo a la papaya por la mosca de
la papaya, Toxotrypana curvicauda Gerstaecker para ayudar a detectar infestaciones
de huevos y de larvas. Se encontr6 una mayor cantidad de huevos en los frutos verdes
que en los frutos maduros, mientras que mas larvas se encontraron en frutos maduros
que en frutos verdes. Bajo condiciones de laboratorio, moscas de la papaya prefirieron
poner los huevos en papayas verdes que en frutos maduros. En un experiment donde
las moscas de la papaya podian escoger, ellas demostraron preferencia para poner los
huevos en cuipulas de cera verde que en cipulas de cera amarilla. Se confirm el color
preferido de la mosca de la papaya.

The papaya fruit fly (PFF) Toxotrypana curvicauda Gerstaecker almost exclusively
uses the papaya Carica papaya L. (Knab & Yothers 1914) and occasionally fruit of
mango Mangifera indica L. (Butcher 1952) as hosts. Toxotrypana occurs only in tropical
America (Wolcott 1933) and in south Florida (Knab & Yothers 1914). Some cosmopolitan
fruit flies, e.g., Dacus dorsalis Hendel, D. cucurbitaceae Coquillet and Ceratitis
capitata (Weideman) also attack papaya in other tropical areas in the world (Seo et al.
1983) but generally they prefer ripe papayas.
Premature fruit maturity and fruit latex exudations have both been related to
papaya fruit fly infestation (Knab & Yothers 1914). These plant reactions, however,
could have been caused by other insects (Sloan 1946) or by pesticides (Sherman &
Tanashiro 1959).
Stimuli eliciting the oviposital response of PFF to its host are unknown. The PFF
has been observed to oviposit in blossom buds and in a wide range of papaya fruits, 1.5
cm diam green fruit to 15 cm-diameter fruit that is almost completely ripe (Landolt &
Hendrichs 1983) and undergoing a change in color from green to yellow. Factors such
as fruit ripeness and color may play a role in the selection of fruits for oviposition.


Pena et al.: Toxotrypana Oviposition


In this study 3 aspects of the relationship between T. curvicauda and the papaya
fruit have been considered:
(1) Are effects of PFF oviposition or larval activity precise measures of PFF popu-
(2) To what extent do females discriminate between fruits with different degrees of
(3) Does color affect oviposition?



Visual observations of the possible effects of PFF infestations were made on 70
fruits collected from an insecticide-free commercial papaya planting (var. Cariflora) at
Homestead, Florida, during August 1984. A visual classification of damaged and undam-
aged fruits was made. A grade scale of 0-6 was devised, based on personal observations
and damage description by Mason (1922) and Knab and Yothers (1914) (Table 1). A RxC
test of independence (Sokal & Rohlf 1969) was used to examine whether the percentages
of fruits with eggs and larvae were independent of the papaya grade scale.


Papaya fruit flies used in color preference tests were obtained from a colony main-
tained in the laboratory at ca. 27C under fluorescent lights that provided a photoperiod
of LD 12:12. The colony was initiated with larvae that were collected in the field and
which pupated in vermiculite. Adults were kept in screen cages (16.5 x 33 x 37 cm)
and were provided with water (65 ml), sucrose (5.8 g) and honey. The effect of the
degree of ripeness of papaya fruit on PFF ovipositional preference was studied. The
ripeness of immature green, quarter-color, half-color, and ripe papayas was determined
visually using the Nickerson Color Fan@. Fifteen females were exposed in each cage


Scale Fruit description

0 = Immature, green with no external symptoms of latex flow.
1 = Immature, green, and with fresh latex flowing from the
fruits, indicative of recent PFF oviposition.
2 = Immature, green, and with dark-coagulated latex on
the fruit, indicative of past PFF oviposition.
3 = Fruit showing premature 1/4 color break, but firmly
attached to the peduncle.
4 = Fruit showing premature 1/4 color break, easily detached
from peduncle, indicative of larval infestation.
5 = Fruit showing 1/2 color break, firmly attached to
the peduncle.
6 = Fruit showing 1/2 color break, soft to the touch and
with larval exit holes, indicative of present
or past larval infestation.

346 Florida Entomologist 69(2) June, 1986

(= one replicate) to the 4 different types of papaya fruits. This multiple choice experi-
ment was replicated 8 times and conducted inside screen cages (16.5 x 22 x 37 cm).
The number of perforations and eggs were recorded after 24 h. Data were analyzed by
ANOVA and the means were separated by Duncan's Multiple Range Test. Later, an
experiment of the multiple-choice type to determine PFF color preference was con-
ducted inside screen cages. Oviposition substrates consisted of wax domes 7 cm in
diameter and 3.5 cm high. To obtain domes of different hues that represented the colors
of papaya fruit, green and yellow paraffin wax and petroleum jelly were mixed. Reflec-
tance spectra of green and yellow colors were measured using the Nickerson Color
Fan. The color values were dark yellowish green = 10 GY 4/5, strong yellow green
= 2.5 GY 6/8, strong greenish yellow = 7.5 Y 7/9, and strong orange yellow = 10 YR
7/10. Ten 6 day old females per cage (= one replicate) were tested. The experiment
was replicated 6 times. The main criterion used for determining oviposition preference
was the number of eggs that were laid in the dome. Counts of eggs were made daily
and the positions of the domes in the cages were re-randomized daily. For the analysis
of egg count data, analysis of variance was used and the means were separated by
Duncan's Multiple Range Test at the P = 0.05 significance level.



The grade-scale study provided information on PFF oviposition and larval infestation
of papaya fruit. Frequency of eggs and larvae in papaya fruit (Table 2) were dependent
(P < 0.05) on the fruit grade scale. More eggs (Mean SE = 2.70 1.87 and 2.04 +
1.05; X2 .005 [6] = 18.548) were obtained from fruit grades 1 and 2 than from any other
types of fruit (0.0 + 0.0). More larvae (P < 0.05; X2 .005, [6] = 18.548 were obtained
from 3, 4, 6, 2 and 1 grade-type fruit than from 0 or 5th grade. By using this grade
scale, it was possible to partially determine egg and larval infestation in the field. The
grade scale could be developed into a tool to help scouts select fruits with the best
chance of containing PFF eggs or larvae.


The results of this multiple choice test (Table 3) demonstrated that females re-
sponded positively (P = 0.05) to a certain degree of fruit development, especially to


% fruit % fruits
Grade scale Mean eggs SEa with eggs Mean larvae + SEb with larvae

0 0.0 +0.0 0 0.0 0.0 0
1 2.701.87 29 0.72+0.41 21
2 2.041.05 18 4.801.29 52
3 0.0 0.0 0 8.75+1.21 100
4 0.0 0.0 0 14.881.05 100
5 0.0 0.0 0 0.0 0.0 0
6 0.0 +0.0 0 5.622.33 62

aX .005 6 = 18.548; g-value (X2 = 22.25) was significant at P < 0.005.
bX2 .00t, 6 = 19.548; g-value (X2 43.43) was significant at P < 0.005.

Pena et al.: Toxotrypana Oviposition


CONDITIONS (27+1C; 75% R.H.).

Fruit No. of perforations/ Mean no. % total
characteristic fruit eggs/fruit eggs

Immature green 10.5 a 55.0 a 68
Quarter color 3.3 b 3.81 b 5
Half color 5.4 b 9.45 b 12
Ripe 2.3 b 12.45 b 15

aNumbers followed by a different letter were significantly different at the P = 0.05


Color of dome Avg. no. of eggsa % of total

dark green 15 a 47
yellow green 6 b 18
greenish yellow 4 b 13
orange yellow 7 b 22

aNumbers followed by a different letter were significantly different at the P = 0.05

immature dark green, and less to quarter color and ripe fruit. This behavior indicates
that physical characteristics such as fruit color may be important for the selection of
the oviposition site. Because changes in the chemical properties of ripening papaya fruit
may also have an equally important effect on ovipositional behavior, the response of
PFF females to color alone was determined.
The results (Table 4) demonstrated a strong color preference of the papaya fruit fly.
About 46% of the eggs were laid in the dark green domes, followed by yellow green
domes (18%) and orange-yellow domes (22%), and less for greenish yellow domes (13%).
For example, the mean number of eggs laid in dark green domes was 15 (P = 0.05)
compared to mean numbers of 6, 4 and 7 eggs laid in the other domes. The strong
preference for the dark green domes indicated that this color constitutes a strong
stimulus to the fruit-seeking papaya fruit fly females.


This research was partially supported by USDA Cooperative Agreement No. 5B-
7B30-9-116. Florida Agricultural Experiment Stations Journal Series No. 6537.


BUTCHER, F. G. 1952. The occurrence of papaya fruit fly in mango. Proc. Florida State
Hort. Soc. 65: 196.

348 Florida Entomologist 69(2) June, 1986

KNAB, F., AND W. W. YOTHERS. 1914. Papaya fruit fly. J. Agric. Res. 2: 447-453.
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.
MASON, A. C. 1922. Biology of the papaya fruit fly Toxotrypana curvicauda in Florida.
U.S. Dept. Agric. Bull. 1081-1-10.
SEO, S. T., C TANG, S. SANIDAD, AND T. TAKENABA. 1983 Hawaiian fruit flies.
(Diptera: Tephritidae): variation of index of infestation with benzyl isothiocyanate
concentration and color of maturing papaya. J. Econ. Entomol. 76: 535-538.
SHERMAN, M., M. TAMASHISO. 1959. Toxicity of insecticides and acaricides to the
papaya, Carica papaya L. Hawaii Agric. Exp. Station, U. of Hawaii. Tech. Bull.
40: 56 p.
SLOAN, W. J. 1946. The fruit spotting bug. Queensland Agric. J. 1: 229-233.
SOKAL, R., AND F. ROHLF. 1969. Biometry: The principles and practice of statistics
in biological research. W. H. Freeman and Co., San Francisco. 776 p.
WOLCOTT, G. N. 1933. Economic entomology of the West Indies. Clay and Sons, Ltd.,
Bengay, Suffolk. 688 p.


Bishop Museum
Honolulu, Hawaii 96817


In and around the coconut plantations in Jamaica, 107 species of thrips were collected
or caught by sticky traps. Prior to this collection, thrips recorded from the island were
highly limited. Species repeatedly collected from coconuts were Franklinella bruneri
Watson, F. kelliae Sakimura, and F. insularis (Franklin) (flower feeders); Anisopilo-
thrips venustulus (Priesner) and Heliothrips haemorrhoidalis (Bouche) (fruit feeders);
Hoplandrothrips flavipes Bagnall (fungal growth feeder); and Karnyothrips merrilli
(Watson) (predator). Of every species enumerated, their extra distributions within the
Caribbean areas were searched through the published data as well as accumulated
holdings in museum collections. Twenty-two species of Frankliniella were collected,
and a key to them is provided. Two new synonymies are designated, and a species is
partly redescribed.


Dentro y alrededor de plantaciones de cocos en Jamaica, 107 species de "thrips"
fueron colectadas o atrapadas en trampas pegajosas. Anteriormente a esta colecci6n, el
registro de "thrips" de la isla era muy limitado. Especies repetidamente colectadas en
cocos eran Frankliniella bruneri Watson, F. kellia Sakimura, y F. insularis (come-
dores de flores); Anisopilothrips venustulus (Priesner) y Heliothrips haemorrhoidalis
(Bouche) (comedores de frutas); Hoplandrothrips flavipis Bagnall (comedores de hon-
gos); y Karnyothrip merrilli (Watson) (depredador). De cada especie enumerada, su
distribuci6n extra dentro del area del Caribe fue indagada a traves de datos publicados,

Sakimura: Jamaican Thrips


lo mismo que en materials acumulados en colecciones de los museos. Se colectaron y
se provee una clave para 22 species de Frankiniella. Se designaron 2 nuevos sin6nimos
y una especie es parcialmente redescrita.

A thrips survey in and around the coconut plantations in Jamaica was conducted in
the whole month of November 1964, upon request of a FAO project on searches among
thrips for possible vector species of the lethal yellowing disease of coconut. About 15
years subsequent to that date, the disease was conclusively demonstrated to be trans-
mitted by Myndus crudus Van Duzee (Homoptera: Cixiidae) (Howard et al. 1983).
Thrips specimens were collected from weedy undergrowth, coconuts, and other trees
and shrubs intermixed within the coconut plantations as well as in the surrounding
areas. Thrips specimens caught by sticky traps, hung among coconut foliage in every
plantation visited were examined also. The plantations visited were aligned along the
north shore of the island from Buff Bay to Negril (Table 1). Supplemental but extensive
general collections were made also in the vicinity of Kingston on the south shore (Table
1). The few specimens previously collected by T. H. Farr, Institute of Jamaica were
examined also. Total numbers of specimen lots were 430 (Acc. No. 4208-48, 4301-4579
and other long non-numbered series), and the number of species recognized was 107.
Names of host plants were provided by the Botany Department, University of the West
The Caribbean fauna of Thysanoptera has long been poorly known, and the only
faunistic work published is the one of Puerto Rico (Medina Gaud 1961) enumerating 78
species. However, over the years since about 1910 many new species have been de-
scribed from the Caribbean areas, and numerous specimens from various islands and
shore countries have been accumulated in some museum collections, particularly at the
United Stated National Museum of Natural History (USNMNH).
This survey fairly well covers the lowland species but is deficient for the highland
indigenous species, since the few cursory collections made at Hardwar Gap (4000') were
all that had been done over the extensive forested highland. The number of species
previously recorded from Jamaica was 16 prior to this survey made in 1964, but it
increased to 28 by the time of this writing in 1985. Among those previously recorded,
the following 6 species, 3 of them being highland species, were not collected during this
survey: Arachisothrips mills Stannard, Kurtomathrips morrelli Moulton, Eurythrips


Coconut Plantation Areas (on north shore):
Buff Bay Area (annual rainfalls: about 100"; 22" during Oct.-Nov. 1964 at Buff Bay):
Woodstock, Kildare, White River, Buff Bay River Estate, Orange Vale, Buff Bay,
Silverstock, Benzenn, Dover, and Gray's Inn.
Montego Bay Area (annual rainfalls: about 50 "; 9" during Oct.-Nov. 1964 at Montego
Bengal, Bryan Castle, Hague, Braco, and Falmouth (east of Montego Bay).
Barnett, Montego Bay, and Irwin (vicinity of Montego Bay).
Round Hill, Hill Top, and Negril (west of Montego Bay).
Non-coconut Plantation Areas (On south shore):
Kingston and Vicinity (annual rainfalls: about 50", higher on highland and lower on
coastal strip; 7" during Oct.-Nov. 1964 at Kingston):
Kingston, Hope Garden, Palisadoes, Hardwar Gap, Bodles, and Barnard Lodge.

350 Florida Entomologist 69(2) June, 1986

batesi (Watson), Orthothrips bilineatus Stannard, Plectothrips pallipes Hood, and
Trisclerothrips hurricanes Stannard. In the following enumeration of the species, 20
species are incomplete in determination to the species level. Many of them are believed
to be undescribed species. All such material will be available for any specialist who may
be interested in it. A set of all the species will be deposited in the USNMNH collection
and others.
For every species to be enumerated later, 1) brief taxonomical remarks, if any; 2)
abbreviated remarks on host range and population status; and 3) extra distribution data
within the Caribbean area that are all based on the published data and accumulated
holdings in museum collections are given. The details on locality, date, host, numerical
data on collected specimens, accession number, and the collector's name are all omitted,
except in a few cases where such complete data are considered significant for future
references. Most of the collections were made by the author, except all the sticky trap
material that was collected by the late Walter Carter or Randall Latta of the FAO
project. Numbers in parentheses in the enumeration of species are the author's acces-
sion numbers.


Ceratothripoides funestus (Hood, 1915:24) (= Taeniothrips martorellorum Medina,
1961:48), Syn. N. See p. 351.
Holopothrips fulvus Morgan, 1929:6 (= Holopothrips anacardii Hood, 1942:581),
Syn. N. See p. 357.



Family Aeolothripidae
Franklinothrips vespiformis (Crawford, 1909): Frequent on various plants
everywhere, particularly in the wet Buff Bay area; predacious on other thrips. Known
from Brazil (Para), Surinam, Trinidad, Grenada, Barbados, St. Vincent, Martinique,
Dominica, Guadeloupe, Antigua, St. Thomas, Puerto Rico, Dominican Republic, Cuba,
Jamaica, Florida, Texas, Mexico, Guatemala, Honduras, Nicaragua, Panama, Colombia,
and Venezuela. F. tenuicornis Hood was not collected during this survey.

Family Heterothripidae
Heterothrips borinquen Hood, 1915:1 Irwin, Eupatorium villosum (4429). Known
only from Puerto Rico.

Family Thripidae
Anisopilothrips venustulus (Priesner, 1923): Frequent on Cyathula prostrata and
young coconut fruits (found breeding there) in the wet Buff Bay area. Known from
Surinam, Trinidad, Grenada, Guadeloupe, Puerto Rico, Dominican Republic, Bermuda,
Jamaica, and Florida.
Aptinothripine sp. (C): Differs from Baileyothrips by head and pronotum striate and
targal seta pair i far apart from each other. 79 9, Barnett, Desmodium sp. (4439a).
Aptinothripine sp. (D): Differs from Chaetanaphothrips by antennal I-III uniquely
modified, mouth cone pointed, and tergite VIII without stippled areas. 16, Round Hill
(Hill Top), Mimosa pudica (4484c).
Baileyothrips limbatus (Hood, 1935): 19, Woodstock, Desmodium sp. (4335b).
Known from Panama. B. arizonensis is not synonymous.

Sakimura: Jamaican Thrips

Bregmatothrips gracilis Hood and Williams, 1915: 1 Woodstock, Commelina dif-
fusa (4306b). Known from Trinidad, Patos Isle, Cuba, Jamaica, Florida, Georgia, and
Caliothrips insularis (Hood, 1927): Abundant and frequent on Seteria palmifolia.
Known from Trinidad, Tobago, Grenada, Martinique, St. Croix, Puerto Rico, Cuba,
Jamaica, Mexico, Panama, and Venezuela.
Caliothrips phaseoli (Hood, 1912): Abundant or frequent on Colocasia esculenta and
Ricinus communis, and occasionally on some leguminous plants; caught by sticky traps.
Known from Brazil (Para), Tobago, Dominica, Puerto Rico, Cuba, Florida, Texas,
Mexico, Panama, and Colombia.
Ceratothripoides funestus (Hood, 1915) (= Taeniothrips martorellorum Medina,
1961, New Synonymy): The type was examined. Abundant on Spigelia anthelmia, and
occasionally on some other plants in the wet Buff Bay area; caught by sticky traps.
Known from Trinidad, Puerto Rico, Texas, and Mexico (Tobasco).
Chaetanaphothrips orchidii (Moulton, 1907): Frequent on Cyathula prostrata; once
from young coconut fruit. Known from Trinidad, Guadeloupe, Puerto Rico, Florida,
Louisiana, Mexico, Honduras, Costa Rica, and Panama.
Chaetisothrips striatus caribeanus Sakimura, 1969: 29 Montego Bay and Negril;
Porana paniculata, sticky trap (4498a, 4500b). Known from Puerto Rico, Cuba, Mexico,
and Honduras.
Chirothrips mexicanus Crawford, 1909: From Eleusine indica; caught by sticky
traps. Known from Martinique, Antigua, St. Thomas, Puerto Rico, Dominican Republic,
Cuba, Grand Cayman, Florida, Mississippi, Texas, Mexico, Costa Rica, Panama, and
Chirothrips texanus Andre, 1939: From several grasses, fairly frequent in the drier
areas; caught by sticky traps. Known from Georgia, Louisiana, Texas, and Mexico.
Apparently the first collection from any of the Caribbean islands.
Corynothrips stenopterus Williams, 1913: 19, Woodstock, sticky trap (4573A).
Known from Brazil (Para), Surinam, Guyana, Trinidad, Tobago, Grenada, St. Vincent,
Barbados, Puerto Rico, Costa Rica, and Panama.
Dendrothripoides innoxius (Karny, 1914) (= D. ipomeae Bagnall): From Ipomoea
fistulosa and a few other plants. Known from Trinidad, Grenada, Barbados,
Guadeloupe, St. Croix, Dominican Republic, and Panama.
Dinurothrips hooker Hood, 1913: Abundant on Leonotis nepetifolia, and also from
Cyathula prostrata in the wet Buff Bay area. Known from Trinidad, Grenada, St.
Lucia, Guadeloupe, Puerto Rico, Cuba, Jamaica, Florida, and Panama.
Echinothrips caribeanus Hood, 1955: Abundant or frequent on Psophocarpus palus-
tris and Cyathula prostrata in the wet Buff Bay area. Known from Trinidad, St. Vin-
cent, Martinique, Guadeloupe, Puerto Rico, and Panama.
Frankliniella spp.: Speciation of the flower-feeding genus Frankliniella apparently
is centered in the Caribbean and Central American region from where more than 65
species have been described, and many more may be discovered in the future. In
Jamaica, 22 species were collected during this survey, and they are the most abundant
thrips there. Many species of this genus are poorly characterized and difficult to iden-
tify. The following key may serve for recognizing the Jamaican species. This key in-
eludes the 3 species that are to be described in another papaer now under preparation,
but not the 4 others that belong to the different subdivisions of the genus.
Abbreviations for the names of setae used in the following key: aa: Anteroangular
pronotal seta. am: Anteromarginal pronotal seta. io: Interocellar seta. IXi,ii, iii and Xi,
ii: Major setae on abdominal IX and X, numbered outwardly from the meson. IXd:

352 Florida Entomologist 69(2) June, 1986

Discal seta on abdominal IX. pai, ii: Posteroangular pronotal setae, numbered outwardly
from the meson. pmii: Posteromarginal pronotal seta, second from the meson. po: Prin-
cipal postocular seta, largest and usually 4th in the series from the meson, directed
posterolaterally. In comparing lengths of 2 setae, the minimal visually recognizable
differential usually is about 15% of the longer seta. In determining the slenderness of
antennal style, numerical index used is a value of the combined length of VII and VIII
including interspace that is divided by the basal width of VII. For details and explana-
tory illustrations, see Sakimura and O'Neill (1979).


1. Both io and po minute or nearly so and also both am and aa minute to
subminute or rarely somewhat larger (Minuta group). Antenna 7-seg-
mented, abdominal IX with all major setae shorter than X ......... jamaicensis
Sakimura & O'Neill
Those setae all developed or only po rarely subminute to small ............... (2)
2. Antennal III with pedicel simple and not dilated (Fig. 1) (Intonsa group) .. (3)
Antennal III with pedicel variously dilated (Fig. 2-6) (Tritici group) ......... (11)
3. Brown to dark brown body ................................................................. (4)
Pale to yellow body ........................................................................... (6)
4. Mid and hind tibiae dark brown; am decidedly smaller than aa. Abdominal
VIII with comb incomplete and short, X shorter than IX (0.8 times of
IX). Male colored same as female ................................. insularis (Franklin)
Mid and hind tibiae totally or partly yellow; am subequal to aa in length ... (5)
5. Mid and hind tibiae totally yellow; abdominal VIII with comb nearly
complete and very short, X as long as IX. Male yellow with brown head...
............................................................................ citripes H ood
Hind tibia with distal half dark grayish brown; abdominal VIII with comb
complete and moderately long, X longer than IX (1.2 times of IX). Male
uniformly yellow ........................................................... varipes M oulton
6. Yellow abdomen with grayish brown shadings and blotches along antecos-
tae; io between posterior ocelli; antennal VI with linear attachment bases
of both major sense cones. Pronotal am decidedly smaller than aa; abdom-
inal VIII with comb incomplete and short ........ schultzei (Trybom) (pale form)
Yellow abdomen without grayish brown shadings and blotches; io before
posterior ocelli; antennal VI with circular attachment bases of sense
cones. .......................................................... ................................ (7)
7. Cephalic po nearly subminute (less than 20 [m long). Antennal III with
a pair of ordinary dorsal setae; mouth cone long and slender; am smaller
than aa ............................................................................... exigua H ood
Cephalic po well developed (more than 30 im) ..................................... (8)
8. Antennal III with a pair of ordinary dorsal setae (about 0.6 of the seg-
ment length) and ordinary sense cone. Antennal style stout (VII + VIII/
VIIw: 2.6-2.9); head transverse (1.4 times as wide as long); abdominal
VIII with comb complete but short ................................... bruneri Watson
Antennal III with a pair of fully developed long dorsal setae (0.75-0.90
of the segment length) and unusually long sense cone ............................ (9)
9. Antennal style fairly stout (VII+VIII/VIIw: 3.3-3.6); head subtrans-
verse (1.3 times as wide as long). Mouth cone very bulky ................ Sp. F
Antennal style very slender (VII + VIII/VIIw: 4.4 4.5); head subquad-
rate (1.2 times as wide as long) .......................................................... (10)

Sakimura: Jamaican Thrips

\ -/ \ / \ / ... ./

I 2 3 4 5 6

Fig. 1-6. Pedicels of antennal III of various Frankliniella spp.: 1, Simple undilated
form (Minuta and Intonsa groups); 2-6, Dilated forms (Tritici group): 2, Mushroom form
(Tritici subgroup); 3, Angulate ring form (Cubensis subgroup); 4, Cup and saucer form
(Cephalica subgroup); 5-6, Miscellaneous forms (Miscellaneous subgroup): 5, F. bre-
vicaulis; 6, F. parvula.

10. Mouth cone moderately long, subvertically directed, strongly recurved,
bulky (often strongly twisted to a side in mounted specimens); abdominal
VIII with comb very long (up to 15 pim) ......................................... Sp. P
Mouth cone very long, subvertically directed, straight and tapered; ab-
dominal VIII with comb short (6-8 pm) ........................................... Sp. C
11. Antennal III with pedicel in mushroom form (Tritici subgroup)(Fig. 2) ...... (12)
Pedicel in angulate ring form (Cubensis subgroup)(Fig. 3) ..................... (15)
Pedicel in cup and saucer form (Cephalica subgroup)(Fig. 4) ................... (16)
Pedicel in specific unique forms (Miscellaneous subgroup)(Fig. 5-6) ............ (17)
12. Pronotal am decidedly smaller than aa. Abdominal VIII with comb incom-
plete and very short .......................................................................... (13)
Pronotal am subequal to aa. Yellow body without any brownish shadings,
head transverse (1.4 times as wide as long), abdominal VIII with comb
complete but very short .................................................................... (14)
13. Grayish brown body with subtransverse head (1.3 times as wide as
long); antennal III 2.4 times as long as wide ......................... salviae Moulton
Yellowish body with transverse head (1.4 times as wide as long); antennal
III 2.8-2.9 times as long as wide (Not yet collected in Jamaica) ..........
.................. ............................ tritici (Fitch)(pale form )
14. Ocellar crescent brownish. Male: abdominal IXd long thick seta, VIII
with no comb .................................................................. kelliae Sakimura
Ocellar crescent bright red. Male: abdominal IXd short thorn, VIII with
comb complete but short (Not yet collected in Jamaica). .......... difficilis Hood
15. Pronotal am decidedly smaller than aa; po small (about 18 jim); abdominal
VIII with comb nearly absent; yellow body with very weak grayish brown
shading throughout; antennal II dark to grayish brown ....... breviseta Moulton
Pronotal am subequal to aa; po well developed (about 31 jim); abdominal
VIII with comb complete and long; yellow body without any grayish
brown shading; antennal II pale to brownish yellow ................ cubensis Hood
16. Antennal II dorsoapically elevated strongly, and produced variably from
slight to as much as about a half of the ventral segment length, always
with a pair of strongly thickened blackish thorn-like setae at its apex.
Yellow body usually with grayish brown median blotches on abdomen .
................................................................... cephalica (Craw ford)
Antennal II dorsoapically elevated weakly but not produced at all, always
with a pair of ordinary thin setae at its apex (in male apical setae slightly

Florida Entomologist 69(2)

June, 1986

thicker than that of female, but far thinner than that of cephalica). Abdo-
men without any grayish brown blotches ............................. borinquen Hood
17. Antennal III with pedicel as illustrated (Fig. 5), III about 2.7 times as
long as wide; io outside of ocellar triangle. Brown body with dark brown
abdomen and all tibiae yellow ......................................... brevicaulis Hood
Antennal III with pedicel as illustrated (Fig. 6). III about 3.5 times as
long as wide; io inside of ocellar triangle. Body color as in brevicaulis;
male yellow with broad median grayish brown blotches on abdomen....
.............................................................................. parvula H ood
Frankliniella borinquen Hood, 1942: Abundant or frequent on Allamanda cathar-
tica, Hibiscus tiliaceus, and Poinciana pulcherrima; also on many other plants in the
drier surroundings, particularly in Kingston area; also rarely caught by sticky traps.
Known from Puerto Rico and Mexico.
Frankliniella brevicaulis Hood, 1937: 2 9 2, 16, Hope Garden, Bauhinia purpurea
(4410b). Known from Trinidad, Cuba, Panama, and Venezuela.
Frankliniella breviseta Moulton, 1948: From Viola sp. at Hardwar Gap (4000');
occasionally caught by sticky traps in the wet Buff Bay area. Known from Trinidad,
Martinique, Cuba, and Florida.
Frankliniella bruneri Watson, 1926: The most common and abundant Franklinielia
sp. throughout Jamaica when this survey was conducted, particularly in the wetter
surroundings. Frequently and abundantly caught by sticky traps, and frequent on
coconut inflorescence; abundant or frequent on Gliricidia sepium, Malvaviscus ar-
boreus, Taraxacum officinale, and Terminalis catappa; also on many other plants.
Known from Cuba, southern Texas, and throughout Mexico along the Gulf coast.
Frankliniella cephalica (Crawford, 1910): F. melanommata Williams is not synony-
mous. One of the common Frankliniella spp. in Jamaica. Abundant or frequent on
Agave sisalana, Bidens pilosa, and Tribulus cistoides, and also on many other plants;
often caught by sticky traps; more abundant in the wet Buff Bay area. Known from
Trinidad, St. Vincent, Barbados, St. Lucia, Dominica, St. Croix, St. Thomas, Puerto
Rico, Cuba, Florida, Alabama, Mississippi, Texas, Mexico, Panama and Colombia.
Frankliniella citripes Hood, 1916: 19, Bryan Castle, Eupatorium odoratum
(4449b); also caught by sticky traps in the wet Buff Bay area. Known from Puerto Rico
and Cuba.
Frankliniella cubensis Hood, 1925: Abundant on Gouania lupuloides, and also some
other plants in Kingston area; often caught by sticky traps in Buff Bay area. Known
from St. Croix, Puerto Rico, Haiti, Cuba, Florida, Mexico, Guatemala, and Panama.
Frankliniella sp. (C): To be described in another paper now under preparation.
From Desmodium sp. and a few others in both Buff Bay and Montego Bay areas
(4398Ad, 4348, 4388, 4433c, 4492).
Frankliniella exigua Hood, 1925: From Eupatorium odoratum and others all in the
dry Montego Bay area (4449d, 4454c, 4463b, 4487c). Known from Georgia, Florida,
Mississippi, Texas, and northeastern Mexico. First collection from any of the Caribbean
Frankliniella sp. (EEE): Differs in male form from F. citripes by body uniformly
grayish brown, mid and hind femora grayish brown, scale brown in basal portion, and
tergite IX with discal seta fully developed. Apparently similar males from Brazil (Para)
and Mexico remain unnamed in the USNM collection. 16, Hope Garden, Rosa sp.
(4417b). Excluded in the key.
Frankliniella sp. (F): To be described in another paper now under preparation.
Acalypha hispida, Hope Garden (4413).

Sakimura: Jamaican Thrips

Frankliniella sp. (G): Differs from F. terminalis and bicolor by stouter antenna,
pronotal chaetotaxy, and incomplete comb of tergite VIII. 19, Hope Garden, Lantana
camera (4414c). Excluded in the key.
Frankliniella insularis (Franklin, 1908) (= F. fortissima Priesner, 1925): Ex-
tremely variable in its body size, and also variable in color and size of antennal IV-V.
One of the common Frankliniella spp. in Jamaica. Abundant or frequent on Cajanus
cajan, Canavalia maritima, Gliricidia sepium, Hibiscus rosa-sinensis, and Malvavis-
cus arboreus; and also on many others everywhere; often and on occasions abundantly
caught by sticky traps, but only occasionally collected from coconut inflorescence in Buff
Bay area. Known throughout the Caribbean areas.
Frankliniella jamaicensis Sakimura and O'Neill, 1979: From Eupatorium villosum,
Round Hill (Hill Top) (4487d); once caught by sticky trap at Woodstock. Known also
from Cuba.
Frankliniella kelliae Sakimura, 1981: Had long been confused in Jamaica with F.
dificilis which was not collected during the present survey. One of the common
Frankliniella spp. there. Abundant or frequent on Bauhinia purpurea, Caesalpinia
coriaria, Calophyllum inophyllum, Cassia siamea, and Tithonia diversifolia; and also
many others; often and on occasions abundantly caught by sticky traps, and also often
collected among coconut inflorescence; more abundant in the dry Montego Bay and
Kingston areas. Known from Puerto Rico, Dominican Republic, Bahamas, Cuba,
Jamaica, and Florida.
Frankliniella sp. (0): Differs from F. deserticola by color of antennal II-V, ex-
tremely slender antennal III, and tergite X 0.65 times as long as IX. 1 Hardwar Gap,
4000', Viola sp. (4513a). This specimen is incomplete. Before being named, complete
specimens are needed. Excluded in the key.
Frankliniella parvula Hood, 1925: Apparently monophagous and abundant on Musa
spp. in dry as well as wet areas; also caught by sticky traps. Known from Trinidad,
Grenada, St. Lucia, Puerto Rico, Dominican Republic, Jamaica, Mexico, Guatemala,
Honduras, Costa Rica, Panama, and Colombia.
Frankliniella sp. (P): To be described in another paper now under preparation.
29 9, Orange Vale, Cassia occidentalis (4324a); also often caught by sticky traps in the
wet Buff Bay area.
Frankliniella salviae Moulton, 1948: 19, Woodstock, Synedrella nodiflora (4303a).
Known only from Mexico.
Frankliniella schultzei (Trybom, 1910) (pale form): From Cajanus cajan, Ipomoea
pes-caprae, Kallstroemia maxima, and Plumbago capensis in the dry Kingston area
only. This pantropic species was observed elsewhere also preferring dry surroundings.
Known from St. Thomas, Puerto Rico, Dominican Republic, Jamaica, Florida, and Col-
ombia. Only the Colombian specimens are dark form.
Frankliniella varipes Moulton, 1933: Abundant on Lantana camera (4374a, 4480a);
often caught by sticky traps in the wet Buff Bay area. Known from southern Brazil and
Peru. The first collection from the Caribbean area.
Frankliniella sp. (X): Differs from F. kelliae by anteromarginal pronotal seta larger
than anteroangular seta, and unusually large interocellar seta and posteromarginal pro-
notal seta ii. 1 Hope Garden, Bauhinia purpurea (4410f). Excluded in the key.
Heliothrips haemorrhoidalis (Bouch6, 1833): Frequent or on occasions abundant on
coconut fruits (often found breeding) and occasionally caught by sticky traps in Buff
Bay and Montego Bay areas; also from some other plants. Known from Brazil (Para),
Surinam, Guyana, Trinidad, Grenada, Barbados, St. Lucia, Martinique, Dominica,
Guadeloupe, Puerto Rico, Cuba, Jamaica, Florida, Alabama, Mexico, Honduras,
Panama, and Colombia.

Florida Entomologist 69(2)

Hercinothrips femoralis (Reuter, 1891): From coconut fruit and Emilia javanica.
Known from Martinique, Bermuda, Puerto Rico, Cuba, Florida, Georgia, Texas, and
Leucothrips theobromae (Priesner, 1923): From Clerodendrum speciosissimum,
Phaseolus lunatus, and Stachytarpheta jamaicensis in the wet Buff Bay area. Known
from Surinam, Panama, and Colombia.
Microcephalothrips abdominalis (Crawford, 1910): From some composite plants in
both wet Buff Bay and dry Montego Bay areas; occasionally caught by sticky traps.
Known from Trinidad, Martinique, Bermuda, Puerto Rico, Cuba, Jamaica, Florida,
Louisiana, Texas, Mexico, Guatemala, Costa Rica, and Panama.
Plesiothrips perplexus (Beach, 1896): From grass sweepings and others. Known
from Guyana, Trinidad, Tobago, Antigua, Puerto Rico, Dominican Republic, Cuba,
Florida, Texas, Panama, and Venezuela.
Psectrothrips interruptus (Hood, 1957) (= Pseudothrips): The present of a median
split on tergite X was considered as a minor local peculiarity. 4 9, 66 6, Woodstock,
sticky traps (4553f, 4556b, 4557, 4559d, 4564e). Known from Panama. An unnamed
allied species from Barro Colorado Island, Panama (Zetek 5178) in the USNM collection
was found not conspecific with this Jamaican species.
Rhamphothrips pandens Sakimura, 1983: From Cassia siamea, Gouania
lupuloides, Lantana camera, Pithecellobium dulce, and sticky traps in the wet Buff
Bay and the dry Montego Bay and Kingston areas. Known from Florida.
Salpingothrips minimus Hood, 1935: 1d, Round Hill, Desmodium sp. (4475Ab).
Known from Panama.
Scirtothrips sp.: Differs from S. manihoti and mutistriatus by wider frontal costa,
hind vein of fore wing with 3 to 4 setae, and male with a prominent pair of drepana.
69 9, ld, Woodstock, Coccoloba uvifera (4365c).
Selenothrips rubrocinctus (Giard, 1901): Abundant on several trees including
Anacardium occidentale, Malpighia punicifolia, Mangifera indica, and Pimenta
dioica, and also on many other plants in both Buff Bay and Kingston areas; occasionally
caught by sticky traps. Known throughout the Caribbean areas (collected in Jamaica as
early as 1921 at Hill Garden).
Sericothrips burungae Hood, 1935: 49 Round Hill, Eupatorium odoratum
(4479a). Known from Panama.
Sericothrips flavicollis Hood, 1954: 19, Kildare, Sida sp. (4326a). Known from
southern Brazil; apparently the first collection from the Caribbean area.
Sericothrips geminus Hood, 1935: 99 9, 66 d, Hope Garden, Cajanus cajan (4407).
Known from Puerto Rico and Panama.
Sericothrips gracilipes Hood, 1924: Differs from S. campestris from Florida. Abun-
dant or frequent on Macroptilium lathyroides, Malvastrum sp. and Sida spp.
everywhere in Jamaica. Known from Jamaica (previously collected in 1970), Texas and
Sericothrips inversus Hood, 1928: Frequent on Desmodium sp. and also from Di-
gitaria sp. Known from Trinidad, Dominica, and Panama.
Sericothrips portoricensis Morgan, 1925: From Psophocarpus palustris, and 2 other
plants. Known from Brazil (Para), Trinidad, St. Lucia, Guadeloupe, Puerto Rico, Cuba,
and Panama.
Sericothrips tricinctus Hood, 1928: 29 Woodstock, Psophocarpus palustris
(4313b). Known from Trinidad, Martinique, Dominica, and Guadeloupe.
Thrips hawaiiensis (Morgan, 1913): The first collection of this African-Oriental-
Pacific species from the Caribbean region. 1 Woodstock, sticky trap (4564f). Known
from Florida, Georgia, and South Carolina (all collected after 1967).


June, 1986

Sakimura: Jamaican Thrips



Family Phlaeothripidae-Subfamily Phlaeothripinae
Adraneothrips decorus Hood, 1938: Abundant on Sporobolus indicus; also from
coconut inflorescence; occasionally caught by sticky traps in the wet Buff Bay area.
Known from Cuba, Florida, Georgia, Texas, and Mexico.
Adraneothrips sp.: Differs from A. imbecillus from Peru by eye ventrally elongate,
tube shorter, fore wing with double fringes, and tergite IX with setae i-ii not dilated.
19, Orange Vale, sticky trap (4541b).
Aleurodothrips fasciapennis (Franklin, 1908): Predator upon scales. From various
plants including coconut fronds. Known from Brazil (Para), Grenada, Barbados, St.
Vincent, St. Lucia, Dominica, Guadeloupe, St. Croix, Bermuda, Puerto Rico, Haiti,
Cuba, Jamaica, Florida, Georgia, Louisiana, Texas, Mexico, Panama, and Venezuela.
Antillothrips cingulatus (Hood, 1919) (= A. graminatus Stannard, 1957): From
Malvastrum sp. and Sida sp. in the dry Montego Bay area. Known from Trinidad,
Puerto Rico, Jamaica, Florida, and Panama.
Carathrips mediamericanus (Hood, 1933): 29 9, Negril, sticky traps (4501c). Known
from Panama.
Eschatothrips reticulotubus (Stannard, 1953): 19, Hardwar Gap, 4000', beatings
(4511b). Collected earlier at this same locality in 1950.
Eurythrips modestus (Bagnall, 1917): 19, Woodstock, sticky trap (4553c). Known
from Brazil (Para), Trinidad, St. Vincent, Cuba, and Panama.
Eurythrips tarsalis Hood, 1925: 2 9 9, 1 Round Hill (Hill Top), Sporobolus indicus
(4486b). Known from Brazil (Para), Florida, Georgia, and Texas.
Eurythrips sp.: Differs from E. longilabris from Florida and Texas by dark antennal
I-II and strongly slenderized V-VIII with lanceolate VIII, dark legs, and tube dimen-
sion. Also differs from E. batesi previously reported from Jamaica by all the major setae
acuminated at apex. 1 Woodstock, sticky trap (4561a).
Haplothrips gowdeyi (Franklin, 1908): From great many weeds, grasses, and orna-
mentals, often abundantly or frequently on many of them; neither seen among coconut
inflorescence nor caught by sticky traps. A general flower feeder. The most abundant
tubuliferan thrips throughout Jamaica, except in the forested high land. Known
throughout the Caribbean areas including Jamaica.
Haplothrips graminis Hood, 1912: From grasses; Woodstock, Negril (4343b, 4460).
Known from Bahama, Cuba, Florida, Alabama, Texas, and Mexico.
Haplothrips sp.: Differs from H. humilis from Panama by head nearly as long as
wide and postocular seta small and dilated. An unnamed species from Cuba (Pinar City,
grass, 12.VII.1940, J. C. Bradley) in the USNMNH collection is, however, not con-
specific because its praepectus, probasisternum, and pelta are more degenerated than
the Jamaican species. 32 9 96d d, Woodstock, Kildare, Negril; Rhynchospora nervosa,
Cyperus diffusus (4340c, 4347, 4400b, 4464b). A common sedge feeder.
Haplothrips (Anchylothrips) sp.: Differs from H. (A.) preeri from Texas and Florida
by pale body, head 1.4 times as long as wide, and all the major setae dilated. Determined
with advice from Kellie O'Neill. 4 9 4d 6, Round Hill (Hill Top), Sporobolus indicus
Holopothrips sp.: Differs from H. fulvus Morgan, 1929:6 (= H. anacardii Hood,
1942: 581, New Synonymy; both types were compared) from Brazil (Bahia) by abdominal
VIII-X dark grayish brown, longer head (1.3 times as long as wide) with well developed
mid-dorsal seta, tergite IX with all major setae pointed, and male sternite VIII with
divided subbasal and undivided subapical transverse glandular strips. 6 9, 7 d,
Woodstock, Coccoloba uvifera (4365a).

Florida Entomologist 69(2)

June, 1986

Holothrips lucyae (Medina 1961:117) (= PII/yIlhnitJI.hi/ps): The type was examined;
its color description is supplemented as follows: A beautiful golden yellow large species
with extensive grayish brown blotches and shadings; apical extremes of mid-hind femora
and entire mid-hind tibiae and tarsi hyaline; abdominal pelta hyaline but from there on
graduating to deep yellow on tube; subcutaneous pigments reddish brown. Grayish brown
blotches and shadings: head with small light spots at both sides of anterior ocellus and
along posterolateral corner of eyes; pronotum lightly shaded; pterothorax deeply shaded
except distal 1 of mesial area; fore femur with small spot subapically along inner mar-
gin, mid-hind femora with 2 ill-defined and broad bands at base and also next to apical
hyaline band leaving the middle yellow, fore tibia and fore-mid coxae lightly shaded;
abdominal II-VIII with broad cross band along fore margin with a small break at middle
and distally distending along lateral margins, more extensively on VII-VIII, IX with
one light spot each on both sides along hind margin, tube with clearly-defined and broad
apical band. Antennal I-II pale brownish yellow, III yellowish brown with distal 1
grayish brown, IV-VIII dark grayish brown; wing pale grayish brown with darker
broad band at middle and narrow band at base; ocellar crescent brownish red; major
setae yellow to light brown. Both sexes concolorous, body length (mm) 2.6-2.7 (d),
3.4-3.5 (2), tube about 2.3 (6), 2.6 (9) times as long as wide at base. A species with a
manuscript name by Hood (2Y ?, 16, Brazil, Para, 1951, J. D. Hood) in the USNMNH
collection was found conspecific with H. lucyae. 39 9, Barnett, Hope Garden, Round
Hill; Lantana camera, Malvaviscus arboreus, Sporobolus indicus (4355b, 4438, 4486d).
Known from Brazil (Para) and Puerto Rico.
Holothrips phaeura (Hood, 1941) (= Polyphemothrips): 1d, Irwin, dead branches
(4474a). Known from Florida. Determined with advice from Kellie O'Neill.
Hoplandrothrips erythrinae Priesner, 1925: 29 9, Bengal, young coconut fruits fal-
len on ground, R. Latta (4427a). Known from Brazil (para), Surinam, Cuba, Mexico,
Panama, and Colombia.
Hoplandrothrips flavipes Bagnall, 1923: Frequent among the calyxes of coconut
fruits on tree or fallen on ground and coconut inflorescence; often caught by sticky traps
in both the wet Buff Bay and the dry Montego Bay areas. No collection was made from
any other plant. Known from Brazil (Para), Venezuela, and Colombia.
Hoplandrothrips reynei Priesner, 1923: 19, d1, Irwin, Buff Bay; dead branches,
sticky trap (4472a, 4570b). Known from Brazil (Para), Trinidad, Puerto Rico, Cuba, and
Hoplandrothrips sp.: Differs from H. pallens Hood from Cuba by subbasal seta iii
of fore wing and lateral seta of tergite VII both dilated, and seta ii of tergite IX pointed.
1 9, 26 d, Round Hill, Woodstock, Buff Bay; all from sticky traps (4502c, 4564g, 4565b).
Hoplothrips angusticeps (Hood, 1908): 19, Kildare, caught in flight in a coconut
grove (4562). Known from Georgia, Florida, and Costa Rica. A common bracket fungus
feeder in the eastern U.S.; probably the first collection recorded from any Caribbean
island (no specimen was found in the USNMNH collection).
Hoplothrips fungosus Moulton, 1928: 29 9, Woodstock, sticky traps (4554p, 4559a).
Known from Brazil (Para) and Puerto Rico. A common bracket fungus feeder among
various Pacific islands.
Hoplothrips moultoni (Hood, 1934): 19, Gray's Inn, sticky trap (4566c). Known
from Panama.
Hoplothrips spp. (B, D): Both are presently unplaceable. B: 3 9 9, Woodstock, White
River, sticky traps (4554q, 4567b). D: 29 9, Buff Bay, sticky traps (4570a).
Karnyothrips flavipes (Jones, 1912): 29 9, Round Hill, Woodstock-Kildare; both by
sticky traps (4502a, 4555); not common. Scale feeder. Known from Trinidad, St. Vincent,
Barbados, Bahamas, Bermuda, Puerto Rico, Dominican Republic, Florida, Georgia,
Louisiana, Texas, Mexico, and Venezuela.

Sakimura: Jamaican Thrips


Karnyothrips melaleuca (Bagnall, 1911): From Panicum maximum, Sporobolus
indicus, and coconut fronds; rarely caught by sticky traps. Scale feeder. Known from
Brazil (Para), Guyana, Trinidad, Patos Isle, Tobago, Barbados, St. Vincent, St. Lucia,
Dominica, St. Croix, Puerto Rico, Bahamas, Haiti, Cuba, Jamaica, Florida, Georgia,
Louisiana, Texas, Mexico, and Panama.
Karnyothrips merrilli (Watson, 1920): Frequent on coconut inflorescence and fruit
feeding on scales, and also occasionally caught by sticky traps; also from great many
weeds, grasses, ornamentals, and crop plants. The most common and abundant Kar-
nyothrips everywhere in Jamaica; scale feeder. Known from Guyana, Trinidad, Tobago,
Grenada, St. Vincent, St. Lucia, Guadeloupe, Puerto Rico, Cuba, Jamaica, Florida,
Georgia, Texas, Mexico, Honduras, Panama, and Colombia.
Karnyothrips ochropezus Hood, 1934: 19, Round Hill, Chloris sp. (4437a). Probably
scale feeder. Known from Panama.
Leptothrips vittipennis Hood, 1938: From Calliandra sp., Lantana camera, Malvas-
trum coromandelianum, Malvaviscus arboreus, and coconut inflorescence; also occa-
sionally caught by sticky traps in Buff Bay and Kingston areas. Probably mite feeder.
Known from Panama.
Liothrips urichi Karny, 1923: 1 Round Hill (Hill Top), Borreria laevis, by R.
Latta (4428). Known from Trinidad.
Liothrips sp.: Differs from L. leucopus Titschack from the western Mediterranean
area, the only other congener with a combination of dark brown body and totally yellow
legs but not closely related, by longer head (1.3 times as long as wide), uniformly dark
grayish brown and more slender tube (3.2 times as long as wide), and pale major setae.
16, Hardwar Gap, 4000', beatings in forest (4511a).
Macrophthalmothrips heinzei Mound, 1972: 29 Woodstock, Gray's Inn; Sticky
traps (4554i, 4566b). The type series was collected subsequently in 1970 6 miles east of
Macrophthalmothrips helenae Hood, 1934: 49 9, Negril, sticky traps; Woodstock,
Saccharum officinarum; Hope Garden, a legume in greenhouse by W. Carter (4501a,
4550c, 4573). Known from Cuba, Florida, Texas, Mexico, and Panama.
Orthothrips sp.: Differs from 0. caudatus Priesner from Surinam by longer head
(1.3 times as long as wide), antennal II totally dark grayish brown, stouter tube (2.6
times as long as wide), and tergite IX with seta i dilated. Also differs from 0. bilineatus
Stannard, described from Jamaica but not collected during the present survey, by dark
tube, lanceolate-pedicellate antennal VIII, and fully developed cepharic and pronotal
setae. 29 9, Kildare, sticky traps (4562c, 4572).
Phrasterothrips sp.: Differs from P. conducans Priesner from Paraguay and Brazil
by antennal III with a single sense cone, inner epimeral seta V1 as long as outer seta,
and tube in different profile. 19, Woodstock, sticky trap among wild sugar cane (4550d).
Pygmaeothrips columniceps Karny, 1920: Abundant on bracket fungus, occasionally
also under bark or on dead branches in Montego Bay area. Apparently the first collection
of this Oriental and Pacific species in the Caribbean area (no specimen was found in the
USNMNH collection).
Sophiothrips squamosus Hood, 1933: 1 Buff Bay, sticky trap (4570c). Known from
Trinidad, Cuba, and Panama.
Stephanothrips occidentalis Hood and Williams, 1925: 19, Hope Garden, Panicum
maximum (4350a). Known from Brazil (Para), Trinidad, Bermuda, St. Croix, Puerto
Rico, Cuba, Jamaica, Caymans, Florida, Mexico, and Panama.
Strepterothrips floridanus (Hood, 1938): 39 9, Woodstock, Dover; sticky traps
(4554k, 4569b). Known from Guyana, Trinidad, Caymans, Cuba, Florida, Texas, and

Florida Entomologist 69(2)

June, 1986

Tylothrips osborni (Hinds, 1902): 29 9, Woodstock, Negril; sticky trap, Sida or
Malvastrum spp. (4468e, 4558). Known from Trinidad, Florida, and Panama.
Family Phlaeothripidae-Subfamily Idolothripinae
Compsothrips graminis (Hood, 1936) (= Oedaleothrips): 39 9, 46 d, Palisadoes
sweepings, 8.XII.1957, by T. H. Farr (4575). Known from Venezuela (Patos Isle),
Trinidad, St. Croix, Tortola, Bahamas, Mexico.
Diceratothrips bicornis Bagnall, 1908 (= D. armatus Bagnall): 19, Buff Bay,
coconut inflorescence (4529a). Known from Brazil (Para), Surinam, Guyana, Trinidad,
Mexico, Panama, and Venezuela.
Diceratothrips picticornis Hood, 1914 (= D. wolcotti Morgan; both type series were
examined): 109 9, 66 d, Irwin, dead branches piled on the ground (4472c); 16, Ben-
zenn, general sweepings (4395). Known from Puerto Rico, Cuba, Bahamas, and Panama.
Elaphrothrips laevicollis (Bagnall, 1910): 9d 6, Bath Mountain, Rock Hall, Christ-
iana, and east of Montego Bay (all high mountainous localities), all by sweepings except
one from Moghania strobilifera, 1955-1959, by T. H. Farr (4574, 76, 77, 79). Known
from Brazil (Para), Surinam, Guyana, Trinidad, Mexico, Honduras, Costa Rica,
Panama, and Venezuela.
Ethirothrips brevis (Bagnall, 1921) (= Dichaetothrips claripennis (Moulton): 26 6,
Woodstock, Bengal; sticky trap, on coconut fruit (4450, 4568e). Known from Trinidad,
Haiti, Bahamas, Jamaica, Florida, and Mexico.
Gastrothrips anolis Morgan, 1925: ld, Round Hill (Hill Top), Solanum corvum
(4485c). Known from Puerto Rico, Cuba, Caymans, and Panama.
Neosmerinthothrips sp.: Differs from N. collaris Bagnall known from St. Vincent,
Dominica, and Puerto Rico, by shorter head (1.1 times as long as wide) with slenderer
tube (2.4 times as long as wide), fore wing clear at base, anteromarginal pronotal seta
well developed, and tergite IX with seta i subequal to seta ii and longer than tube. An
unnamed species in the USNMNH collection (19, Haiti, Port-Au-Prince, dead branches
with leaves, 29.VIII.1951, Hood coll. (2728)) is conspecific with this Jamaican sp. 1S
and larvae, Bengal, coconut fruit, among calyx (4450). Known from Haiti.
Nesothrips lativentris (Karny, 1913) (= Rhaebothrips): Abundant or frequent on
many different plants, particularly various grasses in the wet Buff Bay area as well as
the dry Montego Bay and Kingston areas; never caught by sticky traps and only once
collected on coconut fruit. A fungus spore feeder. Known from Trinidad, Virgin Islands,
Puerto Rico, Dominican Republic, Bahamas, Cuba, Jamaica, Caymans, Florida, and


Throughout the world, no thrips species has been known as a pest of the coconut
crops, and practically nothing is known of the thrips fauna on coconut. Extensive sticky
trappings among coconut fronds and inflorescences and repeated manual collections on
different parts of coconut trees, as well as among the weedy undergrowth in and around
the coconut plantations finally yielded some data (Table 2) on the thrips fauna on
coconuts in Jamaica, and its sources of migrations onto coconuts. The inflorescences are
interspersed among fronds and bloom periodically in short intervals throughout the
year, and most thrips trapped were flower feeding species. Sometimes more than 300
thrips were trapped per week.
Of 56 species enumerated in Table 2, 48 were trapped, and 10 of them were collected
also from coconuts, but the remaining 38 were not; another 8 were collected from
coconuts but not trapped; all together 18 were collected from coconuts. Nineteen are
flower feeders; 8, leaf-fruit feeders; 20, fungal hyphae and fungal decaying breakdown


Sakimura: Jamaican Thrips


Chaetisothrips striatus caribeanus Sakimura 2 a
Chirothrips mexicanus Crawford 2 a
Chirothrips texanus Andre 2 a
Frankliniella borinquen Hood 2 a
Frankliniella breviseta Moulton 2 a
Frankliniella bruneri Watson 1-2 a (inflorescence, frequent)
Frankliniella caphalica (Crawford) 2 a
Frankliniella citripes Hood 2 a
Frankliniella cubensis Hood 2 a
Frankliniella insularis (Franklin) 1-2 a (inflorescence)
Frankliniellajamaicensis Sakimura & O'Neill 2 a
Frankliniella kelliae Sakimura 1-2 a (inflorescence, frequent)
Frankliniella parvula Hood 2 a
Frankliniella sp. (P) 2 a
Frankliniella varipes Moulton 2 a
Microcephalothrips abdominalis (Crawford) 2 a
Psectrothrips interruptus (Hood) 2 b
Rhamphothrips pandens Sakimura 2 a
Thrips hawaiiensis (Morgan) 2 b
Anisopilothrips venustulus (Priesner) 1 a (young fruit, frequent)
Caliothrips phaseoli (Hood) 2 a
Ceratothripoidesfunestus (Hood) 2 a
Chaetanaphothrips orchidii (Moulton) 1 a (young fruit, once)
Corynothrips stenopterus Williams 2 b
Heliothrips haemorrhoidalis (Bouch6) 1-2 a (young fruit, frequent)
Hercinothripsfemoralis (Reuter) 1 a (fruit)
Selenothrips rubrocinctus (Giard) 2 a
Adraneothrips decorus Hood 1-2 a (inflorescence)
Adraneothrips sp. 2 b
Carathrips mediamericanus Hood 2 b
Eurythrips modestus (Bagnall) 2 b
Eurythrips sp. 2 b
Hoplandrothrips erythrinae Priesner 1 b (fallen fruit)
Hoplandrothripsflavipes Bagnall 1-2 b (inflorescence, fruit)
Hoplandrothrips reynei Priesner 2 a
Hoplandrothrips sp. 2 b
Hoplothrips angusticeps (Hood) 2 b
Hoplothripsfungosus Moulton 2 b
Hoplothrips moultoni Hood 2 b
Hoplothrips sp. (B) 2 b
Hoplothrips sp. (D) 2 b
Macrophthalmothrips heinzei Mound 2 b
Macrophthalmothrips helenae Hood 2 a
Orthothrips sp. 2 b
Sophiothrips squamosus Hood 2 b
Strepterothripsfloridanus (Hood) 2 b
Tylothrips osborni (Hinds) 2 a

362 Florida Entomologist 69(2) June, 1986

Diceratothrips bicornis Bagnall 1 b (inflorescence, once)
Ethirothrips brevis (Bagnall) 1-2 b (fruit)
Neosmerinthothrips sp. 1 b (fruit)
Nesothrips lativentris (Karny) 1 a (fruit, once)
Aleurodothripsfasciapennis Franklin la (frond, once)
Karnyothripsflavipes (Jones) 2 b
Karnyothrips melaleuca (Bagnall) 1-2 a (frond)
Karnyothrips merrilli (Watson) 1-2 a (young fruit and
inflorescence, frequent)
Leptothrips vittipennis Hood 1-2 a (inflorescence)

products feeders; 4, fungal spore feeders; and 5, predators. All the flower and leaf-fruit
feeders are Terebrantians, and most of them were trapped frequently and abundantly,
and also were found abundantly living among the undergrowth of coconuts. All the
fungal growth feeders and predators are Tubuliferans, all of them were only sporadically
and sparsely trapped and were mostly not or sparsely present among the undergrowth
in the immediate vicinity.
The flower feeder group was predominated by 12 species of Frankliniella, and the
remainder were incidental transients. F. bruneri, kelliae, and insularis were trapped
abundantly. Frequently, all 3 species were collected from inflorescence, the first species
abundantly. All are polyphagous species and are the most common and abundant species
of the genus in Jamaica. F. breviseta, cephalica, cubensis, and varipes were trapped
occasionally in fair numbers, but they were not collected from inflorescence. Anisopilot-
hrips venustulus and Heliothrips haemorrhoidalis of the leaf-fruit feeder group were
frequently or abundantly trapped as well as often collected from young fruits where
both were found breeding. No infestation of thrips was observed on the mature fronds,
but observations were not made on the developing tender hearts.
Most of the fungal hyphae and fungal decaying breakdown products feeders were
sporadic transients. An exception was Hoplandrothrips flavipes which was trapped
often as well as collected in good numbers from young fruits. Coconut fruits appeared
to be its preferred feeding niches. All the fungal spore feeders were collected rarely
from fruits but only once from inflorescence, and only one was trapped. Among the
predator group, Karnyothrips merrilli, a scale feeder, was trapped occasionally, but it
was often collected from young fruits and among inflorescence. This species also was
found abundantly among the undergrowth of coconuts. The rest of the group are inciden-
tal transients.


Sincere acknowledgments are gratefully made to Kellie O'Neill and Steve Nakahara
of Systematic Entomology Laboratory at Beltsville, U.S. Department of Agriculture,
Paul H. Arnaud, Jr. and W. J. Pulawski of California Academy of Sciences, Richard
zur Strassen of Forschungsinstitut Senckenberg, S. Medina Gaud of University of
Puerto Rico, the Botany Department, University of the West Indies, and the late
Walter Carter of the FAO project all of whom provided much advice and help in various
ways during the progress of this work over the long period of time.

Frank: Florida Staphylinidae


HOOD, J. D. 1915. Descriptions of new American Thysanoptera. Insec. Inscit. Menstr.
3: 1-40.
HOOD, J. D. 1942. A century of new American Thysanoptera. III. Rev. Ent. (Rio de
Janeiro) 12: 547-678.
HOWARD, F. W., R. C. NORRIS, AND D. L. THOMAS. 1983. Evidence of transmission
of palm lethal yellowing agent by a planthopper, Myndus crudus (Homoptera:
Cixiidae). Trop. Agric. (Trinidad) 60: 168-171.
MEDINA GAUD, S. 1961. The Thysanoptera of Puerto Rico. Univ. Puerto Rico, Agric.
Exp. Sta. Tech, Papers 32: 1-159.
MORGAN, A. C. 1929. A new genus and five new species ofThysanoptera foreign to the
United States. Proc. Ent. Soc. Washington 31: ;1-9.
SAKIMURA, K. AND KELLIE O'NEILL. 1979. Frankliniella, redefinition of genus and
revision of Minuta group species (Thysanoptera: Thripidae). United States Dept.
Agric. Tech. Bull. 1572: 1-49.


Entomology and Nematology Department,
3103 McCarty Hall,
University of Florida,
Gainesville, Florida 32611, USA.


A preliminary checklist of the Staphylinidae of Florida (USA) is presented. It in-
cludes 324 species in 122 genera and gives bibliographical information on their original
descriptions, their synonymies, and the publications in which they were recorded as
occurring in Florida. The total size of the staphylinid fauna of Florida is estimated
conservatively as 450 species. The species/area relationship suggests that this fauna is
depauperate relative to the staphylinid fauna of western Europe.


Se present una lista preliminary de las species de Staphylinidae que se encuentran
en Florida (E.U.A.). La list incluye 324 species en 122 g4neros, y ofrece informaci6n
bibliogrAfica sobre las descripciones originales, los sinonimos, y las publicaciones en las
cuales se registraron las species como ocurriendo en Florida. El tamafio total de la
fauna estafilinida de Florida se estima conservativamente en 450 species. La relaci6n
especies/Area indica que esta fauna es depauperada respect de la fauna estafilinida de
Europa Occidental.

In numbers of described species, Coleoptera represent the largest order of the
largest class of the largest phylum of the animal kingdom. At least in western Europe
and in America north of Mexico, in numbers of described species Staphylinidae repre-


364 Florida Entomologist 69(2) June, 1986

sent the largest family of Coleoptera. However, the composition of the staphylinid fauna
of only one large geographical area (western Europe) is at all adequately known.
Only two publications have dealt with the staphylinid fauna of Florida specifically
and in detail. After collecting expeditions in 1875 and 1876, Schwarz (1878) named 99
species as occurring there and mentioned existence of about as many species which
could not then be identified specifically; a section of the text was devoted to descriptions
by LeConte (1878) of new species. Notman (1920) found 101 species represented in the
collections of the American Museum of Natural History, and he described several of
them as new.
In 1969 was published the first of a series of taxonomic revisions of the North
American staphylinid fauna and including the Florida fauna. Several authors have con-
tributed such revisions, usually at the generic level. These modern works differ from
older taxonomic publications in that they are more thorough, are based on examination
of long series of specimens whenever available, and they use reproductive structures
as taxonomic characters. It may be fairly stated that in 1968 all North American
staphylinid genera required modern taxonomic revision, but it has been possible to
revise only a small proportion of the total fauna since then.
Although the number of named species recorded from Florida is now over 300, there
is no single comprehensive treatment of them in the literature, and neither is such
treatment now possible because taxonomic knowledge of the bulk of the species is
inadequate. This checklist is preliminary because it is incomplete. It provides concise
answers to the questions: what staphylinid species are known to occur in Florida, where
in the literature is information available about them, and how did they come to be
recorded as occurring in Florida? It also estimates the total size of the staphylinid fauna
of Florida, and compares the species richness of that fauna with faunas of two areas in
western Europe where the species composition is better known.


A species list was drafted by examination of two North American catalogs (Leng
1920 and supplements, Moore and Legner 1975). Curiously, the earlier catalog attri-
buted many more species to Florida than did the latter. Over a period of about five
years, I examined the entire literature relevant to nomenclature and occurrence of
Staphylinidae in Florida, from which sources the drafted list was augmented and cor-
Names of subfamilies and tribes are capitalized and centered in Table 1. Names of
genera within each tribe, and of species within each genus, are in most cases arranged
alphabetically. Taxa revised since 1968 are treated differently: an asterisk is placed to
the left of the name of each taxon (subfamily, tribe or genus) revised, and names of
species within each genus, and genera within each tribe are arranged in the order used
by the author of the revision.
Synonyms of names of species are in most cases listed below the valid name, follow-
ing an equivalence sign (=). This listing of synonyms is deliberately restricted to names
used for species described from the southeastern United States and the neotropical
region. However, in the case of taxa revised since 1968, and against whose names an
asterisk is placed, no synonyms are listed.
Against the author name of each species (left column) is given in the central column
the date and page of publication of each species name. This provides an index to the
literature cited. In the right column, Table 1 lists the author, date and page of publica-
tion establishing occurrence of each species in Florida. In most cases, the right column
gives the first record of occurrence of a species in Florida, but where a taxonomic

Frank: Florida Staphylinidae 365



Proteinus Latreille
P. thwrruasi Frank

*Eudectoides Campbell
E. crassicornis (LeConte)

*Ephelinus Cockerell
E. notatus (LeConte)

Omalium Gravenhorst
0. foraminosum Miklin
0. humerosum Fauvel
0. repandum Erichson

Hypotelus Erichson
H. hostilis Fauvel

Clavilispinus Bernhauer
C. exiguus (Erichson)
= C. rufescens (LeConte)
Nacaeus Blackwelder
N. tenellus (Erichson)
= N. tenuis (LeConte)
= N. flavipennis (Fauvel)
= N. fauveli (Sharp)
Thoracophorus Motschulsky
T. brevicristatus (Horn)
T. costalis Erichson

Holotrochus Erichson
H. brachypterus Fauvel
= H. minor LeConte
(nec Fauvel
Osorius Latreille
0. brevicornis Notman
0. latipes (Gravenhorst)
0. parviceps Notman
0. planifrons LeConte
0. politus LeConte

*Bledius Leach
B. ceratus Blackwelder
B. politus Erichson
B. basalis LeConte
B. cordatus (Say)
B. dimidiatus LeConte
B. thinopus Herman
B. turbulentus Casey
B. cognatus LeConte


1979: 333 ibid.
subfamily OMALIINAE



subfamily PIESTINAI

Campbell 1978:14

Campbell 1978:69

Leng 1920:94 SM
Notman 1920:693
Leng 1920:94 SM

1865:43 Leng 1920:93
subfamily OSORIINAE

1840:830 Fauvel 1878:180
1863: 59 LeConte 1877: 249

1887: 720

1871: 332

1863: 437)

Fauvel 1878: 179
Schwarz 1878: 442

Blackwelder 1939:22
Schwarz 1878:442


1920:698 ibid.: 699
1806:198 Schwarz 1878:442
1925:12 ibid.
1877:215 Leng 1920: 98 SM
1877:215 ibid.

1889a: 70

Herman 1983:135
Herman 1976:131
Herman 1976:98
Herman 1976:85
Herman 1976:85
ibid.: 88
Herman 1976:94
Herman 1983:14

Florida Entomologist 69(2)

B. wudus Herman
B. mandibularis Erichson
B. rubiginosus Erichson
B. semiferrugineus LeConte
Carpelimus Leach
C. agonus (Casey)
C. basicornis (Notman)
C. convexulus (LeConte)
C. fulvipes (Erichson)
= C. rubripennis (Fauvel)
= C. aequalis (Gundlach)
C. maculicollis (Notman)
C. obesus (Kiesenwetter)
= C. memnonius (Fauvel)
(nee (Erichson)
= C. spectatus (Casey)
C. quadripunctatus (Say)
= C. laticollis (LeConte)
Manda Blackwelder
M. nearctica Moore
*Microbledius Herman
M. litoreus Herman
*Psamathobledius Herman
P. punctatissimus (LeConte)
Thinodromus Kraatz
T. arcifer (LeConte)
Anotylus Thomson
A. exiguus (Erichson)
A. insignitus (Gravenhorst)
A. nanus (Erichson)
A. rugosus (Fabricius)
Apocellus Erichson
A. sphaericollis (Say)
= A. amabilis (Sachse)
= A. longicornis (Sachse)
A. stilicoides LeConte
Oxytelus Gravenhorst
0. convergens LeConte
0. incisus Motschulsky
0. incolumis Erichson
0. pennsylvanicus Erichson
0. sculptus Gravenhorst
Platystethus Mannerheim
P. spiculus Erichson
= P. exiguus Jacquelin du Val
*Oxypomrs Fabricius
0. femoralis Gravenhorst
0. vittatus Gravenhorst
Megalopinus Eichelbaum
M. caelatus (Gravenhorst)
M. rufipes (LeConte)
Stenus Latreille
S. alacer Casey


1889b: 356
1863: 440
1920: 693
1844: 375
1889b: 345


Herman 1972:193
Herman 1983:140
Herman 1972: 205

Leng 1920:96 SM
Leng 1920:96 SM
Schwarz 1878: 442

ibid.: 694

Schwarz 1878: 442

ibid.: 346
Leng 1920:96 SM


1972:129 ibid.

1877:226 Herman 1972:138


1840: 797


1857b: 504

1840: 784

Casey 1889b: 330

Casey 1893:397
Schwarz 1878:442
Notman 1920:695
Leng 1920:96

Schwarz 1878: 442


Frank & Thomas 1981b: 400
Schwarz 1878: 442
Frank & Thomas 1981b: 402
Schwarz 1878:442

Frank 1976:157


1802:196 Campbell 1969:241
1802:195 Campbell 1969:251

1802:197 Schwarz 1878:442
1863:51 Leng 1920:98 SM

1884a: 135 ibid.: 136

June, 1986

Frank: Florida Staphylinidae

= S. fauvelianus Sharp
S. arculus Erichson
S. callosus Erichson
S. carolinae Casey
S. colonus Erichson
= S.floridanus Casey
S. champion Sharp
= S. inermis Sharp
= S. chapini Blackwelder
S. cubensis Bernhauer
S. lutzi Notman
S. megalops (Casey)
S. meridionalis (Casey)
S. nitescens (Casey)
S. rostellifer Puthz
S. sectilifer Casey
= S. milleporus Casey
= S. teter Notman
= S. odius Blackwelder
S. tuberculatus Casey
*Edaphus Motschulsky
E. nitidus Motschulsky
E. americanus Puthz
Euaesthetus Gravenhorst
E. americanus Erichson
E. atomus Casey
E. bicoloratus Casey
E.floridae Casey
E. punctatus Casey
E. similis Casey
Cubanotyphlus Coiffait & Decou
C. largo Frank

Araeocerus Nordmann
A. picipes (Erichson)
Pinophilus Gravenhorst
P. confusus Fall
= P. opacus Casey
(nec LeConte
P. latipes Gravenhorst
P. opacus LeConte
P. parcus LeConte
Palaminus Erichson
P. contortus LeConte
P. cribratus LeConte
P. flavipennis LeConte
P. larvalis LeConte
P. normalis LeConte
P. pumilis LeConte
P. testaceus Erichson

1886: 646
1840: 744 Schwarz 1
1840: 737 Schwarz ]
1884a: 116 Leng 192(
1840:699 Casey 188
1884a: 95 ibid.: 96
1910:364 Puthz 197
1920:700 ibid.: 701
1884a: 161 ibid.
1884a: 185 ibid.: 186
1884a: 170 ibid.: 171
1972:315 ibid.: 316
1884a: 110 Notman 1
1884a: 111 ibid.
1920:699 ibid.: 700
1884a: 129 ibid.: 130

):99 SM
4: 97

4a: 10

920: 700

1857a: 7 Puthz 1974b: 917
1974b: 920 ibid.: 922

1840: 747 Casey 1884:28
1884b: 28 ibid.: 29
1924:150 ibid.
1884b: 21 ibid.: 22
1884b:21 ibid.
1884b:22 ibid.: 23

1984b: 1412 Frank & Thomas 1984a: 1414

1840:671 Schwarz 1878:441






Schwarz 1878:442
Leng 1920: 100 SM
Notman 1920:701


Florida Entomologist 69(2)

Achenomorphus Motschulsky
A. corticinus (Gravenhorst)
= A. rufescens (Nordmann)
Acrostilicus Hubbard
A. hospes Hubbard
Astenus Dejean
A. binotatus (Say)
A. fusciceps (Casey)
A. linearis (Erichson)
A. prolixus (Erichson)
A. spectrum (Casey)
Biocrypta Casey
B. magnolia Blatchley
Dacnochilus LeConte
D. laetus LeConte
Echiaster Erichson
E. brevicornis (Casey)
= E. virginicus (Fall)
Homaeotarsus Hochhuth
H. badius (Gravenhorst)
H. bicolor (Gravenhorst)
H. cinctus (Say)
H. clavicornis (Casey)
H. despectus (LeConte)
H. floridanus (LeConte)
H. lugubris (LeConte)
H. obliquus (LeConte)
= H. parcus (LeConte)
H. pallipes (Gravenhorst)
Lathrobium Gravenhorst (sensu lato)
L. americanum Duvivier
= L. anale LeConte
(nec Lucas
L. collare Erichson
L. dimidiatum Say
L. emarginatum Watrous
L. floridae (Casey)
L. floridanum (Casey)
L. pallidulum LeConte
L. parcum LeConte
L. politum Gravenhorst
L. tetricum (Casey)
= L. nigriceps (Notman)
= L. canoaensis Bierig
L. ventrale LeConte
L. tricolor Casey
Lithocharis Dejean
L. ochracea (Gravenhorst)
Myrmecosaurus Wasmann
M. ferrugineus Bruch
Ophioomma Notman
0. rufa Notman
Paederus Fabricius
P. floridanus Austin
P. littoreus Austin
P. obliteratus LeConte



Schwarz 1878: 441

1896:299 ibid.

1905: 240

Schwarz 1878: 441
Notman 1920:704
Schwarz 1878:441
Schwarz 1878:441
Notman 1920:704

1917:236 ibid.: 237




1849: 117)
1934c: 325

Schwarz 1878:441

Notman 1920:704

Horn 1885:89
Schwarz 1878: 441
Schwarz 1878: 441
Horn 1885: 90
ibid.: 394
Horn 1885:101

Notman 1920:702

Leng 1920:102 SM
Schwarz 1878: 441
Leng 1920:102 SM

1802:59 Notman 1920:702


Frank 1977:31

1920: 705 ibid.


Notman 1920:701

June, 1986

Florida Entomologist 69(2)

Rugilus Samouelle
R. angularis (Erichson)
R. dentatus (Say)
Scopaeus Erichson
S. carolinae Casey
S. exiguus Erichson
S. femineus (Moore & Le
S. macilentus Casey
S. opacus (LeConte)
S. picipes Casey
Stamnoderus Sharp
S. monstrosus (LeConte;
S. pallidus Casey
Stilicopsis Sachse
S. paradoxa Sachse
S. subtropica Casey
Sunius Curtis
S. confluentus (Say)
S. debiliconis (Wollastol
S. minutes (Casey)
S. testaceus (Casey)
Thinocharis Kraatz
T. exilis (Erichson)
= T.fragilis (Sharp)
= T. minute (Sharp)
= T. delicatula (Case:
= T. pertenuis (Casey
T. quadriceps (Notman)

*Diochus Erichson
D. schaumii Kraatz

*Neoxantholinus Cameron
N. cristatus Smetana
*Nudobius Thomson
N. luridipennis Casey
*Phacophallus Coiffait
P. tricolor (Kraatz)
*Microlinus Casey
M. pwsio (LeConte)
*Lithocharodes Sharp
L. ruficollis (LeConte)
L. nigripennis (LeConte
L.floridanus (LeConte)
*Neohypnus Coiffait & Sai:
N. attenuatus (Erichson'
N. bicolor Smetana
N. pusillus (Sachse)
N. fusciceps (LeConte)
N. melanops (Casey)
N. emmesus (Gravenhor

Belonuchus Nordmann
B. pallidus Casey
B. rufipennis (Fabricius

1840:634 Schwarz 1878:441
1834:457 Leng 1920:104 SM

1840: 608
1905: 208

Notman 1920: 703
Schwarz 1878:441
Notman 1920: 703
Schwarz 1878: 441
Casey 1905: 206

1863:48 Schwarz 1878:441
1905:233 ibid.

1852:145 Schwarz 1878:441
1910:191 ibid.

1834: 456

1840:627 Black
y) 1905:159
F) 1910:188
1920: 703 ibid.

Frank 1981a: 388
Blackwelder 1939:23

welder 1939:23

1860:27 Smetana 1982:32

1982: 64

ibid.: 57

1906: 383 Smetana 1982:90

1859:110 Smetana 1982:111

1880:171 Smetana 1982: 115




1839:330 Smetana 1982:204
1982:23 ibid.: 215
1852:124 Smetana 1982:234
1880:173 Smetana 1982:241
1906:390 Smetana 1982:244
t) 1802:176 Smetana 1982:250



June, 1986

Frank: Florida Staphylinidae

= B.formosus (Gravenhorst)
= B. apicalis (Dejean)
= B. moquinus Casey
Cafius Curtis
C. bistriatus (Erichson)
= C. bilineatus (Erichson)
C. caribeanus Bierig
C. rufifrons Bierig
C. subtilis Cameron
Creophilus Samouelle
C. maxillosus (Linn6)
*Erichsonius Fauvel
E. crescenti Frank
E. parcus (Horn)
E. rusticus Frank
E. rosellus Frank
E. alumnus Frank
E. floridanus Frank
E. civicus Frank
Gabrius Curtis
G. nigritulus (Gravenhorst)
Gabronthus Tottenham
G. mgogoricus Tottenham
G. thermarum (Aube)
Hesperus Fauvel
H. apicialis (Say)
H. baltimorensis (Gravenhorst)
Laetulonthus Moore & Legner
L. laetulus (Say)
*Neobisnius Ganglbauer
N. paederoides (LeConte)
N. ludicrus (Erichson)
N. jocosus (Horn)
N. sobrinus (Erichson)
Philonthus Curtis
P. alumnus Erichson
P. cautus Erichson
P. debilis (Gravenhorst)
P. flavolimbatus Erichson
P. fidvipes (Fabricius)
P. gopheri Hubbard
P. hepaticus Erichson
= P. vilis Erichson
= P. orphanus Erichson
= P. pauxillus Solsky
= P. partimanus Sharp
P. lomatus Erichson
= P. georgianus Sachse
P. longicornis Stephens
P.feralis Erichson
= P.fumosus Solsky
P. sericans (Gravenhorst)
= P. brunneus (Gravenhorst)
P. ventralis (Gravenhorst)
*Platydracus Thomson
P. comes (LeConte)

1884b: 125

1934a: 68
1934a: 68

Schwarz 1878:441

Frank et al. 1986

Frank 1985:61
Frank etal. 1986
Frank 1985:62

1758:421 Schwarz 1878:441

1975a: 182
1975a: 194
1975a: 195
1975a: 196
1975a: 198
1975a: 201


Notman 1920: 705
ibid.: 203
Frank 1981c: 100
Frank 1981c: 100
Frank 1981c:100

Fauvel 1889: 116

1955:185 Frank 1984a:476
1850:316 Frank 1984a: 475

1834:451 Frank 1984a: 478
1802:163 Frank 1984a: 478

1834:449 Frank 1984a: 479






Horn 1884:199
Horn 1884:188
Leng 1920:107 SM
Notman 1920:706
Notman 1920:706
Schwarz 1878: 441

Notman 1920:706

Castle & Laurent 1896:303

Leng 1920:107 SM
Frank 1984a: 480

Newton 1973:294

Florida Entomologist 69(2)

P. pinorum (Casey)
P. praelongus (Mannerheim)
= P. cicatricosus (LeConte)
P. tomentosus (Gravenhorst)
P. fossator (Gravenhorst)
P. temporalis (Casey)
P. femoratus (Fabricius)
P. caseyi (Scheerpeltz)#
= P. quadraticeps (Casey)
(nec Menktri&s
P. cinnamopteruj (Gravenhorst)#
(# part of a species complex un
*Heterothops Stephens
H. campbelli Smetana
*Quedius Stephens
Q. peregrinus (Gravenhorst)
Q. compransor Fall
Q. rresomelin'us (Marsham)
Q. laticollis (Gravenhorst)
Q. verres Smetana
*Hemiquedius Casey
H.ferox (LeConte)
*Anaquedius Casey
A. vernix (LeConte)
*Acylophorus Nordmann
A. densus LeConte
A.fallax Smetana
A. princeps Smetana
A. lecontei Duvivier
A. pusillus Smetana
A. longistylus Casey
A. zdenae Smetana
*Atanygnathus Jacobson
A. bicolor (Casey)
BryoporMs Kraatz
B. rufescens LeConte
= B. testaceus LeConte
= B. parvulus Casey
*Lordithon Thomson
L. angularis (Sachse)
L. obsoletus (Say)
Mycetoporus Mannerheim
M. americanus Erichson
= M. lucidulus LeConte
M. brunneus (Marsham)
= M. lepidus Horn
M. flavicollis LeConte

1915: 429
1801: 594
1832: 143)
der revision)

Newton 1973:304
Newton 1973:304




Leng 1920:108 SM


1971:22 Smetana 1981: 632

1971: 174

Smetana 1971: 59
Smetana 1978:823
Smetana 1971: 77
Smetana 1978:826
Smetana 1976: 176

1878:388 Smetana 1971:239

1878:389 Smetana 1978: 831

1878: 387
1971: 255

ibid.: 255
Smetana 1971:261
Smetana 1971:262
Benjamin 1983:347

1915:424 Smetana 1971:272


Schwarz 1878:440

1852:122 Campbell 1982:30
1834:464 Campbell 1982:65

1863: 33

Leng 1920:112

Horn 1877:123

*Coproporus Kraatz
C. ventriculus (Say)
C. pulchellus (Erichson)
C. segnis (Sharp)
C. hepaticus (Erichson)



Campbell 1975:183
Campbell 1975: 187
Campbell 1975: 188
Campbell 1975:190

June, 1986

Frank: Florida Staphylinidae

C. laevis LeConte
*Sepedophilus Gistel
S. macer (Casey)
S. velocipes (Casey)
S. versicolor (Casey)
S. basalis (Erichson)
S. kiteleyi Campbell
S. opicus (Say)
S. scriptus (Horn)
*Tachinus Gravenhorst
T. axillaris Erichson
*Tachyporus Gravenhorst
T. jocosus Say
Trichopsenius Horn
T. depressus (LeConte)
Xenistusa LeConte
X. hexagonalis Seevers
Anacyptus Horn
A. testaceus (LeConte)

*Deinopsis Matthews
D. illinoisensis Klimaszewski
D. franki Klimaszewski
*Adin psis Cameron
A. bicornis Klimaszewski
A. cuspidata Klimaszewski
A. myllaenoides (Kraatz)

*Myllaena Erichson
M. potowatomi Klimaszewski
M. currax Notman
M. koasati Klimaszewski
M. insipiens Casey
M. seminole Klimaszewski
M. cuneata Notman
M. arcana Casey
M. magnolia Klimaszewski

*Aleochara Gravenhorst
A. bimaculata Gravenhorst
A. notula Erichson
A. verna Say
A. lacertina Sharp
A. puberula Klug
A. lustrica Say
A. lata Gravenhorst
A. gracilicornis Bernhauer
A. litoralis (Maklin)
Genosema Notman
G. pulchra (Kraatz)
G. sexualis Notman
Lophomucter Notman
L. laevicollis Notman



Campbell 1975: 196

Campbell 1976:22
Campbell 1976:32
Campbell 1976:42
Campbell 1976:53
Campbell 1976:57
Campbell 1976:60

1839:261 Campbell 1973:38

1834:466 Campbell 1979: 62


Seevers 1938:437

1941:324 ibid.: 326


Schwarz 1878:440

1979:52 Klimaszewski 1985:142
1980:110 ibid.: 113

1857a: 38

1982a: 192
1982a: 196
1982a: 213
1920: 710
1982a: 221


Klimaszewski 1982b: 328
Klimaszewski 1979:73

ibid.: 193
Klimaszewski 1982a: 194
ibid.: 198
Klimaszewski 1982a: 204
ibid.: 211
Klimaszewski 1982a: 217
Klimaszewski 1982a: 221
ibid.: 226



1857a: 6 Schwarz 1878:440
1920:721 ibid.

1920: 722 ibid.: 723

Frank: Florida Staphylinidae

Tinotus Sharp
T. amplus Notman
T. brunnipes Notman
T. planulus Notman

*Myrmecocephalus Macleay
M. cingulatus (LeConte)
M. concinnus (Erichson)
M. gracilis (Casey)
*Falagria Leach
F. dissecta Erichson
*Aleodorus Say
A. partitus (LeConte)
*Lissagria Casey
L. laticeps (Notman)
*Borboropora Kraatz
B. sulcifrons (Casey)

Acrotona Thomson
A. hebeticornis Notman
A. picescens Notman
Asthenesita Casey
A. pallens Casey
Atheta Thomson
A. aspericauda Bernhauer
A. coriaria (Kraatz)
A.fulviceps Notman
A. macrops Notman
Trichiusa Casey
T. ursina Notman

Dexiogyia Thomson
D. tenuicauda (Casey)
Gnypeta Thomson
G. floridana Casey
Meronera Sharp
M. venustula (Erichson)

Apalonia Casey
A. seticornis Casey
Dinocoryna Casey
D. bisinuata Casey
= D. foreli (Wasmann)
Ecitocala Frank
E. rugosa Frank
Ecitoxenidia Wasmann
E. alabamae Seevers
Microdonia Casey
M. nitidiventris (Brues)
M. occipitalis Casey
Zyras Stephens
Z. angustulus (Casey)

Agaricochara Kraatz
A. anomala (Notman)

1920: 723
1920: 724



ibid.: 725



Hoebeke 1985: 970

1866:371 Hoebeke 1985:986

1920:732 Hoebeke 1985: 999

1893:348 Hoebeke 1985:1005


1920:730 ibid.
1920: 729 ibid.

1893:366 ibid.

1907:400 ibid.
1856:282 Fran
1920:726 ibid.:
1920:725 ibid.:

1920:725 ibid.

1911:17 ibid.

1906:195 ibid.

1839:55 Schw

k 1981a: 388

arz 1878:440

1906:324 ibid.


ibid.: 321

1981b: 145 Frank & Thomas 1981a: 145


Frank & Thomas 1981a: 139

1904:250 Frank & Thomas 1981a: 141
1893:319 Frank & Thomas 1981a: 141

1893:323 ibid.: 324

1920:718 ibid.


374 Fl

Bolitochara Mannerheim
B. tenuicornis (Notman)
Coenonica Kraatz
C. puncticollis Kraatz
Elachistarthron Notman
E. ambiguum Notman
Eumicrota Casey
E. atoma Casey
E. corruscula (Erichson)
E. minutissima Casey
E. social (Erichson)
Euvira Sharp
E. debilis Sharp
Gyrophaena Mannerheim
G. keeni Casey
G. laetula (Casey)
Heterota Mulsant & Rey
H. plumbea (Waterhouse)
Homalota Mannerheim
H. plan (Gyllenhal)
Orthodiatelus Notman
0. innotabilis Notman
*Phanerota Casey
P. brunnessa Ashe
P. carinata Seevers
P. cubensis Casey
P. dissimilis (Erichson)
P. fasciata (Say)
Placusa Erichson
P. despecta Erichson
Schistacme Notman
S. obtusa Notman
Thecturota Casey
T. fracta Casey
T. nevadica Casey

Philotermes Kraatz
P. cubitopilis Seevers
P. fuch-ii Kraatz
P. werneri Seevers

Oligota Mannerheim
0. chrysopyga Kraatz
0. parva Kraatz
0. testaceorufa Bernhauer
0. zonata Bierig

orida Entomologist 69(2) June, 1986

1920:714 ibid.

1857b: 46 Frank & Thomas 1984b: 416

1920:715 ibid.: 716


Seevers 1951:738
Seevers 1951:734
Seevers 1951:739
Seevers 1951:736

1883:281 Bierig 1939:25

1911:185 Seevers 1951:682
1906:300 Notman 1920:719

1858:6074 Frank & Thomas 1984b: 412

1810:402 Schwarz 1878:440

1920:716 ibid.: 717


ibid.: 240
Ashe 1986:241
Ashe 1986:238
Ashe 1986:241
Ashe 1986:241

1839:197 Schwarz 1878: 440

1920:712 ibid.: 713

1911:209 Notman 1920:713
1911:209 Notman 1920:713

1957: 255
1857a: 15

1934b: 115

Seevers 1938:433
ibid.: 256

Frank 1975b:280
Notman 1920:710
Frank 1975b:280
Frank 1975b:280

Frank: Florida Staphylinidae 375

revision has been published since 1968, the right column gives the author, date and page
of the revision. Provision of this third column allows verification of each record in the
original literature. Omission of such a third column in most checklists and catalogs
allows errors to be perpetuated in the literature (see Seevers 1978, for comments on
such errors).
This checklist contains no new distributional records. It includes all known records
published in the primary literature. The secondary literature (catalogs by Leng 1920
and supplements; Moore and Legner 1975) posed some difficulties. The Leng (1920)
catalog listed > 20 species whose recording for Florida I could not discover in the
primary literature. Eventually, with the help of D. R. Whitehead (USDA, Washington,
DC) I discovered in the Smithsonian Institution Archives, a reprint of Schwarz's (1878)
paper annotated by Schwarz with most of these species names. This appears to be the
source of records cited by Leng (1920). I have considered this annotated reprint as a
valid publication (Schwarz MS in the REFERENCES CITED) and have accepted these
records (see e.g., 'Leng 1920:98 SM' in the right column for Osorius planifrons). I have
arbitrarily accepted existence of three additional species recorded apparently for the
first time in the Leng (1920) catalog (Hypotelus hostilis, Anotylus rugosus, and
Mycetoporus americanus) as validly recorded for Florida. I have deliberately excluded
Thoracophorus fletcheri Wendeler, apparently recorded for the first time in the Moore
and Legner (1975) catalog, because Moore (in litt.) could not recall how this came to be
recorded for Florida.


The foregoing pages list 324 species belonging to 122 genera as occurring in Florida.
Illustrated keys to the adults of Florida species of most of these genera are lacking.
Autecological studies have not been performed in Florida. This known fauna' is admit-
ted to be only a part of the 'total fauna', so then what is the size of the 'total fauna' and
how does the species/area relationship compare with those of other faunas?
The Florida staphylinid fauna is estimated conservatively as about 450 species as
follows: (1) the taxa Omaliinae (part), Bledius, Oxyporus, Edaphus, Xantholininae,
Erichsonius, Neobisnius, Platydracus, Quediini, Lordithon, Coproporus,
Sepedophilus, Tachinus, Tachyporus, Deinopsini, Myllaenini, Aleochara, Falagriini
and Phanerota have been revised taxonomically since 1968, and they were recordeli as
having 73 species in Florida before revision, 121 after revision; (2) the Florida list in
1968 (before the initiation of a series of modern revisions) contained 249 species, so if
all taxa had been revised instantaneously the list could be estimated to contain 249 X
121/73 = 413 species; (3) even in the revised taxa not all Florida species were recorded
during revision, and I am aware that seven species remain to be added to the 121
mentioned above (a 6% increase); (4) additionally, the presence of 14 species belonging
to genera unrecorded for Florida in 1968 has subsequently been recorded, without
taxonomic revision, a 3% increase (14 as percentage of 413); (5) therefore the estimate
of 413 should be increased by roughly 9% to 450. Possible sources of error may be an
underestimation of diversity within Aleocharinae: by analogy with the European fauna
this subfamily might be expected to have a greater number of species than predicted.
Florida has an area of 152,000 km2 and a conservatively estimated staphylinid fauna
of 450 species. The British Isles have an area of 314,000 kmn and a staphylinid fauna of
990 species [counted in Pope (1977)], whereas "Middle Europe" [Denmark to Austria]
has an area of about 725,000 km2 and a staphylinid fauna of 1,790 species [counted in
Freude, Harde and Lohse (1964, 1974)]. On the basis of the species/area relationships
for the two European areas and the slight slopes of most species/area curves (Williams

Florida Entomologist 69(2)

1964), the staphylinid fauna of Florida is depauperate relative to that of western
About a third of the Florida staphylinid fauna (1-324/450) remains unrecorded, so
an accurate assessment of the origin of its components must await further study. The
largest component is nearctic, with smaller neotropical, cosmotropical, and cosmopolitan
components. Absence of a larger nearctic component probably has to do with absence
of a mountain chain, so that species restricted to high altitudes in the southeastern USA
are excluded from Florida. Absence of a larger neotropical component may well have
to do with the distance of mainland Florida from large land areas to the south and west;
even though a chain of small islands (the Florida Keys) leads southward, these may be
inhospitable to many staphylinid species because they lack fresh water and other sorts
of habitats elsewhere occupied by staphylinids.
It is estimated that about 126 species (450-324) remain to be added to the Florida
list. Verification that this number is a reasonable estimate comes from my collection:
specimens of about 60 species not mentioned in Table 1 await taxonomic treatment
before incorporation. I find it entirely conceivable that a further 66 (126-60) species
await discovery. Future work will consist not merely of adding species names to the
list, but in changes of generic and even of tribal assignments, in recognition of additional
synomymies, in realization that some species names on the list are due to misidentifica-
tion or were recorded from Florida in error, and in preparation of illustrated keys for
identification. All of this forms a necessary background to the mapping of species distri-
butions and to the beginning of an understanding of the ecological roles performed by
these species.


I am indebted to D. R. Whitehead (USDA, Washington, DC) for a copy of an anno-
tated reprint of the Schwarz (1878) paper from the Smithsonian Institution Archives.
Through the kindness of A. F. Newton (Harvard University) I am permitted to cite
Florida records for Platydracus species from his unpublished Ph.D. thesis (it should be
noted that this thesis also contains descriptions of new species of Platydracus, three of
which are recorded from Florida; I have chosen not to list two additional Platydracus
and one Ontholestes species known to A. F. Newton (in litt.) to occur in Florida).
Reviews of this manuscript by J. S. Ashe (Field Museum, Chicago) and A. F. Newton
are gratefully acknowledged. The abstract was translated into Spanish by J. R. Rey
(Vero Beach). Institute of Food and Agricultural Sciences, University of Florida, Jour-
nal Series no. 6748. The author is a Research Associate of the Florida State Collection
of Arthropods.


ASHE, J. S. 1986. Phanerota cubenis and Phanerota brunnessa n. sp., with a key to
the species of Phanerota occurring in Florida (Coleoptera: Staphylinidae). Flor-
ida Ent. 68: 236-245.
AUBE, C. 1850. Description de quelques insects coldopteres appartenant a l'Europe et
A 1'Alg6rie. Ann. Soc. Ent. France (sir. 2) 8: 299-346.
AUSTIN, E. P. 1876. On the species of Sunius and Paederus found in the United States.
Proc. Boston Soc. Nat. Hist. 19: 4-11.
BENJAMIN, R. K. 1983. Comparative morphology of Idiomyces and its possible allies
Diplomyces, Sandersoniomyces, Symplectromyces, and Teratomyces (Ascomy-
cetes: Laboulbeniales). Aliso 10: 345-381.

June, 1986

Frank: Florida Staphylinidae

BERNHAUER, M. 1901. Neue exotische Arten der Gattung Aleochara Gray. Stettiner
Ent. Ztg. 62: 366-373.
1907. Neue Aleocharini aus Nordamerika (Col.) (3. Stuck). Deutsche Ent.
Ztschr. (1907): 381-405.
---. 1910. Beitrag zur Kenntnis der Staphyliniden-Fauna von Zentralamerika. Verh.
Zool.-Bot. Ges. Wien 60: 350-393.
1923. Coleopterologische Beitrage. Ent. Tidskr. 44: 141-146.
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Florida Entomologist 69(2)

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June, 1986

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