Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00029
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 2002
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
 Record Information
Bibliographic ID: UF00098813
Volume ID: VID00029
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text

Wetterer & O'Hara: Ants of the Dry Tortugas




We examined the distribution of ants on the DryTortugas, the outermost of the Florida Keys.
These small islands are important nesting grounds for sea turtles and sea birds. We sampled
ants on the five vegetated islands: Garden Key, Loggerhead Key, Bush Key, Long Key, and
East Key. We found 17 ant species, seven of which had not been recorded from the Dry Tortu-
gas. Paratrechina longicornis was common on four of five islands. Otherwise, the five islands
had strikingly different ant faunas. On Garden Key, Solenopsis geminata was dominant, and
Pheidole megacephala was absent. On Loggerhead Key and Bush Key, Ph. megacephala was
dominant, and S. geminata was absent. On Long Key, vegetated primarily with mangrove, the
most common ant was the arboreal Pseudomyrmex elongatus. On sparsely-vegetated East Key,
all ants were uncommon. Twenty ant species are now known from the Dry Tortugas: eight
New World and 12 Old World species. Only Solenopsis globularia is an undisputed Florida
native. Florida specimens previously identified as C. tortuganus are actually Camponotus zona-
tus; true C. tortuganus are Bahamian. The two dominant ant species, Ph. megacephala and
S. geminata, may pose a threat to native fauna, including sea turtle and sea bird nestlings.

Key Words: Camponotus tortuganus, exotic species, Solenopsis geminata, tramp ants,
Pheidole megacephala


Examinamos la distribuci6n de hormigas en las Dry Tortugas, las islas mas exteriores de las
Florida Keys. Estas islas pequenas son areas importantes para nidos de las tortugas del mar
y de los pdjaros del mar. Muestreamos hormigas en las cinco islas vegetadas: Garden Key,
Loggerhead Key, Bush Key, Long Key, y East Key. Encontramos 17 species hormigas; siete
de estas no fueron registrados de Dry Tortugas. Paratrechina longicornis eran comunes en
cuatro de cinco islas. De otras maneras, las cinco islas tenian llamativo diversos faunas de
la hormiga. En Garden Key, Solenopsis geminata era dominant, y Pheidole megacephala es-
taba ausente. En Loggerhead Key y Bush Key, Ph. megacephala era dominant, y S. gemi-
nata estaba ausente. En Long Key, vegetada sobre todo con el mangle, la hormiga mas co-
mun era la arboreal Pseudomyrmex elongatus. En East Key, una isla sin much vegetaci6n,
las hormigas en general eran raro. Viente species de las hormigas estan conocidas de las
Dry Tortugas ahora: ocho species del mundo viejo y 12 del mundo nuevo. Solamente Sole-
nopsis globularia esta native de la Florida sin disputaci6n. Los especimenes de la Florida
identificada previamente como C. tortuganus es realmente Camponotus zonatus; C. tortug-
anus verdadero es de las Bahamas. Las dos species dominantes de la hormiga, Ph. megace-
phala y S. geminata, pueden plantear una amenaza a la fauna native, incluyendo las
juveniles de la tortuga marinas y del pdjaro del mar.

The Dry Tortugas (2433'-2444'N; 8246-
8315'W) are the outermost of the Florida Keys,
located 100-112 km West of Key West, Florida.
These islands and the surrounding waters were
declared a wildlife refuge in 1908, a national mon-
ument in 1935, and a national park in 1992. Most
biological research in the Dry Tortugas area has
concerned the diverse marine biota in the coral
reefs surrounding the keys (see Schmidt & Pikula
1997 for an annotated bibliography of research).
The Dry Tortugas are also important ecologically
as the feeding and nesting grounds for loggerhead
turtles (Caretta caretta), green sea turtles (Chelo-
nia mydas), and numerous sea birds, including
masked boobies (Sula dactylatra), brown boobies
(Sula leucogaster), sooty terns (Sterna fuscata),

and magnificent frigate birds (Fregata magnifi-
cens) (Robertson 1964; Lenihan 1997). The is-
lands are far from pristine and have numerous
non-indigenous plants and animals (Stoddart &
Fosberg 1981), such as wetland nightshade
(Solanum tampicense; Fox & Bryson 1998) and
the Old World house gecko (Hemidactylus
mabouia; Meshaka & Moody 1996). In the
present study, we examined the distribution of
native and exotic ants on the Dry Tortugas.
Hurricanes periodically reshape the Dry Tor-
tugas, altering the size and even the number of
keys (Stoddart & Fosberg 1981). Currently, there
are seven keys: (from west to east) Loggerhead
Key, Garden Key, Bush Key, Long Key, Sand Key,
Middle Key, and East Key. Garden Key and Log-

Florida Entomologist 85(2)

gerhead Key are the largest and the only inhab-
ited islands. At present, about ten park staff
members live on these two keys. Garden Key is
the site of Fort Jefferson, a 19th century structure
that occupies most of the island. Plans are being
made to expand tourist facilities on Garden Key
to accommodate the large number of people who
visit the Dry Tortugas each year (almost 90,000 in
1999; Dustin 1999). Loggerhead Key has a Coast
Guard Station and lighthouse and is the former
site of Tortugas Laboratory, a research station ad-
ministered by the Carnegie Institute from 1904 to
1939 (Mayer 1902; Schmidt & Pikula 1997). Bush
Key is an important breeding site for many sea-
birds. It is currently connected by a sand spit to
Long Key, which is largely vegetated with man-
groves. East Key is a small, sparsely vegetated is-
land far to the east of the other keys. Finally, the
smallest two islands, Sand Key and Middle Key,
are simple sand bars without vegetation.
The aim of my study was to establish a base-
line of information on the distribution of ants in
the Dry Tortugas, as a first step in evaluating the
potential impact of ants on native species on the


In addition to compiling published records on
ants, we surveyed the ants on the five vegetated
islands of the Dry Tortugas, Loggerhead Key,
Garden Key, Bush Key, Long Key, and East Key,
on 17-19 March 2000 during daylight hours, with
the help of undergraduate assistants from the
Honors College of Florida Atlantic University. We
did not sample on the unvegetated Sand Key and
Middle Key because these islands are periodically
submerged for extended periods, and thus have
no permanent ant populations.
To evaluate the overall variety of ant species
present on each island, we conducted intensive
hand-collecting, searching accessible surfaces
and turning over rocks and logs.
To evaluate the relative dominance of different
ant species on Garden Key, we surveyed ants at
40 bait stations, each with two bait cards. A bait
card consisted of a 7.6 x 12.7 cm index card with
approximately 2 g of tuna bait placed at its center.
We folded each card in half and placed it flush
with the ground about one meter from the road,
trail, transect, or building, using a small rock or
branch to weigh it down. We returned to each sta-
tion 1.5-2.0 hours later and placed the cards,
along with all ants on the bait, in plastic bags.


Published Ant Records

In addition to one questionable record, we found
reports of 14 ant species from the Dry Tortugas

(Table 1; Emery 1895; Mayor 1922; Wheeler 1932;
MacArthur & Wilson 1967; Deyrup et al. 1988).
The questionable record from the Dry Tortugas
is the earliest. Emery (1895) described Campono-
tus tortuganus based on specimens that he be-
lieved were collected by T. Pergande in the Dry
Tortugas. Mackay (pers. comm.), however, found
that the collection locality for Emery's C. tortuga-
nus holotype specimen was labeled in Emery's
handwriting as Egg Island, Bahamas. Also, Mackay
(pers. comm.) has received additional specimens
from the Bahamas that match the C. tortuganus
holotype. In contrast, Mackay (pers. comm.)
found that all "C. tortuganus" specimens from
Florida that he examined are actually Campono-
tus zonatus (formerly Camponotus conspicuous
zonatus). Mackay (pers. comm.) suggested that
Emery erroneously attributed the Dry Tortugas
as the type locality for C. tortuganus and that true
C. tortuganus is Bahamian.
The next mention of ants from the Dry Tortu-
gas come from Cole (1906), who worked at Tortu-
gas Laboratory on Loggerhead Key, and reported
that he "studied the reactions of ants" there. It ap-
pears that this work was never published, and we
could not find any Cole specimens.
Mayor (1922) studied an Old World exotic ant,
Monomorium destructor, on Loggerhead Key, de-
scribing it as "a great pest in the wooden build-
ings of Tortugas Laboratory, making its nests in
crevices of the woodwork. So voracious are these
insects that we are obliged to swing our beds from
the rafters and to paint the ropes with a solution
of corrosive sublimate, while all tables must have
tape soaked in corrosive sublimate wrapped
around their legs if ants are to be excluded from
them. These pests have the habit of biting out
small pieces of skin, and I have seen them kill
within 24 hours rats which were confined in
cages." Mayor (1922) mentioned no other ant spe-
cies on the island.
Wheeler (1932), in his review of ant records
from Florida, listed 91 species, including only two
species from the Dry Tortugas, both collected by
Pergande (again, presumably from Garden Key):
Camponotus tortuganus and Tetramorium
guineense (now Tetramorium bicarinatum).
Wilson's (1964) survey of the ants of the Flor-
ida Keys did not include any records from the Dry
Tortugas. MacArthur & Wilson (1967) stated that
13 ant species "have succeeded in colonizing the
Dry Tortugas," but only mentioned two species by
name, Paratrechina longicornis, and Pseudo-
myrmex elongatus. MacArthur & Wilson (1967)
reported that on the Dry Tortugas, Paratrechina
longicornis was "an overwhelmingly abundant
ant and has taken over nest sites that are nor-
mally occupied by other species in the rest of
southern Florida: tree-boles, usually occupied by
species of Camponotus and Crematogaster, which
are absent from the Dry Tortugas; and open soil,

June 2002

Wetterer & O'Hara: Ants of the Dry Tortugas



Loggerhead Garden Bush Long East Status

Brachymyrmex obscurior Forel f2 NW/N?
*Camponotus tortuganus Emery a* ?
Camponotus zonatus Emery fO f NW/N?
Cardiocondyla ectopia Snelling f f OW/E
Cardiocondyla emery Forel dfO OW/E
Cardiocondyla venustula Wheeler fl f OW/E
Cardiocondyla wroughtoni (Forel) fO OW/E
Hypoponera opaciceps (Mayr) d NW?/N?
Monomorium destructor (Jerdon) b OW/E
Monomorium floricola (Jerdon) f df6 f OW/E
Paratrechina bourbonica (Forel) f df5 f f OW/E
Paratrechina guatemalensis (Forel) e f NW/E
Paratrechina longicornis (Latr.) f cdefl4 f f OW/E
Pheidole megacephala (Fabr.) f de f OW/E
Pseudomyrmex cubaensis Forel f NW/N?
Pseudomyrmex elongatus (Mayr) dfO cf f f NW/N?
Solenopsis geminata (Fabr.) def28 NW/N?
Solenopsis globularia (Smith) f fl f NW/N
Tapinoma melanocephalum (Fabr.) def3 OW/E
Tetramorium bicarinatum Nylander a OW/E
Tetramorium caldarium (Roger) df6 f OW/E
Collectors/Authors: a* = Pergande (in Emery 1895) a = Pergande (in Wheeler 1932); b= Mayor (1922); c = MacArthur & Wilson (1967); d = Winegarner
(in Deyrup et al. 1988); e = Carlin (in Deyrup et al. 1988); f = present study (including # of Garden Key bait stations with the species present); = ap-
parently erroneous record. Status (from Deyrup et al. 1988, 2000): NW = New World origin; OW = Old World origin. N = native; E = exotic; ? = status in

normally occupied by crater nests of Conomyrma
[now Dorymyrmex] and Iridomyrmex [now Fore-
lius], which genera are also absent from the Dry
Tortugas." MacArthur & Wilson (1967) also re-
ported that Pseudomyrmex elongatus was "the
only member of the arboreal assemblage that has
colonized the Dry Tortugas, where it has a red
mangrove swamp on Bush Key virtually to itself.
Yet it is still limited primarily to thinner twigs in
the canopy and, unlike Paratrechina longicornis,
has not increased perceptibly in abundance."
Deyrup et al. (1988), in their review of the ants
of the Florida Keys, included records from recent
collections in the Dry Tortugas by C. Winegarner
(collected 12 April 1983; no island site locality,
presumably from Garden Key) and by N. F. Carlin
(no island site locality, presumably from Garden
Key). These two collections recorded a total of 11
ant species from the Dry Tortugas, bringing the
total number of previously published ant records
to 14 species (Table 1). Although Deyrup et al.
(1988) reported Pheidole dentata specimens col-
lected by Winegarner, these were actually mis-
identified Pheidole megacephala specimens
(M. Deyrup, pers. comm.).
New Ant Collection
We collected 17 ant species on the Dry Tortu-
gas (Table 1). Seven of these were not previously

recorded from the Dry Tortugas. One species, Bra-
chymyrmex obscurior, belongs to a genus in which
species boundaries for Florida specimens have
not been adequately defined (Deyrup 1991). We
did not find four species recorded in previous
studies: Camponotus tortuganus (see discussion
above), Tetramorium bicarinatum, Monomorium
destructor, and Hypoponera opaciceps.
Paratrechina longicornis was common on four
of the five islands. Otherwise, the five islands had
strikingly different ant faunas. On Garden Key,
Solenopsis geminata and Paratrechina longicor-
nis were by far the most common ants at baits,
found, respectively at 28 and 14 of the 40 bait sta-
tions (Table 1). Solenopsis geminata was the dom-
inant ant on the ground, while Paratrechina
longicornis was the most common ant up in trees.
On Loggerhead Key and Bush Key, Pheidole
megacephala and Paratrechina longicornis were
the most common ants. Solenopsis globularia,
which we found only once on Garden Key, was
common on Loggerhead and Bush Key. On Long
Key, vegetated primarily with mangrove, the
most common ant was the mangrove-inhabiting
Pseudomyrmex elongatus. On sparsely vegetated
East Key, all ants were uncommon.
We collected Camponotus specimens on both
Garden Key and on Long Key. On Garden Key, we
found Camponotus workers foraging in trees and

Florida Entomologist 85(2)

we collected dead Camponotus workers and
males from the windowsills of a park warden's
house. On Long Key, we found Camponotus work-
ers inside a bamboo pole near the beach. Mackay
(pers. comm.) examined all my Camponotus spec-
imens and found that they did not match Emery's
C. tortuganus holotype, but instead were Cam-
ponotus zonatus.


Excluding Camponotus tortuganus, which ap-
pears to be an erroneous record, 20 ant species
are recorded from the Dry Tortugas (Table 1), 12
of Old World origin and eight from the New World.
Only one species, Solenopsis globularia, is an un-
disputed Florida native. Of the other seven New
World species, Deyrup et al. (2000) considered
Paratrechina guatemalensis as an exotic in Flor-
ida and listed Brachymyrmex obscurior, Cam-
ponotus tortuganus (= Camponotus zonatus),
Solenopsis geminata, Pseudomyrmex cubaensis,
and Pseudomyrmex elongatus as "dubious na-
tives." In addition, although Deyrup et al. (1988)
considered Hypoponera opaciceps as native to the
Florida Keys, Dlussky (1994) has proposed that
this species is actually of Old World origin.
We found a strikingly different ant fauna on
each of the five vegetated keys of the Dry Tortu-
gas. On Garden Key, Solenopsis geminata was the
dominant ant species, occurring at 70% of the bait
stations, usually with several hundred workers
on each bait card. Their presence on the island
was noted by visitors who were often stung on the
feet by these fire ants. Although Pheidole mega-
cephala may have occurred at one time on Garden
Key, we found none. In contrast, we found no
S. geminata on any other key in the Dry Tortugas.
Instead, on Loggerhead Key and Bush Key,
Ph. megacephala was the dominant ant.
Wheeler (1908) similarly observed mutually
exclusive distributions of Ph. megacephala and
S. geminata on two Caribbean islands: "I devoted
ten days to a careful study of the ant-fauna of the
little island of Culebra off the eastern coast of
Porto Rico without seeing a single specimen of
Ph. megacephala. This island is, however, com-
pletely overrun with a dark variety of the vicious
fire-ant (Solenopsis geminata). One day, on visit-
ing the island of Culebrita, which is separated by
a shallow channel hardly a mile in width from the
eastern coast of Culebra, I was astonished to find
it completely overrun with Ph. megacephala. This
ant was nesting under every stone and log, from
the shifting sand of the sea-beach to the walls of
the light-house on the highest point of the island.
The most careful search failed to reveal the pres-
ence of any other species, though the flora and
physical conditions are the same as those of Cul-
ebra. It is highly probable that Ph. megacephala,
perhaps accidentally introduced from the island

of St. Thomas a few miles to the east, had exter-
minated all the other ants which must have pre-
viously inhabited Culebrita. The absence of
megacephala on Culebra is perhaps explained by
the presence of the equally prolific and pugna-
cious fire-ant" (Wheeler 1908). Mutually exclu-
sive distributions between pairs of dominant ant
species has also been noted for many other spe-
cies, such as between Ph. megacephala and Line-
pithema humile on Madeira (Schmitz 1896) and
Bermuda (Haskins & Haskins 1965).
Paratrechina longicornis coexisted at high den-
sities with both Solenopsis geminata (on Garden
Key) and Ph. megacephala (on Loggerhead Key
and Bush Key). An Old World native,P longicornis
now has a pantropical distribution and commonly
occurs at high densities in disturbed habitats (e.g.,
see Wetterer 1998, Wetterer et al. 1999).
Remarkably, Monomorium destructor, found in
such overwhelming abundance on Loggerhead
Key by Mayor (1922), was not found on any island
in the Dry Tortugas by later collectors. It is possi-
ble that M. destructor only dominated the Tortu-
gas Lab buildings which have subsequently been
torn down. Deyrup et al. (1988) wrote that M. de-
structor "is not common in Florida, and may be on
the decline in the Keys, except in Key West where
it is a dominant urban ant." Deyrup (1991) wrote
that M. destructor "is spectacularly common on
Key West." It is possible that Ph. megacephala
and/or S. geminata competitively excluded M. de-
structor from the Dry Tortugas.
The two dominant ant species of the Dry Tortu-
gas, S. geminata and Ph. megacephala, may pose a
threat to the native fauna of the Dry Tortugas.
Pheidole megacephala, an African native, is now a
pantropical pest. In lowland Hawaii, it has been
implicated in the extermination of much of the en-
demic fauna (Zimmerman 1948). Zimmerman
(1948) reported that the "voracious immigrant
ant," Ph. megacephala, eliminates most endemic
insects throughout its range. Solenopsis geminata
is an important exotic pest on Pacific islands (Wet-
terer 1997). On the Dry Tortugas, this species may
prey on hatchling sea turtles and birds (e.g., see
Kroll et al. 1973; Wetterer & Wood 2001).
Two highly-destructive exotic ant species
known from the Florida Keys that also pose im-
portant threats to native species (e.g., see Vinson
1994; Jourdan 1997; Wetterer & Wood 2001) are
the red imported fire ant, Solenopsis invicta, and
the little fire ant, Wasmannia auropunctata (Dey-
rup et al. 1988, Horvitz 1997). These species are
not yet known from the Dry Tortugas. Construc-
tion of new tourist facilities on the Dry Tortugas
may result in the introduction of these and other
exotic ant species, arriving with building material
or heavy machinery during the building stage, or
accompanying visitors in their camping supplies
and picnic baskets. On the Dry Tortugas, a rela-
tively small effort in preventing the further inva-

June 2002

Wetterer & O'Hara: Ants of the Dry Tortugas

sion of exotic ants could have a great effect in
protecting the native inhabitants of these ecolog-
ically important islands.


We thank S. Paquette, N. McClain, L. Eurich, J. Chang,
and L. Arencibia (students from Florida Atlantic Univer-
sity's Honors College) for field assistance; S. Cover (Har-
vard University) for ant identification; W. Mackay (Texas
Tech) for unpublished information concerning Campo-
notus; A. Wetterer, M. Wetterer, and M. Deyrup (Archbold
Field Station) for comments on this manuscript; S. Bass,
P. Taylor, and W. Meshaka (U.S. National Park Service)
for logistic support; Florida Atlantic University and the
National Save the Sea Turtle Foundation for financial


COLE, L. J. 1906. Ant Studies. Carnegie Inst. Washing-
ton, Year Book 5: 110.
DEYRUP, M. A. 1991. Exotic ants of the Florida Keys.
Proc. 4th Symp. Nat. Hist. Bahamas. pp. 15-22.
1988. A review of the ants of the Florida Keys. Flor-
ida Entomol. 71: 163-176.
Deyrup, M., L. Davis, and S. Cover. 2000. Exotic ants in
Florida Trans. Amer. Entomol. Soc. 126: 293-326.
DLUSSKY, G. M. 1994. Zoogeography of southwestern
Oceania, pp. 48-93. In Y. G. Puzatchenko, S. I. Golo-
vatch, G. M. Dlussky, K. N. Diakonov, A. A. Zakharov,
and G. A. Korganova [eds.], Animal population of the
islands of Southwestern Oceania. Nauka Publishers,
Moscow, Russia.
DUSTIN, D. 1999. The curious history of Dry Tortugas
National Park. Parks Recreat. 34: 126-133.
EMERY, C. 1895. Beitage zur Kenntniss der nordameri-
kanischen Ameisenfauna. Zoolog. Jahr. Abtheil. Sys-
tem. Geogr. Biol. Tiere 8: 257-360.
Fox, A. M., AND C. T. BRYSON. 1998. Wetland night-
shade (Solanum tampicense): a threat to wetlands in
the United States. Weed Technol. 12: 410-413.
HASKINS, C. P., AND E. F. HASKINS. 1965.Pheidole mega-
cephala and Iridomyrmex humilis in Bermuda-equi-
librium or slow replacement? Ecology 46: 736-740.
HORVITZ, C. C. 1997. The impact of natural distur-
bances, pp. 63-74. In D. Simberloff, D. C. Schmitz,
and T. C. Brown [eds.], Strangers in Paradise. Impact
and management of nonindigenous species in Flor-
ida. Island Press, Washington, D.C. 467 pp.
JOURDAN, H. 1997. Threats on Pacific islands: the spread
of the tramp ant Wasmannia auropunctata (Hymenop-
tera: Formicidae). Pacific Conserv. Biol. 3: 61-64.

KROLL, J. C., K. A. ARNOLD, AND R. F. GOTIE. 1973. An
observation of predation by native fire ants on nest-
ling Barn Swallows. Wilson Bull. 85: 478-479.
LENIHAN, D. J. 1997. The Tortuga Triangle. Natural
Hist. 106: 36-41.
MACARTHUR, R. H., AND E. 0. WILSON. 1967. The The-
ory of Island Biogeography. Princeton Univ. Press.
MAYER, A. G. 1902. The Tortugas, Florida as a station for
research in biology. Science 17: 190-192.
MAYOR, A. G. 1922. The tracking instinct in a Tortugas
ant. Papers Tortugas Lab. 18: 101-107.
MESHAKA, W. E. JR., AND B. A. MOODY. 1996. The Old
World tropical house gecko (Hemidactylus mabouia)
on the Dry Tortugas. Florida Scientist 59: 115-117.
ROBERTSON, W. B., JR. 1964. The terns of the Dry Tor-
tugas. Bull. Fla. State Mus. 8(1): 1-93.
SCHMIDT, T. W., AND L. PIKULA. 1997. Scientific studies
on Dry Tortugas National Park: an annotated bibli-
ography. Atoll Res. Bull. 449: 2-9.
SCHMITZ, E. 1896. As formigas da Madeira. Ann. Sci.
Nat. 3: 55-58.
STODDART, D. R., AND F. R. FOSBERG. 1981. Topographic
and floristic change, Dry Tortugas, Florida, 1904-
1977. Atoll Res. Bull. 253: 1-55.
VINSON, S. B. 1994. Impact of the invasion of Solenopsis
invicta (Buren) on native food webs, pp. 240-258. In
D. F. Williams [ed.], Exotic ants. Biology, impact, and
control of introduced species. Westview Press, Boul-
der, CO. 322 pp.
WETTERER, J. K. 1997. Alien ants of the Pacific islands.
Aliens 6: 3-4.
WETTERER, J. K. 1998. Ants on Cecropia trees in urban
San Jos6, Costa Rica. Florida Entomol. 81: 118-121.
1999. Ecological dominance by Paratrechina longicor-
nis (Hymenoptera: Formicidae), an invasive tramp
ant, in Biosphere 2. Florida Entomol. 82: 381-388.
WETTERER, J. K., AND L. D. WOOD. 2001. Distribution
and impact of ants on a sea turtle nesting beach in
Palm Beach County, Florida. Proc. 21st Ann. Symp.
Sea Turtle Biol. Conserv. NOAA Tech. Mem. NMFS-
SEFSC, in press.
WHEELER, W. M. 1908. The ants of Porto Rico and the Vir-
gin Islands. Bull. Am. Mus. Nat. Hist. 24: 117-158.
WHEELER, W. M. 1932. A list of the ants of Florida with
descriptions of new forms. J. New York Entomol. Soc.
40: 1-17.
WILSON, E. 0. 1964. Ants of the Florida Keys. Breviora
210: 1-14.
ZIMMERMAN, E. C. 1948. Insects of Hawaii. Volume 1.
Introduction. Univ. Hawaii Press, Honolulu, HI.

Florida Entomologist 85(2)

June 2002


1U.S.D.A. Forest Service, Forestry Sciences Laboratory, 320 Green St., Athens, GA 30602

2U.S.D.A. Forest Service, Department of Forest Resources, Clemson University, Clemson, SC 29634


As a result of human activity, longleaf pine (Pinus palustris Miller) forests in the southern
United States have been lost or drastically altered. Many of the plant species that histori-
cally occupied those forests now persist only as remnants and are classified as threatened or
endangered. In order to safeguard such species, a better understanding of their pollination
ecology is needed. We identified insect visitors and potential pollinators of Harperocallis
flava (McDaniel) (Amaryllidaceae), Macbridea alba Chapman (Lamiaceae) and Scutellaria
floridana Chapman (Lamiaceae) that occur in longleaf pine habitat on the Apalachicola Na-
tional Forest in Florida. We observed that potential pollinators of H. flava were Halictidae,
of M. alba were bumble bees (Apidae: Bombus), and of S. floridana were Megachilidae and
Halictidae. However, the rates at which these insects visited the flowers were very low. Our
results raise important concerns about how forest management practices affect the survival
of rare plants, as well as their pollinators.

Key Words: Harperocallis flava (McDaniel), Macbridea alba Chapman, Scutellaria floridana
Chapman, Pinus palustris Miller, threatened species, endangered species


Como resultado de la actividad humana, los bosques de pino de hoja larga (Pinus palustris
Miller) del sureste de los Estados Unidos han desaparecido o han sido drasticamente altera-
dos. Muchas de las species de plants que historicamente ocupaban estos bosques persisten
en la actualidad como restos y estan clasificadas como amenazadas o en peligro de extinction.
Para salvaguardar estas species es necesitario entender la ecologia de su polinizacion. Para
ello, identificamos los insects visitadores y polinizadores potenciales de Harperocallis flava
(McDaniel) (Amaryllidaceae), Macbridea alba Chapman (Lamiaceae) y Scutellaria flori-
diana Chapman (Lamiaceae) que existen en el habitat del pino de hoja larga del Bosque Na-
cional de Apalachicola en Florida. Observamos que los polinizadores potenciales de H. flava
eran Halictidae, de M. alba eran abejorros (Apidae: Bombus), y de S. floridiana eran Mega-
chilidae y Halictidae. Sin embargo, los niveles de visitacion de estos insects a las flores era
muy bajo. Nuestros resultados crean dudas importantes acerca de como practices de mante-
nimiento de los bosques afectan a la supervivencia de plants amenazadas, asi como a sus

The longleaf pine ecosystem is a conservation
priority area within the U.S. Department of Agri-
culture Conservation Reserve Program (Food Se-
curity Act of 1985, Title XII). Longleaf pine (Pinus
palustris Miller) forests once occupied >24 million
ha in the southern United States. Today, <1.3 mil-
lion ha remain as small isolated parcels (Outcalt
& Sheffield 1996). The diversity of groundcover
plants per unit area (e.g., 140 vascular plant spe-
cies/1000 m2 in mesic longleaf woodlands) illus-
trates the remarkable species richness of longleaf
pine ecosystems (Peet & Allard 1993). At least 30
endangered or threatened plant species now re-
side in the few remnant understory communities
of longleaf pine forest, and populations of at least
191 taxa of vascular plants have been reduced to
low levels (Hardin & White 1989; Walker 1993).
Thus, it is important to understand the physiolog-

ical and ecological requirements of these plants in
order to develop appropriate recovery plans.
Studies of potential management effects on rare
plants in some fire-disturbed habitats have consid-
ered the response of plants to management prac-
tices e.g., Hessl & Spackman 1995; Brewer 1999;
Lesica 1999), but not the effects on their pollinator
systems. One reason for this lack of information is
simply that the effective pollinators are unknown.
Pollinators are critical to the long-term survival of
many flowering plants because they provide a
mechanism for ensuring seed set and develop-
ment, and often facilitate gene flow between plants
and plant populations. In return for pollination
services, the flowers provide vital floral resources
for the foraging insects (Proctor et al. 1996).
A worldwide pollination crisis is at hand, and
environmental degradation from habitat destruc-

Pitts-Singer et al.:Pollinators of Rare Plants

tion, modification, or fragmentation can disrupt
plant-pollinator interactions and jeopardize their
existence (Rathcke & Jules 1993; Kearns et al.
1998). Tepedino et al. (1997) express a need for
"extended care" in conservation. When rare
plants are imperiled, their extended families of
pollinators also must be considered. It is impor-
tant to "maintain the integrity of ecosystems by
preserving interactions between plants and their
pollinators" (Tepedino et al. 1997).
Our study investigates the pollinator-plant
relationships of Harper's beauty, Harperocallis
flava (McDaniel) (Amaryllidaceae), white birds-
in-a-nest, Macbridea alba Chapman (Lamiaceae),
and Florida skullcap, Scutellaria floridana Chap-
man (Lamiaceae). All of these plants are federally
listed species and are endemic to the longleaf pine
ecosystem in the Apalachicola lowlands of the
Florida panhandle (Kral 1983; Walker 1993). They
are fire-dependent species (Kral 1983; Walker
1993), and the local USDA Forest Service uses
prescribed fires to help maintain their habitats.
For H. flava pollination, insects may not be as
important as for the other species in this study, if
they are important at all. Allozyme studies have
indicated that H. flava individuals and popula-
tions are genetically uniform (Godt et al. 1997).
Further, a preliminary study by Wagner & Spira
(1996) indicated that H. flava flowers are self-
compatible and capable of selfing. Mature fruits
were produced from flowers that were open-polli-
nated, cross-pollinated, and self-pollinated.
Harperocallis flava was first listed as an en-
dangered species in 1979 (U.S. Fish & Wildlife
Service 1992). It occurs in Franklin and Liberty
Counties, Florida, in open pineland bogs and
along moist roadside ditches. This perennial spe-
cies has a single yellow flower with 6 tepals (each
being 9-15 mm in length) produced atop a stalk
that emerges from stiff, grasslike leaves. Flower-
ing occurs from mid-April through May and fruits
mature in July. Kral (1983) reported that the den-
sity of H. flava has declined since its discovery in
1965 primarily because of lack of fire in the area.
Presently, the USDA Forest Service manages a
Franklin County location and carries out periodic
controlled burns to help maintain the open habi-
tat required by this species (U.S. Fish & Wildlife
Service 1992).
Macbridea alba was first listed as a federally
threatened species in 1992 (U.S. Fish & Wildlife
Service 1992). It occurs in open savannahs as well
as in drainage areas in pine stands in Bay, Gulf,
Franklin, and Liberty Counties, Florida (Kral
1983; U.S. Fish & Wildlife Service 1992). This pe-
rennial herb usually has only one stem, which
may be branched. Brilliant white flowers are clus-
tered in terminal compressed thyrses on erect
stems. Each flower has a green calyx (1 cm in
length) and a white, 2-lipped corolla (3 cm in
length). Flowering of M. alba is stimulated by fire

during the growing season and usually occurs
from May to July (U.S. Fish & Wildlife Service
1992, Madsen 1999). Some information available
concerning this flower species relies on insect pol-
linators. Madsen (1999) found that when insects
were excluded from M. alba flowers, almost no
seeds were produced.
Scutellaria floridana was also placed on the
federal threatened species list in 1992 (U.S. Fish
& Wildlife Service 1992). It is found in Franklin,
Liberty, and Gulf Counties, Florida. The stem is
simple or sparingly branched, and its solitary
purple flowers are well separated in the axils of
short, leafy bracts. The corolla (2.5 cm in length)
has 2 lips, the lower one being white in the mid-
dle. The preferred habitat of S. floridana is simi-
lar to that of M. alba, although it is more
restricted (Kral 1983; U.S. Fish & Wildlife Service
1992). This perennial herb is reported to flower in
May or June (Kral 1983).
We monitored arthropod visitors to flowers of
these plants, identified them, and evaluated their
importance as pollinators. The results of our
study are an important first step towards under-
standing the interactions between these rare
plants and their pollinators in this Florida
longleaf pine ecosystem.


All study sites were located on the Apalachi-
cola National Forest near Sumatra and Wilma,
Florida. Field observations were made to deter-
mine which insects frequently visited flowers and
how they behaved. In the summers of 1999 and
2000, we selected and tagged 10-25 flowers (or in-
florescences) at each site, and then monitored the
flowers for 3- or 5-min periods throughout the day
for 2-5 days. Arthropod activity and the time they
were present on the flowers were recorded during
the observation period. The visits of arthropods to
flowers were counted only when they occurred
during the observation period for that flower, al-
though visits to other tagged flowers in the vicin-
ity could be seen during this same time.
Observations were not made at the same time
each day. Because the 1999 sites did not produce
enough flowers for study in 2000, sites chosen in
2000 were not the same as those used in 1999. In
summer 2000, we also used a video camcorder to
record insect activity on focal flowers.
The video equipment consisted of a tripod-
mounted Sony CCD-VX3 Video Recorder that was
auto-controlled via a Dell Latitude Xpi Laptop
computer with custom-developed "VideoSpy" soft-
ware (Mark Evans, John Deere Commercial Prod-
ucts, Inc., unpublished). The custom software
allowed us to set a filming schedule for each day
using "record" times (3 or 5 min) and "wait" times
(10-15 min) so that up to 2 h of video could be re-
corded throughout the day. The system was pow-

Florida Entomologist 85(2)

ered by a 12-volt marine deep cycle battery. Video-
imaging allows pollinator visits to be reviewed re-
peatedly or in slow-motion. Furthermore, this
technique eliminates any effect of human pres-
ence (e.g., scent and motion) on the behavior of
the visitors. From both researcher and video mon-
itoring, we recorded the identity of a visitor, the
duration of its visit, and its behavior on a flower.
From field observations, we identified insects
that were potentially pollinators, herbivores, nec-
tar robbers, or incidental visitors. We also deter-
mined the time of day potential pollinators were
present. Visitation rates were calculated from data
recorded by the field observer while watching focal
flowers and also from data recorded on individual
flowers by the camcorder. Rate of visitation (visits
per flower per min) was calculated as number of
arthropod visits to flowers per number of flowers
observed per total observation time. When flowers
were observed over more than one day, a visitation
rate was calculated for each day, and then an aver-
age rate for those days was determined.
On two separate occasions in 1999, a halictid
bee was collected after it had visited an H. flava
flower. Each bee was washed in a vial containing
deionized water to remove pollen from its body,
and then it was released unharmed. The pollen
was cleaned and slide-mounted according to a po-
tassium hydroxide-acetolysis procedure provided
by Jean Porter at the University of Georgia Paleo-
ecology Laboratory, Athens, Georgia (see also Erdt-
man 1969). Jean Porter also prepared a reference
slide ofH. flava pollen from pollen extracted from
a specimen in the University of Georgia Herbar-
ium. We used the reference slide for determining
if H. flava pollen was present on the slides we
made from field collections.


Harperocallis flava

In 1999, 25 H. flava flowers at one roadside
study site and 20 flowers at a nearby site (1 km
north of other site, approximately 20-40 m off the
road) were tagged and then monitored at 3-min
periods for a total of 20 h of observation over 3
days (May 18-20; range 0830-1555 h EDT). Al-
though many bees, wasps, and beetles were seen
on nearby Ilex glabra L. (Aquifoliaceae), Hyperi-
cum sp. (Clusiaceae), and Iris sp. (Iridaceae), only
5 different insect species visited H. flava (Table 1),
resulting in 8 floral visits. Of all the insect visitors,
halictid bees (n = 3) were the only ones that spent
time on the flowers gathering pollen (Fig. 1). One
of the halictids was collected, and later identified
by T. L. Pitts-Singer as Dialictus sp. (using key in
Mitchell 1960). The specimen was retained for ref-
erence at the U.S. Department of Agriculture For-
est Service, Forestry Sciences Laboratory, Athens,
Georgia. While on a flower, each bee focused its at-

tention on one or two anthers for pollen collection,
occasionally crawling across the centrally located
stigma. Duration of visits by the halictids ranged
from 30 s to 7 min on a single flower.
In summer 2000 at a different site located ap-
proximately 5.1 km from the 1999 site, an ob-
server monitored 12 H. flava flowers on May 16,
and 13 flowers on May 17, for a total of 6 h 15 min
(range 0830-1656 h EDT). Observations were
made for 5-min periods (instead of 3-min). The
video camcorder recorded 3 h 56 min of videotape
over the same 2 days noted above. Four visits (2
by halictids, 1 by a mordellid, and 1 by a syrphid)
were recorded on one flower using the video cam-
corder. Arthropods that were recorded visiting
H. flava by both the human observer and by video
camcorder are shown in Table 1. Again, halictid
bees were the only insects that gathered pollen
from the flowers (Fig. 1), and their visits lasted
from 1 s to 10 min (i.e., some visits exceeded the 5-
min period). Halictid visits occurred throughout
the day (0945-1345 h EDT), but not in the early
morning or late afternoon (Fig. 1). Another small
yellow flower, Xyris baldwiniana Roemer &
Schultes (Xyridaceae), was also present at this
H. flava site and was visited by halictid bees.
Xyris baldwiniana flowers were only open during
the morning, and their presence could have af-
fected the rate at which bees visited H. flava. No
halictids were collected in 2000 for identification
to the generic level.
In both 1999 and 2000, average insect visita-
tion rates to H. flava were very low (Table 2).
Based on human observations, the rate was
higher in 2000 than in 1999. Rates from the video
camcorder used in 2000 on a focal flower were
higher than that recorded by the human observer.

Macbridea alba

Twenty inflorescences of M. alba at each of two
sites (distance between sites = approx. 4.4 km)
were observed for 34 h over 5 days (June 24-25,
June 29-July 1; range 0724-1805 h EDT). Nine in-
sect and spider species made 70 visits (Table 1),
but only bumblebees (Bombus) were large enough
to contact the reproductive structures of the
flower. No bumblebees were collected in order to
determine their specific scientific identity during
this study. Bumblebee visits to each flower were
very short (1-9 s), but they usually visited more
than one flower per inflorescence (Fig. 2) and vis-
ited throughout the day (0800-1645 h EDT). The
bees landed on the lower lip (petal) of the flower
and immediately pushed their heads down into
the corolla, presumably to obtain nectar. With its
head in the flower corolla, the bee's thorax con-
tacted anthers and stigma. Occasionally, just be-
fore leaving the flower, the bee appeared to probe
the anthers with its proboscis for ~1 s. The purpose
of the latter behavior is not clear. Overall, visita-

June 2002

Pitts-Singer et al.:Pollinators of Rare Plants


Plant species;
observation time and no. visits

H. flava
obs. time = 20 h
visits = 8

human obs. = 8 h 25 min
video = 3 h 56 min
visits = 8

M. alba
obs. time = 34 h
visits = 70

S. floridana
obs. time = 6 h
visits = 9

S. floridana
human obs. = 6 h
video = 2 h 43 min
visits = 10

Visitor scientific name

Halictidae: Dialictus*
Apidae: Bombus
Halictidae: sp. #1*
Halictidae: sp. #2*

Apidae: Bombus*
Formicidae: Paratrechina

Apidae: Bombus*
Halictidae: Dialictus
Halictidae?: sp. #2


Common name

sweat bee
bumble bee
tumbling flower beetle
katydid nymph
sweat bee
sweat bee
tumbling flower beetle
bee fly

bumble bee
false darkling beetle
gossamer-winged butterfly
lynx spider
crab spider

leafcutter bee
sweat bee
syrphid fly
crab spider

leafcutter bee
sweat bee
small katydid
crab spider

*Indicates insects that were seen to collect pollen and are probably important pollinators
?Researcher is not positive of family-level identification.

tion rates were higher for M. alba than for H. flava
(Table 2). Bumble bees also visited Rhexia alifa-
nus Walter (Melastomaceae) and Hibiscus aculea-
tus Walter (Malvaceae) that occurred at the same
sites. The bees spent more time collecting pollen
on these common flowers than they did onM. alba
collecting nectar.
In 2000, very few M. alba flowers were pro-
duced, presumably because of drought and other
unfavorable environmental conditions. Thus, we
were unable to make further observations of visi-
tors to this flower species.

Scutellaria floridana

In 1999, 10 inflorescences ofS. floridana at one
site were monitored over 2 days (September 14-
15; range 0825-1632 h EDT) for 6 h of observa-
tion, during which 7 insect and spider species
made 9 visits (Table 1). Visitations occurred be-
tween 1000 and 1530 h (Fig. 3). Of the visitors
recorded during observation periods, megachilid
bees and possibly halictid bees displayed behav-
ior that may have resulted in pollination of S.
floridana flowers (Fig. 3). Megachilids landed on

Florida Entomologist 85(2)

Harperocallis flava


1600 -
n =
1500 25 flowers

1400 -

1300 B

1200 -





20 flowers

n =
25 flowers

' I*',t

MAY 18 MAY 19
1999 1999

MAY 20

n = 13 flowers
(+ 1 w/ video)

= 12 flowers
1 w/ video)


MAY 16

MAY 17

= Halictid:

= Halictid:

Dialictus sp.

sp. #1 observer

*= Halictid: sp. #1 video

*= Halictid: sp. #2 observer

Fig. 1. Visitation by halictids on H. flava flowers recorded by human observer and by video camcorder in May
1999 and 2000 at the Apalachicola National Forest, Florida. Bars represent time frame in which 3- or 5-min obser-
vation periods occurred. Symbols represent pollinator visits.

the flower, clung to the flower hood in an upside-
down position, and then kicked into the hood
through the seam so that pollen fell onto the
scopa on the venter of the gaster. Durations of
megachilid visitations were 21-23 s. Although
halictids (Dialictus sp.) also visited these flowers
(Fig. 3), they did not appear to come into contact
with the stigma and, therefore, may have been
pollen "robbers." The small halictids entered into
the hood of the flower, out of the observer's view.
The halictids presumably collected pollen before
exiting the flower. Their visits lasted from 2 s to 1
min. In addition to the megachilid and halictid
bees, the observer noted that one bumblebee vis-
ited a flower before the study began (Fig. 3). This
bee spent a few seconds on a flower and used flo-
ral sonication to dislodge pollen from the anthers.
A few lepidoptera (not on marked flowers) and a
fly also were observed on the flowers, although

they appeared to collect only nectar and did not
contact the anthers. Compared to the other 2
plants in this study, the overall visitation rate
was high (Table 2).
In 2000 at a site located approximately 16.25
km from the 1999 site, 10 flowers of S. floridana
were monitored by a human observer for 6 h on
April 20-21 (range 0850-1754 h EDT) in 2000. The
video camcorder captured 2 h 43 min of tape, re-
vealing visits by 4 megachilids and 1 halictid. Al-
together, 4 insect species made 10 visits to flowers
(Table 1). Megachilids (visit duration = 10-15 s)
and halictids (visit duration = 5-31 s) were the
most frequent visitors to S. floridana (Fig. 3). Be-
havioral observations of megachilids and halic-
tids, which were larger halictids than those seen
in 1999, indicated that these bees probably are ef-
fective pollinators of S. floridana. These bees
clung to the outside of the flower hood, sometimes


June 2002

Pitts-Singer et al.:Pollinators of Rare Plants

MERS OF 1999 AND 2000.

Visitation rate (x 102)

Flower species: visitors-method 1999 2000

H. flava: halictids-observer 0.01 (n = 3) 0.06 (n = 3)
H. flava: all visitors-observer 0.03 (n = 8) 0.06 (n = 3)
H. flava: halictids-video n.a. 0.85 (n = 2)
H. flava: all visitors-video n.a. 2.1 (n = 5)
M. alba: bumblebees-observer 0.07 (n = 21) n.a.
M. alba: all visitors-observer 0.18 (n = 70) n.a.
S. floridana: bees*-observer 0.14 (n = 5) 0.06 (n = 3)
S. floridana: all visitors-observer 0.25 (n = 9) 0.15 (n = 5)
S. floridana: bees*-video n.a. 3.72 (n = 5)
S. floridana: all visitors-video n.a. 3.72 (n = 5)

*"Bees" includes megachilids and halictids.

touching the stigma, as they forced their heads
and thoraces into the hood. Pollinator visitations
occurred from morning until late afternoon (Fig.
3). Visitation rates were lower in 2000 than they
were in 1999. Carpenter bees (Xylocopa sp.) were
seen to rob nectar by piercing the base of the co-
rolla. Other flowers such as Aletris lutea Small
(Liliaceae), Balduina uniflora Nuttall (Aster-
aceae), and Pityopsis graminifollia (Michaux)
(Asteraceae) were also at this site, attracting
bumblebees and butterflies.

Pollen from Halictid Bees

We found pollen on only 1 of the 2 slides pre-
pared from washes of the 2 halictid bees captured
on H. flava flowers. There were 7 pollen grains of
H. flava, 1 grain of 3 other types, and 3 grains of
a fourth type. The absence of pollen on the second
slide does not preclude that pollen was not
present on the bee. The pollen may have been lost
in the insect net during collection, the water wash
may have not removed pollen from the bee, or
maybe the pollen was lost in the process of ex-
tracting pollen out of water in the vial.


We found that native bees were the likely pol-
linators of all 3 of the threatened or endangered
flowers we studied in the Apalachicola longleaf
pine forest. Bumblebees are probably important
pollinators of M. alba and possibly of S. floridana,
but megachilids and halictids may be the primary
pollinators of S. floridana. Halictids may play a
significant role in the pollination system of
H. flava. Although honeybee (Apis mellifera L.)
hives are present near the forest (Louise Kirn,

U.S.D.A. Forest Service, Bristol, FL, pers. comm.),
we observed no honeybees on these flowers.
Arthropod visits to the rare flowers were quite
infrequent (Table 2) (Figs. 1-3). Unfortunately, we
found no other pollinator visitation data in the lit-
erature for flowers in the Apalachicola National
Forest or vicinity for making comparisons. How-
ever, pollinator studies performed on flowers from
other habitats may offer an idea of potential bee
visitation rates. For example, a study on Hibiscus
moscheutos in Maryland revealed average visita-
tions by bumblebees and anthophorid bees of 27-
13 visits per flower per min x 10-2 (Spira et al.
1992). Another study in Michigan on catnip,
Nepeta cataria L., found average visitations by
bumblebees to be 8.6 visits per flower per minx 10-
2 and by Halictidae to be 7.4 visits per flower per
min x 10-2 (Sih & Baltus 1987). Compared to these
studies, bee visitation rates to the flowers we stud-
ied are indeed quite low (range = 0.01 x 10-2 to 3.72
x 10-2 per min) (Table 2). However, we saw these
same insects visiting other flowers in the area.
Regardless of whether or not these rare flow-
ers are imperiled because of low pollinator fre-
quency, our studies showed that certain insects
collect pollen and nectar from them and may be
important in their pollination ecology. Bumble-
bees were the only insects observed whose size
and behavior was efficient for pollinatingM. alba.
Observations of M. alba flowers in the South
Carolina Botanical Garden suggest that bumble-
bees are very frequent visitors and efficient polli-
nators (J. L. Walker, pers. obs.). Continued studies
of pollinators of this flower species should deter-
mine how many individual bumble bees are visit-
ing the rare flowers or other flowers in the area
and if these bees are important for out-crossing in
this species.

Florida Entomologist 85(2)

Macbridea alba

n = 20 flowers






.E 1300-

1200 n = 20 flowers

1100- *W*


n = 20 flowers

1999 1999

*= Bombus

n = 20 flowers




* 20 flowers



Fig. 2. Visitation by bumblebees on M. alba inflorescences recorded by human observer in June-July 1999 at the
Apalachicola National Forest, Florida. Bars represent time frame in which 3-min observation periods occurred.
Symbols represent pollinator visits.

Although some evidence has indicated that
H. flava flowers are self-compatible and undergo
selfing, we observed several halictid bees collect-
ing pollen from H. flava flowers, especially in 2000
(Tables 1 and 2) (Fig. 1). The pollen-collecting be-
havior of halictids on H. flava flowers could have
facilitated pollination. For a hermaphroditic spe-
cies like H. flava, visitation by bees may improve
out-crossing; or the activity of bees on anthers
may dislodge pollen for better wind dispersal
(Cane et al. 1992). Nonetheless, if out-crossing
were evolutionarily important for this species in
the past, it may now be ineffectual as a result of
the low genetic variability in H. flava. Though pol-
linator services may not benefit H. flava, the
availability of this pollen may be a useful resource
for the solitary bees that harvest it.
Our study provides the first documentation of
the pollination ecology of S. floridana. The behav-
iors of megachilids and halictid bees on the flow-
ers were appropriate for pollination. We also

noted that the newly opened flowers were more
readily approached and subsequently visited
than 2-day-old flowers. In 2000, the flowers
bloomed at the beginning of the summer (May)
and not at the end of the summer (September) as
in 1999. These different flowering dates may have
had an effect on the diversity and abundance of
insects that visited the flowers. Further investi-
gation of pollinators of this species should include
insect exclusion experiments on S. floridana and
comparative studies with another common skull-
cap, S. integrifolia L., which occurs in the area.
Another useful outcome from this study is in
the methodology. Using the video camcorder to
collect visitation data may have had an advan-
tage over human observation (Table 2). By focus-
ing on only 1 flower throughout the day, up to 2
hours of observable data could be gathered with-
out human intervention. A higher visitation rate
was recorded for both H. flava and S. floridana
when this method was used.


0900 -

0800 -

0700 -

June 2002

Pitts-Singer et al.:Pollinators of Rare Plants

Scutellaria floridana

1800 -
,7An n = 10 flowers


1600 -



1300 -

1200 -


1000 -

0900 -



SEPT. 14

n = 10 flowers
(+ 1 w/ video)

= 10 flowers


SEPT. 15



n = 10 flowers
(+1 w/ video)



S= Bombus A = Megachilid 0 = Halictid: Dialictus sp.
*Bee observed outside of designated time range.

A = Megachilid observer H = Halictid observer
A = Megachilid video U = Halictid video
Fig. 3. Visitation by bumblebees, megachilids, and halictids on S. floridana flowers recorded by human observer
and video camcorder in September 1999 and April 2000 at the Apalachicola National Forest, Florida. Bars repre-
sent time frame in which 3-min observation periods occurred. Symbols represent pollinator visits.

Early in the study, we planned to collect in-
sects that visited the rare flowers in order to cre-
ate a reference collection and to identify pollen
samples from the insects. However, because visi-
tation rates were low, we decided that removal of
potential pollinators might have been detrimen-
tal to the reproductive success of the plants, and
we abandoned this effort.
Each of the plants we studied responds to fire
by increased growth and flowering (Madsen 1999;
Louise Kirn, USDA Forest Service, Apalachicola
Ranger Station, Bristol, FL, pers. comm.). This
response suggests that fire timing is important to
ensure that flowering occurs when pollinators are

available. Thus, we must also understand the life
cycle of important pollinators, and, further,
whether fire has an effect on the availability of
food and nesting materials.
At least three requirements must be met for ef-
fective conservation of pollinators in any ecosys-
tem and of bee pollinators in the Apalachicola
ecosystem. They require proper nesting sites,
nest-building material, and a sufficient amount of
food at appropriate times both for adults (who
need nectar) and for larvae (who need pollen)
(Westrich 1996). Thus, it is imperative to identify
the specialized needs of each pollinator. Such in-
formation will help in determining which, if any,

Florida Entomologist 85(2)

forest management practices affect availability of
these resources; or if improvements can be made
in certain areas to maintain suitable habitat.
The pollination ecology of some communities
has been neglected, and more information concern-
ing the plant-pollinator interactions of rare and
endangered plants is needed (Kevan 1975; Kevan
et al. 1993; Buchmann & Nabhan 1996; Kearns et
al. 1998). Researchers must first have basic knowl-
edge of a system before making hypotheses at an
ecosystem level and designing experiments to test
them. Researchers must perform the necessary
studies to determine the importance of insect (and
other animal) visitors in pollination (Kwak et al.
1996). Conservation plans need to be backed by
solid, scientific evidence. Our results are a starting
point for understanding insect pollinators of three
rare plants in a longleaf pine ecosystem and pro-
vide a basis for future examination and conserva-
tion of these plants and their pollinators.

We thank personnel of the Apalachicola National
Forest Ranger Station for their support of this project,
especially Louise Kirn (USDA Forest Service) who kept
us informed of when flowers were in bloom, guided us to
field sites, and helped us identify various flowers at field
sites. David Jenkins (University of Georgia) was a valu-
able field and laboratory assistant in summer 1999, and
his observations and suggestions are much appreciated.
We thank Robert W. Matthews, Christopher J. Fettig,
and James P. Pitts (Department of Entomology, Univer-
sity of Georgia, Athens, Georgia), Kerry 0. Britton
(USDA Forest Service, Athens, Georgia), and Paul
Smith (USDA Forest Service, Asheville, North Caro-
lina) and 3 anonymous reviewers for their critical as-
sessments of this manuscript.

BREWER, J. S. 1999. Effects of competition, litter, and
disturbance on an annual carnivorous plant (Utricu-
lariajuncea). Plant Ecol. 140: 159-165.
BUCHMANN, S. L., AND G. P. NABHAN. 1996. The Forgot-
ten Pollinators. Island Press/Shearwater Books:
Washington, D.C.
1992. The solitary bee Melissodes thelypodii thelypo-
dii Cockerell (Hymenoptera: Anthophoridae) collects
pollen from wind-pollinated Amaranthus palmeri
Watson. Pan-Pacific Entomol. 68: 97-99.
ERDTMAN, G. 1969. Handbook of Palynology. Hafner
Publishing Co., New York, NY.
Genetic diversity in the endangered lily Haperocal-
lis flava and a close relative, Tofieldia racemosa.
Conserv. Biol. 11: 361-366.
HARDIN, E. D., AND D. L. WHITE. 1989. Rare vascular
plant taxa associated with wiregrass (Aristida stricta)
in the southeastern United States. Nat. Areas J. 9:
HESSL, A. AND S. SPACKMAN. 1995. Effects of fire on
threatened and endangered plants: an annotated bib-
liography. Information and Technology Report 2. U.S.
Department of the Interior, Washington, DC. 55 pp.

Endangered mutualisms: the conservation of plant-
pollinator interactions. Ann. Rev. Ecol. Syst. 29: 83-
KEVAN, P. G. 1975. Pollination and environmental con-
servation. Environ. Conserv. 2: 293-298.
sects and plants in the pollination ecology of the bo-
real zone. Ecol. Res. 8: 247-267.
KRAL, R. 1983. A report on some rare, threatened, or en-
dangered forest-related vasular plants of the south.
U.S.D.A. Forest Service Tech. Pub. R8-TP, Vols. 1&2.
1996. Insect diversity and the pollination of rare plant
species, pp. 116-124. In A. Matheson, S. L. Buch-
mann, C. O'Toole, P. Westrich, and I. Williams [eds.],
The Conservation of Bees. Academic Press, London.
LESICA, P. 1999. Effects of fire on the demography of the
endangered, geophytic herb Silene spaldingii
(Caryophyllaceae). Amer. J. Bot. 86(7): 996-1002.
MADSEN, D. L. 1999. Seed production and germination
studies ofMacbridea alba. Master's Thesis, Clemson
MITCHELL, T. B. 1960. Bees of the Eastern United
States, Vol. I. North Carolina Agricultural Experi-
ment Station, Tech. Bul. No. 141.
PEET, R. K., AND D. J. ALLARD. 1993. Longleaf pine-
dominated vegetation of the southern Atlantic and
eastern Gulf Coast region, USA. Proc. Tall Timbers
Fire Ecol. Conf. #18, pp. 45-81.
PROCTOR, M., YEO, P. AND A. LACK. 1996. The Natural
History of Pollination: Timber press, Portland, OR.
longleaf pine forest: trends and current conditions.
U.S. Forest Service Resource Bulletin SRS-RB-9.
RATHCKE, B. J. AND E. S. JULES. 1993. Habitat fragmen-
tation and plant-pollinator interactions. Current
Science 65: 273-277.
SIH, A. AND M.-S. BALTUS. 1987. Patch size, pollinator
behavior, and pollinator limitation in catnip. Ecol-
ogy 68: 1679-1690.
1992. Flower visitation, pollen deposition, and pol-
len-tube competition in Hibiscus moscheutos (Mal-
vaceae). Am. J. Bot. 79: 428-433.
HICKERSON. 1997. The need for "extended care" in
conservation: examples from studies of rare plants
in the Western United States, pp. 245-248. In K.W.
Richards [ed.], Seventh International Symposium on
Pollination, Acta Hort 437, ISHS.
and Threatened Species of the Southeastern United
States (The Red Book), Vol. 2., Fish & Wildlife Ser-
vice, Atlanta, GA.
WAGNER, L. K. AND T. P. SPIRA. 1996. Germination be-
havior of Harperocallis flava, an endangered Florida
endemic. Botanical Society of America Annual Meet-
ing, University of Washington, Seattle.
WALKER, J. 1993. Rare and vascular plant taxa associ-
ated with the longleaf pine ecosystems: patterns in
taxonomy and ecology. Proc. Tall Timbers Fire Ecol.
Conf. #18, pp. 105-125.
WESTRICH, P. 1996. Habitat requirements of central
European bees and the problems of partial habitats,
pp. 1-16. In A. Matheson, S. L. Buchmann, C. O'Toole,
P. Westrich, and I. Williams [eds.], The Conserva-
tion of Bees. Academic Press, London.

June 2002

Fedorka & Mousseau: Nuptial Gifts in a Cricket


Department of Biological Sciences, University of South Carolina, 700 Sumter Street, Columbia, SC 29208


Many male insects provide somatic nuptial gifts that may strongly influence reproductive
fitness by insuring an effective copulation or by increasing paternal investment. In the
striped ground cricket, Allonemobius socius (Scudder), females receive a nuptial gift by
chewing on a specialized spur on the male's hind tibia during copulation. Using a series of
no-choice trials, we attempted to quantify gift magnitude and to determine the relationships
between male size, gift contribution, and male mating success. Tibial spur chewing duration
was a significant predictor of gift contribution (F,,,17 = 17.02, P < 0.001) and the magnitude of
the gift ranged between 0.2% and 8% of the male's body mass, implying that females receive
mostly hemolymph. Large males produced bigger gifts than small males (2.52 0.59 mg vs.
1.33 + 0.28 mg, ti, = 1.88, P < 0.05, respectively) and females were more likely to mate with
larger males (F,,3 = 4.76, P < 0.05). If gift size is shown to influence female reproductive fit-
ness, then nuptial gifts may play a large role in the evolution of male body size.

Key Words: nuptial gift, tibial spur, body size, Allonemobius socius


El macho en muchos insects provee un regalo nupcial somatico que puede influenciar fuer-
temente la adaptabilidad 6ptima reproductive (fitness) al asegurar una c6pula eficaz o al in-
crementar la inversion paternal. En el grillo Allonemobius socius, las hembras reciben un
regalo nupcial al morder un espol6n especializado en la tibia posterior del macho durante la
c6pula. Utilizando una series de pruebas de no alternatives, tratamos de cuantificar la mag-
nitud del regalo y determinar la relaci6n entire el tamano del macho, la contribuci6n del re-
galo, y el 6xito de la c6pula del macho. La duraci6n de las mordidas del espol6n de la tibia fue
un predictor significativo de la contribuci6n del regalo (F,17 17.02, P < 0.001) y la magnitude
del regalo abarc6 entire el 0.2% y el 8% del peso del cuerpo del macho, indicando que las hem-
bras reciben principalmente hemolinfa. Los machos grandes produjeron regalos mas grandes
que los machos pequenos (2.52 0.59 mg vs 1.33 0.28 mg,t,7 =1.88, P < 0.05) y era mas pro-
bable que las hembras copularan con los machos mas grandes. Si el tamano del regalo in-
fluencia la adaptabilidad 6ptima reproductive de las hembras, entonces los regalos nupciales
pueden actuar un papel important en la evoluci6n del tamano del cuerpo de los machos.

In many birds and insects, males offer nuptial
gifts to females prior to and/or during copulation
(Wiggins and Morris 1986; Gwynne and Brown
1994; Simmons 1995; Neuman et al. 1998). These
offerings include captured prey, somatic tissue,
synthesized secretions and suicidal food transfers
(see Andersson 1994 and references therein). The
function of these gifts may vary, serving as a form
of male mating effort by increasing fertilization
success (Alexander & Borgia 1979; Sakaluk 1984)
and/or as paternal investment by increasing repro-
ductive fitness (e.g., increasing egg size, offspring
number or offspring viability; Gwynne 1984; Rein-
hold 1999). For instance, in the hangingfly, Hylo-
bittacus apicalis (Byers), nuptial prey functions to
increase mating effort by increasing copulation du-
ration, which is in turn associated with the volume
of sperm transferred (Thornhill 1976). In contrast,
nuptial feeding in the osprey, Pandion haliaetus
(Linnaeus), acts as paternal investment since it
was positively associated with offspring growth

rate (Green and Krebs 1995). Regardless of the
precise functional significance, these associations
suggest that variation in nuptial gift mass may
have strong fitness implications for both sexes.
Nuptial gifts are generally transferred to fe-
males via external, intermediate packages such
as prey items or self-contained somatic secretions
(Andersson, 1994), permitting an easy quantifica-
tion of their mass. However, in some animal gen-
era the gift is internally transferred making the
magnitude and the fitness implications of such
gifts difficult to assess. For instance, in the cricket
genus Allonemobius (Orthoptera: Gryllidae),
males internally deliver a nuptial gift through a
specialized spur on each hind tibia that is chewed
by the female during copulation (Fig. 1). Although
chewing damages the spur, males can mate mul-
tiple times and females do not discriminate
among males based on the spur's condition (un-
published data). This somatic gift has previously
been described as a limited glandular contribu-

Florida Entomologist 85(2)

Fig. 1. The tibial spur. This specialized spur delivers a somatic nuptial gift directly to the female through court-
ship feeding. The spur on the right has been chewed by a female whereas the left is still intact indicating a male's
previous mating success.

tion contained within the spur that is exuded once
the tip is removed (Fulton 1931; Mays 1971; For-
rest 1991). However, a preliminary study sug-
gested that, when the spur tip is artificially
amputated and a capillary tube attached, the vol-
ume of the extract exceeds the volume of the spur
(unpublished data). Thus, the spur may provide
females with direct access to the male's hemo-
lymph, making male mass a potential limiting
factor in gift contribution. This is important since
the evolutionary trajectory of male body size and
sexual size dimorphism may be modified if posi-
tive selection on the gift exists, coupled with a ge-
netic correlation between body and gift size.
Unfortunately, no mass measure of this or any
other internally transferred nuptial gift exists.
Using the striped ground cricket,Allonemobius
socius (Scudder), we attempted to quantify the
magnitude of an internally transferred gift. We
predicted that male A. socius donate more than a
limited glandular contribution, and may offer fe-
males a continuous supply of hemolymph. Fur-
thermore, we predicted that male contribution is
constrained by male mass, with the largest males
capable of offering the largest gift. Therefore, fe-
males should prefer large males if gift mass is pos-
itively related to male mass and to female fitness.


A. socius is a small chirping ground cricket
found throughout the southeastern United
States, with closely related sister taxa ranging
throughout North America (Alexander & Thomas
1959; Howard & Furth 1986; Mousseau & Roff

1989). All crickets used in this experiment were
second-generation lab-reared individuals origi-
nating from a single population near Asheville,
North Carolina. Crickets were maintained in 10 x
10 x 8 cm plastic cages containing ground cat
food, a carrot slice, water vial and strips of brown
paper towel for cover. Every three days the food,
carrot and paper towel were replaced. Cages were
kept in a constant environment at 28C and a
12:12 [L:D] photoperiod provided by a Percival in-
cubator. The age of all experimental crickets was
held constant at 12 + 2 days post eclosion.
A. socius males perform a calling song used to
attract distant females. Once a potential mate is
encountered, males switch to a courtship song
and dance that culminates with the male orient-
ing his abdomen toward the stationary female. If
the female is receptive, she will briefly mount the
male in a "mock copulation" lasting only a few sec-
onds. Once an effective mock copulation is
achieved (this may take several attempts) the
male will cease courting and begin to form a sper-
matophore (approximately 20 min). When com-
plete, he will renew his courtship behavior, again
enticing the female to mount. At this time, the
couple will adjoin abdomens as the male adheres
the spermatophore to the female's genitalia. The
male will then bring his hind tibia forward allow-
ing the female to chew on his spur until the couple
separates (upwards of 30 min). Once apart, the fe-
male will remove and consume the spermato-
phore. If a spermatophore is formed but not
attached, no spur chewing will take place and the
spermatophore will be removed and consumed by
the male (Alexander & Thomas 1959; Mays 1971).

June 2002

Fedorka & Mousseau: Nuptial Gifts in a Cricket

Virgin males and females were randomly paired
(n = 41 pairs), placed into a mating arena (6 cm di-
ameter petri dish) and allowed to mate once. Al-
though crickets mate multiply in the wild (Walker
1980; Sakaluk and Cade 1983), only one mating
bout is needed to detect the mass changes used in
estimating the size of the nuptial gift. Two groups
were created depending on the outcome of the mat-
ing trial. Males who attached their spermatophore
to the female were grouped as successful. Males
who formed a spermatophore, but failed to re-at-
tract a female for copulation were grouped as un-
successful. In both cases, spermatophores were
collected after removal to avoid consumption.
Male, female and petri dish mass were recorded be-
fore and after each mating trial. The mass gain in
the petri dish gave us a measure of mass loss due to
defecation. In addition, spermatophore mass, trial
duration, spermatophore attachment duration, tib-
ial spur chewing duration, and post-chewing sper-
matophore attachment duration (i.e., the amount
of time it takes the female to remove the spermato-
phore after chewing the tibial spur) were recorded.
Since by nature of the mating ritual, a positive as-
sociation between chewing duration and spermato-
phore attachment seems inherent, post-chewing
spermatophore attachment duration was exam-
ined to assess whether larger gifts satiate females
to where they will postpone removal and consump-
tion of the spermatophore. Mass was measured us-
ing a Sartorius scale (Bohemia, NY) accurate to
0.01 mg. Trial duration was defined as the time
elapsed from the onset of male courtship to the re-
moval of the spermatophore by the female (suc-
cessful) or the male (unsuccessful).
The mass of the gift contribution was esti-
mated for each successful trial by taking the total
change in male mass (Am) and subtracting out
the mass lost to defecation (Ap), spermatophore
mass (s), and respiration (r), such that,

Gm] = Ami (Ap/2) s r (eq. 1)

The term Ap was halved since we assumed that
the rate of defecation was equal between the
sexes. To estimate r, we first estimated the aver-
age mass lost per minute, R, from the unsuccess-
ful male data while controlling for defecation and
spermatophore production,

y(|AmI (Ap/2) s)/t
R 1 (eq. 2)

where t is the duration of the trial. We assumed
that R was equal for successful and unsuccessful
males. Thus, for each successful trial, r was esti-
mated by multiplying R by the duration of the

mating trial, t. A second independent estimate of
the nuptial gift, G,, was based on female mass
gain and obtained by substituting the change in
female mass, (Af), for (Am) into equations one and
two and dropping s. All data analyses were per-
formed using SAS 8.1.


Since the ejaculate contained within the sper-
matophore that is transferred directly to the fe-
male genitalia may confound our estimate of gift
mass, we compared the mass of the successful
male's depleted spermatophore (mean+ se: 1.03 +
0.01 mg; n = 19) with the unsuccessful male's in-
tact spermatophore (0.99 0.07 mg; n = 22) to es-
timate ejaculate contribution. No difference
existed between these two groups (one tailed t-
test: t39 = -0.93, NS), suggesting that the ejaculate
mass was negligible. There was no effect of male
body size on spermatophore mass (F,39 = 2.07, NS)
or estimated respiratory mass loss (F,7 = 0.17,
NS). Average male mass was 77.55 + 1.24 mg and
the average GEm; was 1.89 0.33 mg. Thus, the av-
erage nuptial gift was approximately 2.44% of the
male's initial body mass. When coupled with aver-
age spermatophore size (1.03 + 0.01 mg) males in-
vested approximately 3.77% of their total mass
into a single copulation. This is a large invest-
ment considering that males are promiscuous
and that a substantial proportion of their mass is
attributable to exoskeleton.
When we replaced Am with Af in equations one
and two, average G. was estimated to be 1.01 +
0.45 mg, providing a second, independent mea-
sure of gift size. Although this estimate is lower
than the male estimate, they are not significantly
different (two tailed t-test: t6 = 1.53, NS). Six G,
estimates were negative (Fig. 2), and were most
likely the result of the error inherent in the meth-
odology used to estimate the mass lost to respira-
tion, r. All six negative G, estimates were well
within one standard deviation in r away from a
positive estimate. As a consequence, our estimate
of the average G, is conservative.
Since no difference existed between the suc-
cessful and unsuccessful groups with regard to
spermatophore mass (s), mass lost to defecation
(0.58 0.02 mg and 0.67 0.02 mg, respectively;
Ap two tailed t-test: t3, = 0.52, NS), and the dura-
tion of the trial (two tailed t-test: t,= -0.72, NS),
we calculated two additional estimates of gift
mass, GRmi and Gf,, using the least squares means
difference in Am and Af between the two groups,
respectively. These estimates were calculated be-
cause they are free of the error attributed to esti-
mating mass lost to respiration and defecation.
Since initial male mass was significantly associ-
ated with Am (F,39 = 4.41, P < 0.05), we used an
analysis of covariance with initial male mass as
the covariate (ANCOVA: F2,M = 14.75, P < 0.001;

Florida Entomologist 85(2)


B 7



0 10 20 30


E *

0 10 20 3C
Spur Chewing (min)

Fig. 2. Nuptial gift contribution as a function of spur
chewing duration. A) As females chewed on the tibial
spur, male gift contribution increased (y = 0.1496x +
0.2444, R2 = 0.50). B) The relationship between female
mass gain and courtship feeding also provided a rate of
male contribution (y = 0.1547x 0.6922, R2 = 0.28). Both
male (Gem9) and female (G, ) estimates of the nuptial gift
were controlled for mass lost to defecation, respiration
and spermatophore production. The arrows indicate a
potential outlier in the female G, estimate and the cor-
responding GEm] estimate.

no interaction between covariate and group). Us-
ing this method Ge,.] was estimated to be 1.75 mg.
Likewise, G1, was estimated to be 1.28 mg. Con-
sidering that the tibial spur weighs less than
0.001 mg, these estimates suggest that the gift far
exceeds the capacity of the spur. Moreover, these
gift estimates exceed the weight of the entire male
tibia (0.75 + 0.05 mg), implying that the gift is
comprised mostly or entirely of hemolymph.


*o 0


20 -

. 10-


Male Mass (mg)

Fig. 3. Spur chewing duration as a function of male
mass. Large males were chewed longer than small
males (y = 564.18x 35.419, R2 = 0.20) suggesting that
male size may ultimately constrain gift size.






0 0


June 2002

Chewing duration covaried with both the male
estimate (Fig. 2a; GEm: F1,,, = 17.02, P < 0.001) and
the female estimate (Fig. 2b; G.: F1,1 = 6.67, P <
0.05) of the nuptial gift implying that females
who chewed longer received a larger gift contribu-
tion. The size of the gift ranged from 0.2% to 8% of
initial male mass. Although, the rate of male gift
contribution and female gift gain (Fig. 2) were not
significantly different (F1,16 = 2.12, NS), G, ex-
ceeded GEm. in four trials. In three of these trials
the differences were small and, as with the nega-
tive estimates of Gf, fell well within 1 standard
deviation of our estimate of r. The remaining trial
discrepancy showed female mass gain to be 3.94
mg greater than the male contribution, and can-
not be reconciled with the previous argument
suggesting that it may be the result of measure-
ment error. However, when this observation was
removed, the relationship between chewing dura-
tion and GEmi and G, remained unchanged (F1,,
=15.68, P < 0.001, R2 = 0.50 and F,1,= 10.21, P <
0.01, R2 = 0.40, respectively).
In addition, male mass was significantly asso-
ciated with spur chewing duration (Fig. 3: F1, =
4.24, P = 0.05) implying that larger males provide
a larger gift. In turn, spur chewing duration was
significantly associated with spermatophore at-
tachment duration (F1,1 = 4.51, P < 0.05), which is
expected since the spermatophore is attached at
the onset of spur chewing. However, chewing du-
ration was not associated with post-chewing sper-
matophore attachment duration (F,1, = 0.06, NS),
implying that females do not delay removal of the
spermatophore if a larger gift is received.

Fedorka & Mousseau: Nuptial Gifts in a Cricket

Successful males were significantly larger
than unsuccessful males (F,,, = 4.76, P < 0.05).
This may be the result of female choice or a me-
chanical constraint of small male size. In crickets,
copulation may be impeded if the size difference
of a mating pair is too great, making proper geni-
talic alignment and spermatophore transfer diffi-
cult (Sakaluk, pers. comm.). However, there was
no distinction between successful and unsuccess-
ful males with regard to male and female size dif-
ferences (F,,, = 1.28, NS), suggesting that large
male success was not due to small male inability
in passing a spermatophore, but perhaps to fe-
male preference for larger mates.
Although chewing duration was related to
both male mass and estimated gift mass, male
mass and gift mass were not directly related. To
further investigate the relationship between body
and gift mass, we separated the successful males
into two groups, large and small, based on their
mass relative to the mean (82.30 1.27 mg). The
ten males who fell below the mean were placed
into the 'small male' group, and the nine males
who fell above the mean were placed into the
'large male' group. On average, larger males were
chewed upon twice as long and provided nearly
twice as much gift as did smaller males (chewing
duration: 15.46 + 2.38 min vs. 7.01 + 1.20 min, t17
= 3.27, P < 0.01; gift donation: 2.52 0.59 mg vs.
1.33 0.28 mg, t1, = 1.88, P < 0.05, respectively T-
tests were one tailed). Using this analysis, male
mass was significantly associated with gift mass,
suggesting that the magnitude of the nuptial gift
may be constrained by male size.


In ground crickets (Neonemobius sp.), females
are attracted to larger males and this preference
was previously speculated to be based on the
male's ability to provide a larger nuptial gift (For-
rest et al. 1991). In this study, we have shown that
larger males are more successful at attracting
and copulating with females. More importantly,
we have shown that male mass was positively as-
sociated with the magnitude of gift contribution,
providing a mechanism for the maintenance of fe-
male preference. This body size and gift size rela-
tionship coupled with sexual selection on gift size
via female choice may have profound evolution-
ary implications.
In insects, the common pattern of female-
biased size dimorphism is usually attributed to the
reproductive advantages of being large (Shine
1988). However, if gift mass is related to female
fitness, then the strength of size selection on males
may surpass females, eventually increasing male
size and modifying degree of dimorphism (e.g.,
Leimar et al. 1994). An association between gift
mass and female fitness is common in Orthoptera
(Gwynne 1983, 1984; Brown 1997; Calos & Saka-

luk 1998; Reinhold 1999). However, an associa-
tion between male mass and gift mass is not
(Wedell 1997), though an association between
male mass and a spermatophylax gift/sperm am-
pulla complex has also been shown in some spe-
cies (Gwynne 1982; Sakaluk 1985). To elucidate
the implications of the nuptial gift on male size
evolution in this system, we are presently exam-
ining the impact of gift mass on female fitness
along with the selective pressures and underlying
genetic architecture surrounding these traits.
Currently, it is unknown whether gift mass is
shaped through male mating effort (e.g., by in-
creasing sperm transfer) or paternal investment
(i.e., by increasing the number and fitness of the
gift givers offspring). Our data suggest that mat-
ing effort is a plausible hypothesis since gift size
(i.e., chewing duration) was highly associated
with spermatophore attachment duration. These
results run contrary to a previous study by Bi-
dochka and Snedden (1985) that examined
mating behavior in a closely related sister spe-
cies, Allonemobius. fasciatus. In this study female
access to the tibial spur was manipulated through
three treatments (spur covered, surrounding spur
area covered-spur uncovered, and spur uncov-
ered) and subsequent copulation duration (an ap-
proximation of chewing duration and hence
nuptial gift size) and spermatophore attachment
duration recorded. They found that females who
were denied access to the spur had a significantly
shorter copulation duration than the other two
treatments. In addition, they concluded no associ-
ation between treatment and spermatophore at-
tachment duration. However, if we compare the
published summary statistics for the covered
(16.72 3.59 min, n = 30) and uncovered treat-
ments (29.26 + 5.35 min, n = 19) only, they are sig-
nificantly different (two tailed t-test: t, = -2.023,
P < 0.05), suggesting that spermatophore attach-
ment duration may be related to copulation dura-
tion in A. fasciatus, supporting our results.
In the wild, male and female ground crickets
are promiscuous and mate with numerous indi-
viduals throughout the breeding season, exceed-
ing the mating rate necessary to continually
produce offspring prior to senescence. Promiscu-
ous behavior often carries associated costs includ-
ing increased time and energy expenditure
(Thornhill & Alcock 1983), increased predation
risk (Arnqvist 1989), increased disease suscepti-
bility (Hurst et al. 1995), and/or caustic seminal
fluids that potentially reduce fitness (Fowler &
Partridge 1989; Rice 1996). The large size of the
nuptial gift in A. socius described here provides
the opportunity for these costs to be offset by in-
creasing female fitness through increasing repro-
ductive rate, reproductive longevity or fecundity.
Associations between gift size and female repro-
ductive fitness components are common in insects
(Gwynne 1984; Andersson 1994).

Florida Entomologist 85(2)

Currently, the contents of the gift inA. socius
are unclear though our data suggest it is mostly
hemolymph and not simply a limited glandular
secretion. Other systems have shown that nuptial
gifts may contain oviposition inducing hormones
(Friedel & Gilliot, 1977), or act as indicators of
important male chemical resources that may af-
fect offspring fitness (Eisner et al. 1996). Consid-
ering the relationship between gift size, male size
and large male mating success in our data, gift
quantity may be more important than gift quality.


We thank Jacqueline Litzgus, Scott Sakaluk and
Wade Winterhalter for their insightful comments on this
manuscript. This research was supported in part by a
grant from the National Science Foundation (NSF
0090177) to T.A.M. and by a GAANN fellowship to K.M.F.


ALEXANDER, R. D., AND THOMAS, E. S. 1959. Systematic
and behavioral studies on the crickets of the Nemo-
bius fasciatus group (Orthoptera: Gryllidae: Nemo-
biinae). Ann. Entomol. Soc. Amer. 52: 591-605.
ALEXANDER, R. D., AND BORGIA, G. 1979. On the origin
and basis of the male-female phenomenon, pp. 417-
440. In M. S. Blum and N. A. Blum (eds.), Sexual se-
lection and reproductive competition in insects. New
York: Academic Press.
ANDERSSON, M. 1994. Sexual Selection. Princeton, NJ:
Princeton University Press.
ARNQVIST, G. 1989. Multiple mating in a water strider:
mutual benefits or intersexual conflict? Animal Be-
hav. 38: 749-756.
BIDOCHKA, M. J., AND W. A. SNEDDEN. 1985. Effect of
nuptial feeding on the mating behaviour of female
ground crickets. Canadian J. Zool. 63: 207-208.
BROWN, W. D. 1997. Courtship feeding in tree crickets
increases insemination and female reproductive life
span. Animal Behav. 54: 1369-1382.
CALOS, J. B., AND SAKALUK, S. K. 1998. Paternity of off-
spring in multiply mated female crickets: the effect
of nuptial food gifts and the advantage of mating
first. Proc. Royal Soc. London, 265: 2191-2195.
ROACH, AND J. MEINWALD. 1996. Chemical basis of
courtship in a beetle (Neopyrochroa flabellata):
cantharidin as precopulatory 'enticing' agent. Proc.
Nat. Acad. Sci., USA, 93: 6499-6503.
AND M. EUBANKS. 1991. Mate choice in ground crick-
ets (Gryllidae: Nemobiinae). Florida Entomol. 71:
FOWLER, K., AND L. PARTRIDGE. 1989. A cost of mating
in female fruit flies. Nature 338: 760-761.
FRIEDEL, T., AND C. GILLOTT. 1977. Contribution of
male-produced proteins to vitellogenesis in Melano-
plus sanguinipes. J. Insect Physiol. 23: 145-151.
FULTON, B. B. 1931. A study of the genus Nemobius.
(Orthoptera: Gryllidae). Ann. Entomol. Soc. Amer. 24:

GREEN, D. J., AND E. A. KREBS. 1995. Courtship feeding
in ospreys Pandion hallaetus-a criterion for mate
assessment. Ibis. 137: 35-43.
GWYNNE, D. T. 1982. Mate selection by female katydids
(Orthoptera: Tettigoniidae, Conocephalus nigropleu-
rum). Animal Behav. 30: 734-738.
GWYNNE, D. T. 1983. Male nutritional investment and
the evolution of sexual differences in Tettigoniidae
and other Orthoptera, pp. 337-366. In D. T. Gwynne
and G. K. Morris (eds.), Orthopteran mating sys-
tems: Sexual competition in a diverse group of in-
sects. Boulder, Colorado: Westview Press.
GWYNNE, D. T. 1984. Courtship feeding increases female
reproductive success in bushcrickets. Nature 307:
GWYNNE, D. T., AND W. D. BROWN. 1994. Mate feeding,
offspring investment, and sexual differences in katy-
dids (Orthoptera, Tettigoniidae). Behavioral Ecology
5: 267-272.
HOWARD, D. J., AND D. G. FURTH. 1986. Review of the
Allonemobius faciatus (Orthoptera: Gryllidae) com-
plex with the description of two new species sepa-
rated by electrophoresis, songs, and morphometrics.
Ann. Entomol. Soc. Amer. 79: 472-481.
M. E. N. MAJERUS. 1995. Sexually transmitted dis-
ease in a promiscuous insect, Adalia bipunctata.
Ecol. Entomol. 20: 230-236.
predictable food and sexual size dimorphism in in-
sects. Proc. Royal Soc. London, 258: 121-125.
MAYS, D. L. 1971. Mating behavior of nemobiine crick-
ets Hygronemobius, Nemobius and Pteronemobius
(Orthoptera: Gryllidae). Florida Entomol. 54: 113-
MOUSSEAU, T. A., AND D. A. ROFF. 1989. Adaptation to
seasonality in a cricket-patterns of phenotypic and
genotypic variation in body size and diapause ex-
pression along a cline in season length. Evolution 43:
Courtship feeding and reproductive success in black-
legged kittiwakes. Colonial Waterbirds 21: 73-80.
REINHOLD, K. 1999. Paternal investment in Poecilimon
veluchianus bushcrickets: beneficial effects of nup-
tial feeding on offspring viability. Behav. Ecol. Socio-
biol. 45: 293-299.
SAKALUK, S. K., AND W. CADE. 1983. The adaptive sig-
nificance of female multiple matings in house and
field crickets, pp. 319-336. In D. T. Gwynne and G. K.
Morris (eds.), Orthopteran mating systems: Sexual
competition in a diverse group of insects Boulder,
Colorado: Westview Press.
SAKALUK, S. K. 1984. Male crickets feed females to en-
sure complete sperm transfer. Science 223: 609-610.
SAKALUK, S. K. 1985. Spermatophore size and its role in
the reproductive behaviour of the cricket, Gryllodes
supplicans (Orthoptera: Gryllidae). Canadian J. Zool.
63: 1652-1656.
SHINE, R. 1988. The evolution of large body size in fe-
males: a critique of Darwin's 'fecundity advantage'
model. American Naturalist 131: 124-131.
SIMMONS, L. W. 1995. Correlates of male quality in the
field cricket, Gryllus campestris L: Age, size and
symmetry determine pairing success in field popula-
tions. Behav. Ecol. 6: 376-381.

June 2002

Fedorka & Mousseau: Nuptial Gifts in a Cricket

THORNHILL, R. 1976. Sexual selection and paternal in-
vestment in insects. American Naturalist 110: 153-
THORNHILL, R., AND J. ALCOCK. 1983. The evolution of
insect mating systems. Harvard University Press,
VAHED, K. 1998. The function of nuptial feeding in insects:
a review of empirical studies. Biol. Rev. 73: 43-78.

WALKER, T. J. 1980. Reproductive behavior and mating
success of male short-tailed crickets: differences
within and between demes. Evol. Biol. 13: 219-260.
WEDELL, N. 1997. Ejaculate size in bushcrickets: the
importance of being large. J. Evol. Biol. 10: 315-325.
WIGGINS, D. A. AND R. D. MORRIS. 1988. Courtship feed-
ing and copulatory behavior in the common tern
Sterna hirundo. Ornis Scandinavica 19: 163-165.

Florida Entomologist 85(2)


1Dept. Biological Sciences, Oral Roberts University, Tulsa, OK 74171

2USDA, ARS, Yakima Agriculture Research Laboratory, 5230 Konnowac Pass Rd., Wapato, WA 98951


In a flight tunnel, mated female codling moths, Cydia pomonella L., were attracted (upwind
flight with zigzagging flight patterns) to cold-stored thinning apples. Greater numbers of cod-
ling moths were attracted to apples infested with codling moth larvae than to uninfested apples.
However, codling moth response to piped odor from cold-stored thinning apples infested with
larvae was not significantly greater than that of moths to piped odor from uninfested apples.
In a flight tunnel, significant numbers of mated female codling moths were captured in traps
baited with fresh-picked immature apples or in traps through which odor from such apples
was piped. Also, more codling moths were captured in traps baited with infested versus un-
infested apples, and more were captured in traps with odor from infested apples compared
to odor from un-infested apples. These studies demonstrate upwind attraction by flying fe-
male codling moths to apple fruit and odors from apple fruit and show increased response by
moths to odors of fruit that are infested with codling moth larvae. It is suggested that this
heightened response to infested apples may be due to increased apparency of infested fruit
that may release greater amounts of volatile odorants.

Key Words: codling moth, host-finding, attraction, apple, kairomone


En un tunel de vuelo, hembras apareadas de Cydia pomonella L. fueron atraidas (vuelo con-
tra el viento en un patron zigzag) a manzanas inmaduras y refrigeradas. Un mayor numero
de palomillas fueron atraidas a manzanas infestadas con larvas que a manzanas no infesta-
das. Sin embargo, la respuesta de las palomillas al olor de manzanas inmaduras refrigeradas
e infestadas con larvas introducido por un tubo no fu6 significativamente mayor que al de
manzanas no infestadas. En el tunel de vuelo, un numero significativo de hembras apareadas
fueron capturadas en trampas cebadas con manzanas inmaduras reci6n cosechadas o en
trampas con el olor introducido de tales manzanas. Tambi6n se capturaron mas palomillas en
trampas cebadas con manzanas infestadas que no infestadas, y se capturaron mas palomillas
en trampas con el olor de manzanas infestadas que con el olor de manzanas no infestadas.
Estos studios demuestran la atracci6n de las palomillas hembra de Cydia pomonella en
vuelo contra el viento hacia manzanas y hacia el olor de manzanas y demuestra una res-
puesta mayor de las palomillas hacia los olores de frutos infestados con larvas de C. pomone-
lla. Esta respuesta mayor a manzanas infestadas puede ser debido a una mayor apariencia
de las frutas infestadas por que tal vez emiten mayores cantidades de olores volatiles.

Adult codling moths are attracted to host fruit
and codling moth attraction responses to apple
odorants are thought to be important host-finding
behavior (Wearing et al. 1973, Wearing and
Hutchins 1973, Sutherland et al. 1974, Hern and
Dorn 1999, Yan et al. 1999). Although Wearing et
al. (1973) could not show upwind movement by
moths in an olfactometer in response to apple
odor, they and others (Wearing and Hutchins
1973, Sutherland et al. 1974) speculated that ap-
ple odor, as well as the apple odorant a-farnesene,
may be an attractant for the adult female codling
moth. An ambulatory upwind response to apple
odor by adult codling moths was recently demon-
strated by Yan et al. (1999). In that study, both
virgin and mated female moths moved farther up-

wind in tubes with apple odor compared to tubes
without apple odor. Using a Y-tube olfactometer
design, Hern and Dorn (1999) demonstrated an
ambulatory orientation response by codling moth
females to the apple odorant (E,E)-a-farnesene.
Attraction of flying codling moths to apple odor
has yet to be shown, although Light et al. (2001)
trapped male and female codling moth with the
pear chemical ethyl (E,Z)-2,4-decadienoate. Un-
derstanding such behavior is critical to the devel-
opment of appropriate assays to explore codling
moth host-finding and to isolate and identify host
plant kairomones that codling moths use to locate
and select oviposition sites.
Adult moth host-finding behavior can include
chemoanemotactic flight in response to host odor

June 2002

Reed & Landolt: Attraction of Codling Moth to Apple

(Phelan and Baker 1987, Landolt 1989, Tingle et
al. 1989, Rojas and Wyatt 1999). Such responses
appear to be similar to those documented for male
moth responses to female pheromone, a combina-
tion of upwind chemoanemotaxis with self-
steered countering (Baker 1989). We investigated
whether codling moths, like other moths, respond
to host (apple) odor with zigzagging upwind
flights that lead them to the odor source.
Attraction or orientation responses of phyto-
phagous insects to host plant odor may be en-
hanced or increased with injury to the plant. This
has been noted in flight tunnel studies of cabbage
looper moth, Trichoplusia ni Hfibner, attraction
to cotton plants, but not to cabbage or potato
plants (Landolt 1993, 2001), and in both wind
tunnel and olfactometer ambulatory assays of
Colorado potato beetle, Leptinotarsa decemlin-
eata Say, responses to potato plants (Bolter et al.
1997, Landolt et al. 1999), and larval codling
moth attraction to apple fruit (Landolt et al. 1998,
2000). Particularly because of the increased re-
sponse of codling moth larvae to apple fruit in-
fested with other codling moth larvae (Landolt et
al. 2000), adult codling moths may also respond
more strongly to the odors of infested apple fruit,
compared to un-infested fruit. Additional experi-
ments were conducted to test this hypothesis, us-
ing a flight tunnel assay.



Moths used in assays were obtained as pupae
from a laboratory colony maintained on an artifi-
cial diet (Toba and Howell 1991) at the Yakima
Agricultural Research Laboratory, Wapato, Wash-
ington. Pupae were sorted by sex and were placed
in plastic-screened cages in a controlled environ-
ment room. The room was at 23C, 50-70% RH,
with both a red incandescent light on 24 h per day
and white fluorescent lamps on a reversed 14 h
light:10 h dark photocycle. As adults emerged, pu-
pae were moved to new cages daily to provide
males and females of known ages. Cages of moths
were provided water on cotton.
To obtain mated female codling moths for bio-
assays, 15-20 females (2-3 day old) were placed in
one cage with 20-25 males (2-5 day old) for one
scotophase (dark period of the light cycle). Fe-
males were removed during the following photo-
phase (light period) and were then used in flight
tunnel tests during the next 2 scotophases. Fe-
males used in bioassays were dissected to confirm
the presence of spermatophores in the bursa cop-
ulatrix. Data sets for groups of females that were
<90% mated were rejected (one group).
For flight tunnel experiments involving obser-
vation of moth responses to cold-stored thinning
apples, Red Delicious apples were obtained from a

commercial apple orchard in June of 1999 and
were placed in cold storage at 2C until used in
experiments from December 1999 through Febru-
ary 2000. Apples were infested by placing 2 neo-
nate codling moth larvae with individual apples
in paper cups with lids for 5 to 12 days. Infesta-
tion was evident by physical damage to the fruit
and the presence of frass. Apples were 30 to 45
mm in diameter.
Fresh-picked Granny Smith apples were ob-
tained from commercial apple orchards in June
and July of 2000 and in July of 2001. Apples were
picked at 800 to 900 h the morning of the day they
were used in flight tunnel experiments. Apples
were infested by confining 3rd instar codling
moth larvae with sleeved fruit on trees for 5 days
prior to picking of fruit and their use in assays.
Again, successful infestation of fruit was con-
firmed by the presence of physical damage and
frass. Fresh-picked apples were 30 to 50 mm in di-
ameter, depending on the date.

Observational Experiments

A set of experiments was conducted as obser-
vational assays of moth responses in a flight tun-
nel. The flight tunnel was a 1 m wide x 1 m tall x
2 m long plexiglas box with a blower motor push-
ing air into the upwind end and a second blower
motor pulling air through the downwind end. Air-
flow was balanced to produce a slight positive
pressure in the tunnel, with an in-tunnel air speed
of 22 cm/sec. Charcoal-coated filters were installed
at both ends of the flight tunnel. Moths were
tested one at a time, released from a horizontal
polystyrene vial near the center of the downwind
end of the tunnel, during the 2nd and 3rd h of the
scotophase. Moths were observed for 3 min and
were scored for upwind oriented flight and con-
tact with the odor source (apples or airflow vent)
at the center of the upwind end of the tunnel. Up-
wind oriented flights were zigzagging upwind
flights within the likely odor plume downwind of
the fruit or pipe vent. These flights are here re-
ferred to as attraction, or upwind oriented flights.
The first experiment compared moth re-
sponses to 3 cold-stored thinning apples that were
not infested with a codling moth larva, 3 cold-
stored thinning apples infested with a codling
moth larva, and a control (no apples). Apples were
placed on a petri plate on a ring stand at the cen-
ter of the upwind end of the flight tunnel. On each
of 11 days, five moths were tested per treatment,
with the 3 treatments rotated in the treatment
sequence each day. Totals of 55 moths were tested
per treatment in this experiment. Treatment
means were separated by Tukey's test following a
significant ANOVA F value (DataMost 1995).
The second experiment compared mated fe-
male codling moth responses to 1) airflow that
was passed over 6 un-infested cold-stored thin-

Florida Entomologist 85(2)

ning apples in a glass jar, 2) airflow passed over 6
such apples each infested by a codling moth larva,
or 3) airflow through a "system blank" consisting
of the jar and plumbing but with no apples. On
each of 10 days, 5 females were tested per treat-
ment, with the 3 treatments rotated in the treat-
ment sequence each day. Totals of 50 moths were
tested per treatment. Treatment means were sep-
arated by Tukey's test following a significant
ANOVA F value (DataMost 1995).

Flight Tunnel Trapping Assays

Two sticky traps containing a bait or odor
source were placed near the center of the upwind
end of the flight tunnel at the beginning of the
scotophase. Traps were triangular cardboard tent
traps (10 cm wide, 15 cm tall and 10 cm deep)
painted yellow and coated on all 3 inside panels
with adhesive. Two traps were set up for each as-
say, with one serving as a control and the other
containing an apple treatment. Groups of 20-25
female codling moths were released from five
20-ml clear plastic vials hung horizontally at the
center of the downwind end of the flight tunnel at
the end of the second hour of the scotophase. Vials
were open on the upwind end and screened on the
downwind end to permit moth escape upwind into
the tunnel. Traps were 20 cm apart and 30 cm
from the tunnel walls. Four hours after the re-
lease of codling moths into the flight tunnel, traps
were checked to count codling moth captured in
the traps Three experiments were conducted to
evaluate codling moth responses to two or three
fresh-picked, immature apples, using this experi-
mental design. Data were analyzed as mean per-
centages of released moths recaptured in traps.
Treatment and control means were compared by
a paired t-test (DataMost 1995) to determine if
they were significantly different.
The first trapping experiment evaluated moth
response to two un-infested apples in a trap com-
pared to a control trap (no apples). The assay was
conducted six times, with releases of moths in the
flight tunnel made on each of six days. The second
trapping experiment evaluated moth response to
two infested apples in a trap compared to a con-
trol trap (no apples). This assay was conducted
five times, with releases of moths in the flight
tunnel made on each of five days. The third trap-
ping experiment compared moth response to two
un-infested apples and to two infested apples, in a
trap. This assay was conducted six times, with re-
leases of moths in the flight tunnel made on each
of six days.
Three additional trapping tests evaluated
moth response to airflow passed over fresh picked
immature apples. For each test replicate, three
apples were placed in a glass jar and air was
pumped through the jar and into a trap in the
flight tunnel. Airflow was also pumped through

an empty jar and into a second trap that served as
an experimental control. Twenty to 25 female cod-
ling moths were released at the downwind end of
the tunnel at the end of the second hour of the sco-
tophase and traps were checked 4 h later to count
moths captured on the sticky insides of the traps.
The fourth trapping experiment compared moth
response to airflow with and without uninfested
apples in the jar. This test was replicated five
times over five days. The fifth trapping experi-
ment compared moth response to airflow with
and without apples infested with codling moth in
the jar. This test was also replicated five times
over five days. The sixth trapping experiment
compared moths trapped in response to infested
apples compared to un-infested apples. This test
was replicated 10 times over 10 days. Apples were
infested in the field by placing 3rd instar codling
moth larvae on fruit in sleeves on the tree, five
days prior to assays.
Data for moths captured in traps were ana-
lyzed as percentages of released moths recap-
tured in traps. Differences between treatments or
between treatments and controls were deter-
mined using a paired t-test (DataMost 1995) at p


Flight Tunnel Observational Assays

Mated female codling moths were attracted to
and contacted cold-stored thinning apples pre-
sented in the flight tunnel (Table 1). Percentages
of moths exhibiting upwind oriented flights to-
wards either un-infested or infested apples were
significantly greater than percentages of moths
responding to the control. Also, percentages of
moths contacting un-infested or infested apples
were significantly greater than percentages of
moths contacting the control. Percentages of
moths attracted to infested apples were signifi-
cantly greater than percentages of moths at-
tracted to un-infested apples (Table 1), but source
contact was similar on infested and un-infested
apples in this test.
Mated female codling moths were also at-
tracted to odors of cold-stored thinning apples.
Percentages of moths exhibiting upwind oriented
flights towards the pipe that vented odor of un-
infested or infested apples were significantly
greater than the percentages of moths responding
to the control airflow (Table 1). Also, percentages
of moths contacting the odor source (pipe vent)
were greater in response to odor from over un-
infested or infested apples compared to the system
control (Table 1). Percentages of moths respond-
ing (attraction or contact) to odor of infested
apples were not statistically different than per-
centages of moths responding to odor of un-
infested apples.

June 2002

Reed & Landolt: Attraction of Codling Moth to Apple


Treatment n % Upwind oriented flight % Source contact

Experiment 1
Blank 55 0.0 + 0.0 a 0.0 + 0.0 a
Un-infested apples 55 18.2 + 3.3 b 12.7 + 3.0 b
Infested apples 55 38.2 + 8.3 c 18.2 + 6.3 b
Experiment 2
Blank airflow 50 0.0 + 0.0 a 0.0 + 0.0 a
Un-infested apple airflow 50 10.0 + 3.3 b 8.0 + 0.0 b
Infested apple airflow 50 8.0 + 4.4 b 6.0 + 4.3 b

Within a column and within an experiment, means followed by a the same letter are not significantly different by Tukey's test, at p < 0.05.

Flight Tunnel Trapping Assays


Significantly more moths were captured in
flight tunnel traps baited with immature apples,
compared to moths captured in un-baited traps,
when those apples were infested with a codling
moth larva (Table 2). In a direct comparison of un-
infested and infested apples, significantly more
moths were captured in traps baited with infested
apples (Table 2).
Female codling moths were captured in traps
into which odor (as airflow) from over fresh-
picked apples was introduced, indicating attrac-
tion to the apples (Table 2). This response was sig-
nificant in comparison to traps without odor from
apples, both using apples infested with codling
moth and using apples that were not infested. In
a direct comparison of odor from infested apples
and odor from un-infested apples, significantly
more moths were captured in traps baited with
odor of infested apples (Table 2).

These results demonstrate attraction re-
sponses of flying mated female codling moths to
apple fruit and to apple fruit odor. Previous stud-
ies had shown attraction by walking codling moth
adult to apple fruit, apple odor, and the apple
odorant E,E-a-farnesene (Hern and Dorn 1999,
Yan et al 1999), but not flight responses. Flight re-
sponses by moths to host plants have been shown
for the noctuids T ni (Landolt 1989), Heliothis
subflexa Tingle et al. 1989), and Mamestra bras-
sica L. (Rojas and Wyatt 1999), and the plutellid
Plutella xylostella (L.) (Palaniswamy et al. 1986,
and Pivnick et al. 1990). Additionally, the codling
moth has been captured in traps baited with the
pear volatile ethyl-(Z, E)-2,4-decadienoate (Light
et al. 2001). Such flight responses indicate a po-
tential for long distance host-finding by the co-
dling moth, with a significant role of host odor in
host finding.


% Upwind oriented flight

Treatment N Apples in trap N Apple airflow into trap

Experiment 1
Blank 6 1.3 + 0.8 a 5 2.0 + 1.2 a
Un-infested apples 3.3 + 1.0 a 7.0 + 1.2 b
Experiment 2
Blank 5 1.6 + 1.6 a 5 1.4 + 1.4 a
Infested apples 18.0 + 2.7 b 9.0 + 2.1 b
Experiment 3
Un-infested apples 5 0.9 + 0.9 a 10 2.0 + 0.8 a
Infested apples 7.5 + 1.6 b 5.5 + 1.6 b

Means within an experimental comparison that are followed by the same letter are not different by a paired t-test at p < 0.05.

Florida Entomologist 85(2)

These results also indicate that host-attraction
responses of adult codling moth, like that of neo-
nate larvae, are enhanced by prior infestation of
fruit by the codling moth. Both anemotactic and
klinotactic responses by neonate larvae of the cod-
ling moth are enhanced when larvae are in the air-
stream with odor from apple fruit infested with
other codling moth larvae, compared to un-in-
fested fruit (Landolt et al. 2000). Similar enhanced
orientation responses by herbivores to damaged or
induced plant odors have been shown for the cab-
bage looper moth and for the Colorado potato bee-
tle (Landolt 1993, Landolt et al. 1999). Although it
might appear disadvantageous for a phytopha-
gous insect to respond to an infested plant as an
oviposition site, because the moth's offspring
would likely face immediate competition for food,
there are other explanations for the observed be-
havior. If host plants occur in patches, locating an
infested plant or fruit may provide proximity to
other, un-infested, plants or fruits. In flight tunnel
studies of cabbage looper attraction to cotton
plants, moths were likely to fly to plants damaged
by conspecific larvae but were also likely to then
oviposit on undamaged adjacent plants, when
given that choice (Landolt 1993). Plants, including
apple and pear, that are injured or induced, often
produce, or produce increased amounts of, particu-
lar odorants (Boeve et al. 1996, Landolt et al. 2000,
Scutareanu et al. 1997) and may be more chemi-
cally apparent to a plant-seeking insect compared
to an un-injured plant. Perhaps the codling moth
cannot easily detect apple fruit from a distance un-
less it is injured or infested.
The attraction response rates of codling moths
in these flight tunnel assays were not high (up to
35%), compared to male moth responses to phero-
mones, but are not out of line with studies of other
moths and their responses to host plants. Rates of
attraction to cabbage plants by cabbage looper
moths in a flight tunnel ranged from 22% for
males up to 41% for females (Landolt 1989). At-
traction responses of the same moth to potato
plants were 20-26% (Landolt 2001). Rojas (1999)
reported attraction (upwind flight) response rates
of ca 20-55%) for female Mamestra brassicae (L.)
moths to cabbage plants in a flight tunnel.
This information should assist efforts to iso-
late and identify additional host attractants for
codling moth and to understand the role of vola-
tile chemicals in host finding and host selection
behavior of this insect. If moths use flight re-
sponses to apple odor as a means of locating ovi-
position sites, then flight attraction assays might
be useful in studying codling moth host finding
behavior and also in isolating those chemicals
emitted from apple fruit that attract codling
moth. Additionally, if infested fruit are more
strongly attractive to codling moth adults and lar-
vae, then the odors of these fruits might best be
used in studies to isolate and identify improved

host attractants for the codling moth. Qualitita-
tive and quantitative comparisons of the odor of
un-infested and infested apple fruit may provide
clues as to which volatile chemicals are important
to codling moth host finding and host selection.


Technical assistance was provided by J. F. Alfaro,
L. L. Biddick, J. A. Brumley, J. R. Dedlow, and D. Lovelace.
Helpful comments to improve the manuscript were made
by W. L. Yee and R. S. Zack. This work was supported by
funding from the Washington State Tree Fruit Research
Commission, by a USDA, NRI Research Career Enhance-
ment grant to Oral Roberts University, and by a sabbati-
cal leave from Oral Roberts University to H. C. Reed.


BAKER, T. C. 1989. Sex pheromone communication in the
Lepidoptera: New research progress. Experientia 45:
T. C. J. TURLINGS. 1996. Volatiles emitted by apple
fruitlets infested by larvae of the European apple
sawfly. Phytochemistry 42: 373-381.
SER, AND M. A. POSTHUMUS. 1997. Attraction of Col-
orado potato beetle to herbivore-damaged plants
during herbivory and after its termination. J. Chem.
Ecol. 23: 1003-1023.
DATAMOST. 1995. StatMost statistical analysis and
graphics. DataMost Corporation. Salt Lake City, UT.
HERN, A., AND S. DORN. 1999. Sexual dimorphism in the
olfactory orientation of adult Cydia pomonella in
response to alpha farnesene. Entomol. Exp. Appl. 92:
LANDOLT, P. J. 1989. Attraction of the cabbage looper to
host plants and host plant odor in the laboratory. En-
tomol. Exp. et Appl. 53: 117-124.
LANDOLT, P. J. 1993. Effects of host plant leaf damage
on cabbage looper moth attraction and oviposition.
Entomol. Exp. et Appl. 67: 79-85.
LANDOLT, P. J. 2001. Moth experience and not plant in-
jury affected female cabbage looper moth (Lepidop-
tera: Noctuidae) orientation to potato plants. Florida
Entomol. 84: 243-249.
1998. Neonate codling moth larvae (Lepidoptera:
Tortricidae) orient anemotactically to odor of imma-
ture apple fruit. Pan-Pacific Entomol. 74: 140-149.
1999. Attraction of Colorado potato beetle (Coleop-
tera: Chrysomelidae) to damaged and chemically in-
duced potato plants. Environ. Entomol. 28: 973-978.
BIDDICK, AND R. W. HOFSTETTER 2000. Apple fruit
infested with codling moth are more attractive to ne-
onate codling moth larvae and possess increased
amounts of (E,E)-alpha farnesene. J. Chem. Ecol. 26:
CAMPBELL. 2001. A pear-derived kairomone with
pheromonal potency that attracts male and female

June 2002

Reed & Landolt: Attraction of Codling Moth to Apple

codling moth, Cydia pomonella (L.). Naturwissen-
schaften 88: 333-338.
Attraction of diamondback moth, Plutela xylostella
(L.) (Lepidoptera: Plutellidae), by volatile com-
pounds of canola, white mustard, and faba bean.
Can. Entomol. 118: 1279-1285.
PHELAN, P. L., AND T. C. BAKER. 1987. An attracticide for
control of Amyelois transitella (Lepidoptera: Pyral-
idae) in almonds. J. Econ. Entomol. 80: 779-783.
AND E. W. UNDERHILL. 1990. Attraction of the dia-
mondback moth (Lepidoptera: Plutellidae) to vola-
tiles of oriental mustard: the influence of age, sex
and prior exposure to mates and host plants. Envi-
ron. Entomol. 19: 704-709.
ROJAS, J. C. 1999. Influence of age, sex and mating sta-
tus, egg load, prior exposure to mates, and time of
day on host-finding behavior of Mamestra brassicae
(Lepidoptera: Noctuidae). Environ. Entomol. 28:
ROJAS, J. C., AND T. D. WYATT. 1999. Role of visual cues
and interaction with host odour during the host-find-
ing behavior of the cabbage moth. Entomol. Exp. et
Appl. 91: 59-65.
MUS, AND M. W. SABELIS. 1997. Volatiles from Psylla
infested pear trees and their possible involvement in

attraction of anthocorid predators. J. Chem. Ecol. 23:
WEARING. 1974. The role of the hydrocarbon a-farne-
sene in the behavior of codling moth larvae and
adults, pp. 249-263. In Browne, L. B. (ed.), Experi-
mental Analysis of Insect Behaviour. Springer-Ver-
lag, New York.
Flight responses of Heliothis subflexa (Gn.) (Lepidop-
tera: Noctuidae) to an attractant from ground cherry,
Physalis angulata L. J. Chem. Ecol. 218: 168-170.
TOBA, H. H., AND J. F. HOWELL. 1991. An improved sys-
tem for mass-rearing codling moths. J. Entomol. Soc.
B. C. 3: 625-631.
WEARING, C. H., AND R. F. N. HUTCHINS. 1973. a-farne-
sene, a naturally occurring oviposition stimulant for
the codling moth, Laspeyresia pomonella. J. Insect
Physiol. 19: 1251-1256.
Olfactory stimulation of oviposition and flight activ-
ity of the codling moth Laspeyresia pomonella, using
apples in an automated olfactometer. N. Z. J. Science
16: 697-710.
ioral response of female codling moths, Cydia
pomonella, to apple volatiles. J. Chem. Ecol. 25:

Florida Entomologist 85(2)

June 2002


1USDA, ARS, U.S. Horticultural Research Laboratory, 2001 South Rock Road, Fort Pierce, FL 34945

2Visiting scientist, Biological Control Research Institute, Fujian Agricultural University, Fuzhou, Fugian 35002 China

'Visiting scientist, USDA, ARS, U.S. Horticultural Research Laboratory, 2001 South Rock Rd., Fort Pierce, FL 34945


A commercial preparation of the microbial entomopathogen, Bacillus thuringiensis subsp.
tenebrionis (Btt) was evaluated for biological activity against the Diaprepes root weevil, Dia-
prepes abbreviatus (L.). A reduction in survival was observed for neonatal larvae exposed to
insect diet incorporated with Btt and in potted citrus treated with a Btt soil application. A
treatment-induced, weight gain reduction for neonates was indicated only in the diet assay.
Larvae exposed at 5 weeks old to diet treated with Btt demonstrated a dose-dependent mor-
tality response. The mean ages for larval death ranged from 111 to 128 days among treat-
ments. The LC5s for larvae in this age group was 6.2 ppm [AI] and the slope of the probit line
was 2.29. The mortality response of larvae exposed at 12 weeks old also was dose dependent
and the mean ages for larval death ranged from 130 to 141 days among treatments. The LC5s
for larvae in this age group was 25.4 ppm [AI] and the slope of the probit line was 2.75. The
delayed patterns of mortality that we observed among larvae treated at 5 and 12 weeks old
indicates that disease is slow to develop in older larvae but that death occurs before matura-
tion is completed.

Key Words: Diaprepes abbreviatus, Bacillus thuringiensis, entomopathogen, citrus, Diaprepes
root weevil


Se evalu6 una formulaci6n commercial del entomopat6geno microbial, Bacillus thuringiensis
subsp. tenebrionis (Btt) por su actividad biol6gica en contra del gorg6jo Diaprepes abbrevia-
tus (L.). Se observe una reducci6n en la sobrevivencia de larvas neonatas expuestas a una
dieta artificial que contenia Btt, y de larvas en potes con arboles tratados con Btt. Una re-
ducci6n del peso ganado por las larvas neonatas fue inducida por el tratamiento solamente
en el caso del ensayo con dieta artificial. Larvas expuestas a dieta tratada a las 5 semanas
de edad manifestaron una mortalidad proporcional a la dosis. El rango del promedio del nu-
mero de dias hasta la muerte de las larvas fue de 111 a 128 dias entire tratamientos. La DL50
para las larvas de este grupo fue de 6.2 ppm (IA) y el pendiente de la linea probit fue de 2.29.
La mortalidad de larvas expuestas a las 12 semanas de edad fue dependiente de la dosis y
la edad promedia de la muerte vari6 entire 130 y 141 dias entire tratamientos. La DL50 para
las larvas de este grupo fue de 25.4 ppm (IA) y el pendiente de la linea probit fue de 2.75. El
patron de mortalidad que observamos entire larvas tratadas a las 5 y a las 12 semanas de
edad indica que la enfermedad se desarroll6 lentamente en las larvas mayores pero la
muerte ocurri6 antes de la maduraci6n.

The most economically important insect pest ranean larvae extensively damage host plant root
of citrus in Florida is the Diaprepes root weevil, systems on which they feed. Larval feeding in cit-
Diaprepes abbreviatus (L.). Losses to citrus grow- rus predisposes root tissues to infections by
ers are estimated in excess of $75 million yearly pathogens such as Phytophthora spp. (Rogers et
(Anonymous 1997). This root weevil also attacks al. 1996), induces a decline in tree health and
sugar cane, vegetable crops, and ornamental fruit production, and may eventually kill the tree
plantings in areas that are infested. The subter- (Schroeder & Sutton 1977). In severe cases of un-
controlled weevil populations, entire citrus groves
are destroyed (pers. obs.).
Mention of a trademark or proprietary product does not con- Entomopathogens have received considerable
stitute a guarantee or warranty of the product by the U.S. De-
partment of Agriculture and does not imply its approval to the attention as potential biocontrol agents of root
exclusion of other products that may also be suitable. weevil larvae. Beavers et al. (1983) and Roman

Weathersbee et al.: Susceptibility of D. abbreviatus to B. thuringiensis

and Beavers (1983) initially demonstrated the
pathogenicity and seasonal prevalence of several
naturally occurring fungi and nematodes attack-
ing D. abbreviatus. Since then, many others have
demonstrated the potential for biological control of
root weevil larvae by entomopathogens (Duncan et
al. 1996; Figueroa & Roman 1990; Quintella & Mc-
Coy 1997, 1998; Schroeder 1987, 1994). Nematode
applications were recommended for control as re-
cently as 1999, but the insecticidal compound,
bifenthrin, is currently the only material listed in
the Florida Citrus Pest Management Guide for
control of larval stages (Futch et al. 2001).
Bacterial entomopathogens have not been pre-
viously investigated as potential biocontrol
agents for Diaprepes root weevil. The microbial
pathogen, Bacillus thuringiensis subsp. tenebrio-
nis (Brr, was shown to have activity against rep-
resentatives of the order Coleoptera including the
families Chrysomelidae, Bostrichidae, and Cur-
culionidae (Herrnstadt et al. 1986; Krieg et al.
1983, 1987; Beegle 1996; Saade et al. 1996). The
bacterium produces a crystalline 6-endotoxin dur-
ing the process of sporulation (Hofte & Whiteley
1989). Upon ingestion by a susceptible insect
host, the endotoxin is proteolytically activated in
the midgut and causes feeding inhibition, septi-
caemia, and eventual death (Knowles 1994). We
investigated a commercially available Btt prod-
uct, Novodor (Novo Nordisk, North Chicago, IL),
to determine if it had biological activity against
D. abbreviatus, envisioning that a demonstration
of activity may provide the impetus to develop
this and possibly other B. thuringiensis subsp. for
field application.


Insect Rearing

Neonatal, 5-week-old, and 12-week-old larvae
ofD. abbreviatus were obtained from a laboratory
colony maintained by the U.S. Horticultural Re-
search Laboratory, Fort Pierce, FL. Larvae were
reared on a commercially-prepared insect diet
(Product No. F1675, Bio-Serv, Frenchtown, NJ)
placed within sealed diet cups (PC100, 30 ml cups
and lids, Jet Plastica, Harrisburg, PA). Prepara-
tion of insect diet and rearing of larvae were sim-
ilar to the methods described by Beavers (1982)
with temperature and moisture content of diet op-
timized for larval development (Lapointe 2000,
Lapointe & Shapiro 1999). Weevil larvae used in
the bioassays were maintained in diet cups stored
in plastic trays in a growth chamber at 26C and
a 24-h dark cycle.

Diet Incorporation of Btt

Prepared insect diet was heated to 90C for 15
min., covered with foil, and allowed to cool to 56C

in a heated water bath before incorporating treat-
ments. A commercial preparation of B. thuring-
iensis subsp. tenebrionis (Novodor 3% [Al], [30 mg
spores and 6-endotoxin crystals per ml product])
was incorporated into the diet at rates of 0, 0.3,
3.0, 30, and 300 ppm (pg AI/ml diet). Treatments
were incorporated into the agar-laden diet with
the aid of a heated, stirrer plate. The resulting
mixtures were pipetted into diet cups (15 ml diet
per cup), allowed to solidify and cool to room tem-
perature, and then were closed with a lid. All
steps after heating of the diet were performed in a
laminar flow, clean bench to avoid contamination.

Evaluation of Btt Activity

The biological activity of Novodor was evalu-
ated initially against neonatal larvae exposed to
treated insect diet. Neonates were briefly surface
sterilized in a 5% bleach solution and rinsed with
sterile, deionized water prior to being placed in
diet cups. The treatments included the 5 rates of
treatment-incorporated diet described above and
there were 3 replications of each treatment. A
treatment comprised 18 diet cups, each infested
with 5 neonatal larva. The numbers and weights
of surviving larvae in each diet cup were assessed
after 6 weeks of exposure to treated diet.
The activity of Novodor also was evaluated
against 5-week old weevil larvae. Larvae used in
the experiment weighed ~110 mg before exposure
to treatments. Treatments were prepared as
above and replicated 6 times. Each treatment
comprised 18 diet cups, each containing 1 larva.
Mortality, age at death, and weights of closed
adults were recorded twice weekly. The experi-
ment was terminated after 5 months when most
larvae had either died or matured to adults.
Novodor activity also was evaluated against
12-week old larvae that weighed ~510 mg before
exposure to treatments. The treatments were in-
corporated in diet at the rates described above.
There were 3 replications of treatments, each
comprising 30 larvae confined individually to diet
cups. Mortality, age at death, and weights of
closed adults were recorded twice weekly. The
experiment was terminated after 3 months when
most larvae had either died or matured to adults.
Another experiment was conducted to deter-
mine the effect of Novodor suspensions applied as
soil treatments against neonatal larvae feeding
on potted citrus roots. The citrus plants used in
the study were 1 year old Cleopatra Mandarin
(Citrus reshni Hort., ex Tan.) rootstock potted in
473 cm3 containers of potting soil (Metromix 500,
Scotts, Marysville, OH). The treatments included
4 rates of Novodore (0, 3.0, 30, and 300 ppm [(g AI/
ml DI water]) applied as 50 ml suspensions to the
soil of each container. Two drench applications
were made 14 days apart. There were 6 replica-
tions of each treatment. A treatment comprised 1

Florida Entomologist 85(2)

potted citrus plant infested with 20 neonates im-
mediately after the first drench application. All
treatments were maintained in a growth chamber
at 26C with a photoperiod of 14:10 (L:D) h. The
numbers and weights of surviving larvae were as-
sessed after 6 weeks.

Data Analyses and Statistics
Percentage data were adjusted for control mor-
tality using the Abbott (1925) formula and trans-
formed (arcsine) before analyses. Data were
analyzed by the General Linear Models Proce-
dure, and differences among treatment means
were determined by Tukey's studentized range
test (SAS Institute 1990). Differences among
means were considered significant at a probabil-
ity level of 5 percent (P < 0.05). Probit analyses
were conducted using the Probit Procedure (SAS
Institute 1990) to generate LC5o values and slopes
of probit lines for larval mortality due to treat-
ments. Untransformed means were presented in
the data tables.


Neonatal Larvae
The survival of root weevil larvae exposed as
neonates to diet treated with Novodor was signif-
icantly reduced (F = 9.36; df = 4, 262; P < 0.0001)
after 6 weeks. While 2 of 5 larvae survived after 6
weeks in the control group, only 1 larvae survived
on average in the 300 ppm treatment (Table 1).
The low survival rate observed for control larvae
is addressed in the discussion section. The fresh
weights of surviving larvae were significantly re-
duced (F = 8.46; df = 4, 262; P < 0.0001) by treat-
ments indicating that larval feeding or nutrient
assimilation may have been inhibited. Surviving
larvae in the 300 ppm treatment weighed 53%
less than those in the control group (Table 1).

5-Week Old Larvae
Insect diet treated with Novodor caused a sig-
nificant increase (F = 76.57; df = 4, 20;P < 0.0001)

in mortality of larvae exposed at the age of 5
weeks. Mortality of larvae exposed to the 3 ppm
treatment (44%) was significantly (P < 0.05)
greater than that of the controls (15%). Larval
mortality in the 30 ppm treatment exceeded 50%
(Table 2). The calculated LCs0 for larvae in this
age group was 6.2 (95% FL = 2.7-13.2) ppm [AI].
The slope of the probit line was 2.29 (SE = 0.31)
(X2 = 43.07; df = 1; P < 0.0001). Treatments also
caused a significant reduction (F = 6.25; df = 4,20;
P = 0.0020) in the mean death ages of larvae. The
death ages for larvae maintained on treated diets
ranged from 111 to 128 d while that of control lar-
vae was 154 d. Larvae in the 30 and 300 ppm
treatments died significantly (P < 0.05) earlier
than the control larvae (Table 2). The mortality
response we observed occurred later than is typi-
cal for insects susceptible toB. thuringiensis; nev-
ertheless, the response was consistent, dose-
dependent, and occurred earlier than that ob-
served for the controls. We observed that disease
and death occurred in treated insects during the
later stages of larval development and early
phases of pupation (Fig. 1). The presence of Btt
was confirmed by isolating the bacteria from in-
ternal tissues of dead larvae. Bacterial isolates
were cultured in 40 ml LB broth in a 125 ml baf-
fled flask maintained in an incubator shaker at
28C and 200 rpm until sporulation phase. Cul-
tures were examined using Hoffman contrast
microscopy at 1000x magnification to identify Btt
endospores and 8-endotoxin crystals. The mean
fresh weights of adults that closed on treated di-
ets also were significantly reduced (F = 3.39; df =
4, 20; P = 0.0286). Adult weevils that survived the
300 ppm treatment weighed 15% less than con-
trol adults, indicating that inhibition of feeding or
disruption of nutrient assimilation may have oc-
curred among survivors of treatments (Table 2).

12-Week Old Larvae

A significant increase (F = 8.82; df = 4, 8; P <
0.0050) in mortality was observed for 12-week old
larvae fed Novodor treated diet. Mortality in the
30 ppm treatment (53%) was significantly (P <


Treatment Number of surviving Weight (mg) of surviving
(ppm AI)" larvae SE (n = 3)b larvae SE (n = 3)b

0.0 2.0 + 0.1 a 306.3 + 22.4 a
0.3 1.9 + 0.1 a 251.8 + 16.7 a
3.0 1.8 + 0.1 a 236.1 + 21.5 a
30.0 1.6 + 0.1 a 231.1 + 20.5 a
300.0 1.0 + 0.1 b 144.0 + 20.9 b

aAI refers to the concentration (pg/ml) of active ingredient, comprising spores and 5-endotoxin of B. thuringiensis var. tenebrionis, in prepared diet.
'Means within a column sharing the same letter were not significantly different (P > 0.05, Tukey's studentized range test [SAS Institute 1990]).

June 2002

Weathersbee et al.: Susceptibility of D. abbreviatus to B. thuringiensis


Treatment % Mortality + SE Death age (d) + SE Adult wt. (mg) + SE
(ppm AI)" (n = 6)b,' (n = 6)b,' (n = 6)b,'

0.0 15.3 + 4.4 a 154.3 + 14.5 a 315.0 + 12.0 a
0.3 30.3 + 4.3 a 127.9 + 10.0 ab 329.2 + 23.8 ab
3.0 43.7 + 5.8 b 126.9 + 8.6 ab 323.2 + 15.2 ab
30.0 58.1 + 4.9 c 114.4 + 8.1 b 294.3 + 9.0 ab
300.0 71.0 + 4.4 d 110.6 + 2.9 b 266.9 + 19.5 b

"AI refers to the concentration ((g/ml) of active ingredient, comprising spores and 8-endotoxin of B. thuringiensis var. tenebrionis, in prepared diet.
'Mortality, age at death, and fresh weights of closed adults were assessed twice weekly during the experiment.
'Means within a column sharing the same letter were not significantly different (P > 0.05, Tukey's studentized range test [SAS Institute 1990]).

0.05) greater than that for the control group (6%)
(Table 3). The calculated LC50 for larvae in this
age group was 25.4 (95% FL = 12.5-60.0) ppm
[AI]. The slope of the probit line was 2.75 (SE =
0.44) (x2 = 47.37; df = 1;P < 0.0001). The mean age
at which mortality occurred also was significantly
reduced by treatments (F = 4.54; df = 4, 8; P =
0.0330). The mean death age for larvae in the 300
ppm treatment (130 d) was significantly (P < 0.05)
less than that for control larvae (153 d) (Table 3).
Treatments did not significantly affect the fresh
weights of closed adults in this age group (F =
1.34; df = 4, 8; P = 0.3341), likely because larvae
used in the experiment had attained near maxi-
mum weights before exposure to treatments.

Soil Treatments

Survival of neonates on potted citrus was sig-
nificantly reduced (F = 4.87; df = 3, 15;P < 0.0146)
by soil treatments with Novodor. Larval survival
was significantly (P < 0.05) less in all treatments
as compared to the controls after 6 weeks expo-
sure to treated soils and citrus roots (Table 4).
The low rate of survival for control larvae is ad-
dressed in the discussion section. In contrast to
the results of the diet bioassay against neonates,
the fresh weights of surviving larvae were not sig-
nificantly reduced (F = 1.02; df= 3, 15;P = 0.4098)
in the potted citrus bioassay (Table 4). Some lar-
vae may have been able to avoid exposure to Btt
where soil drenches resulted in unequal disper-
sion of treatments.


4Y -

Fig. 1. Healthy (left) and diseased (right) larvae and
pupae of D. abbreviatus due to feeding on diet incorpo-
rated with Novodor.

The results of our experiments indicate that
neonatal, 5-week-old, and 12-week-old larvae of
D. abbreviatus were susceptible to the biological
effects of Novodor. Activity for B. thuringiensis
against older larval stages of insects is not nor-
mally expected; nevertheless, the dose-dependent
mortalities we observed for larvae exposed at 5
and 12 weeks old were reproducible.
The low survival rates we observed for neo-
nates in our controls are typical in experiments
such as this, where multiple larvae are collec-
tively grouped to challenge plants or generate
data for bioassays of diet-incorporated materials.
Low survival rates for neonatal D. abbreviatus
have been reported by others (Schroeder & Sie-
burth 1997, Quintella & McCoy 1997), and are
due to natural mortality factors including aggres-
sive interactions among larvae confined together
(Lapointe & Shapiro 1999).
The response we observed for older larvae to
Novodor may be attributed to the finding that
midgut trypsin activity increases to a maximum
in D. abbreviatus larvae at the age of approxi-
mately 7 weeks (Yan et al. 1999). Yan et al. also
inferred that the midgut of D. abbreviatus larvae

Florida Entomologist 85(2)


Treatment % Mortality SE Death Age (d) + SE Adult Wt. (mg) + SE
(ppm AI)" (n = 3)b,' (n = 3)b,' (n = 3)b,'

0.0 5.6 + 5.6 a 153.1 + 7.4 a 353.0 + 6.5 a
0.3 21.1 +13.1 ab 132.0 + 14.9 ab 331.2 + 22.2 a
3.0 31.1 + 21.5 abc 141.3 + 7.8 ab 317.6 + 32.3 a
30.0 53.3 + 15.4 bc 134.8 + 10.8 ab 338.5 + 1.3a
300.0 67.8 + 12.5 c 130.1 + 9.4 b 374.6 + 13.5 a

"AI refers to the concentration (pg/ml) of active ingredient, comprising spores and 8-endotoxin of B. thuringiensis var. tenebrionis, in prepared diet.
'Mortality, age at death, and fresh weights of closed adults were assessed twice weekly during the experiment.
'Means within a column sharing the same letter were not significantly different (P > 0.05, Tukey's studentized range test [SAS Institute 1990]).

is probably alkaline, as shown for many Lepi-
doptera and Diptera, since the enzyme activity
observed was most active at a pH of 10.4. Alkaline
solubilization and proteolytic cleavage by serine
proteases, such as trypsin, are required for acti-
vation of many B. thuringiensis 6-endotoxins (Pie-
trantonio et al. 1993) including those produced by
Btt. The Cry3a toxin of Btt may undergo substan-
tial cleavage at the N-terminus without loss of
biological activity (Carroll et al. 1989). It has been
suggested that these additional protease cleavage
sites may facilitate insertion of a portion of the
protein into the target membrane of the midgut
epithelium (Knowles 1994) and subsequent pore
formation. The mortality responses we observed
for larvae treated at 5 and 12 weeks old may have
been a product of appropriate levels of pH and
serine proteases in midgut tissues during those
stages of development. The delayed period to lar-
val death may have been a function of the ability
of D. abbreviatus to survive for long periods with-
out food or water (personal observation) and the
slow development of disease. Any future evalua-
tion of Btt on larval D. abbreviatus, and possibly
other insects, should consider the importance of
suitable midgut environments for endotoxin acti-
vation and the general hardiness of the insect.
We observed dose-dependent responses for
mortality, age at death, and fresh weights of adult
survivors from larvae treated at the age of 5
weeks. Although the mortality response devel-

oped slowly, the mean ages at death were consis-
tently earlier among treated as compared to
control larvae. Also, the fresh weights of adults
that survived treatments were reduced compared
to those of the control, indicating that midgut
damage or feeding inhibition may have occurred
among larvae that survived treatments.
A dose-dependent mortality response also was
observed for larvae treated at the age of 12 weeks.
Larvae treated at this stage of development often
died as pupae, further supporting the contention
that disease was slow to develop. The fresh
weights of the adults that developed from larvae
treated at 12 weeks old were not affected by treat-
ments. Lapointe (2000) reported that larval
weights of D. abbreviatus were maximum at 12
weeks old, and then declined during further de-
velopment, thus explaining the lack of a weight
response for larvae in this age group.
There is much yet to be learned about the acti-
vation and binding patterns of B. thuringiensis
endotoxins, and the subsequent onset of disease
in susceptible insects. We observed that disease
was slow to develop inD. abbreviatus. The results
of our experiment indicate that there may be age-
related factors that influence larval D. abbrevia-
tus response to Btt endotoxins. There may also be
other endotoxins awaiting discovery that are bet-
ter adapted to activation and binding by larval
D. abbreviatus. This report provides the impetus
to further explore these processes in the Dia-


Treatment Number of surviving Weight (mg) of surviving
(ppm AI)" larvae SE (n = 6)" larvae SE (n = 6)b

0.0 4.2 + 1.1 a 52.0 + 22.5 a
3.0 1.7 + 0.3 b 40.7 + 9.9 a
30.0 1.5 + 0.4 b 19.8 + 6.6 a
300.0 1.3 + 0.3 b 41.2 + 12.1 a

aAI refers to the concentration (pg/ml) of active ingredient, comprising spores and 5-endotoxin of B. thuringiensis var. tenebrionis, in prepared diet.
'Means within a column sharing the same letter were not significantly different (P > 0.05, Tukey's studentized range test [SAS Institute 1990]).

June 2002

Weathersbee et al.: Susceptibility of D. abbreviatus to B. thuringiensis

prepes root weevil and other insects. Our demon-
stration of Novodore activity against different
larval stages of D. abbreviatus is encouraging
enough to warrant additional testing.


The authors thank Ginny Schmit and Karin Crosby
(USDA, ARS, U.S. Horticultural Research Laboratory,
Fort Pierce, FL) for technical support and assistance
with insect rearing and preparation of insect diets.


ABBOTT, W. S. 1925. A method for computing the effective-
ness of an insecticide. J. Econ. Entomol. 18: 265-267.
ANONYMOUS. 1997. Diaprepes Task Force Minutes, July
17, 1997. University of Florida. Lake Alfred, FL.
BEAVERS, J. B. 1982. Biology of Diaprepes abbreviatus
(Coleoptera: Curculionidae) reared on artificial diet.
Florida Entomol. 65: 263-269.
BEAVERS, J. B., C. W. McCOY, AND D. T. KAPLAN. 1983.
Natural enemies of subterranean Diaprepes abbre-
viatus (Coleoptera: Curculionidae) larvae in Florida.
Environ. Entomol. 12: 840-843.
BEEGLE, C. C. 1996. Efficacy of Bacillus thuringiensis
against Lesser Grain Borer, Rhyzopertha dominica
(Coleoptera: Bostrichidae). Biocontrol. Sci. Tech. 6:
CARROLL, J., J. LI, AND D. J. ELLAR 1989. Proteolytic
processing of a coleopteran-specific 6-endotoxin pro-
duced by Bacillus thuringiensis var. tenebrionis. Bio-
chem. J. 261: 99-105.
1996. Estimating sample size and persistence of en-
tomogenous nematodes in sandy soils and their effi-
cacy against the larvae of Diaprepes abbreviatus in
Florida. J. Nematol. 28: 56-67.
FIGUEROA, W., AND J. ROMAN. 1990. Biocontrol of the
sugarcane rootstalk borer, Diaprepes abbreviatus
(L.) (Coleoptera: Curculionidae), with entomophilic
nematodes. J. Agric. Univ. P.R. 74: 395-404.
FUTCH, S. H., J. L. KNAPP, AND C. W. McCoY. 2001. Fact
Sheet ENY-611, 2001 Florida Citrus Pest Manage-
ment Guide, FCES, IFAS, University of Florida. Re-
vised: October 2000.
EDWARDS. 1986. A new strain of Bacillus thuringien-
sis with activity against coleopteran insects. Bio-
technology. 4: 305-308.
HOFTE, H., AND H. R. WHITELEY. 1989. Insecticidal crys-
tal proteins of Bacillus thuringiensis. Microbiol. Rev.
53: 242-255.
KNOWLES, B. H. 1994. Mechanism of action of Bacillus
thuringiensis insecticidal 6-endotoxins. Adv. Insect.
Physiol. 24: 275-308.
"Bacillus thuringiensis var. San Diego" strain M-7 is
identical to the formerly in Germany isolated B. thu-
ringiensis var. tenebrionis strain BI 256-82. J. Appl.
Entomol. 104: 417-424.

W. SCHNETTER. 1983. Bacillus thuringiensis var.
tenebrionis, a new pathotype effective against larvae
of Coleoptera. Z. Angew. Entomol. 96: 500-508.
LAPOINTE, S. L. 2000. Thermal requirements for the de-
velopment of Diaprepes abbreviatus (Coleoptera:
Curculionidae). Environ. Entomol. 29: 150-156.
LAPOINTE, S. L., AND J. P. SHAPIRO. 1999. Effect of soil
moisture on development of Diaprepes abbreviatus
(Coleoptera: Curculionidae). Florida Entomol. 82:
1993. Interaction of Bacillus thuringiensis endotox-
ins with the insect midgut epithelium, pp.55-80. In
N. E. Beckage, S. N. Thompson, and B. A. Federici
(eds.), Parasites and Pathogens of Insects, Vol 2. Ac-
ademic Press, New York.
QUINTELLA, E. D., AND C. W. MCCOY. 1997. Pathogenic-
ity enhancement ofMetarhizium anisopliae and Beau-
veria bassiana to first instars of Diaprepes abbreviatus
(Coleoptera: Curculionidae) with sublethal doses of
imadacloprid. Environ. Entomol. 26: 1173-1182.
QUINTELLA, E. D., AND C. W. McCoY. 1998. Synergistic
effect of imadacloprid and two entomopathogenic
fungi on the behavior and survival of larvae of Dia-
prepes abbreviatus (Coleoptera: Curculionidae) in
soil. J. Econ. Entomol. 91: 110-122.
ROGERS, S., J. H. GRAHAM, AND C. W. MCCOY. 1996. In-
sect-plant pathogen interactions: Preliminary stud-
ies of Diaprepes root weevil injuries and
Phytophthora infections. Proc. Fla. State Hort. Soc.
109: 57-62.
ROMAN, J., AND J. B. BEAVERS. 1983. A survey of Puerto
Rican soils for entomogenous nematodes which at-
tack Diaprepes abbreviatus (L.) (Coleoptera: Curcu-
lionidae). J. Agric. Univ. P.R. 67: 311-316.
Response of the Carrot weevil, Listronotus oregonen-
sis (Coleoptera: Curculionidae), to strains of Bacillus
thuringiensis. Biological Control. 7: 293-298.
SAS INSTITUTE. 1990. SAS/STAT user's guide for per-
sonal computers, vol. 6. SAS Institute, Cary, NC.
SCHROEDER, W. J. 1987. Laboratory bioassays and field
trials of entomogenous nematodes for control of Di-
aprepes abbreviatus (Coleoptera: Curculionidae) in
citrus. Environ. Entomol. 16: 987-989.
SCHROEDER, W. J. 1994. Comparison of two steinerne-
matid species for control of the root weevil Diaprepes
abbreviatus. J. Nematol. 26: 360-362.
SCHROEDER, W. J., AND P. J. SIEBURTH. 1997. Impact of
surfactants on control of the root weevil Diaprepes
abbreviatus larvae with Steinernema riobravis. J.
Nematol. 29: 216-219.
SCHROEDER, W. J., AND R. A. SUTTON. 1977. Citrus root
damage and the spatial distribution of eggs of Di-
aprepes abbreviatus. Florida Entomol. 60: 114.
LOCK, AND D. BOROVSKY. 1999. Sequencing and char-
acterization of the citrus weevil, Diaprepes
abbreviatus, trypsin cDNA: Effect of Aedes trypsin
modulating oostatic factor on trypsin biosynthesis.
Eur. J. Biochem. 262: 627-636.

Florida Entomologist 85(2)


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

Through laboratory choice tests involving 19 plant species, we assessed the host selection
behavior of six grasshopper species: Stenacris vitreipennis (Marschall) (glassywinged tooth-
pick grasshopper), Leptysma marginicollis (Serville) (cattail toothpick grasshopper),
Gymnoscirtetes pusillus Scudder (little wingless grasshopper), Paroxya clavuliger (Serville)
(olivegreen swamp grasshopper), Paroxya atlantica Scudder (Atlantic grasshopper), and
Romalea microptera (Beauvois) (eastern lubber grasshopper). This grasshopper assemblage
is commonly associated with semi-aquatic habitats in the southeastern United States. These
poorly studied species display both graminivorous (S. vetreipennis and L. marginicollis) and
mixed graminivorous-forbivorous feeding habits (the remaining species), the nature of
which are fairly predictable based on examination of mouthpart morphology, but not entirely
consistent with the tendency of cyrtacanthacridine species to feed on forbs.

Key Words: host preference, plant acceptance, mouthpart morphology


Por medio de pruebas de selecci6n de laboratorio utilizando 19 species de plants, evalua-
mos el comportamiento de selecci6n de hospederos de seis species de saltamontes: Stenacris
vitreipennis (Marschall), Leptysma marginicollis (Serville), Gymnoscirtetes pusillus Scudder,
Paroxya clavuliger (Serville), Paroxya atlantica Scudder, y Romalea microptera (Beauvois).
Este grupo de saltamontes, esta comunmente asociado con habitats semiacuaticos, en el su-
reste de los Estados Unidos. Estas species poco estudiadas demuestran habitos de alimen-
taci6n graminivora (S. vetreipennis y L. marginicollis) y de alimentaci6n mixta graminivora
y de hierbas (las species restantes), la naturaleza de los cuales son suficientemente prede-
cibles basandose sobre la examinaci6n de la morfologia del aparato bucal, pero no totalmente
consistentes con la tendencia de las species cirtacanthacridines para alimentarse de plan-
tas herbaceas.

Although often viewed as polyphagous herbi-
vores, most grasshoppers are selective to some de-
gree, exhibiting definite plant preferences
(Mulkern 1967). In previous studies, it has been
shown that grasshoppers are conveniently classi-
fied as grass-feeders (graminivorous), forb-feed-
ers (forbivorous), or a mix of the two (ambivorous
or mixed feeders) (Isely 1944). Phylogenetic dif-
ferences exist among grasshoppers in relation to
host plant preferences (Dadd 1963, Joern 1979).
For example, members of the acridid subfamily
Gomphocerinae tend to have a preference for
grasses, Cyrtacanthacridinae (Melanoplinae in
part) prefer forbs, and Oedopodinae eat both
grasses and forbs (Dadd 1963, Joern & Lawlor
1980, Otte 1981). Joern (1986) points out that
most monophagous and polyphagous species are
forb-feeders while oligophagous species are grass
Information on dietary habits of Florida's
grasshoppers is growing, though still far from
complete. Preference tests (Capinera 1993,
Scherer 1997) have been conducted for some of
Florida's upland plants and crops, though infor-
mation on most of Florida's grasshoppers is lack-

ing. Such tests commonly are used to construct
preference hierarchies (Lewis and van Emden
1986), but are constrained by experimental de-
sign. The investigator must have the wisdom to
present the correct array of plants, which should
be based on the habitat in which the insect is
found. Nevertheless, even good designs are sub-
ject to faulty interpretation, as lack of a "pre-
ferred" host among the array of choices may force
a hungry individual to feed on non-preferred
plants which, in nature, might be accepted only if
faced with starvation.
A very general method to determine the diet of
a grasshopper is by the morphology of the grass-
hopper's mandibles (Mulkern 1967, Patterson
1984). The morphological characters of the man-
dibles, incisor and molar surfaces are useful in la-
beling grasshoppers as grass- or forb-feeders
(Chapman 1964, Bernays & Barbehenn 1987,
Kang et al. 1999) though most species with forb-
feeding mandibles feed on a mixture of grasses
and forbs. Isley (1944) suggested that the study of
mandibular morphology would aid in under-
standing grasshopper ecology and their role in
terrestrial communities.

June 2002

Squitier & Capinera: Grasshopper Host Selection

We evaluated host selection behavior by grass-
hoppers among the plant species abundantly
found inhabiting semi-aquatic habitats. Most re-
search on grasshopper feeding behavior has been
conducted in arid environments, where grasshop-
pers most often attain high and damaging levels
of abundance. Very little is known concerning spe-
cies that inhabit moist or wet environments be-
cause such species usually do not become pests.
The host selection behavior displayed by grass-
hoppers in choice tests was compared with host
preference predictions based on mouthpart (man-
dible) morphology.


Grasshoppers were collected from several wet
habitats: freshwater marshes, lakeside, flat-
woods and ditchbanks. The grasshopper species
included in this study were Stenacris vitreipennis
(Marschall) (glassywinged toothpick grasshopper),
Leptysma marginicollis (Serville) (cattail tooth-
pick grasshopper), Gymnoscirtetes pusillus Scud-
der (little wingless grasshopper) Paroxya clavuliger
(Serville) (olivegreen swamp grasshopper), Par-
oxya atlantica Scudder (Atlantic grasshopper),
and Romalea microptera (Beauvois) (eastern lub-
ber grasshopper). The grasshoppers were field-
collected as large nymphs or adults and main-
tained in the laboratory during the experimental
period. The insects and choice tests were held in
an insect growth room at 30-32C with a photope-
riod of 14:10 (L:D) and a relative humidity of 40+
10%. Between study repetitions, the grasshop-
pers fed Romaine lettuce, a plant that seems to
have nearly universal acceptance by acridids.
The plant species were presented to the grass-
hoppers in a series of four-choice tests. The plants
were clipped into equal sized portions, the stems
wrapped in cotton, and the cuttings were placed
into vials filled with water in order to maintain
plant turgidity. The 19 plant species were ran-
domly assembled into 6 clusters of 4 plants each,
with each cluster containing cattail as a common
plant, and the components not varying among
grasshoppers species or replicate tests. Each clus-
ter was replicated 7 times, with replicates occur-
ring on different days. Each of the four plants was
randomly placed at equal distances from the oth-
ers and from the sides of screen cages measuring
0.3 m on each side.
The grasshoppers were exposed to the host
plants for approximately 24 h followed by estima-
tion of the amount consumed. The scale for deter-
mining grasshopper consumption was taken from
Capinera (1993): the consumption was deter-
mined by a visual estimate of the remaining plant
material and assigning it a value of 1 to 5. A value
of 1 would be assigned when 0-20% of the plant
was eaten, 2 when 21-40% consumed, 3 when 41-
60% consumed, 4 when 61-80% consumed and 5

when 81-100% consumed. The feeding trials were
conducted under the same environmental condi-
tions mentioned earlier. The temperature and hu-
midity maintained in the laboratory during this
experiment was approximately the optimal feed-
ing range for most grasshoppers (Chapman 1957,
Mulkern 1967).
The number of grasshoppers per cage was ad-
justed to reflect the individual appetites of the
grasshoppers, thereby allowing measurable con-
sumption without exhausting any of the plant
material. Thus, there were 5 Romalea microptera,
6 Paroxya clavuliger, 6 Leptysma marginicollis, 6
Stenacris vitreipennis, 10 Paroxya atlantica, and
12 Gymnoscirtetes pusillus per cage. These grass-
hopper populations resulted in relatively the
same amount of plant material eaten between
species over the 24-h test period, an average of
about 30%.
We took steps to assure that the grasshoppers
had ample opportunity to explore the cages and
host plants before registering acceptance of hosts
by prolonged feeding. We maintained relatively
low densities, and in no case was consumption
high enough on one plant species to influence con-
sumption of another plant. The cages were small,
allowing the grasshoppers to encounter most or
all plant species with relatively little movement.
Therefore, we believe that the grasshoppers were
fully capable of assessing the host options, and
registered "preference" by their host consumption
behavior. Observation of the insects confirmed
that grasshoppers moved freely and often sam-
pled plants without continuing to feed.
The 19 plants that were evaluated in this
study were collected from the same habitats as
the grasshoppers, and represented the most
abundant floral elements in the semi-aquatic
habitats sampled. The study plants included
Typha spp. (cattail) (Typhaceae); Eichhornia
crassipes (floating water hyacinth), and Ponte-
deria cordata (pickerel weed) (Pontederiaceae);
Urochloa mutica (para grass), Leptochloa spp.
(sprangletop), Panicum repens (torpedograss),
Sacciolepis striata (American cupscale) and
Chasmanthium sessiliflorum (long leaf spike-
grass) (Gramineae); Polygonum punctatum (dot-
ted smartweed) and Polygonum hirsutum (hairy
smartweed) (Polygonaceae); Hydrocotyle spp.
(pennywort) and Cicuta mexicana (water hem-
lock) (Umbelliferae); Ludwigia octovalvis (long
fruited primrose willow) and Ludwigia suffruti-
cosa (headed seedbox) (Onagraceae); Sagittaria
latifolia (common arrowhead) (Alismataceae);
Cyperus compressus (poorland flatsedge) and
Cyperus surinamensis (tropical flatsedge) (Cyper-
aceae); Juncus efusus (softrush) (Juncaceae); and
Sesbania macrocarpa (hemp sesbania) (Legumi-
nosae). For the purposes of this study the monocot
families of Typhaceae, Pontederiaceae, Gramineae,
Alismataceae, Cyperaceae and Juncaceae were

Florida Entomologist 85(2)

considered grasses and the dicot families Poly-
gonaceae, Umbelliferae, Leguminosae and Ona-
graceae were considered forbs.
The mean consumption values for each plant
from the 6 plant clusters were calculated for each
grasshopper species using Graph Pad Software
(Instat 1993) one-way analysis of variance
(ANOVA). The individual means were then com-
pared in each grasshopper species trial using the
Tukey-Kramer multiple comparison test. Simul-
taneously, mandibles of the grasshoppers were ex-
amined visually from preserved specimens and
classified as forb- or grass-feeding types according
to the descriptions and drawings of Isley (1944).


Of the 6 species collected from the wetland
plant habitats, 5 were in the subfamily Cyrtacan-
thacridinae whereas Romalea microptera, though
closely related to Cyrtacanthacridinae, is placed
in the subfamily Romaleinae or in the family
Romaleidae. Based on subfamily taxonomy, such
cyrtacanthacridines might be expected to be forb-
Examination of the mandibles revealed that not
all species were morphologically equipped to feed
on forbs (Fig. 1). Three of the cyrtacanthridines,
Paroxya clavuliger, Paroxya atlantica, and Gym-
noscirtetes pusillus possessed toothed mandibles,
suggesting a tendency to feed on forbs. However,
Leptysma marginicollis and Stenacris vitreipenni
were found to possess blunt-toothed mandibles,
characteristics of grass-feeding species. Romalea
microptera displayed mandibles suitable for forb
feeding (Isley 1944, Patterson 1984).
The grouping of grasshoppers by mandible
type to predict host preference was, for the most
part, confirmed with the food choice experiments
(Table 1). The cyrtacanthridine species with forb-
feeding mandibles proved to be mixed feeders, ac-
cepting both grasses and forbs. Gymnoscirtetes
pusillus displayed a mixed preference by readily
consuming 2 grasses and 2 forbs. Paroxya atlan-
tica also displayed a mixed preference by selecting
3 grasses and 2 forbs, and Paroxya clavulger chose
2 grasses and 2 forbs. Romalea microptera also
displayed a mixed preference, preferring 2 forbs
and 4 grasses. The two species with grass-feeding
mandibles, Stenacris vitreipennis and Leptysma
marginicollis, displayed strong preference for
grasses. Although in a small number of cases
forbs were consumed, in no case was a forb con-
sumed by these latter species more than a grass.
Typha spp. (cattail) was used as a standard in
each trial because it is a very common aquatic
plant and relative consumption of this species
would allow comparison to the other plants. Con-
sumption values for cattail ranged from 1.3 to 5.0
in the various tests. Overall, cattail is one of the
most readily accepted hosts for the grasshopper






Fig. 1. Diagrams of the left mandible of the grass-
hopper species: (a) Romalea microptera Beauvois, (b)
Paroxya clavuliger (Serville), (c) Paroxya atlantica
Scudder, (d) Leptysma marginicollis (Serville), (e) Sten-
acris vitreipennis marshallll, (f) Gymnoscirtetes pusil-
lus Scudder.

species tested. Other plant species were often
consumed about as readily, or more readily, than
cattail. Gymnoscirtetes pusillus had low con-
sumption values but showed preferences for cat-
tail, long fruited primrose willow, headed seedbox
and sprangletop. Paroxya atlantica showed pref-
erence for cattail, pennywort, common arrow-
head, poorland flatsedge and water hemlock.
Paroxya clavuliger showed preference for cattail,
water hyacinth, dotted smartweed, pennywort,
water hemlock and sprangletop. Stenacris vit-
reipennis showed preference for cattail and poor-
land flatsedge. Leptysma marginicollis showed
preference for cattail, pickerel weed, long fruited
primrose willow and softrush. Romalea mi-
croptera showed preference for cattail, common
arrowhead, poorland flatsedge, headed seedbox,
sprangletop and hairy smartweed.
This study provides the first documentation of
the host selection behavior of the grasshopper as-
semblage commonly associated with semi-aquatic
habitats in the southeastern United States. These
poorly studied species display both graminivorous
and forbivorous feeding habits, the nature of
which is fairly predictable based on examination
of mouthpart morphology, but not entirely consis-
tent with the tendency of cyrtacanthridine species
to feed on forbs. The graminivorous feeding behav-
ior of Stenacris vitreipennis and Leptysma mar-
ginicollis is also reflected in modified body form.
Both species display unusually long, thin bodies
that allow them to blend with narrow emergent
grass vegetation. This crypsis undoubtedly makes

June 2002

Squitier & Capinera: Grasshopper Host Selection

xxxxxx xxxxxx xxxxxx xxxxxx xxxxxx xxxxxx

'- 1 0 CO 1:- (0
0 ~' (0 C-^ ~' cO
0 '-I t- 00 0

0L-i Cq 00i C

'~ClC -(01

01 Cq 01 x CO (0
(0 (C CO 0 (0 1 -
CO IC 0 Cl It

i-il IC 01 Iil x
0 L- 1 C1 00
000 n 0C 0
0 ij 0 00 0

L- -i -i (0 -i Cl
Cq 00 Cq 00
000 000

O= C L-7 OD Q;^ O U L-7i 1o CO 1 CI q 1q L-7 -! C -! L-7 C0 Q; -! C0 C t 0 L O Oq Iq lC CO
T--l T--l T--I ~~~C'I T-- C11T- -l0 T- T- T- -IT-


-2 c< -2 -2 -2 -2
a' 0 co 0
'T 'T 'ct-o 00
T- -! L-7 C^ :=T

-2 -2 -2 -2 c< -2
0 ^' 0 0 10
0 'a 00 CO .-

c3 c3 c3 -2 c3 c3
0 I0 0 0 In 0

0 C 00 0

-a ct-a co -a -a


, m *, ,- co .

010C~0 CO^ CO
Ca 00 c
C^ 00 Co '-c

-t -t -t -t -t -t

bp- bfcp m b p m b p m b m cj m

,C m bj) z m bj) zc, mc tQbcjcc, c) z ) cc cc t Qc c2 cm t) z c mc b) z mc c, cc)
o ^- as ^ as as as ^ga 'g
-^ p t o+ o t c + ' r o+ o '^ Pir o co o ir co +J c o .r o J

+1 +1 +1 +1 +1 +1

0 e -I
0C IC In o "C
0 00 '-i 'i 0
+1 +1 +1 +1 +1 +1
0C CO CQ (

+1 +1 c+ +1 c+ +1

+1 +1 +1 +1 +1 +1
t- 0 [0 [-0

+1 +1 +1 +1 +1 +1

-ac~c co c~c
0 'C t0 0 'D
+1 +1 +1 +1 +1 +1
0 CO C It In

r-; 0 r^ Ci -2 r-2
0 ^00 i'j i' 0
+1 +1 +1 +1 +1 +1

b bp X 1c bf 0c1 0cc 0cc x

0i) -z) b) i cc i
?-| ~ ~ 0^ CF B 0 ccc ?- F OC F OC F OC F OC F OC

+1 +1 +1 +1 +1 +1
0 ij o o o

10 CO 1000 C

+1 +1 +1 +1 +1 +1
11 ; 0i-!O
X- O T- Cq

+1 +1+1+1+1 +1
'-1 0 0 '-l CO

0 0 00 In In

+1 +1 +1 +1 +1 +1

-2 ca ca ca -2 ca
(0 01 C<1 (0 In In

+1 +1 +1 +1 +1 +1

Cl C0 x CO x

ca ca -2 -2 -2 ca

+1 +1 +1 +1 +1 +1
00 CO C (0 00
oi^ : i i-i iA-i C

0 C0 *0 C0 C0 bCO
C j) -j) C j) C j) C j) bl)
) 5 bt bj 5 o g 5 o

.P z OP. z OP. z
" o0^ "g^co0^ "g^c g- ~Q cc$~ 0' o0^ "g^c '

In Iq Iq OqIn
00000 + 1
[- CO^ ;f

Z 0TI z 1

+1 +1 +1 +1 +1 +1

x Cq Cq Cq x; Cq

' I 0 In CO C

+1 +1 +1 +1 +1 +1

x0 Cq CO x Cq Cq

C)i C)i C)i C) C)i C)

0 O0 ICnC
cqi TicqiTiT-

[ +1 (0C 00


c; -; q~ cq c^ q
-IC Z- TI z- z- ^

0x x cq I

+11. 1. 1. m. m1 m1 . . +. . .



9 p


o b


o) a

o V

ov V

a s
s. &

r1 r


+ l


Florida Entomologist 85(2)

them difficult to detect and presumably safer from
predation. Interestingly, these grass-feeding cyrt-
acanthacridine species have very slanted faces,
thereby physically resembling grass-feeding gom-
phocerines more than their close relatives, the
forb-feeding cyrtacanthacridines.


This research was supported by Florida Agricultural
Experiment Station, and published as Journal Series
number R-08174.


BERNAYS, E. A., AND R. BARBEHENN. 1987. Nutritional
ecology of grass foliage-chewing insects, pp. 147-175.
In F. Slansky and J. G. Rodriguez (eds.), Nutritional
Ecology of Insects, Mites, Spiders, and Related In-
vertebrates. John Wiley & Sons, New York.
CAPINERA, J. L. 1993. Host-plant selection by Schisto-
cerca americana (Orthoptera: Acrididae). Environ.
Entomol. 22: 127-133.
CHAPMAN, R. F. 1957. Observations on the feeding of
adults of the red locust (Nomadacris septemfasciata
iS... .. .. Brit. J. Anim. Behav. 5: 60-75.
CHAPMAN, R. F. 1964. The structure and wear of the
mandibles in some African grasshoppers. Proc. Zool.
Soc. London 142: 107-121.
DADD, R.H. 1963. Feeding behavior and nutrition in grass-
hoppers and locusts. Adv. Insect Physiol. 1: 47-109.
INSTAT. 1993. Graph Pad Instat Mac Instat Statistics.
Graph Pad Software, San Diego, CA. 110 pp.

ISLEY, F. B. 1944. Correlation between mandibular mor-
phology and food specificity in grasshoppers. Ann.
Entomol. Soc. Am. 37: 47-67.
JOERN, A. 1979. Resource utilization and community
structure in assemblages of arid grassland grasshop-
pers (Orthoptera: Acrididae). Trans. Am. Entomol.
Soc. 105: 253-300.
JOERN, A. 1986. Resource partitioning by grasshopper
species from grassland communities. Proc. Triennial
Mtg. Pan Am. Acrid. Soc. 4: 75-100.
JOERN, A., AND L. R. LAWLOR 1980. Food and microhab-
itat utilization by grasshoppers from arid grass-
lands: comparisons with neutral models. Ecology 61:
KANG, L., Y. GAN, AND S. L. LI. 1999. The structural ad-
aptation of mandibles and food specificity in grass-
hoppers on Inner Mongolian grasslands. J. Orth.
Res. 8: 257-269.
LEWIS, A. C., AND H. F. VAN EMDEN. 1986. Assays for in-
sect feeding, pp. 95-119. In J. R. Miller and T. A.
Miller (eds.), Insect-Plant Interactions. Springer-
Verlag, New York.
MULKERN, G. B. 1967. Food selection by grasshoppers.
Annu. Rev. Entomol. 12: 59-78.
OTTE, D. 1981. The North American Grasshoppers. Vol-
ume I: Acrididae. Gomphocerinae and Acridinae.
Harvard Univ. Press, Cambridge. 275 pp.
PATTERSON, B. D. 1984. Correlation between mandibu-
lar morphology and specific diet of some desert
grassland Acrididae (Orthoptera). Am. Midl. Nat.
11: 296-303.
SCHERER, C. W. 1997. Response of grasshoppers (Orthop-
tera: Acrididae) to different forest restoration tech-
niques in a Florida sandhill community. Unpublished
M.S. Thesis. Univ. of Florida.

June 2002

Sourakov & Mitchell: Laboratory Biology of C. scutellaris


1Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611

2Center for Medical, Agricultural and Veterinary Entomology, U.S. Department of Agriculture
Agricultural Research Service, P.O. Box 14565, Gainesville, FL 32604, USA

The tachinid parasitoid Chetogena scutellaris (Wulp) was reared from southern armyworm,
Spodoptera eridania (Cramer), a new host record. When reared in cabbage looper, Tricho-
plusia ni (Htibner), (a new host) and in fall armyworm, Spodoptera frugiperda (J. E. Smith),
in the laboratory, C. scutellaris developed successfully ca. 30% of the time. Success of para-
sitism depended on the numbers of eggs laid per host, host age, and host species. Parasitoid
development was not synchronized with host development. C. scutellaris developed mostly
as a solitary parasitoid. Female flies preferred fifth instar hosts for oviposition. Cabbage
looper was a better host than fall armyworm for mass rearing of this parasitoid.

Key Words: biological control, parasitoid development, southern armyworm, Spodoptera fru-
giperda, Spodoptera eridania, Tachinidae, Trichoplusia ni

El parasitoide taquinido Chetogena scutellaris (Wulp) fu6 criado de Spodoptera eridania
(Cramer), un nuevo registro de hospedero. Cuando fu6 criado en el gusano medidor de repo-
11o, Trichoplusia ni (Hlibner), (un nuevo registro de hospedero) y en el gusano cogollero, Spo-
doptera frugiperda (J. E. Smith), en el laboratorio, C. scutellaris se desarroll6 exitosamente
ca. 30% del tiempo. El 6xito de parasitismo dependio del numero de los huevos puestos en
cada hospedero, la edad del hospedero, y la especie del hospedero. El desarrollo del parasi-
toide no fu6 sincronizado con el desarrollo del hospedero. C. scutellaris se desarroll6 princi-
palmente como un parasitoide solitario. Las moscas hembras prefirieron ovipositar en el
quinto estadio larval de los hospederos. El gusano medidor de repollo fu6 un mejor hospedero
para criar el parasitoide en masa que el gusano cogollero.

The tachinid fly Chetogena scutellaris (Wulp)
is largely known by its synonym Euphorocera
floridensis Townsend (Aldrich & Webber 1924).
Two families of Coleoptera (Cerambycidae and
Coccinellidae) and 11 families of Lepidoptera
(Arctiidae, Citheroniidae, Ctenuchidae, Geometri-
dae, Hesperiidae, Noctuidae, Notodontidae, Sphin-
gidae, Zygaenidae, Pierdae, and Saturniidae) are
listed as hosts for this species (Arnaud 1978).
C. scutellaris was recorded from seven noctuid
species that are crop pests: velvetbean caterpillar,
Anticarsia gemmatalis Hfibner; in Georgia, cot-
ton leafworm, Alabama argillacea (Hfibner), true
armyworm, Pseudoletia unipuncta (Hawitson.),
and corn earworm, Helicoverpa zea (Boddie); in
North Carolina, green cloverworm, Plathypena
scabra (F.); in S. Carolina, Florida and Maryland,
fall armyworm, Spodoptera frugiperda (J. E.
Smith); in Mississippi, soybean looper, Pseudo-
plusia includes (Walker). It is also known from
Costa Rica and Peru (Arnaud 1978).
In June, 2000, we collected ca. 100 late-instar
larvae of southern armyworm, Spodoptera erida-

nia (Cramer), from lambsquarters, Chenopodium
album (L.), in a corn field near Bunnell, Florida.
Approximately 25% of late instar larvae had one or
more tachinid eggs on their surface, and laboratory
colony was established from the resultingC. scutel-
laris adults. Two major noctuid pests were used as
hosts: cabbage looper, Trichoplusia ni (Hfibner),
and fall armyworm. This study was conducted to
assess the biology of C. scutellaris in the laboratory
and evaluate its potential for mass rearing.


Host third-fifth instar larvae were maintained
on a pinto bean diet (Guy et al. 1985) and offered
in groups of 20-30 larvae to C. scutellaris main-
tained inside 20-cm3 cages with netting sides.
C. scutellaris tend to parasitize larger larvae, thus
to obtain parasitism in younger larvae they had to
be offered separately from the old ones. The time
that larvae were exposed to flies varied, though
usually flies attack larvae right away, so within an
hour larvae were removed and checked for eggs.

Florida Entomologist 85(2)

Initially, flies were maintained in groups on
honey and water, and then some were randomly
chosen for the experiments (freshly emerged cop-
ulating pairs were carefully removed from the
communal cage into solitary ones, insuring that
the female is young, mated and that it has a male
partner with it for further matings). Parasitized
larvae were identified by observing the eggs that
were laid on them. Host larvae with one or more
tachinid eggs were transferred into individual
200-ml waxed paper cups with plastic lids, sup-
plied with a diet cube, and checked daily for par-
asitoid emergence. Thus, the development rates
from egg to pupa, and from pupa to adult were
recorded. The study was repeated three times,
using flies and hosts of three consecutive genera-
tions. Overall, 705 parasitized larvae were reared
To estimate host age preference, 20 fall army-
worm larvae (10 fourth and 10 fifth instar) were
exposed for a period of five minutes to several 10-
d-old female flies in a similar cage. Larvae were
then checked for parasitoid eggs, easily noticeable
on the larval surface, to compare egg numbers
laid on hosts of different stages. This experiment
was repeated 12 times and was analyzed using
ANOVA and t-tests ("JMP", SAS Institute 1995).
Standard error values are provided for all means.


Substantial variation was observed among in-
dividual C. scutellaris females in their parasitism
(emergence of maggot(s)) in hosts of different
stages and species (Table 1). Higher percentage of
successful parasitism was observed when fifth-
instar versus fourth-instar cabbage loopers were
attacked (46.5 5% vs. 22.7 4%, P < 0.05 (Means
+ SD)). In fall armyworm, differences in parasit-
ism were not statistically significant among lar-
val stages. The decline in success of parasitism
towards the end of the fourth instar probably
should be attributed to moulting prior to hatching
of parasitoid eggs. Parasitized third-instar hosts
produced no parasitoids in either host species.
Thus, majority (close to 70%) of attacked larvae

escaped parasitism and developed into healthy
adult moths.
In 10 trials of 12, flies parasitized older larvae
at a higher rate (fifth instars, 6.6 0.4%; fourth
instars, 5.0 + 0.5%, P < 0.05), which suggests that
older larvae are preferred for oviposition. The to-
tal number of eggs laid also was higher in older
larvae (fifth instars, 11.5 1.1 eggs; fourth in-
stars, 6.5 0.7 eggs, P < 0.05). The probability of
successful parasitism increased from 29 8.2% to
53.1 4.1% (P < 0.05), when number of eggs ovi-
posited was more than one per host (Fig. 1). A sin-
gle maggot emerged from 81% (N = 228) of the
hosts, though two-thirds of these hosts had multi-
ple parasitoid eggs laid on them. In 16% of the
hosts, two parasitoids per host emerged and, only
in 3% of the hosts did three or more (maximum of
six) parasitoids emerge. The latter parasitoids
were smaller (e.g., the dry weight of an average
fly was 10.7 mg, exceeding eight times the weight
of a fly that developed gregariously in a group of
six (1.3 mg)). When five or six parasitoids came
from a single host, half of the flies never emerged
from their puparium. Our data concerning influ-
ence of host age and number of eggs laid on suc-
cess of parasitoid development correspond with
similar studies on other species of Tachinidae
(Konotie & Paolo 1992).
The host stage from which the parasitoid
emerged depended on the stage at which it was
parasitized. When fourth-instar host larvae were
attacked, 85% of maggots emerged from larvae
and 15% emerged from pupae (N = 65). In con-
trast, when hosts were attacked as fifth instars,
13% of maggots emerged from larva while 87%
emerged from pupae (N = 139). Thus, this parasi-
toid's development does not appear to be synchro-
nized with development of its host. C. scutellaris'
development time from egg to pupa at 21C was
12.3 + 0.2 days (N = 145) with no significant dif-
ference (P > 0.05) found between different hosts.
Flies emerged from pupae in 13.3 + 0.6 days.
C. scutellaris reproduce easily in the labora-
tory on hosts fed artificial diet. Cabbage looper, a
species that has not been previously recorded as a
host of C. scutellaris, would probably be attacked


Successful parasitism (% + SE)

Stage Attacked Cabbage looper N Fall armyworm N

Early 4th 24.0 + 5.0 23 49.0 + 8.4 92
Mid-4th 30.5 + 4.5 38 0
Late 4th 13.5 + 8.5 30 2.9 + 1.5 68
Early 5th 48.3 + 16.3 56 50.0 + 0.0 16
Mid-5th 47.3 + 3.9 89 15.0 + 8.0 93
Late 5th 44.0 + 12.5 164 31.0 + 11.6 36
Mean 34.5 8.5 400 29.5 5.9 305

June 2002

Sourakov & Mitchell: Laboratory Biology of C. scutellaris



o 50- ---


0 -- -- -- -----
1 egg 2 eggs 3 eggs 4 eggs >4 eggs

Fig. 1. Success of development of one or more Cheto-
gena scutellaris flies in relation to the number of eggs
oviposited on a host larva. Increase was significant (P <
0.05) when one (29 8.2%) and more than one (53.1 +
4.1%) egg per host was laid.

in the field should both the host and the parasi-
toid occur synchronously (cabbage looper is
mostly a winter pest in the southeastern United
States). Field trials are needed to evaluate poten-
tial of this species as a biological control agent of
noctuid pests. Cabbage looper also proved to be a
more convenient host for mass rearing C. scutel-
laris. Cabbage looper larvae make cocoons on top
of the cage, while maggots pupate on the bottom
where they can be easily collected. In contrast,
southern and fall armyworms pupate inside the
diet cakes in the laboratory, as in nature they pu-

pate in soil. When these hosts were used, collect-
ing pupae of C. scutellaris was labor intensive
because they had to be extracted from the diet.


We would like to thank Dr. Susan Webb and Dr. Rob
Meagher for critical reviews of this manuscript. Dr.
James O'Hara and Dr. Gary J. Steck identified the par-
asitoids. This article reports the results of research only.
Mention of a proprietary product does not constitute an
endorsement or the recommendation for its use by


ALDRICH, J. M., AND R. T.WEBBER 1924. The North
American species of parasitic two-winged flies be-
longing to the genus Phorocera and allied genera.
Proc. U.S. Natl. Museum No. 2486, Vol. 63, Art. 17,
pp. 1-90.
ARNAUD, P. H., JR. 1978. A host-parasite catalog of
North American Tachinidae (Diptera), pp. 1-860.
USDA, Misc. Pub. No. 1319, Washington, DC.
GuY, R. H., N. C. LEPPLA, J. R. RYE, C. W. GREEN, S. L.
BARRETTE, AND K. A. HOLLIEN. 1985. Trichoplusia
ni, pp. 487-494 In Pritam Singh and R. F. Moore
[eds.], Handbook of Insect Rearing, Vol. 2, Elsevier
Science Publishers B. V., Amsterdam.
KONOTIE, C. A., AND F. PAOLO. 1992. Eucelatoria bryani
Sabr. (Diptera Tachinidae) rearing on the factitious
host Galleria mellonella L. (Lepidoptera Galleri-
idae): Effect of host age at exposure to the parasitoid
females. Bolletino dell'Istituto di Entomologia
"Guido Grandi" della Univesita degli Studi di Bolo-
gna. 46(0): 229-238.
SAS INSTITUTE. 1995. JMP, Version 3, SAS Institute
Inc., Cary, NC.

Florida Entomologist 85(2)


1USDA-ARS, Invasive Plant Research Laboratory, 3205 College Ave., Ft. Lauderdale, FL 33314

2Fort Lauderdale Research and Education Center, University of Florida
3205 College Ave., Ft. Lauderdale, FL 33314

Podisus mucronatus Uhler is a generalist predator found in Florida and the islands of the
Caribbean. Adult P. mucronatus were observed preying on larvae of the Australian weevil
Oxyops vitiosa (Pascoe), a biological control agent of Melaleuca quinquenervia (Cav.) S.T.
Blake. To facilitate field-based identification of this predator, we present descriptions of eggs,
nymphal stages, and adults. Life history traits of P. mucronatus when held with no food or
either of two prey species (0. vitiosa and Tenebrio molitor (L.) larvae) are also reported. The
potential use of this species as a biological control agent of arthropods and its interference
with weed biological control are discussed.
Key Words: Podisus mucronatus, Pentatomidae, developmental rates, predatory stinkbug,
Oxyops vitiosa, Melaleuca quinquenervia, Tenebrio molitor

Podisus mucronatus Uhler es un depredador generalista que se encuentra en Florida y en el
Caribe. Se observaron adults de P. mucronatus alimentandose de larvas de Oxypos vitiosa
(Pascoe), un gorgojo australiano y un agent de control biol6gico de Melaleuca quinquenervia
(Cav.) S. T. Blake. Se presentan descripciones morfol6gicas de los huevos, estadias ninfales
1-5, y los adults, para facilitar la identificaci6n de este depredador en el campo. Se reportan
tambi6n caracteristicas de la historic natural de P. mucronatus mantenidas sin alimento o
sin las larvas de cualquier de las dos species presa (0. vitiosa y Tenebrio mollitor (L.)). Se
discute el uso potential de esta especie como un agent de control biol6gico de artr6podos y
su interferencia con el control biol6gico de malezas.

Species in the genus Podisus (Pentatomidae) are
generalist predators, that attack primarily lepi-
dopteran and coleopteran larvae (Aldrich et al.
1991). Because prey species include important pests
in agroecosystems, some Podisus species have re-
ceived attention as potential biological control
agents for agricultural pests (Drummond et al.
1984, Stamopoulos & Chloridis 1994, De Clercq
2000). Podisus maculiventris (Say), for instance, has
been the focus of various life history and compara-
tive development studies as well as morphological
descriptions of eggs, nymphs and adults (Aldrich
1986, Decoursey & Esselbaugh 1962, Legaspi &
O'Neil 1993, Legaspi & O'Neil 1994). This attention
is due, in part, to the use ofP maculiventris as a bi-
ological control agent of the Colorado potato beetle,
Leptinotarsa decemlineata (Say) (Drummond et al.
1984, Stamopoulos & Chloridis 1994).
Only about 10% of the 300 known asopine spe-
cies have been studied in detail (De Clercq 2000).
For instance, the native Floridian and Caribbean
species Podisus mucronatus Uhler has rarely been
reported in the literature (J. Eger, pers. comm.).
Occurrences of P mucronatus attacking agricul-
tural and horticultural pests were reported by Ge-
nung (1959) and Genung et al. (1964). Aldrich et

al. (1991) described the attractant pheromone pro-
duced by male Podisus species, including that of
P mucronatus. Unfortunately, little else is known
concerning this species.
We recently observed adultP mucronatus prey-
ing on larvae of the weed biological control agent
Oxyops vitiosa (Pascoe) (Coleoptera: Curculion-
idae) that feed on Melaleuca quinquenervia (Cav.)
S. T. Blake in Lee Co., FL. We wished to quantify
the population dynamics of both 0. vitiosa and its
newly acquired generalist predator, P mucrona-
tus. To date, no morphological descriptions of im-
mature stages or life history traits exist for this
predator, making accurate identification of eggs
and nymphs difficult. To facilitate proper identifi-
cation and future understanding of its ecological
relationship with 0. vitiosa, we describe herein
the basic morphological characteristics and life
history traits of P mucronatus.


Rearing and Maintenance of P. mucronatus Colonies

Podius mucronatus adults, found in associa-
tion with 0. vitiosa, were collected from coppicing

June 2002

Costello et al.: Description and Life History ofPodisus mucronatus

M. quinquenervia stumps at a site near Estero,
Florida (N2625.530' W8148.620'). The biological
control agent was originally released at this site
in 1998 (Center et al. 2000). Adult P. mucronatus
were maintained in 30 x 25 x 9 cm diameter plas-
tic containers provisioned with paper towels lin-
ing the interior of the container. The paper towels
were used to provide seclusion and oviposition
sites. Podisus mucronatus colonies were sus-
tained by feeding nymphs and adults mixed lar-
val stages of Tenebrio molitor (L.) and 0. vitiosa.
Paper towels and dead prey were removed
weekly. Adult colonies were examined every 12 h
at which time newly deposited eggs were removed
and placed into 10 x 1.5 cm diameter petri dishes.
Aged cohorts were held separately. Eggs and
first-instar nymphs were reared in 10 x 1.5 cm di-
ameter petri dishes on filter paper with a small
M. quinquenervia terminal vegetative bud (tip)
for moisture. After the eggs hatched, decapitated
T. molitor third and fourth-instar larvae were
placed in the petri dish. Tenebrio molitor were de-
capitated to facilitate predation byP. mucronatus
nymphs. Second and third-instar nymphs were
then transferred into a 15 x 4 cm diameter plastic
container with filter paper, several M. quinquen-
ervia tips, and live larvae of T. molitor. Fourth
and fifth-instar nymphs were reared in 18 x 8 cm
diameter plastic containers and maintained in
the same manner. Colonies were held under labo-
ratory conditions at 25 (5) C, a photoperiod of
16:8 (L:D), and 70 (10)% relative humidity. All
stages were monitored every 12 h and exuvia
were removed to accurately track life stages.

Morphological Descriptions and Measurements

Each nymphal and adult stage was examined
using a Nikon dissecting microscope (10-50x).
Descriptions were based on live individuals
whereas measurements were based on specimens
preserved in ethyl alcohol. The number of individ-
uals measured ranged from 11-35, depending on
the number available from each respective cohort.
Size and number of micropylar processes were
measured on 44 eggs. Length of each nymph and
adult was measured from the tip of the tylus to the
tip of the abdomen. The width of the head was
measured from the outer margins of the com-
pound eyes. The pronotal width of adults was mea-
sured between the humeral spines. Lengths of the
metathoracic leg and femur were also measured.

Life History Parameters

Life history parameters were assessed in ex-
perimental arenas that consisted of 10 x 1.5 cm
plastic petri dishes, provisioned with a slightly
moistened filter paper. Newly hatched (<10 h old)
P mucronatus first-instar nymphs were collected
from colonies and transferred using a camel's hair

brush to individual arenas. Twenty randomly se-
lected first-instar nymphs were assigned to each
of 4 diets: 1) moistened filter paper only, 2) diet 1
plus the addition of aM. quinquenervia tip, 3) diet
2 plus a single 0. vitiosa larva (second-third in-
star) or 4) diet 2 plus a single T molitor larva (sec-
ond-fourth instar). Melaleuca quinquenervia tips
consisted of the terminal 4 cm of a newly devel-
oped, succulent shoot. All petri dishes were
stacked with an additional dish at the top, which
contained only a moistened filter paper. Stacks of
petri dishes were placed in an environmental
chamber at 25 (+1) C, a photoperiod of 16:8 (L:D),
and 65 (10)% relative humidity. Developmental
stages and survivorship were assessed every 24 h.
The presence of exuvia was used to assess molting
between stages. Melaleuca quinquenervia tips
were replaced every 48 h. Oxyops vitiosa survivor-
ship was assessed daily by probing each larva
with a camel's hair brush, those not responding
were considered dead and were replaced. Tenebrio
molitor larvae were replaced every 48 h.

Nymph and Adult measurements are presented in Table
Eggs (Fig. 1A)

Egg shape elliptical, height 0.91 mm (0.08;
mean (SD), and width 0.80 mm (0.05), laid in
clusters. Color opaque initially then silver or ma-
roon with a shiny surface before eclosion. An av-
erage of 8.25 (0.90) micropylar processes
occurring circularly around the operculum, gen-
erally curving outward. Micropylar processes
white terminating with a spherical structure,
ranging from white to black.

First-Instar Nymph

Shape oval, convex, widest at second or third
abdominal segment. Head convex dorsally, widest
basally, narrowing to tip of tylus. Head brown and
eyes red with a metallic appearance. Antennae 4-
segmented, segments brown with white annuli.
Antennae covered with increasingly more setae
towards apex. Rostrum 4-segmented and brown,
apex extending slightly beyond metacoxae.
Thorax narrowest anteriorly, width increasing
posteriorly. Length of pro-, meso-, and metatho-
rax medially equal. Dorsal sutures well defined.
Dorsal color brown. Thorasic median line visibly
lighter in color. Coxa, femur, and tibia also brown
with tarsi and tarsal claws gray. As stadium pro-
ceeds, first two tarsal segments turn brown and
tarsal claws elongate. Leg segments setose. Brown
scent glands present on pro-epimeron and meta-
epimeron, becoming obscured after sclerotization.
Abdominal sutures between segments distinct.
Lateral margins of abdominal segments I and II

Florida Entomologist 85(2)


Nymphal stage
characteristic First Second Third Fourth Fifth Adult male Adult female

Body length 1.24 (0.23)d 2.00 (0.22) 4.01 (0.92) 4.77 (0.35) 8.46 (0.80) 10.13 (0.88) 11.46 (0.94)
Pronotal widthb 0.91 (0.16) 1.22 (0.17) 2.34 (0.34) 2.73 (0.26) 4.91 (0.35) 6.51 (0.47) 7.05 (0.47)
Head width' 0.56 (0.13) 0.73 (0.11) 1.33 (0.23) 1.34 (0.14) 2.16 (0.13) 2.22 (0.12) 2.28 (0.14)
Antennal segment
1 0.07 (0.01) 0.08 (0.03) 0.15 (0.03) 0.12 (0.04) 0.21 (0.05) 0.27 (0.11) 0.20 (0.00)
2 0.20 (0.08) 0.38 (0.08) 0.83 (0.18) 1.02 (0.12) 1.69 (0.24) 1.20 (0.19) 1.27 (0.20)
3 0.18 (0.06) 0.30 (0.07) 0.62 (0.12) 0.71 (0.08) 1.14 (0.14) 0.97 (0.25) 0.99 (0.16)
4 0.28 (0.06) 0.38 (0.05) 0.64 (0.08) 0.67 (0.06) 0.95 (0.12) 1.14 (0.29) 1.19 (0.17)
5 0.00 (0.00) 0.00 (0.00) 0.00 (0.00) 0.00 (0.00) 0.00 (0.00) 0.95 (0.21) 1.00 (0.12)
Total length 0.74 (0.19) 1.13 (0.18) 2.25 (0.37) 2.48 (0.19) 3.94 (0.40) 4.51 (0.80) 4.46 (0.43)
Rostrum segment
1 0.19 (0.03) 0.30 (0.03) 0.44 (0.09) 0.51 (0.05) 1.07 (0.12) 1.03 (0.16) 1.19 (0.14)
2 0.18 (0.06) 0.35 (0.07) 0.64 (0.09) 0.66 (0.11) 1.13 (0.17) 1.22 (0.16) 1.28 (0.16)
3 0.16 (0.06) 0.22 (0.04) 0.35 (0.05) 0.43 (0.10) 0.89 (0.16) 0.96 (0.11) 1.06 (0.14)
4 0.19 (0.04) 0.27 (0.06) 0.49 (0.10) 0.51 (0.04) 0.76 (0.11) 0.93 (0.17) 0.94 (0.11)
Total length 0.72 (0.16) 1.15 (0.14) 1.92 (0.27) 2.11 (0.13) 3.86 (0.29) 4.14 (0.30) 4.46 (0.31)
Femur length 0.42 (0.08) 0.63 (0.10) 1.19 (0.28) 1.33 (0.24) 2.73 (0.25) 3.22 (0.81) 3.37 (0.36)
Total leg length 1.09 (0.20) 1.58 (0.22) 2.92 (0.42) 3.81 (0.40) 7.23 (0.48) 7.80 (0.89) 8.22 (0.88)

"Length of each nymph and adult individual was measured from the tip of tylus to tip of abdomen.
'The pronotal width of adults was measured between the humeral spines.
'The width of the head was measured from the outer margins of the compound eyes.
'mm (+SD).

curve anteriorly, segments III-X curve posteriorly.
Lateral margin of segments V and VI longest, lat-
eral length of segments narrowing anteriorly and
posteriorly. Segments IX and X not distinguish-
able on every individual. Abdomen orange with
brown medial and lateral plates. Dorsal lateral
plates semi-circular in shape (see Fig. 1C). Ven-
tral lateral plates similar except one spiracle
present on segments II-XIII, centrally located be-
tween midline and anterior margin, resulting in a
small emargination in each lateral plate. One tri-
chobothria present on segments III-VII, caudad
to spiracle, resulting in an additional emargin-
ation in lateral plates on venter. Intersegmental
sulcus, originating from center of each lateral
plate on venter, extends transversely. Lateral
plates decreasing in size posteriorly, those of seg-
ment I minute. Dorsal medial pattern varied.
Most common pattern: brown transverse central
band on suture between segments I and II, hour-
glass shaped plate centered on segments II and
III, rectangular plate centered on segments III-V,
semi-circle plate with arc towards apex centered
on segment VI extending slightly onto segment V,
a central oval macule on segments VII and VIII,
segments IX and X completely brown (see Fig.
1B). Mediodorsal plates with three pairs of scent
gland orifices and a visible connecting suture
curving posteriorly (see Fig. 1B). Mediodorsal
pattern size increases with each exuviation to

next nymphal stage. Degree of expression of hour-
glass shaped plate decreases with each molt. In
fully sclerotized nymphs, orange and brown colors
change to red and black.

Second-Instar Nymph (Fig. 1B)

Oval, convex, widest at third abdominal seg-
ment. Head black, eyes dark red, similar in ap-
pearance to first-instar nymph. Antennal seg-
ments I,IV black, II,III gray, annuli white. Head
shape and setae same as first-instar nymph.
Shape and coloration of thorax similar to first
instar except as follows: Legs from coxa to tibia
gray or brown with white annuli. Coxae black
basally. Tarsal claws white, pulvilli gray to black.
Length of pro- and mesothoracic segments equal
medially, metathorax medially shorter than pro-
and mesothoracic segments.
Abdominal shape and dorsal pattern same as
first-instar nymph. Second ventral trichobothria
present on segments III-VII lateral to first tricho-
bothria. Abdominal sutures less distinct than in
first instar and intersegmental sulci not visible.
Medioventral plates vary by size, darkness, and
number of black oval plates. Predominant medio-
ventral plates dark red as follows: single large
oval on segments I-IV, smaller oval on segments V
and VI, third smaller oval on segments VII and
VIII. Segments IX and X dark red.

June 2002

Costello et al.: Description and Life History ofPodisus mucronatus

sc -ME

ep =



~a-~ .4,


^^ -w
*-t -* *
t ,',. Sfr. *




Fig. 1. Podisus mucronatus adults and instars. A, egg clutch; B, second instar dorsal view; C, third instar dorsal
view; D, fourth instar dorsal view; E, fifth instar dorsal view; F, fifth instar ventral view; G, adult dorsal view; H,
adult ventral view. dp, dorsal abdominal pattern; edp, expanded dorsal abdominal pattern; ep, eye pattern; sc, semi-
circle plate; vp, ventral abdominal pattern.



1( D

Florida Entomologist 85(2)

Third-Instar Nymph (Fig. 1C)
Oval, convex, widest at third abdominal seg-
ment. Head shape, setae, and color same as sec-
ond-instar nymph. Thorax color similar to second-
instar nymph, except tarsi vary in color from
brown to gray.
Abdominal shape and patterns slightly different
from second-instar. Longest lateral margin varies
from segment V-VII. Dorsal abdominal sutures no
longer visible, otherwise, dorsal and ventral pat-
tern and coloration similar to second-instar.

Fourth-Instar Nymph (Fig. 1D)
Oval, convex, widest at second abdominal seg-
ment. Head shape and color similar to third-
instar nymph. Last segment of rostrum dark
brown, all others black. Ocelli dark red. Thorax
similar to third-instar nymph except wing pads
present on mesothoracic segment and median
length of metathoracic segment shortened.
Abdominal morphology similar to third-instar
nymph with few exceptions. A ventral interseg-
mental sulcus reappears, similar to first-instar
nymph. Segment VIII with black ventral lateral
pattern. Coloration of some nymphs vary from or-
ange and black to red and black.

Fifth-Instar Nymph (Fig 1E, 1F)
Oval, convex, widest at third abdominal seg-
ment. Head shape consistent with fourth-instar
nymph. Head color varying from red to black,
base of head usually black. Eyes and ocelli red.
Antennal segment I red to black depending on
maturity, segment II gray to black, segments III
and IV black, annuli white. Posterior portion of
eye white in color and possibly lacking ommatidia
(see eye pattern, Fig. 1E). Last rostral segment
brown, remaining segments black.
Thorax color variable black, red or orange.
Legs typically with coxa, trochanter, and femur
red, tibia, tarsi and tarsal claws black, annuli
white. Legs all black if entire thorax black. Ven-
tral pattern as in fourth-instar nymph. Dorsal
pattern differing from previous stages. Scutellum
and wing pads evident. Wing pads extending
approximately to end of abdominal segment III.
Lateral margins of thorax serrate, black and flat-
tened dorso-ventrally.
Abdominal shape and color same as fourth-
instar nymph. All sutures and patterns more
defined. Ventral patterns and dorsal patterns
similar to fourth-instar nymph, but central dorsal
pattern further expanded (see expanded dorsal
pattern, Fig. IE).

Adult Male
Shape elliptical, wider anteriorly, dorsally flat-
tened, ventrally convex, widest between humeral
spines. Head dorsally punctate. Punctures becom-

ing more dense towards thorax, except for small
white lateral area anterior to thorax. Head ven-
trally white to green with small punctures. Eyes
red to dark red and similar to fifth-instar. Anten-
nae 5-segmented, segments I-IV light brown, seg-
ment V brown, annuli tan. Rostrum 4-segmented,
each segment, base to apex, increasingly darker
from white and green to medium brown.
Thorax dorsally covered with dark brown
punctation. Humeral spines dark brown and pro-
jecting forward (Thomas 1992). Anterolateral
margins of pronotum serrate and green to yellow.
Pronotum with 2 yellow cicatrices. Scutellum tri-
angular, apex and basal angles white to yellow.
Corium and clavus red with same dark punctures
as thoracic dorsum, claval suture white. Hemely-
tral membrane light brown. Hindwing iridescent
with brown venation. Thorax gray to brown punc-
tured ventrally. Coxa, trochanter, femur, and tibia
green with terminus of tibia brown. Three tarsal
segments brown, tarsal claws dark brown, and
pulvilli brown to dark brown. Protibia with brown
spur two-thirds distance from apex to base. Legs
setose, setae denser toward apex. Scent gland
ostiole located on intercoxal region of meta-
thoracic epimera. Ostiolar ruga white, yellow or
green. A white, elliptical callus also occurs on
sternum of mesothorax, anterior to mesocoxae.
Abdomen eight-segmented, length of segments
medially equal except first, which is much shorter.
Lateral margin of segment IV usually longest,
lateral length of segments narrowing anteriorly
and posteriorly. Ventrally, single spiracle on each
side of segments III-VII. Sternum white to green,
concolorously punctate laterally, punctures lack-
ing medially. Tergum brown to red with small
punctures. Connexiva green with larger punctures
than tergum. Dorsal and ventral sutures light in
color. Basal spine occurring ventrally on second
abdominal segment and extending to middle of
metasternum. Ventral posterior margin of male
pygophore setose, setae most dense mesially.

Female Adult (Fig. 1G and 1H)
Female similar in shape but generally larger
than male. Apex of eighth and ninth paratergites
moderately setose, apex of first gonocoxae
sparsely setose. Other characteristics similar to
adult male.

Life History Parameters
Eggs collected from laboratory colonies were
laid in parallel rows, creating circular or elliptical
masses of 28.18 (+10.25) eggs per group. Eye-
spots were apparent under the pseudoperculum
1-2 days before eclosion. Rates of nymphal devel-
opment differed significantly among diets (Table
2). When held with only moistened filter paper
(Diet 1), three individuals (15%) survived to the
second nymphal stage but did not develop further.

June 2002

Costello et al.: Description and Life History ofPodisus mucronatus

When held with moist filter paper and a freshly
excised melaleuca tip (Diet 2), development of
P mucronatus was extended: 90% of those tested
developed to the second nymphal stage and 15%
of these successfully molted to the third nymphal
stage (Table 2). Developmental times were simi-
lar for most within stage comparisons when held
with prey, except second-instar nymphs held with
0. vitiosa, which required approximately one ad-
ditional day to molt into third-instar nymphs.
When compared over the entire developmental
period, individuals feeding on T molitor devel-
oped faster than those on 0. vitiosa (Table 2). Sur-
vivorship was similar among both prey diets, with
5% mortality occurring among fifth-instar
nymphs regardless of prey. Differences in sex ra-
tios were also observed between diets (P = 0.04,
F = 4.16, df = 1,37). Nymphs reared with T moli-
tor larvae were male biased (74:26; J: 9 ; X2 = 4.26,
P = 0.039) whereas those reared with 0. vitiosa
were not (53:47; x2 = 0.053, P = 0.819). The inter-
val between last nymphal molt and first mating
for females was 2.67 (0.71) days. Under labora-
tory conditions, females began ovipositing 4.75
(+0.39) days after mating.


Generalist predators are integral components
of most natural ecosystems due to their abilities
to regulate population densities of lower trophic
levels in the absence of specialist predators
(McMurtry 1992, Chang & Kareiva 1999, Hagen
et al. 1976). However, rarely have trophic level in-
teractions of generalist predators and their prey
been reported for unmanaged systems (Chang &
Kareiva 1999). One explanation for the paucity of
studies concerning generalist predators may be
related to a limited knowledge of their identity
and life history traits, specifically in immature
stages. The predaceous pentatomid P. mucrona-
tus, for instance, is widely distributed in south

Florida. To date the immature stages have never
been described. The descriptions provided herein
should facilitate future research on predator-prey
interactions with P. mucronatus.
Morphological characteristics described herein
are useful for distinguishing various life stages of
P mucronatus from other commonly occurring
Podisus species, specifically P. maculiventris and
P sagitta (Fab.). For instance, P mucronatus eggs
possess 7-11 (x = 8.25, 0.90) micropylar pro-
cesses compared to 13-16 on eggs ofP maculiven-
tris and P sagitta (De Clercq & Degheele 1990).
Nymphal stages 1-4 of both P sagitta and P mac-
uliventris exhibit distinctive orange and white
patterns on the abdominal tergum, which are not
present in P mucronatus (De Clercq & Degheele
1990, DeCoursey & Esselbaugh 1962). Unlike
P sagitta or P. maculiventris, adult P. mucronatus
possess 3 distinctive white callused spots on the
scutellum and forward projecting humeral spines
(Thomas 1992).
Consistent with other predatory pentatomids,
P mucronatus successfully completed the first
nymphal stage in the absence of prey, although
individuals were observed imbibing free water
from the filter paper or from condensation in the
petri dish. Without prey, survivorship from sec-
ond to third nymphal-instars was 0% when reared
with only filter paper and 15% when reared with
a fresh M. quinqueneruia bud. These findings sug-
gest that the presence of M. quinquenervia foliage
increased nymphal survivorship but it remains
unclear if this is related to facultative plant feed-
ing or to an increase in relative humidity from the
vegetation within the arena. Not surprisingly,
survivorship increased to 95% in the presence of
either coleopteran diet (Table 2).
When compared between prey diets, total de-
velopment time was slower with 0. vitiosa larvae
than with T molitor larvae (Table 2). One expla-
nation for this may pertain to the antipredatory
activity of the viscous coating that covers imma-


Duration (days) of nymphal stages: mean (SD)

Diet" 1 2 3 4 5 Total Survivorship

1 1.00 (0.00) bb 0.00 a
2 0.95 (0.22) ab 2.33 (0.58) ab 0.00 a
3 0.70 (0.47) a 3.65 (1.18) a 2.45 (0.76) 2.55(0.69) 3.87(0.69) 13.60 (1.88) a 0.95 b
4 0.90 (0.45) ab 2.70 (0.66) b 2.35 (0.59) 2.35(1.18) 4.00 (0.56) 12.30 (1.59) b 0.95 b

P-value 0.04' 0.01 0.64 0.52 0.80 0.02 <0.00

aDiets consisting of: 1) moistened filter paper only, 2) diet 1 plus the addition of a M. quinquenervia tip, 3) diet 2 plus a single 0. vitiosa larva (2-3 in-
star) or 4) diet 2 plus a 2-4'" T molitor larval instar.
bStudent-Newman-Keuls all pairwise multiple comparison procedure (SigmaStat 1995).
'One-way ANOVA.

Florida Entomologist 85(2)

ture stages of 0. vitiosa (Purcell & Balciunas
1994). Montgomery & Wheeler (2000), for in-
stance, demonstrated that larvae covered with
this coating were repellent to the red imported
fire ant, Solenopsis invicta Buren. While P mucr-
onatus readily attacks 0. vitiosa larvae under
field conditions, it is unclear whether this larval
coating influences development of nymphs.
When comparing immature developmental
rates with those of other Podisus species, P mucr-
onatus (11-18 d at 25C) appears to have a shorter
developmental time than P. maculiventris (25-31
d at 27C), P placidus (33.1 d at 27C) and P. sag-
itta (18-32 d at 23C) (De Clercq & Degheele
1990). DeBach (1964) suggests that a short devel-
opmental time is a desirable characteristic of
insect biological control agents. Assuming differ-
ences in developmental times are not due to prey
suitability or experimental design, rates of popu-
lation increase for P mucronatus may be higher
than those Podisus species commonly introduced
for pest suppression in other agroecosystems.
These findings suggest that P mucronatus may
be a useful biological control agent of pest co-
leopteran and lepidopteran larvae.
Podisus mucronatus has been observed as the
most common predator associated with the weed
biological control agent, 0. vitiosa. Our labora-
tory-based data suggested that P. mucronatus
readily feeds and completes development when
provided 0. vitiosa prey. However, it remains un-
clear if P. mucronatus regulates populations of
this weed biological control agent under field con-
ditions, and what impact this predation has on
weed suppression. Future studies will focus on
assessing the association, predator-prey interac-
tions and population dynamics of this native pen-
tatomid and the introduced herbivore.


The authors would like to thank J. Eger for com-
ments on earlier versions of the manuscript. We appre-
ciate Susan E. Halbert of the Florida Department of
Agriculture (Division of Plant Industries) for the initial
identification of P. mucronatus. We also thank Willey
Durden for assistance with the photography of P. mucr-
onatus. This is Journal Article R-08322 of the Florida
Agricultural Experiment Station.

ALDRICH, J. R. 1986. Seasonal variation of black pig-
mentation under the wings in a true bug (Hemi-
ptera:Pentatomidae): a laboratory and field study.
Proc. Entomol. Soc. Wash. 88(3):409-421.
M. BORGES. 1991. Pheromone blends of predaceous
bugs (Heteroptera: Pentatomidae: Podisus spp.) Z.
Naturforsch. 46c: 264-269.

CELL, AND P. D. PRATT. 2000. Field colonization of
the melaleuca snout beetle (Oxyops vitiosa) in South
Florida. Biological Control 19: 112-123.
CHANG, G. C., AND P. KAREIVA. 1999. The case for indig-
enous generalists in biological control, pp. 103-115.
In B. Hawkins and H. V. Cornell [eds.], Theoretical
Approaches to Biological Control. Cambridge Univ.
Press, London 412 pp.
DEBACH. 1964. Biological Control by Natural Enemies.
Cambridge University Press, Cambridge 323 pp.
DE CLERCQ, P., AND D. DEGHEELE. 1990. Description
and life history of the predatory bug Podisus sagitta
(FAB.) (Hemiptera: Pentatomidiae). Can. Ent. 122:
DE CLERCQ, P. 2000. Podisus online. http://allserv.rug.ac.
be/~-padclerc/ Last updated: May, 4 2000.
scriptions of the nymphal stages of some North
American Pentatomidae (Hemipera-Heteroptera).
Ann. Ent. Soc. Amer. 55: 323-342.
H. FAUBERT. 1984. Development and survival of Po-
disus maculiventris (Hemiptera: Pentatomidae), a
predator of the Colorado potato beetle (Coleoptera:
Chrysomelidae). Environ. Entomol. 13: 1283-1286.
GENUNG, W. G. 1959. Notes on the syntomid moth
Lymire edwardsi (Grote) and its control as a pest of
Ficus in south Florida. Florida Entomol. 42: 39-42.
Inter-relationship of stinkbugs and diseases to Ever-
glades soybean production. Proc. Soil Crop Sci. Soc.
Florida 24: 131-137.
1976. The biology and impact of predators, pp. 93-
142. In C. B. Huuaker and P. S. Messenger [eds.],
Theory and Practice of Biological Control. Academic
Press, New York 788 pp.
LEGASPI, J. C., AND R. J. O'NEIL. 1993. Life history of
Podisus maculiventris given low numbers of Epil-
anchna varivestis as prey. Environ. Entomol. 22(5):
LEGASPI, J. C., AND R. J. O'NEIL. 1994. Developmental
response of nymphs of Podisus maculiventris (Het-
eroptera: Pentatomidae) reared with low numbers of
prey Environ. Entomol. 23(2): 374-380.
McMURTRY, J. A. 1992. Dynamics and potential impact
of 'generalist' phytoseiids in agroecosystems and
possibilities for establishment of exotic species. Exp.
Appl. Acarol. 14: 371-382.
predatory activity of the weevil Oxyops vitiosa: a bi-
ological control agent of Melaleuca quinquenervia. J.
Ins. Behav. 13 (6): 915-926.
PURCELL AND BALCIUNAS. 1994. Life history and distri-
bution of the Australian weevil Oxyops vitiosa, a
potential biological control agent for Melaleuca quin-
quenervia. Ann. Entomol. Soc. Amer. 87: 867-873.
SIGMASTAT 1995. Version 2.0, Jandel Corporation.
tion rates, survivorship and development of Podisus
maculiventris (Het.: Pentatomidae) on larvae of Lep-
tinotarsa decemlineata (Col.: Chrysomelidae) and
Pieris Brassicae (Lep.: Pieridae), under field condi-
tions. Entomophaga 39(1): 3-9.
THOMAS, D. B. 1992. Taxonomic Synopsis of the Asopine
Pentatomidae (Heteroptera) of the Western Hemi-
sphere. Entomol. Soc. Amer., Lanham, MD. 156 pp.

June 2002

Liang & Jiang: First Record and a New Species of Punana


Department of Entomology, Institute of Zoology, Chinese Academy of Sciences
19 Zhongguancun Road, Beijing 100080, P.R. China

Punana sinica Liang sp. nov. (Hemiptera: Fulgoroidea: Delphacidae) is described and illus-
trated from Sichuan, southwest China. This represents the first record of the genus Punana
Muir from China and the fifth known species of Punana. The new taxon extends the range
of the genus Punana northward considerably, previously known only from southeast Asia
and south India. A key for separation of the species of Punana is included.

Key Words: Punana, new species, Delphacidae, Fulgoroidea, China

Se describe y se ilustra Punana sinica sp. nov. (Hemiptera: Fulgoroidea: Delphacidae) de Si-
chuan, en el suroeste de China. Este represent el primer registro del g6nero Punana Muir
en China y la quinta especie de Punana conocida en el mundo. Este nuevo tax6n extiende la
distribuci6n geografica del g6nero Punana hacia el norte considerablemente, que antes se co-
nocia solamente en el suroeste de Asia y en el sur de la India.

The Delphacidae is the largest family of the
Fulgoroidea, comprising more than 2000 de-
scribed species in approximately 300 genera and
six subfamilies worldwide (Asche 1985, 1990).
Members of the group are predominantly mono-
cot-feeders and a few are major agricultural pests
on grasses, such as rice, maize, and sugarcane
(Wilson & O'Brien 1987, Wilson et al. 1994).
The delphacid fauna of China remains inade-
quately studied. The only comprehensive treat-
ment of Chinese Delphacidae was that of Kuoh et
al. (1983) in their Delphacidae volume of the Eco-
nomic Insect Fauna of China, which deals with
123 species distributed in 47 genera, 2 tribes and
2 subfamilies. The number of described species
likely represents only a small fraction of the ac-
tual diversity of the whole Chinese delphacid
fauna considering the vast territory and various
complex habitats of China.
The basal delphacid taxa, the Asiracinae and
Vizcayinae, of Delphacidae in the Chinese fauna
have received very little attention. To date, only
six species in four genera from the Asiracinae
(Asiracini, Ugyopini) and Vizcayinae are de-
scribed or recorded from China, e.g., Asiraca
choui (Yuan & Wang) (Shaanxi in central China),
A. clavicornis (F.) (Xinjiang in northwest China),
A. granulipennis (Kato) (Manchuria and Jinlin in
northeastern China), and Ugyops zoe Fennah
(Hainan Island in south China), all belonging in
the Asiracinae, and Vizcaya longispinosa Liang
(Yunnan in southwest China) andNeovizcaya sin-
ica Liang (Yunnan in southwest China) of the
subfamily Vizcayinae (Fennah 1956, Asche 1985,
1990, Liang 1996, 1998, 2002).

The genus Punana was described by Muir
(1913) for P brunnea Muir from Borneo. Asche
(1983) correctly separated Punana from
Neopunana (8 species, the Caribbean) and Equa-
systatus (monotypic, Ecuador). Four species are
currently included in the genus from Borneo,
Philippines (Luzon, Negros), and south India
(Muir 1913, 1916, Distant 1916, Metcalf 1943,
Asche 1983, 1985). Asche (1985) placed Punana
Muir in the tribe Ugyopini Fennah 1979 of the
subfamily Asiracinae Motschulsky 1863. How-
ever, Emeljanov (1995) upgraded the Ugyopini to
the subfamily rank and recognized three tribes in
the Ugyopinae, e.g., Neopunanini Emeljanov
1995, Eodelphacini Emeljanov 1995 and Ugyo-
pini. Punana Muir was placed by Emeljanov
(1995) in the tribe Eodelphacini together with Eo-
delphax Kirkaldy 1901, Ostama Walker 1857,
Paranda Melichar 1903, Melanesia Kirkaldy
1907, Livatiella Fennah 1956 and Prolivatis
Emeljanov 1995. Eodelphacini is characterized by
the distal part of the aedeagal shaft arched clock-
wise (from the base curved to the left); the pres-
ence of a row of teeth on the metatarsomere II, in
which the marginal teeth are considerably longer
than all others; the presence of the sinus on the
hindwings opposite CuAP; forewings without
postnodal transverse veins; the presence of a
well-defined bend of the membrane when the
wings are folded; and the intermediate carinae of
mesonotum straight (Emeljanov 1995).
In the present paper the senior author de-
scribes a new species of the genus Punana Muir
which was recently found in Sichuan, southwest
China. The new species represents the first record

Florida Entomologist 85(2)

of Punana in China, and its discovery has broad-
ened our knowledge of the morphology and bio-
geography of the genus, as well as that of the
primitive delphacid taxa in the Chinese fauna.

The specimen used in this study is from the In-
sect Collection of the Institute of Zoology, Chinese
Academy of Sciences, Beijing, China (IZCAS).
Morphological terminology follows that of
Kramer (1950) and Asche (1985).

Punana Muir
Punana Muir, 1913: 249. Type species:P, brun-
nea Muir, 1913, by original designation.
Onkelos Distant, 1916: 137. Type species:
0. annulatus Distant, 1916, by original designation
and monotypy. [Synonymized by Muir, 1919: 6.]
Head (Figs. 1, 3, and 4) relatively broad, broadly
roundly produced anteriorly. Vertex (Figs. 1 and 3)
very short, broader than long, disk foveate with a
median carina (very faint in P sinica sp. nov.) and
two sublateral carinae which meet anteriorly with
the extreme apical part incised posteriorly. Frons
(Fig. 4) longer than broad, anterior margin trun-
cate, convex medially, lateral marginal areas de-
pressed, centrally finely carinate. Postclypeus (Fig.
4) more than half length of frons, convex medially,
lateral marginal areas depressed, with median
longitudinal carina. Anteclypeus (Fig. 4) very nar-
row and convex, with median carina. Rostrum
long, reaching between hind trochanters, basal
segment relatively long, apical segment relatively
short, greater than 1/2 length of basal segment.
Antennae (Figs. 1, 3, and 4) moderately long, scape
and pedicel with many sturdy bristles on surface,
scape shorter than pedicel, with base distinctly
narrow, strongly broadening toward apex, apex
distinctly broad; pedicel cylindrical with base
slightly flattened, with about 18 sensory fields.
Pronotum (Figs. 1 and 3) centrally slightly shorter
than head, disk slightly sloping anteriorly, ante-
rior lateral areas strongly sloping laterad, hind
margin centrally slightly arched anteriorly, with
median longitudinal carina. Mesonotum (Figs. 1
and 3) longer than head and pronotum combined,
with five longitudinal carinae on disk, extreme
apex somewhat lobately produced. Legs moder-
ately long, hind tibiae with three lateral spines on
outer edge, five apical spines (the outer one largest
and the middle one smallest) and a long apical con-
ical mobile spur, metatarsomere I with 5 apical
spines: four spines in a row and the fifth, middle
spine, shifted proximally from the row, metatar-
somere II with 3 apical spines (outer side 2, inner
side 1). Forewings (Figs. 1 and 2) relatively narrow
and elongate, about 3 times as long as broad, ob-
liquely depressed laterally, apices broadly angu-


Figs. 1-2. Punana sinica Liang sp. nov. 1. male holo-
type, dorsal view; 2. same, lateral view.

late, veins distinctly prominent and thickly
covered with fuscous granules with long setae, a
more or less continuous series of transverse veins
before apical area. For forewing and hindwing
venation see Asche (1985: fig. 204).
Male genitalia. See description of Punana sin-
ica Liang sp. nov. below and Asche (1985: figs.
393, 441,442).
Remarks. The above generic description was
based on the specimen of the new species de-
scribed below.
Punana is distinguished from other genera in
the tribe Eodelphacini by the shape and length of
the antennae, the shape of vertex and frons, the
number of longitudinal carinae on mesonotum,
the number of lateral spines on hind tibiae, the
wing venation, and the minutiae of the male gen-
italia, as noted above.
Punana can be distinguished from Melanesia
Kirkaldy (7 species, Borneo, Philippines and Fiji)
and Paranda Melichar (monotypic, Sri Lanka) by
the hind tibiae having 3 lateral spines (2 in
Melanesia and Paranda) and the antennal pedicel
distinctly short (distinctly elongate in Melanesia
and Paranda). It can be distinguished from Eodel-
phax Kirkaldy (2 species, Sri Lanka) by the an-
tennal scape and pedicel subcylindrical (both
scape and pedicel compressed and scape obliquely
triangular in Eodelphax). Punana differs from
Prolivatis Emeljanov (monotypic, Vietnam) in the
antennal pedicel short (relatively long in Proliva-
tis) and the male parameres relatively broad
(very slender and narrow in Prolivatis). It differs
from Ostama Walker [2 species, Borneo, Mentawi
Island (East Indies)] in the antennae short (very
elongate in Ostama), mesonotum with 5 carinae
(3 in Ostama) and forewings with about 12 closed
apical cells (about 15 in Ostama). Punana can be
separated from Livatiella Fennah (2 species, east-
ern Caroline Island) by the forewing with Sc+R
with two branches before subapical transverse

June 2002

Liang & Jiang: First Record and a New Species of Punana

nodal line (one branch in Livatiella) and the hind-
wing with the apical cell enclosed by M, and M,
relatively long and large (very short and small in
Included species and distribution. Five spe-
cies; P annulata (Distant, 1916) (south India),

P brunnea Muir, 1913 (Borneo), P. negrosensis
Muir, 1916 [Philippines (Negros)], P philippina
Muir, 1916 [Philippines (Luzon)], andP sinica sp.
nov. [southwest China (Sichuan) (new record)].
A key to the known species of Punana is pro-
vided below.


1. Antennae with three dark rings (Muir 1913); Borneo ......................... ........... brunnea Muir
Antennae without dark rings . . ........................................................ 2
2. Forewings with distinct stramineous suffusions, especially on basal, subapical and apical areas ............ 3
-Forewings without distinct stramineous suffusions . . ........ ............................ 4
3. Vertex (Figs. 1-3) shorter and broader with anterior margin shorter and more rounded; male pygofer (Figs. 5 and
6) with a large median triangular process on ventrocaudal margin; and parameres (Figs. 5, 8, and 9) dis-
tinctly angulately produced laterad near midlength in ventral aspect; southwest China (Sichuan)
...................................................................... sin ica L ian g sp nov.
-Vertex relatively long, relatively distinctly produced anteriorly (Distant 1916: Fig. 98; Asche 1985: Figs.
11c, 161); male pygofer without distinct process on ventrocaudal margin (Asche, 1985: Fig. 393); parameres
not angulately produced laterad near midlength in ventral aspect (Asche 1985: Fig. 441); south India
....................................................................... an n ulata (D instant)
4. Male pygofer with median process on ventrocaudal margin angular (Muir 1916); fore and middle coxae and fem-
ora very light brown; clavus without fuscous spots; Philippines (Luzon). ............... philippina Muir
-Male pygofer with median process on ventrocaudal margin square (Muir 1916); fore and middle coxae
and femora dark brown; clavus with a small fuscous spot; Philippines (Negros) ........ negrosensis Muir

Punana sinica Liang sp. nov.
(Figs. 1-10)

Holotype male, CHINA: Sichuan, Wan County
(3008'N, 10803'E), 1200 m, 27.v.1994 (Jin Yao) (IZ-
Description. Length (from apex of vertex to tip
of forewings): male 5.8 mm.
Vertex, frons, postclypeus and anteclypeus
brownish, the pit on frons in front of vertex
darker, frons (Fig. 4) with two columns of small
rounded pale spots (one next to median longitudi-
nal carina and the other on lateral margin), epis-
tomal suture ochraceous, genae and lora
ochraceous with the latter suffused with fuscous.
Antennae ochraceous with apex of scape and
pedicel suffused with brown, scape and pedicel
covered with long brownish sturdy bristles. Ros-
trum with basal segment ochraceous, apical seg-
ment brown with extreme apex black. Pronotum
ochraceous with anterior lateral depressed areas
blackish brown. Mesonotum (Figs. 1 and 3) fus-
cous or blackish brown, with longitudinal carinae
(excluding the median carina), posterior lateral
margins and extreme apex ochraceous. Thorax
ventrally ochraceous, pleurae with fuscous spots.
Legs ochraceous with dark brown or fuscous suf-
fusion, femora dark brown or fuscous (hind fem-
ora much paler), tibiae with two broad fuscous
rings basally and medially respectively (those on
hind tibiae much paler), tarsi and claws brown
(hind tarsi and claws much paler), tips of apical
spines on hind tibiae and tarsi black. Forewings

(Figs. 1 and 2) fuscous brown, with irregular stra-
mineous suffusions mainly on basal, subapical
and apical areas, granules on veins fuscous, setae
in granules on veins pale yellowish brown; hind-
wings pale fuliginous, veins brown. Abdomen
with tergites brown and sternites ochraceous
with anterior margin broadly brown.
Structural characters as in generic description,
but vertex with disk short and transversely broad,
with very faint median longitudinal carina, frons
with a very small shallow rounded pit dorsally,
and pronotum shorter than vertex medially.
Male genitalia with pygofer (Fig. 5) in ventral
view with apical 2/5 broad and nearly parallel-
sided, narrowing to base over basal 3/5, ventrocau-
dal margin broadly excavated with a large median
triangular process. Pygofer (Fig. 6) in lateral view
elongate, very short anteriorly and very high pos-
teriorly, dorsal margin very short and ventral mar-
gin very long. Parameres (Figs. 5, 6, 8, and 9)
slender and elongate, distinctly angulately pro-
duced laterad near midlength in ventral aspect,
base relatively broad, tapered to apex, somewhat
constricted medially, apical 1/2 directed inwardly
and covered with long hairs on surface. Connective
(Fig. 9) relatively short, slender, parallel-sided,
apex and base expanded laterad. Aedeagal shaft
(Figs. 5, 6, and 10) nearly C-shaped with basal 1/2
compressed and apical 1/2 spinous; gonopore at
midlength on dorsal surface. Suspensorium (Fig.
10) long, compressed, base ring-shaped, embrac-
ing base of aedeagus, apical part curved and broad
with apex bluntly forked. Anal tube (Figs. 6 and 7)

Florida Entomologist 85(2)

- -tt ** ..


1- -




N <4

,I / \.;\
P' \
4,(Y vj1)

"8 i. 9


Figs. 3-10. Punana sinica Liang sp. nov. 3. head, pronotum and mesonotum, dorsal view; 4. head, ventral view;
5. pygofer, ventral view; 6. pygofer, lateral view; 7. anal segment, caudal view; 8. genital styles, ventral view; 9. gen-
ital style and connective, ventral view; 10. aedeagus, lateral view. Scale bars = 0.5 mm in Figs. 3, 4 and 0.2 mm in
Figs. 5-10.

short and broad, gradually widening from base to
apex in dorsal aspect, apex without appendages,
anal style slender, small and short.
Female. Unknown.
ErK \. ..1.....,'. This new species is named sinica
after the Latin sinicus, adjective ("of China"), re-
ferring to its occurrence in China.
Distribution. Southwest China (Sichuan).
Remarks. This new species is externally simi-
lar to P annulatus (Distant) from south India
(Kodaikanal), which was illustrated by Distant
(1916) and Asche (1983, 1985), but differs from
the latter in having the antennal pedicel subcy-
lindrical and in the shape of the vertex, male gen-
italia and pygofer, as noted in the above key.

We thank Dr. Manfred Asche, Museum ffir Natur-
kunde der Humboldt-Universitat zu Berlin, Germany
and Dr. A. F. Emeljanov, Zoological Institute, Russian
Academy of Sciences, Leningrad, Russia, for several dis-
cussions during the preparation of this manuscript. We
also thank Dr. Manfred Asche, Dr. Steve Wilson (Depart-
ment of Biology, Central Missouri State University,
Warrensburg, MO, USA), Dr. Lew Deitz (Department of
Entomology, North Carolina State University, Raleigh,
NC, USA), Dr. R. M. Baranowski (University of Florida,
TREC, Homestead, FL, USA) and Dr. Gary Steck (Divi-
sion of Plant Industry, Florida Department of Agriculture

& Consumer Service, Gainesville, FL, USA) for review-
ing the manuscript and suggesting improvements. The
work on which this paper is based was supported by the
National Natural Science Foundation of China grant
number 39925006 and the Chinese Academy of Sciences
Biological Innovation Fund A2999084.

ASCHE, M. 1983. Aufgliederung der Asiracinen-Gat-
tung Punana Muir, 1913; Equasystatus gen. nov. aus
Equador und Neopunana gen. nov. von den Karibis-
chen Inseln (Homoptera Auchenorrhyncha Fulgoro-
morpha Delphacidae). Marburger Entomol. Pub. 1(8):
ASCHE, M. 1985. Zur Phylogenie der Delphacidae
Leach, 1815 (Homoptera Cicadina Fulgoromorpha).
Marburger Entomol. Pub. 2(1-2): 1-912.
ASCHE, M. 1990. Vizcayinae, a new subfamily of Delpha-
cidae with revision of Vizcaya Muir (Homoptera:
Fulgoroidea)-a significant phylogenetic link. Bishop
Mus. Occ. Papers 30: 154-187.
DISTANT, W. L. 1916. The fauna of British India, includ-
ing Ceylon and Burma. Rhynchota Vol. 6 (Homoptera:
Appendix). Taylor & Francis, London. viii + 248 pp.
EMELJANOV, A. F. 1995. On the question of the classifi-
cation and phylogeny of the Delphacidae (Homop-
tera, Cicadina), with reference to larval characters.
Entomol. Obozr. 74: 780-794 [In Russian with Eng-
lish summary; Russian summary separately pagi-
nated, pp. 944-945. English translation in Entomol.
Rev. 75(9): 134-150, 1996.]


'K ___




June 2002

Liang & Jiang: First Record and a New Species of Punana

FENNAH, R. G. 1956. Fulgoroidea from southern China.
Proc. California Acad. Sci. (4)28: 441-527.
KRAMER, S. 1950. The morphology and phylogeny of
auchenorhynchous Homoptera (Insecta). Illinois
Biol. Monog. 20: 1-109, pls. 1-15.
1983. Economic Insect Fauna of China. Fasc. 27.
Homoptera: Delphacidae. Science Press. Beijing, 166
pp., 13 pls. [In Chinese, English abstract p. 137.]
LIANG, A.-P. 1996. Taxonomic changes in Chinese Lopho-
pidae with a check list of Chinese species (Homoptera:
Fulgoroidea). Pan-Pacific Entomol. 72: 145-151.
LIANG, A.-P. 1998. On the Eurasian planthopper genus Asi-
raca Latreille (Homoptera: Auchenorrhyncha: Fulgoro-
morpha: Delphacidae). Reichenbachia 32:187-196.
LIANG, A.-P. 2002. New taxa of Vizcayinae (Hemiptera:
Auchenorrhyncha: Delphacidae), including a remark-
able new genus from China. J. Nat. Hist. 36: 601-616.
METCALF, Z. P. 1943. General Catalogue of the Hemip-
tera. Fasc. IV. Fulgoroidea, Part 3. Araeopidae (Del-
phacidae). Smith College, Northampton, MA. 552 pp.

MUIR, F. A. G. 1913. On some new Fulgoroidea. Proc.
Hawaiian Entomol. Soc. 2: 237-269, pl. 6.
MUIR, F. A. G. 1916. Additions to the known Philippine
Delphacidae (Hemiptera). Philippine J. Sci. 11: 369-
MUIR, F. A. G. 1919. Notes on the Delphacidae in the Brit-
ish Museum collection. Canadian Entomol. 51: 6-8.
SON. 1994. Evolutionary patterns of host plant use
by delphacid planthoppers and their relatives, pp. 7-
45. In R. F. Denno and T. J. Perfect (eds.), Planthop-
pers: their ecology and management. New York:
Chapman Hall.
WILSON, S. W., AND L. B. O'BRIEN. 1987. A survey of
planthopper pests of economically important plants
(Homoptera: Fulgoroidea), pp. 343-360. In M.R. Wil-
son and L.R. Nault (eds.), Proc. 2nd Int. Workshop on
Leafhoppers and Planthoppers of Economic Impor-
tance. CAB International Institute of Entomology,

Florida Entomologist 85(2)

June 2002


1University of Florida, NFREC-Monticello, Route 4, Box 4092, Monticello, FL 32344

2Metropolitan Life Insurance Co., Inc., Tampa, FL

'Undergraduate, Junior, University of Florida, Gainesville, FL 32611


Trolling, a novel trapping method, was developed and tested for deer flies, Chrysops spp.,
and other Tabanidae. The trap is a plastic plant container coated with Tanglefoot that is
mounted upside down on a rod in an apparatus attached to a vehicle. The vehicle is then
driven "trolled" to attract Tabanidae. Trap movement, color, shape, dimension and size were
evaluated to improve trap catch. Response by Chrysops vittatus Wiedemann/C. pikei Whit-
ney and C. macquarti Philip to the trap's parameters are reported. The most effective trap
is a 15 cm diameter pot (Lerio C-360, B-6, The Lerio Corp. Mobile, AL) painted bright blue,
placed 1-2 m above the ground and moved at a speed of less than 3.13 m/sec. Response of
Chrysops spp. to the trap indicated the hierarchy of behavioral stimuli to Chrysops spp. in
order of importance to be height, movement, speed, dimensions, color, size, and contrast. No
known tabanid attractants tested including CO2, acetone or octenol increased trap captures
and the insect repellent, DEET, N,N-diethyl-3-methylbenzamide, did not reduce trap cap-
tures. Other biological information derived during the trapping experiments is reported. The
trolling trap appears very valuable to detect and monitor certain Chrysops spp. and other ta-
banid populations for scientific purposes. In addition the trap or modifications of it, mounted
either on or near humans (hat or walking stick) or on vehicles, may be useful to reduce or
eliminate attacks from Chrysops spp. or to suppress their activity for short time periods in
small areas such as dooryards.

Key Words: Chrysops vittatus, C. pikei, C. macquarti, trap, Tabanidae, trolling


Trolear, un novedoso m6todo de capture, fue desarrollado y probado para moscas del genero
Chrysops spp., y otros Tabanidae. La trampa consiste en un envase plastico para plants
recubierto con Tanglefoot que esta montado el rev6s sobre una barra en un aparato que va
adherido a un vehiculo. El vehiculo posteriormente es manejado "troleado" para atraer Ta-
banidae. El movimiento de las trampas, color, forma, dimension y tamano fueron evaluados
para mejorar la capacidad de capture de la trampa. La respuesta por parte de Chrysops
vittatus Wiedemann/C. pikei Whitney y C. macquarti Philip a los parametros de la trampa
son reportados. La trampa mas efectiva es un envase de 15 cm de diametro (Lerio C-360, B-
6, La Corp. Lerio, Mobile, AL) pintado de color azul brillante, colocado a 1-2m sobre el suelo
y movida a una velocidad menor a 3.13 m/seg. La respuesta de Chrysops spp. a la trampa,
indicaba lajerarqufa del estimulo de comportamiento a Chrysops spp. en orden de importan-
cia el cual era altura, movimiento, velocidad, dimensions, color, tamano, y contrast. Nin-
gun atrayente para tabanidos conocido que se haya probado incluyendo CO, acetona o
octenol, aument6 el numero de captures en la trampas, y el repelente de insects, DEET, N,
N-dietil-3-metilbenzamida, no redujo el numero de captures en la trampa. Otra information
biol6gica obtenida durante los experiments de trampeo se reportan. La trampa de troleado
parece ser muy valiosa para detectar y monitorear ciertos Chrysops spp. y otras poblaciones
de tabanidos para prop6sitos cientificos. ademas la trampa o modificaciones de esta, coloca-
das tanto sobre o cerca de humans (sombreros o bastones para caminar) o sobre vehiculos,
pueden ser tiles para reducir o eliminar ataques de Chrysops spp. o para suprimir su acti-
vidad por cortos periods de tiempo en pequenas areas tal como patios de casa.

Detection and monitoring of tabanids many types of adult traps for monitoring taban-
(Diptera: Tabanidae) began when Hansens (1947) ids, especially horse flies have been developed.
demonstrated that Tabanus nigrovittatus Mac- The Manitoba fly trap was described and used by
quart could be captured with black shingles Thorsteinson (1958) and Thorsteinson et al.
coated with an adhesive material. Since then, (1965). Black and red spheres and two- and three-

Mizell et al.: Trolling: A Novel trap for Chrysops spp.

dimensional silhouettes were used by Bracken et
al. (1962) and Browne and Bennett (1980). Mal-
aise traps (Smith et al. 1965, Wilson et al. 1966,
Roberts 1971a,b), silhouette traps simulating
cows (Wilson et al. 1966), red and black helium-
filled spheres tethered 1.2-1.8 m above the ground
(Snoddy 1970), modified Manning traps (Granger
1970), canopy traps (Catts 1970, French & Kline
1989, Hribar et al. 1991a), sticky traps (Hansens
et al.1971), rigid canopy traps (Axtell et al. 1975),
aerial netting (Tallamy et al. 1976, Cilek and
Schreiber 1996), box traps (Wall & Doane 1980,
Ailes et al. 1992), panel traps (Allan & Stoffalano
1986), two-tiered box traps (Jackson et al. 1993,
French and Hagan 1995), and brown boards on
the ground and white buckets in the air (Moore et
al. 1996) have been used to successfully monitor
many tabanid species. However, different trap
types vary in their ability to capture deer flies
(e.g., Chrysops spp.) or other tabanid subgroups
(horse flies), but can provide diverse information
about the behavior of individual species. Tallamy
et al. (1976) compared Malaise traps to aerial net-
ting and reported that the latter favored
Chrysops spp. while Malaise traps were better in-
dicators of tabanid diversity and species even-
ness. Neither trap alone should be used in
tabanid community studies.
Trap color is important to tabanid attraction
and landing. Black and red spheres (Bracken et al
1962, Snoddy 1970) were found to be superior to
green and yellow spheres and white dappling or
striping of black traps dramatically reduced at-
traction of Tabanus illotus (O.S.). Browne and
Bennett (1980) reported that both Chrysops spp.
and Hybomitra spp. were attracted to blue or red,
but were consistently not attracted to black, yel-
low or white. Tabanids landed in the areas
painted the attractive color on striped traps with
alternating attractive and unattractive colors. Al-
lan and Stoffalano (1986) reported that T nigro-
vittatus preferred blue, black and red panel traps.
Increased trap capture occurred when trap inten-
sity was increased or decreased against a back-
ground. Wall and Doane (1980) used box traps of
green, black or blue to trap T nigrovittatus from
1970-1979 and indicated a measurable decrease
in the flies over the trapping period. Moore et al.
(1996) used brown boards on the ground and
white buckets on poles to determine emergence,
concentrations and flight periodicity of T abactor
Philip. Male flies preferred the boards while fe-
male flies preferred buckets (Moore et al. (1996).
Hribar et al. (1991b) showed that reduction of
ultraviolet (UV) light increased catch of certain
tabanids. However, Hribar and Foil (1994) re-
tested the effect of UV reflectance of canopy traps
on tabanids in Louisiana and concluded that UV
had little effect on most species.
Odors have also been used to enhance trap re-
sponse. Wilson et al. (1966) used carbon dioxide

(CO2) in silhouette traps and captured 23 species
of female tabanids. Wilson (1968) using sticky
traps with CO2 on a pasture periphery signifi-
cantly reduced tabanids on cattle in the pasture.
Red and black helium-filled spheres tethered 1.2-
1.8 m above the ground captured large numbers
of Chrysops niger taylori Philip, but Snoddy
(1970) determined that CO2 was a poor attractant
for deer flies. Using the canopy trap with CO2,
Catts (1970) reported capturing ca. 1000 tabanids
per hour in Delaware. Roberts (1971a and b) used
Malaise traps and elucidated a species-specific re-
sponse to CO2 rates by tabanids. Roberts (1976a)
compared six types of traps with and without CO2
for collection of tabanids including the Malaise,
plexiglass and canopy traps. Based on the num-
ber of species captured, the Stoneville Malaise
(Roberts 1971a) with CO2 was the most efficient.
Six Tabanus spp. and C. flavidus Wiedemann
were captured in numbers suitable for statistical
analysis (Roberts 1976a). French and Kline
(1989) investigated the response of tabanids to
canopy traps with CO2 plus the tsetse fly attrac-
tant (Hall et al. 1984) 1-octen-2-ol (octenol) iso-
lated from cattle. Octenol alone increased canopy
trap catch three fold over traps without attrac-
tant. Octenol with CO2 enhanced trap catch in to-
tal specimens and number of species captured.
Both Chrysops spp. and Tabanus spp. were af-
fected. Schreck et al. (1993) evaluated configura-
tions of inflated vinyl beach balls in Malaise and
canopy traps as possible insecticide-impregnated
visual targets. Beach balls attracted 2x and 2-5x
more flies respectively when used alone or when
treated with octenol. Leprince et al. (1994) inves-
tigated tabanid response in Louisiana to Jersey
bullocks and canopy traps baited with ammonia,
octenol and CO2. Odors did not significantly in-
crease trap catch. The results of Leprince et al.
(1994) were contrary to the findings of French and
Kline (1989) and Hribar et al. (1991a), and sug-
gest that there may be environmental, geograph-
ical and species differences in response to
attractants by tabanids. Foil and Hribar (1995)
tested the tsetse fly attractants acetone, and
octenol + 3-n-propylphenol and 4-methylphenol
(4:1:8 ratio) in canopy traps in Louisiana. Four-
teen species or species groups of tabanids were
captured in equal numbers to octenol and the
mixture, response to acetone was similar to the
control. Jackson et al. (1993) and French and
Hagan (1995) used a two-tiered box trap to collect
T nigrovittatus, C. fuliginosus Wiedemann and
C. atlanticus Pechuman. Octenol significantly in-
creased the catch of C. atlanticus.
Despite the development of traps and the
knowledge of tabanid behavior, effective monitor-
ing and management methods, and even basic un-
derstanding of the behavior of the thousands of
species of tabanids remain unknown. The objec-
tives of this investigation were to develop and test

Florida Entomologist 85(2)

a novel trapping method and to use the trap to
characterize the behavior of selected tabanid spe-
cies. The trap described herein is most effective
against Chrysops spp. and the responses of three
species trapped in large numbers near Monti-
cello, Florida, C. vittatus Wiedemann, C. pikei
Whitney and C. macquarti Philip, are reported.


Trap development progressed over a series of
experiments in an effort to exploit the observation
of large numbers of deer flies (Chrysops spp.)
chasing and landing on the side mirrors of a vehi-
cle. Experiments were conducted during August-
September, 1994, April-August, 1995, May-Au-
gust, 1996, and May-August, 1998. All trap sur-
faces were covered with a thin layer of heated
Tanglefoot (The Tanglefoot Co., Grand Rapids, MI
49504) applied with a small paint brush. Because
C. vittatus and C. pikei are not possible to sepa-
rate in the field, their counts were combined.
In preliminary experiments we determined
that two-dimensional black silhouette traps
against a completely white background and
mounted on the top or on the sides of a moving ve-
hicle would not induce tabanid species flying
along side to land. Therefore, we developed the
described "trolling" apparatus used for the exper-
iments to mount traps on a vehicle.
All experiments were conducted using a 1992
Dodge Dakota Clubcab pickup, white with gray-
striped side panels and black mirrors. An appara-
tus was mounted on the front and/or back of the
truck to support the traps and enable rotation of
traps among 7 different positions. The apparatus
(Fig. 1) consisted of a 2.5 x 5.0 cm wooden lath
base 1.8 m long with 2, 30 cm long pieces nailed
perpendicularly on the bottom to serve as support
braces. On the front of the vehicle each end of the
base was strapped tightly to the truck wheel wells
using bungi cords. On the back of the vehicle the
apparatus was mounted into the standard holes in
the truck bed. Along the length of the base at ca.
30 cm intervals from each end, two 10 penny nails
were driven through the bottom of the wood lath
ca. 2 cm apart so as to orient with their points ver-
tically through the top side. Small wooden blocks
each with a top centered hole and with 2 bottom
holes to fit snugly over the nails were attached.
Metal rods 30 cm long of ca. 5 mm wire were
formed at the top into a L shape and the straight
bottom was placed into the top hole in the wooden
block. A small binder clip was fastened over the
wire's L-shaped end for trap support. Plant pot
traps were placed on the rods upside down by
drilling 7 mm hole in the center of the bottoms. Po-
sition of the traps was changed by moving the
traps from rod to rod after each replicate.
Experiments were replicated 5-15 times with
each replicate consisting of driving the vehicle for

1-5 min. through an area containing deer flies. Af-
ter each replication the captured flies were re-
moved and recorded by species and trap. Trap
position was rotated clockwise to the next adja-
cent position for the succeeding replication. Traps
in the right end position were returned to the op-
posite left-end position. Replication generally was
conducted such that each trap type was posi-
tioned in each test location 1-3 times.
Paints used for small pyramid and square
shapes were Glidden Rust Master Enamel. Paints
used for the plastic pots were TRU-TEST HI-G,
high gloss enamel of the following colors, 105 Chi-
nese red, U-48 Royal blue (darkest), U-9 Dutch
blue (lightest), and FL-5 Neon blue (brightest), a
TRU-TEST "Easy Color" fluorescent spray. Black
was used as the natural color of the plastic pots.
Spectral reflectance of the blue traps (Fig. 2)
was measured using a USB UV-VIS S2000 spec-
trometer with a DT-1000-MINI tungsten light
source (Ocean Optics, Dunedin, FL). Percent re-
flectance was determined in comparison to a
white Spectralon standard.
Traps were baited with candidate chemicals
using the methods of Foil and Hribar (1995).
Data were analyzed using the SAS Proc GLM
procedures. Means were separated by least signif-
icant difference test when a significant F was indi-
cated by analysis of variance (SAS Institute 1999).
Experiment 1: Pyramid-shaped traps (Tedders
& Wood 1994) ca. 30 cm in height and 15 cm wide
at the base, made of clear plexiglass, unpainted
cardboard or cardboard painted black, red, yellow,
or white and coated with Tanglefoot were used.
Eighteen replicates were conducted with one pyr-
amid of each color mounted on both the front and
back of the truck. Nine replicates were conducted
by driving the truck forward and 9 by driving the
truck backwards for a one minute bioassay. Cap-
tures were processed as described above. The data
were analyzed as a three-way analysis of variance
with direction (driving forward or backwards), po-
sition on the truck (front or back), and trap color
as the factors.
Experiment 2: Black plant pots (15 cm) (Lerio
C-650, B-6, The Lerio Corp., Mobile, AL) were com-
pared in six replicates to the cardboard pyramid
traps of plain brown cardboard, black or white.
Experiment 3: Black unpainted plant pots
(Lerio) of 15, 17.5, and 20 cm were compared to
flat squares of cardboard of 15, 17.5, and 20 cm on
a side and painted black on both sides in eight
Experiment 4: Trap size was tested in this two
part experiment. Black Lerio pots of 5, 7.5, 10,
12.5, 15, 17.5, and 20 cm in diameter were used.
In part one trap sizes from 5-15 cm were tested in
15 replicates. In part two the trap sizes of 15,
17.5, and 20 cm were tested with 8 replicates.
Experiment 5: Red, U-48 Royal blue and black
15-cm pots were tested with 8 replicates. Two

June 2002

Mizell et al.: Trolling: A Novel trap for Chrysops spp.

Fig. 1. The "trolling" deerfly trap: A. truck-mounted apparatus used in experimentation, B. cup version mounted
on cap for personal protection and C. individual trap that can be mounted on a lawnmower, 4-wheeler or other ve-
hicle to suppress deer flies from dooryards and other locations.

traps of each color were used and the pots of the 3
colors were grouped in two sets to begin the first
replicate then randomized as described above.
Experiment 6: Plant pots (15 cm) were painted
with one of three different blue paints. In ascend-
ing order of intensity (authors' vision, Fig. 2) the
colors were: U-9 Dutch blue, FL-5 Neon blue, U-
48 Royal blue. Ten replicates were conducted with
2 traps of each color randomized by position on
the apparatus.
Experiment 7: Two 15 cm black pots were
painted with 2.5 cm alternating vertical stripes of

white and black. The striped pots were compared
to two solid black pots in five replicates.
Experiment 8: The "size" test (expt. 4) was re-
peated using pots painted FL-5 Neon blue. Pots
similar to those described in experiment 3 of sizes
5, 10, 15, 20, and 25 cm were compared in ten rep-
Experiment 9: Background color under the
trap. Cardboard fruit harvesting boxes (25 x 30 x
45 cm) were stapled to form a cup shape. The in-
side of one box each was painted white, FL-5
Neon blue or covered with aluminum foil. To pro-

Florida Entomologist 85(2)


: Neon *

/ Royal
'" '* * \"4 1~


300 400 500 600 700 800

Wavelength (nm)

Fig. 2. Spectral reflectance patterns of three blue paints tested on traps for deer fly response. FL-5 Neon and U-
48 Royal blues captured significantly more deer flies than U-9 Dutch blue (see text).

vide a trap background under the trap, the center
of the cupped box was placed over the nails (fitted
into the wood for holding the rods) on the truck-
mounted apparatus which held the box in place.
FL-5 Neon bluel5 cm traps were placed on the
rods as described above in the center of the back-
ground boxes. Fifteen replicates were conducted
and trap position was changed after 5 replicates
were completed so that each trap was similarly
located next to the other 2 traps for 5 replicates,
i.e., three blocks in time.
Experiment 10: Height above ground: A
wooden lath of 5 x 5 cm was used to construct a
3.7 m pole that fit snugly into the stake holder on
the truck bed. Another piece of lath drilled to hold
the metal rod supporting the trap was nailed per-
pendicular to the pole at 3 heights: 150, 300 and
450 cm. A trap was also placed in the truck bed on
a rod which was at 75 cm above the ground. Traps
of 15 cm diameter painted FL-5 Neon blue were
mounted on the perpendicular stakes at 150, 300,
and 450 cm and 15 replicates were conducted.
Experiment 11: Silhouettes of 30 x 60 cm made
of blue chloroplast sheeting (BBE Graphics Ware-
house, Tampa, FL) were mounted under a trap
placed at 150 cm above ground on the bed of the
truck as described for the height experiment. Two
silhouette traps were compared for each replicate.
The silhouettes were placed either vertically or
horizontally to mimic an animal body under the
trap "head". Replicates were conducted with the
silhouettes oriented differently or the same for
side by side comparison of the effect on deer fly re-

Experiment 12: Attractants and repellents.
Using the methodology of Hribar and Foil (1994)
for elution devices and concentration ranges, CO2
(2.5 1/min open box of dry ice), acetone (500mg/
hr), and octenol (0.5-5.0 mg/hr) were evaluated to
determine the attraction and repellence to C. vit-
tatus/C. pikei. CO2 was investigated using con-
trolled elution rates and dry ice in a variety of
containers to produce a range of low to high vol-
umes both on and around the traps. Ten repli-
cates were also conducted using cannister CO2
eluded through a flowmeter at the rate of 2.5 1/
min. For octenol, 3 FL-5 Neon blue 15 cm pot
traps were used with one trap containing the at-
tractant and 2 traps serving as controls. Traps
were separated on the apparatus by 50 cm (2 sta-
tions) and position was randomized after each of
the 11 replications. The octenol with 98% purity
was purchased from Sigma Chemical Co. Histo-
logical grade acetone was purchased from Fisher
Scientific and was bioassayed as described for
octenol with 7 replicates. N,N-diethyl-3-methyl-
benzamide, DEET, was tested using several elu-
tion methods: vials as with octenol after Hribar
and Foil (1994) and larger dental wicks to provide
higher concentrations. Results from the 8 repli-
cates conducted with dental wicks are reported.
C. macquarti: Experiments (1) with color, size
and background were conducted in April- May,
1997 when this early-emerging species was ob-
served in relatively high numbers. Color and size
were investigated together using 2 Lerio pots
each, one black and the other FL-5 Neon blue, of
diameter 7.5, 15, and 22.5 cm. The six pots were


June 2002

Mizell et al.: Trolling: A Novel trap for Chrysops spp.

initially arranged on the truck apparatus with
the traps of similar size and color adjacent and
then randomized as described above for each of
the 12 replicates. Experiment 2 testing back-
ground colors was conducted as described above
for C. vittatus/C. pikei with 16 replicates. Popula-
tions of C. macquarti were much lower than C.
vittatus/C. pikei and also more variable in space.
Thus, the numbers of flies captured by traps in
each replicate were more variable and trap counts
often were zero. Therefore, the data were trans-
formed before analysis of variance by taking the
square root of trap capture number +1. The un-
transformed means are reported.


Trap efficiency was ca. 95% or better as long as
the Tanglefoot was fresh. Deer flies occasionally
escaped from the Tanglefoot. Therefore, at the be-
ginning and during the experiments as needed,
the Tanglefoot was refreshed by rubbing the sur-
faces of two traps together.
The seasonal occurrence and emergence dates
by years were: first detection of C. vittatus/C
pikei-20 April 1995, 3 May 1996, 29 March 1997,
and 9 April 1998. The last collection date for C.
vittatus/C. pikei was 9 October 1998. We did not
evaluate the diurnal activity of specific Chrysops
spp, but we did collect C. vittatus/C. pikei from
dawn to dusk and occasionally just after dark. C.
macquarti emerged first in Spring and was
present only from late March to early May in
1998. C. macquarti is also much less aggressive
towards humans and smaller in size than C. vit-
tatus/C. pikei. These data agree generally with
Jones (1953) who reported two generations of
C. vittatus in north Florida.
Experiment 1, colored pyramid traps: Colored
pyramid traps mounted on the front on rear of the
vehicle captured deer flies inefficiently relative to
the numbers of C. vittatus/C. pikei that were fly-
ing around them. However, they indicated that
dark colors were more attractive (F = 17.4; df =
5,216; P = 0.0001) (Table 1). Comparison by anal-
ysis of variance of the trap catch on the front and
rear of the vehicle driven either forwards or back-
wards indicated no significant differences: direc-
tion (F = 0.92; df = 1, 216; P = 0.33) vehicle end
(F = 1.39; df = 1,216; P = 0.24) but the interaction
of direction x vehicle end was significant (F =
8.07; df = 1, 216; P = 0.005). The interaction was
significant because the number of deer flies cap-
tured on the front (engine end) of the vehicle was
larger when the vehicle was driven forward or
backward. This result may indicate that heat
from the engine or other characteristic of the ve-
hicle front (white hood providing more contrast)
may have enhanced attraction and/or landing.
Experiment 2, black plant pot vs colored pyra-
midal traps: Because the pyramidal traps were


Experiment 1A Experiment 2
Trap Type Mean + SE Mean + SE

Black pyramid 2.1 + 0.33 A 3.5 + 0.8 A
Tan pyramid 1.5 + 0.29 AB 5.2 + 0.6 AB
Red pyramid 1.8 + 0.27 B -
Clear pyramid 0.2 + 0.07 C -
White pyramid 0.2 + 0.06 C 0 + 0 C
Yellow pyramid 0.2 + 0.05 C -
Black plant pot 8.2 + 2.3 A
aMeans in columns not followed by the same letter are significantly
different as determined by LSD.

inefficient, but "trolling" appeared promising, we
compared other potential trap types. Significant
differences were determined by analysis of vari-
ance with F = 7.3; df= 3, 20;P = 0.0017 and LSD
= 3.7. Black 15 cm plant pots captured 8.2 + 2.3
(mean + SEM) C. vittatus/C. pikei, in comparison
to 5.2 0.6 or less for the other pyramid traps (Ta-
ble 1).
Experiment 3, two-dimensional squares vs
plant pots of 3 sizes: Shape was significantly dif-
ferent (F = 42.3; df = 1,42; P = 0.0001), size was
not significant (F = 2.8; df = 2,42; P = 0.074) and
the shape x size interaction was significant (F =
4.92; df = 2,42; P = 0.012) indicating that the
three-dimensional 15 cm pot trap captured signif-
icantly more (13.9 2.2) than the 15 cm two-di-
mensional trap (1.4 0.6). In this test the 15 cm
black pot trap also captured significantly more
C. vittatus/C. pikei than the 17.5 cm (7.5 + 1.0)
and 20 cm black pot traps (7.3 + 1.5), LSD = 2.34.
Experiment 4, size and color interaction: This
experiment was conducted in two parts. In evalu-
ating black pot traps varying in size from 5-15 cm
we found significant differences (F = 3.84; df =
4,70; P = 0.007) LSD = 3.44. In part two, evaluat-
ing traps of 15, 17.5, and 20 cm, we also found a
significant size effect (F = 6.72; df = 2,21; P =
0.006) LSD = 2.2 (Table 2). The 15 cm pot trap
captured significantly more C. vittatus/C. pikei in
both experiments.
Experiment 5, 15 cm pots of red, black and
blue: Blue traps captured significantly more C.
vittatus/C. pikei than the red and black traps, (F
= 21.5; df = 2,135; P = 0.0001) LSD = 1.56 (Table
Experiment 6, 15 cm pots of 3 intensities of
blue: FL-5 Neon blue and U-48 Royal blue pots
captured significantly more C. vittatus/C. pikei
than the U-9 Dutch blue traps, (F = 4.68; df =
2,59; P = 0.01) LSD = 1.54 (Table 3).
Experiment 7, 15 cm black pots with white
stripes: Striped traps captured significantly less

Florida Entomologist 85(2)


Trap Color
Size (cm) Black Black Neon Blue Neon Blue

5.0 2.4 + 0.7 B 1.9 + 0.59 B -
7.5 3.5 + 0.7 B -
10.0 4.2 + 0.8 B 4.5 + 0.93 A -
12.5 4.3 + 0.6 B -
15.0 8.7 + 2.3 A 8.0 + 1.4 A 4.3 + 0.94 A 7.1+ 1.2 A
17.5 3.5 + 0.8 B -
20.0 4.0 + 0.5 B 3.7 + 0.68 AB 7.4 + 1.6 A
22.5 -
25.0 3.8 + 0.87 AB 4.9 + 1.2 A

"Means in columns not followed by the same letter are significantly different as determined by LSD.

C. vittatus/C. pikei 1.72 0.22 (total = 34) than
the solid black traps (2.84 0.39, total = 94) (F =
6.33; df = 1,18; P = 0.022) (Table 3). The over-
whelming majority of the deer flies captured on
the striped trap landed on the black areas and
avoided the white.
Experiment 8, repeat of the size test with FL-
5 Neon blue pot traps: Part one indicated a signif-
icant size effect (F = 1.59; df = 4, 45; P = 0.019)
LSD = 2.32 (Table 2). Part two evaluating 15, 20,
and 25 cm traps did not indicate significant size
differences (F = 1.02; df = 2,29; P = 0.37). Trap
captures in relation to FL-5 Neon blue trap size
differed from the results obtained with the black
traps in experiment 4 (Table 2).
Experiment 9, background contrast: A FL-5
Neon blue trap on a white background captured
significantly more C. vittatus/C. pikei (16.7 2.2)
than traps with a FL-5 Neon blue background
(11.6 1.1) which captured significantly more
than the traps with a background of aluminum
foil (6.7 1.0) (F = 10.49; df = 2,42; P = 0.0002)
LSD = 4.44. In the blue trap with blue back-
ground treatment, some deer flies landed on the
background which decreased trap capture.
Experiment 10, trap height. All C. vittatus/
C. pikei deer flies were captured on the traps

placed at 75 and 150 cm above ground. No deer
flies were captured on traps placed higher.
Experiment 11: Blue silhouettes of 60 x 120 cm
had no effect on fly response in any orientation
relative to trap placement.
Experiment 12, response to odors: Response by
C. vittatus/C. pikei to octenol (5.1 + 1.5) was not
significantly different from the unbaited control
(5.2 0.8) (F = 0.01; df = 1,31;P = 0.93). Similarly,
no differences were found in response by C. vitta-
tus/C. pikei in these bioassays to FL-5 Neon blue
traps baited with acetone (11.9 3.0 vs 10.6 +
1.2), CO2 (5.9 + 1.1 vs 5.2 0.9) or DEET (control
-10.0 + 2.8 vs DEET 8.4 3.1).
C. macquarti: Experiment 1, size and color in-
teraction: The two-way analysis of variance indi-
cated that color (F = 14.38; df = 1,66; P = 0.0003)
had a significant effect on C. macquarti response
with FL-5 Neon blue trap collection (11.4 1.1)
greater than black traps (6.5 0.7). Size (10, 15,
22.5 cm) was not significant (F = 1.16, df = 2, 66;
P = 0.32) nor was the color x size interaction (F =
1.81, df= 2, 66;P = 0.17).
Experiment 2, background contrast: Back-
ground color had a significant effect on trap re-
sponse by C. macquarti, (F = 3.55; df = 2,45; P =
0.037) LSD = 0.57 with FL-5 Neon blue (6.81 +


Trap Type (15 cm) Experiment 5 Experiment 6 Experiment 7

Black 3.43 + 0.46 B 2.84 + 0.39 A
Red 3.87 + 0.68 A -
Neon Blue 8.10 + 0.51 B 4.20 + 0.60 A
Royal Blue -3.95 + 0.62 A
Dutch Blue -2.05 + 0.40 B
Black & White Stripes -1.72 + 0.22 B

aMeans in columns not followed by the same letter are significantly different as determined by LSD.

June 2002

Mizell et al.: Trolling: A Novel trap for Chrysops spp.

0.93) and white (6.69 1.46) equal, but greater
than response to traps with an aluminum foil
background (3.63 0.97).


The specimens of C. vittatus/C. pikei and C.
macquarti captured on traps in this study were
almost all females with only a rare male. Parity of
the females was not tested. Occasional samples of
captured flies dissected for gut contents found
about equal numbers with and without clear gut
contents on any date. No dissected deer flies con-
tained evidence of blood feeding.
The truck-mounted experimental apparatus
containing the entire array of traps presented a
moving attraction at a distance (ca. 10-15 m =
maximum distance from the woods, most repli-
cates were closer to vegetation) to responding flies
which then selected a single trap on which to
land. The distance from the woods in these bioas-
says is the distance that Phelps and Vale (1976)
suggested was the maximum from which taban-
ids could detect a moving target. The truck-
mounted trap design is analogous to the cluster
design of Hribar et al. (1991b) and Hribar and
Foil (1994) and provides more precise comparison
of fly treatment preference for activation and
landing cues than other designs that offer single
trap replicates in separate locations. The trolling
trap does not fully measure fly attraction, as at-
traction could occur without landing and capture.
The flight orientation pattern and degree of
aggression exhibited by Chrysops spp. to the
traps during trolling was variable, and while the
causes were not measured in these experiments,
these behaviors were most likely related to the
time since emergence, feeding history and the
prevailing weather conditions. During periods of
relatively warm temperatures, deer flies at the
time of the experiments predominantly flew
straight into the trap from any direction attack-
ing the trap in a manner similar to Vespidae de-
fensive behavior. Some deer flies oriented from
behind the trap and followed it for a short period
of time before landing. This behavioral response
could be induced by increasing the vehicle speed.
Other orientation behavior included circling
around several traps and then landing. Often cap-
tured deer flies tended to be in clusters on the
traps which may have been the result of short
range visual orientation to other flies that had
been captured previously. However, an experi-
ment placing either live or dead tabanids of dif-
ferent sizes and species on the traps in different
places as bait prior to trolling (data not shown)
had no effect on deer fly landing patterns. Gener-
ally, the first arriving deer flies landed on the trap
positions at random, but tended to land on the top
portions first and filled in other areas as avail-

Movement of the traps was necessary to elicit
deer fly landing. Both quality and quantity of trap
movement were important. The method of trap
suspension on the metal rods allowed the traps to
rotate and to shake back and forth during troll-
ing. Trolling on the moving truck provided angu-
lar displacement and velocity. A crude
determination indicated that a speed of ca. 3.16
m/sec (7 mi/h) was the maximum vehicle velocity
at which flying C. vittatus/C. pikei could land on
the traps. This figure is within the range of the
flight speed reported for tsetse flies (2.5-5m/s)
(Gibson and Brady 1985). Deer flies, as do many
hematophagous Diptera (Galun 1977), will follow
a moving vehicle and will continue to circle it for
a brief period after it stops moving. Occasionally,
deer flies would land on traps after the truck be-
came stationary. Manually rotating or shaking
the traps after stopping the truck had little effect
on deer fly landing on traps. Hanging traps such
as blue balloons or pots swaying in the wind in
the same area (as the stationary truck) also only
occasionally induced deer flies to land.
No significant response by the Chrysops spp. to
any behavioral chemicals tested (acetone, octenol,
CO2, DEET) were observed in these experiments.
Trap movement, color and size either singularly
or in combination were perhaps such strong stim-
uli to the Chrysops spp. tested that odorant ef-
fects were not detectable. For example, we were
only able to detect the effect of trap size by using
black traps that are less preferred than blue. Al-
ternatively, because the traps measure landing
behavior, the effects of attractants in our design
perhaps were undetectable. Other species of ta-
banids including large horse fly species were reg-
ularly observed flying near the traps or the truck.
However, very few were captured which may indi-
cate that the trolling traps provided some of the
primary attractive stimuli, (i.e., movement and
color), but not the secondary or final landing cues
(Gibson and Torr 1999). Nevertheless, the repel-
lent DEET must ultimately affect fly landing be-
havior which was measurable in these bioassays.
No repellent affect of DEET was observed. More-
over, true attractants if operative would in part
serve to activate and draw flies from a greater dis-
tance to the trap vicinity while the landing behav-
iors would likely remain constant for the test
species. As a result, if attractants were operative,
higher numbers of flies in the test replicates us-
ing attractants (in comparison to other experi-
ments without chemicals) would be expected.
Higher numbers were not observed, however, the
appropriateness of the trolling trap bioassay for
determining the effects of chemicals on deer fly
behavior remains for future research.
Many times during the course of the experi-
ments, C. vittatus C. pikei apparently were elim-
inated from local areas that had been repeatedly
trolled. Lack of deer flies made it necessary to find

Florida Entomologist 85(2)

populations at new locations. During these
searches we trolled the traps through different
landscape habitat types. We found that habitat
characteristics affected deer fly presence and
abundance as shown previously by Dale and Ax-
tell (1976). C. vittatus/C. pikei was found almost
exclusively on the edges of open areas adjoining
woodlands (within 25-30 m of the borders) and in
narrow road corridors through forests. C. mar-
quarti was found further from the forest edge in
more open habitats such as old fields with sparse
tree density. The two species were found together
in road corridors through areas adjacent to habi-
tats described above.
The collective results of these experiments in-
dicate the importance of habitat context and the
hierarchy of stimuli determining the behaviors
used in the orientation to and landing on hosts by
C. vittatus/C. pikei, C. macquarti and presum-
ably related Chrysops spp. Habitat characteris-
tics such as amount of sunlight and vegetation
pattern that may affect micro habitats of perches
where the species were found during trolling were
different and deserve further research enabled by
trolling traps. Height of deer fly flight above
ground as indicated by orientation to traps was
always less than 3.0 m. Roberts (1976b) captured
70% of tabanids in Malaise traps with openings at
less than 1.8 m above ground and Snoddy (1970)
and Schulze et al. (1975) reported similar flight
patterns by other tabanids.
C. vittatus/C. pikei and C. macquarti were
strongly attracted to movement of 3-dimensional
objects and would rarely land on stationary tar-
gets. However, other horse fly species have been
captured effectively using flat panels (Allan and
Stoffalano 1986, Moore et al. 1996). Moving tar-
gets may be more attractive to most tabanids
(Phelps and Vale 1975). Quality of movement is im-
portant to the captured Chrysops spp. such that
angular displacement is required to elicit landing.
Movement of the trap without angular displace-
ment-stationary placement with rotation or
shaking-often attracted deer flies in low num-
bers, but induced landing only in a few cases when
the deer flies had just followed the moving vehicle.
Deer flies preferred the 3-dimensional traps
presumably due to the increased reflectiveness of
the surface as shown by Thorsteinson et al. (1966).
Deer fly response to traps was also strongly af-
fected by color and size and the interaction of color
with size was evident in the results of experiments
1A, 2, 4, 6, and 8 (Table 2). Blue was preferred over
black and red, which were preferred over traps of
clear plexiglass, yellow and white. Trap spectral
reflectance patterns indicated that intensity and
ultraviolet (UV) light may be important determi-
nants of Chrysops spp. behavior (Fig. 2). FL-5
Neon blue captured the highest numbers of deer
flies and was of medium intensity, the most satu-
rated (pure) blue and reflected no UV. Royal blue

had the lowest intensity and also reflected a low
level of UV. Dutch blue reflected a high percentage
of UV and was the most intense (brightest). Color
is primary and size is secondary in the behavioral
hierarchy as indicated by the significant interac-
tion in the behavioral response to size of the less
preferred black traps (Table 2).
Contrast was also an important determinant
of deer fly behavior. A white background provid-
ing the most contrast increased trap catch signif-
icantly. Aluminum foil which provides high
contrast but reflects ca. 80% UV (Allan and Stof-
folano 1986) significantly decreased trap catch for
C. vittatus/C. pikei and C. macquarti and these
results agree with reports by Allan and Stoffolano
(1986) for T nigrovittatus and the results of the
blue series (Exp. 6, Fig. 2) above. The addition of
white stripes to black traps also significantly re-
duced trap collection (95 C. vittatus/C. pikei on
the black traps vs 34 on black and white striped
Although silhouettes have been shown to be im-
portant cues for tabanids (Phelps & Vale 1976) in-
cluding Chrysops spp. in the presence of CO2
(Browne & Bennett 1980), we found no effect when
traps were trolled with silhouettes without CO2.
A Malaise trap run concurrently with these
studies and placed along the edge of a forest with
central swampy areas (the edges and road
through the middle of this ca. 2 ha area were used
for trolling) indicated that the trolling trap did not
catch all of the Chrysops spp. present in our trap/
trolling locations. However, we apparently cap-
tured most Chrysops spp. present that generally
attack humans in the area. Diaclorus ferrugatus
(F.) was not captured by trolling, although this
species also attacks humans usually around the
lower legs. Other species regularly captured in-
cluded C. geminata Wiedemann, C. dorso-vittatus
Hine, and the C. flavidus Wiedemann complex. C.
davisus Walker, Tabanus trimaculatus Palisot de
Beauvois, T pumilus MacQuart, and T fuluulus
Wiedemann were captured occasionally. Approxi-
mately 30 Chrysops spp. have been captured by
using the Malaise trap in Monticello, Florida (Mi-
zell & Roberts 1997-1999, unpublished).
Many humans are allergic to deer fly bites and
often develop painful boil-like areas in response to
tabanid bites. Even in the absence of allergic reac-
tions, deer flies are a nuisance and bites are pain-
ful. Our results provide an inexpensive method to
reduce attacks by certain deer fly species and to
suppress deer fly populations in small areas such
as dooryards. Placing a small blue trap covered
with Tanglefoot such as a plastic drinking cup at-
tached to a hat either worn or carried on a walk-
ing stick, exploits the deer fly's tendency to attack
the highest point on the target animal, and sup-
presses or eliminates most deer fly species that
land on the body. Because deer flies are ambush
predators, placement of traps on a lawnmower, 4-

June 2002

Mizell et al.: Trolling: A Novel trap for Chrysops spp.

wheeler, or other vehicle will enable trapping-out
in small areas such as dooryards for a few days or
so until new deer flies re-colonize the area.
As with all tabanid traps developed to date, the
trolling trap has limitations. The trap catches pre-
dominately female Chrysops spp., but only a few
other tabanids. The trap requires specific move-
ment (angular displacement) in order to attract
deer flies and has the added disadvantage of re-
quiring a sticky substance to trap landing insects.
However, cleanup of Tanglefoot is easy with soaps
containing d-limonene. During this study many
other unidentified species of biting flies were cap-
tured on the trolling trap. As such our trolling
trap offers a novel method to detect and collect
Chrysops spp. and perhaps many other Dipteran
species, and may be useful to investigate a variety
of scientific questions heretofore unattainable. In
addition, the simple, economical methodology has
great potential for demonstration of basic insect
behavior to science students of all ages.


We especially recognize, thank and remember Dr. Ri-
chard Roberts, USDA-ARS, (deceased) for identifying
the Tabanidae and for many enjoyable conversations
about horse fly behavior and research. We thank Pat Mi-
zell, wife and mother, for her kind patience and encour-
agement during the course of these studies. We thank
Stephanie Bloom, Peter Andersen, Richard Roberts,
Frank French, Lane Foil and two anonymous reviewers
for helpful comments on an earlier draft of the manu-
script. Special sentiment and thanks are sent to Dr.
Robert Combs, Mississippi State University, whose jo-
vial humor and wit were often of great amusement and
encouragement to the senior author during graduate
school. This is the University of Florida Agricultural Ex-
periment Station Journal Series No. R-07188.


AND W. GALE. 1992. Mechanical control of greenhead
flies (Diptera: Tabanidae) in a marsh environment.
J. Med. Entomol. 29: 160-164.
ALLAN, S. 1984. Studies on vision and visual attraction
of the salt marsh horse fly, Tabanus nigrovittatus
Macquart. Ph.D. Dissertation. University of Mass.,
Amherst. 178 pp.
ALLAN, S., AND J. G. STOFFALANO, JR 1986. The effects
of hue and intensity on visual attraction of adult Ta-
banus nigrovittatus (Diptera: Tabanidae). J. Med.
Entomol. 23: 83- 91.
ALLAN, S. A., J. F. DAY, AND J. D. EDMAN. 1987. Visual
ecology of biting flies. Ann. Rev. Entomol. 32: 2 97-
Rigid canopy trap for Tabanidae (Diptera). J. GA En-
tomol. Soc. 10: 64-67.
SON. 1962. The orientation behavior of horse flies
and deer flies (Tabanidae: Diptera) II. The role of
some visual factors in the attractiveness of decoy sil-
houettes. Canadian J. Zool. 40: 685-695.

BROWNE, S. M., AND G. F. BENNETT. 1980. Color and
shape as mediators of host-seeking responses of Sim-
uliids and Tabanids (Diptera) in the Tantramar
marshes, New Brunswick, Canada. J. Med. Entomol.
17: 58-62.
CATTS, E. P. 1970. A canopy trap for collecting Taban-
idae. Mosq. News. 30: 472-474.
CILEK, J. E., AND E. T. SCHREIBER. 1996. Diel host-seek-
ing behavior of Chrysops celatus (Diptera: Taban-
idae) in northwest Florida. Florida Entomol. 79: 520-
DALE, W. E., AND R. C. AXTELL. 1976. Salt marsh Ta-
banidae (Diptera): comparisons of abundance and
distribution in Spartina and Juncus habitats. J.
Med. Entomol. 12: 671- 678.
FOIL, L. D., AND L. J. HRIBAR 1995. Evaluation of tsetse
attractants as baits for horse flies and deer flies
(Diptera: Tabanidae) in Louisiana. Florida Entomol.
78: 129-133.
FRENCH, F. E., AND D. L. KLINE. 1989. 1-Octen-3-ol, an
effective attractant for Tabanidae (Diptera). J. Med.
Entomol. 26: 459-461.
FRENCH, F. E., AND D. V. HAGAN. 1995. Two-tiered box
trap catches Chrysops atlanticus and C. fulginosus
(Diptera: Tabanidae) near a Georgia salt marsh. J.
Med. Entomol. 32: 197-200.
GALUN, R. 1977. Responses of blood-sucking arthropods
to vertebrate hosts, pp. 103-115. In H. H. Shorey and
J. J. McKelvey [eds.]. Chemical control of insect be-
havior, theory and application. John Wiley and Sons.
N.Y. 414 pp.
GIBSON, G., AND J. BRADY. 1985. 'Anemotactic' flight
paths of tsetse flies in relation to host odors: a pre-
liminary video study in nature to host odor. Physiol.
Entomol. 10: 395-406.
GIBSON, G., AND S. J. TORR. 1999. Visual and olfactory
responses of haematophagous Diptera to host stim-
uli. Med. and Vet. Entomol. 13: 2-23.
GRANGER, C. A. 1970. Trap design and colors as factors
in trapping salt marsh greenhead fly. J. Econ. Ento-
mol. 63: 1670-1672.
HANSENS, E. J. 1947. Greenhead flies (Tabanus nigro-
vittatus) like dark colors. New Jersey Agr. 29: 3-4.
1971. Use of traps for study and control of saltmarsh
greenhead flies. J. Econ. Entomol. 64: 1.
HRIBAR, L. J., AND L. D. FOIL. 1994. Color and UV reflec-
tance of canopy traps for collecting horse flies
(Diptera: Tabanidae) in Louisiana. Bull. Soc. Vector
Ecol. 19:49-52.
Design for a canopy trap for collecting horse flies
(Diptera: Tabanidae). J. Am. Mosq. Contr. Assoc. 7:
HRIBAR, L. J., D. J. LEPRINCE, AND L. D. FOIL. 1991b. In-
creasing horse fly (Diptera: Tabanidae) catch in can-
opy traps by reducing Ultraviolet light reflectance. J.
Med. Entomol. 28: 974-877.
Two-story French box traps for Tabanids (Insect:
Diptera). J. GA. Science. 51: 24-25.
JONES, C. M. 1953. Biology of Tabanidae in Florida. J.
Econ. Entomol. 46:1108-1109.
NEWTON. 1996. Effect of trap design and color in
evaluating activity of Tabanus abactor Philip in
Texas rolling plains habitats. Southwest. Entomol.
21: 1-11.

Florida Entomologist 85(2)

PHELPS, R. J., AND G. A. VALE. 1976. Studies on the local
distribution and on the methods of host location of
some Rhodesian Tabanidae (Diptera). J. Entomol.
Soc. Southern Africa. 39: 67-81.
ROBERTS, R. H. 197 la. The seasonal appearance of Ta-
banidae as determined by Malaise trap collections.
Mosq. News. 31: 509-512.
ROBERTS, R. H. 1971b. Effect of amount of CO2 on collec-
tion of Tabanidae in Malaise traps. Mosq. News. 31:
ROBERTS, R. H. 1976a. The comparative efficiency of six
trap types for the collection of Tabanidae (Diptera).
Mosq. News 36: 530-535.
ROBERTS, R. H. 1976b. Altitude distribution of Taban-
idae as determined by Malaise trap collections.
Mosq. News. 36: 518-520.
SAS INSTITUTE. 1996. SAS User's Guide v. 6.12. SAS In-
stitute, Cary, NC.
Some environmental factors affecting the daily and
seasonal movements of the salt marsh greenhead, Ta-
banus nigrovittatus. Environ. Entomol. 4: 965-971.
The Malaise trap-a survey tool in medical entomol-
ogy. Mosq. News. 25: 398-400.
SNODDY, E. L. 1970. Trapping deer flies with colored
weather balloons (Diptera: Tabanidae). J. Ga. Ento-
mol. Soc. 5: 207-209.
A comparison of Malaise trapping and aerial netting

for sampling a horsefly and deer fly community. En-
viron. Entomol. 5: 788-792.
TEDDERS, W. L., AND B. W. WOOD. 1994. A new tech-
nique for monitoring pecan weevil emergence. J. En-
tomol. Sci. 29: 18-30.
THORSTEINSON. A. J. 1958. The orientation of horse flies
and deer flies (Diptera: Tabanidae). I. The attraction of
heat to Tabanids. Entomol. Expt. et Appl. 1:1 91-196.
1965. The orientation behavior of horse flies and
deer flies (Tabanidae: Diptera). III. The use of traps
in the study of orientation of Tabanids in the field.
Entomol. Exp. et Appl. 8: 189-192.
WARYK. 1966. The orientation behavior of horse flies
and deer flies (Tabanidae: Diptera) V. The influence
of the number and inclination of reflecting surfaces
on attractiveness to Tabanids of glossy black polyhe-
dra. Can. J. Zool. 44: 275-279.
WALL, W. J., AND 0. W. DOANE, JR 1980. Large scale
use of box traps to study and control saltmarsh
greenhead flies (Diptera: Tabanidae) on Cape Cod,
Massachusetts. Environ. Entomol. 9: 371-375.
WILSON, B. H. 1968. Reduction of Tabanid populations
on cattle with sticky traps baited with dry ice. J.
Econ. Entomol. 61: 827-829.
Attraction of Tabanids to traps baited with dry ice
under field conditions in Louisiana. J. Med. Entomol.
3: 149-149.

June 2002

Scientific Notes


USDA-ARS, U.S. Horticultural Research Laboratory, 2001 South Rock Road, Ft. Pierce, FL 34945

The population dynamics of the sweetpotato
whitefly, Bemisia tabaci (Gennadius) biotype B
(= silverleaf whitefly, Bemisia argentifolii Bel-
lows & Perring) are driven by seasonal climatic
conditions, natural enemies (including entomo-
pathogens), host-plant interactions, and IPM
practices (Byrne & Bellows 1991; Coudriet et al.
1985; Nava-Cameros et al. 2001; Tsai & Wang
1996; Yee and Toscano 1996). Physiological disor-
ders (Schuster et al. 1990, 1991) and the spread of
geminiviruses (Blair et al. 1995; Polston & Ander-
son 1997) associated with whitefly infestations in
Florida vegetables are becoming increasingly im-
portant limitations to grower profitability. The ef-
fect of plant viruses on the reproductive potential
of the vector is key to understanding geminivirus
epidemiology and developing effective control
measures. The objective of this study was to de-
termine the effect of tomato mottle virus (ToMoV)
on whitefly oviposition and survival rates on
healthy tomato.
Adult B. tabaci biotype B were obtained from
laboratory colonies maintained by the U.S. Horti-
cultural Research Laboratory, Ft. Pierce, FL.
Whiteflies used in these experiments were origi-
nally obtained from Dr. Lance Osborne, Univer-
sity of Florida, Apopka, FL and have been
maintained on dwarf cherry tomato (Lycopersicon
esculentum cv. Florida Lanai) since 1996 by serial
transfer. In 1997, a ToMoV whitefly colony was es-
tablished by obtaining tomato plants infected
with ToMoV from Dr. Philip Stansly, University of
Florida, Immokalee, FL and infesting with white-
flies from the healthy colony. Whitefly biotyping
was based on RAPD PCR analysis using primers
developed by De Barro and Driver (1997). Nonvir-
uliferous and viruliferous whitefly colonies were
housed separately in screened Plexiglass cages lo-
cated in separate growth chambers at 25 + 1C
and a 16:8 L:D photoperiod. Whiteflies from the
viruliferous colony were confirmed to be infected
with ToMoV prior to infestation by PCR analysis
(Sinisterra et al., 1999).
In each experiment, one male and one female
whitefly of unknown age from healthy or ToMoV-
infected whitefly colonies were confined in clip
cages attached to the terminal leaf of the 3rd fully
expanded leaflet of a healthy cv. Florida Lanai
plant. Clip cages were made from clear plastic
cups (PC100 30 ml cups, Jet Plastica, Harrisburg,
PA) fitted with a foam seal on the bottom and an
organdy window on top for ventilation. The foam
bottom was backed with a thin square of balsa

wood, and an aluminum hair clip was glued to the
balsa wood and cup portion. Whiteflies were intro-
duced through a small hole in the side of the cup.
After a 48-h access period, adult whiteflies were
removed and eggs were counted. Treatments were
maintained separately at 25 + 1C and a photo-
period of 16:8 L:D. For each experiment, 20 test
plants were typically used for each treatment;
however, the final number of replicates (= clip
cages) per treatment varied when leaves of test
plants died or were severed during the experi-
ment. A minimum of 8 replicates was used for all
treatments. Experiments were repeated five
times. There were no significant interactions be-
tween experiment*treatment (F = 2.0; df = 4,80;
P = 0.10) or treatment*clip cage (F = 0.66; df =
19,80; P = 0.85) so results were pooled over exper-
iments for mean comparison (Tukey option in
SAS GLM procedure, SAS Institute 1998). In the
last two experiments, progeny from cohorts used
in the oviposition clip cage experiments (n = 15)
were used to include survival to adult emergence
which was evaluated 30 days after egg lay to en-
sure that all viable whiteflies had emerged.
Whiteflies infected with ToMoV deposited sig-
nificantly more eggs (F = 19.51; df = 1,80;P < 0.01)
on healthy tomato leaves than nonviruliferous
whiteflies (Table 1) and supports earlier findings
(McKenzie et al. 2002). There was no significant
difference between virus-infected and nonvirulif-
erous whiteflies for the number of adults emerged
or the proportion of those adults surviving from
the egg stage. There was no significant correlation
between the number of eggs deposited per female
and progeny survival rates on healthy tomato for
whitefly infected with or without the virus (SAS
CORR procedure, SAS Institute 1998).
A report by Costa et al. (1991) demonstrated
that whitefly maintained on pumpkin for ~ 6
years had a higher rate of survival on virus-in-
fected pumpkin compared to healthy pumpkin
out of six virus-plant hosts evaluated, including
tomato. In those experiments, researchers did not
take host-plant adaptation time into consider-
ation. For example, whitefly survival rates from
egg to adult on tomato were 8% on virus-infected
plants and 17% on noninoculated control plants,
but the parental generation of whiteflies used in
those experiments were maintained on pumpkin.
In our experiments, high survival (Table 1) of
both healthy and ToMoV-infected whitefly reflect
this host-plant adaptation when compared to the
previous work by Costa et al. (1991). Plants from

Florida Entomologist 85(2)

June 2002


WF colony Egg SE' Adult + SE2 % Survival + SE3

Healthy 8.80 + 0.70 a 7.28 + 1.90 a 70.56 + 14.11 a
ToMoV 14.69 + 1.11 b 11.09 + 2.16 a 77.44 + 6.37 a

Means within a column followed by the same letter are not significantly different, P < 0.05, using Tukey studentized range test.
'F = 19.51; df = 1,80; P = < 0.01.
'F = 3.16; df= 1,11; P = 0.10.
'Proportion of whitefly surviving from egg to adult stage; F = 0.44; df = 1,8; P = 0.53.

the virus treatment exhibited characteristic To-
MoV symptoms 30 days after clip cages were re-
moved. This suggests adaptation to the host plant
and virus by the vector could override any ad-
verse effect the virus had on host plant physiol-
ogy. McKenzie et al. (2002) found healthy plants
infested with ToMoV-infected whiteflies consis-
tently had 2.5-fold more eggs and 4.5-fold more
nymphs than plants with nonviruliferous white-
flies (P < 0.05) 56 days after infestation with the
same number of whitefly. In the present study,
whiteflies were well adapted to the host plant, ei-
ther with or without ToMoV.


Whiteflies carrying ToMoV deposited signifi-
cantly more eggs than nonviruliferous whiteflies
when provided a healthy tomato host. Insect ad-
aptation to the host-plant is a critical factor that
should be considered on a host-by-host basis
when evaluating insect biology and vector-host-
plant interactions for polyphagous insect species.
Mention of a trademark or proprietary product
does not constitute a guarantee or warranty of the
product by the U.S. Department of Agriculture
and does not imply its approval to the exclusion of
other products that may also be suitable.


AND M. LAMBERTS. 1995. Occurrence of bean golden
mosaic virus in Florida. Plant Disease 79: 529-533.
BYRNE, D. N., AND T. S. BELLOWS. 1991. Whitefly biol-
ogy. Ann. Rev. Entomol. 36:431-457.
COSTA, H. S., J. K. BROWN, AND D. N. BYRNE. 1991. Life
history traits of the whitefly, Bemisia tabaci (Homop-
tera: Aleyrodidae) on six virus-infected or healthy
plant species. Environ. Entomol. 20: 1102-1107.

D. E. MEYERDIRK. 1985. Variation in developmental
rate on different hosts and overwintering of the
sweetpotato whitefly, Bemisia tabaci (Homoptera:
Aleyrodidae). Environ. Entomol. 14: 516-519.
2000. Effect of leaf age and silverleaf symptoms on
oviposition site selection and development of Bemi-
sia argentifolii (Homoptera: Aleyrodidae) on zuc-
chini. Environ. Entomol. 29:220-225.
DE BARRO, P. J., AND F. DRIVER 1997. Use of RAPD
PCR to distinguish the B biotype from other biotypes
of Bemisia tabaci (Gennadius) (Hemiptera: Aley-
rodidae). Aust. J. Entomol. 36: 149-152.
D. LEE, M. INBAR, AND R. T. MAYER (2002). Effect of
geminivirus infection and Bemisia infestation on ac-
cumulation of pathogenesis-related proteins in to-
mato. Arch Insect Biochem and Physiol. 49: 203-214.
POLSTON, J. E., AND P. K. ANDERSON. 1997. The emer-
gence of whitefly-transmitted geminiviruses in to-
mato in the Western Hemisphere. Plant Disease 81:
SAS INSTITUTE, INC. 1998. SAS/STAT@ User's guide
version 7. Cary, NC, USA: SAS Institute, Inc.
PRICE. 1990. Relationship of the sweetpotato white-
fly to a new tomato fruit disorder in Florida. HortSci.
25: 1618-1620.
SCHUSTER, D. J., J. B. KRING, AND J. F. PRICE. 1991. As-
sociation of the sweetpotato whitefly with a silver-
leaf disorder of squash. HortSci. 26: 155-156.
E. HIEBERT. 1999. Tobacco plants transformed with
a modified coat protein of tomato mottle begomovi-
rus show resistance to virus infection. Phytopath.
89: 701-706.
TSAI, J. H., AND K. WANG. 1996. Development and re-
production of Bemisia argentifolii (Homoptera: Aley-
rodidae) on five host plants. Environ. Entomol. 25:
YEE, W. L., AND N. C. TOSCANO. 1996. Ovipositional
preference and development of Bemisia argentifolii
(Homoptera: Aleyrodidae) in relation to alfalfa. J.
Econ. Entomol. 89: 870-876.

Scientific Notes


1Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611-0620

2Nutkin's Nest Wildlife Rehabilitation Center, 14107 NW 61st Lane, Gainesville, FL 32653-2570

Cuterebra bot fly larvae are obligate, subcuta-
neous parasites of rodents (mice, rats, tree squir-
rels, chipmunks, etc.) and lagomorphs (rabbits
and hares) in the Americas (Sabrosky 1986).
Infestation is revealed by lumps (referred to as
warbles) on an animal caused by the presence of
second and third instar larvae under its skin
(Cogley 1991, Slansky & Kenyon 2000, 2001). Ev-
idence from a limited number of studies indicates
that Cuterebra flies do not oviposit directly on
their hosts but rather on foliage, twigs or other
substrates, often in the vicinity of an animal's
nest (Catts 1967, Baird 1974, 1997). A potential
host thus risks exposure to newly hatched Cutere-
bra larvae when leaving its nest and moving
about in its habitat (Catts 1982).
Once the tiny, infective-stage larvae transfer
to a potential host, they enter an orifice or wound
and begin their approximately week-long journey
through the animal's body, eventually settling un-
der its skin (Gingrich 1981). They then use their
pointed mouth hooks to cut through the host's
hide to create a warble pore through which they
respire and excrete fluid. A larva typically re-
mains in its warble, which increases in size as the
larva grows, until completing development in
about 3 to 10 weeks, depending on the species of
Cuterebra and host (Baird 1975, Jacobson et al.
1978). The mature larva exits through the warble
pore and falls to the ground, where it burrows
into the soil to pupate (Catts 1982).
Over the past several years, we have been
studying the relationship between Cuterebra
emasculator Fitch and its tree squirrel hosts. This
bot fly parasitizes eastern gray (Sciurus carolin-
ensis Gmelin) and fox (S. niger L.) squirrels, as
well as eastern chipmunks (Tamias striatus (L.)),
throughout the eastern and midwestern regions
of North America, from southern Canada to Flor-
ida (Sabrosky 1986); American red (Tamiasciurus
hudsonicus (Erxleben)) and flying Gi..........
spp.) squirrels seem to be rarely infested (Dorney
1965, Slansky & Kenyon 2000). Cuterebra emas-
culator is apparently univoltine throughout its
range, spending some ten months underground in
the pupal stage (Bennett 1972a). Infested ani-
mals are observed from mid-late July to the end of
October (Bennett 1955, Jacobson et al. 1981,
Slansky & Kenyon 2000). While there are many
published records of C. emasculator infesting tree
squirrels and chipmunks active outside the nest,
to our knowledge this note is the first documenta-

tion of infestation of nest-bound infants by larvae
of this bot fly.
As part of a wildlife rehabilitation program in
northcentral (Alachua Co.) Florida, over the last
18 years we have encountered 10-15 cases of infant
S. carolinensis infested by bot fly larvae (out of sev-
eral hundred infants brought in for rehabilitation),
including a record high of five in the summer, 2000
squirrel breeding season (no cases were reported
in 2001). This rarity contrasts with the situation in
adult squirrels, where the incidence of Cuterebra
parasitization often exceeds 50% (unpubl. obs.). In-
festations of the infants typically consisted of one
or two, and rarely three, larvae per individual (Fig.
1). All of these animals would have been nest-
bound at the time of parasitization; most were
young enough to have not yet opened their eyes
(i.e., they were less than five weeks of age) and
even those with opened eyes were too young to
have begun exploring outside the nest. Juvenile
S. carolinensis typically do not begin extra-nest ac-
tivity before about eight weeks of age (unpubl. obs.).
In that exposure to Cuterebra larvae is be-
lieved to occur when an animal moves about out-
side of its nest, infestation of nest-bound infants
seems highly unusual. We are aware of only a few
anecdotal reports of Cuterebra-infested infant an-
imals (e.g., a 6-day-old rabbit and a litter of nurs-
ing wood rats; Dalmat 1943). Kittens and
puppies, both of which are incidental hosts, can
also be parasitized (Hall 1925, Rosenthal 1975,
McKenzie et al. 1978). How infestation of nest-
bound animals occurs has not been determined.
Assuming that female Cuterebra did not deposit
eggs directly on these infants, it is possible that a
mother animal that had acquired infective-stage
larvae while out foraging inadvertently brought
these back to her nest and some transferred to
her nursing offspring. Another possibility is that
larvae from eggs laid on nest material entered the
nest and parasitized the animals within. It is not
known where C. emasculator females oviposit.
Parasitism by Cuterebra typically has been
studied in juvenile, subadult and adult animals
(Bennett 1955, 1972b, 1973, Jacobson et al. 1981,
Manrique-Saide et al. 2000). Individuals in these
age classes typically exit their nests on a daily ba-
sis to forage in the habitat and thus risk exposure
to infective-stage larvae; it is also these individu-
als that are sampled through the commonly used
techniques of trapping, hunting and collection of
roadkills. However, as documented here, seden-

Florida Entomologist 85(2)

Fig. 1. Upper panel: Nest-bound infant gray squirrel (4-5 weeks old, eyes not yet opened) with a moderate-sized
bot fly warble on its side behind its right front leg; the warble contained a second instar larva. Lower left panel: Sec-
ond instar C. emasculator larva removed from its warble; bands of dark spines are visible ('head' end at left; length
= 10.0 mm; original photomicrograph, 25x). Lower right panel: Ventral view of anterior portion of second instar bot
fly larva showing paired mouth hooks (toward top of image) and bands of dark, posterior-pointing spines (the inter-
nal cephaloskeleton to which the mouth hooks are attached is faintly visible through the translucent cuticle; orig-
inal photomicrograph, 60x).

tary, nest-bound infants may also become in-
fested. Therefore, when evaluating the effects of
parasites on their host populations (Grenfell &
Gulland 1995, Milton 1996), it would seem to be
important to study not just animals moving about
in their habitat, but nest-bound individuals as
well, even though locating nests to sample the in-
fants will typically be more difficult in a field set-
ting than trapping mobile animals. Associated

with their small size and the high energy and
nutrient demands of their rapid growth, it is
likely that infants will be deleteriously affected
by parasites to a greater extent than adults with
a comparable intensity of infestation. Thus, when
only older, mobile animals are studied, the impact
of parasites on the host population could be sig-
nificantly underestimated, especially if the inci-
dence of infant infestation is substantial.

June 2002

Scientific Notes

We thank Lance Durden, Heidi Bissell, and
Shelley L. Miller for reviewing this manuscript,
and three anonymous reviewers for their helpful
comments. This research was supported by the
Florida Agricultural Experiment Station, and ap-
proved for publication as Journal Series No. R-


Although infestation of tree squirrels and chip-
munks active outside the nest (i.e., juveniles, sub-
adults and adults) by larvae of the bot fly Cuterebra
emasculator has been well documented, this is ap-
parently the first report of nest-bound infant
eastern gray squirrels (Sciurus carolinensis) par-
asitized by this species.


BAIRD, C. R. 1974. Field behavior and seasonal activity
of the rodent bot fly, Cuterebra tenebrosa, in central
Washington (Diptera: Cuterebridae). Great Basin
Naturalist 34: 247-253.
BAIRD, C. R. 1975. Larval development of the rodent bot
fly, Cuterebra tenebrosa, in bushy-tailed wood rats
and its relationship to pupal diapause. Canadian J.
Zool. 53: 1788-1798.
BAIRD, C. R. 1997. Bionomics of Cuterebra austeni
(Diptera: Cuterebridae) and its association with
Neotoma albigula (Rodentia: Cricetidae) in the south-
western United States. J. Med. Entomol. 34: 690-695.
BENNETT, G. F. 1955. Studies on Cuterebra emasculator
Fitch 1856 (Diptera: Cuterebridae) and a discussion
of the status of the genus Cephenemyia Ltr. 1818.
Canadian J. Zool. 33: 75-98.
BENNETT, G. F. 1972a. Observations on the pupal and
adult stages of Cuterebra emasculator Fitch (Diptera:
Cuterebridae). Canadian J. Zool. 50: 1367-1372.
BENNETT, G. F. 1972b. Further studies on the chipmunk
warble, Cuterebra emasculator (Diptera: Cuterebri-
dae). Canadian J. Zool. 50: 861-864.
BENNETT, G. F. 1973. Some effects of Cuterebra emascu-
lator Fitch (Diptera: Cuterebridae) on the blood and
activity of its host, the eastern chipmunk. J. Wildlife
Diseases 9: 85-93.
CATTS, E. P. 1967. Biology of a California rodent bot fly
Cuterebra latifrons Coquillett. J. Med. Entomol. 4:
CATTS, E. P. 1982. Biology of New World bot flies:
Cuterebridae. Annu. Rev. Entomol. 27: 313-338.

COGLEY, T. P. 1991. Warble development by the rodent
bot Cuterebra fontinella (Diptera: Cuterebridae) in
the deer mouse. Veterinary Parasitol. 38: 276-288.
DALMAT, H. T. 1943. A contribution to the knowledge of
the rodent warble flies (Cuterebridae). J. Parasitol.
29: 311-318.
DORNEY, R. S. 1965. Incidence of botfly larvae (Cutere-
bra emasculator) in the chipmunk (Tamias striatus)
and red squirrel (Tamiasciurus hudsonicus) in
northern Wisconsin. J. Parasitol. 51: 893-894.
GINGRICH, R. E. 1981. Migratory kinetics of Cuterebra
fontinella (Diptera: Cuterebridae) in the white-
footed mouse, Peromyscus leucopus. J. Parasitol. 67:
GRENFELL, B. T., AND F. M. D. GULLAND. 1995. Intro-
duction: Ecological impact of parasitism on wildlife
host populations. Parasitology 111: s3-s14.
HALL, M. C. 1925. The occurrence of cuterebrid larvae in
dogs and cats, and the possible modes of infection. J.
Econ. Entomol. 18: 331-335.
1978. Bot fly myiasis of the cottontail rabbit, Sylvil-
agus floridanus mallurus in Virginia with some biol-
ogy of the parasite, Cuterebra buccata. J. Wildlife
Diseases 14: 56-66.
1981. Prevalence of Cuterebra emasculator in squir-
rels in Mississippi. J. Wildlife Diseases 17: 79-87.
M. T. QUINTERO. 2000. First record of Cuterebra sp.
(Diptera: Cuterebridae) infection in Ototylomys
phyllotis (Rodentia: Muridae). Florida Entomol. 83:
1978. Intracerebral migration of Cuterebra larva in a
kitten. JAVMA 172: 173-175.
MILTON, K. 1996. Effects of bot fly (Alouattamyia baeri)
parasitism on a free-ranging howler monkey (Alou-
atta palliata) population in Panama. J. Zool., London
239: 39-63.
ROSENTHAL, J. J. 1975. Cuterebra infestation of the con-
junctiva in a puppy. Veterinary Medicine/Small Ani-
mal Clinician 70: 462-463.
SABROSKY, C. W. 1986. North American Species of Cute-
rebra, the Rabbit and Rodent Bot Flies (Diptera:
Cuterebridae). Entomol. Soc. Amer. Thomas Say
Foundation Monograph, College Park, MD.
SLANSKY, F., AND L. R. KENYON. 2000. Lumpy squir-
rels-bugged by bot flies. Wildlife Rehab. Today
11(3): 24-31.
SLANSKY, F., AND L. R. KENYON. 2001. Warbles of the
tree squirrel bot fly. http://botfly.ifas.ufl.edu/cutrwrb/

Florida Entomologist 85(2)


1University of Florida, IFAS, Mid-Florida Research and Education Center
2725 Binion Road, Apopka, FL 32703-8504

2University of Florida, IFAS, Department of Fisheries and Aquatic Sciences
7922 NW 71st Street Gainesville, FL 32653-3071

Large swarms of non-biting midges (Diptera:
Chironomidae) emanating from some central
Florida lakes can cause severe nuisance and eco-
nomic problems for businesses, residents, and vis-
itors within the dispersal range of these insects
(Ali 1995). Midges are also a cause of allergies to
humans (Cranston 1995). Because of these prob-
lems, a systematic research program on the bio-
nomics and management possibilities of midge
populations in central Florida has continued for
the past two decades (Ali 1996). As a part of this
program, a preliminary investigation of fish pre-
dation on chironomid midge larvae for the biolog-
ical control perspective of midges was conducted.
Midge predatory fish (bluegill,Lepomis macrochi-
rus) were collected from two lakes on four occa-
sions to elucidate any relationships between
consumption of midge larvae by these fish and the
associated larval composition and distributions in
the lakes. Information concerning fish predation
on midge larvae, species or habitat specific, would
be useful in devising new control strategies.
Fish were collected (May 1999, July, Septem-
ber and December 2000) from Lakes Dora and
Yale (Lake County, Florida) by electrofishing un-
der permit from Florida Fish and Wildlife Conser-
vation Commission. Collections were made between
0830 and 1200 h and up to twenty fish were col-
lected from near-shore areas. Fish were identified
and killed immediately, maintained on ice while
transported to the laboratory, and stored at -10C
until examined. For examination, each fish was
thawed and the foregut was dissected (Bowen
1996), and the contents transferred to 4-dram
vials containing 70% ethanol. Gut contents were
examined under variable magnification of a dis-
secting microscope and enumerated. Chironomi-
dae head capsules and associated fragments were
wet mounted on slides and examined at 400x
magnification using a phase-contrast microscope
and identified to lowest possible taxonomic level
using the keys of Epler (1995). Only head cap-
sules with sufficient morphological features re-
maining for identification were counted for gut
content enumeration, other fragments were used
to improve identification where possible.
Five to 20 fish were successfully collected per
sampling occasion (Table 1). Midge larvae of the
tribe Tanytarsini (>90% Cladotanytarsus spp.)

were most numerous in gut contents of fish from
both lakes (Table 1), comprising 55.9-62.8% of
total consumed midge larvae from Lake Dora and
4.8-48.1% from Lake Yale. Geoldichironomus spp.
larvae were the next most common in the gut con-
tents of fish from Lake Dora (0.0-27.5% of total
larval chironomids). Other midge larvae consumed
by fish from Lake Dora included Chironomus
crassicaudatus, Glyptotendipes paripes, Crypto-
chironomus spp., Pseudochironomus spp. and
Tanypodinae. Seasonal mean number of total
midge larvae in fish gut contents ranged from 4.7
to 44.0. Midge larvae were present in the gut con-
tents of all fish from Lake Dora, except those col-
lected in December 2000, when 40% of collected
fish had empty guts. This was likely due to low
water temperatures reducing feeding activity, as
suggested for bluegill during winter months by
Gilinsky (1984). Pseudochironomus spp. larvae
were the second most prevalent midge larvae in
the fish gut contents of Lake Yale, forming up to
46.2% of total midge larvae, followed by C. crassi-
caudatus (collected only during May 1999),
G. paripes, Cryptochironomus spp., and Tanypod-
inae. Seasonal mean number of larvae per fish in
Lake Yale ranged between 1.0 and 18.9. Bluegill
feeding on midge larvae in Lake Yale was also re-
duced during December 2000, though only one
fish had an empty gut. Other food items identified
from fish in these two lakes included immature
Insecta (Odonata, Ephemeroptera and Trichop-
tera), Crustacea (Decapoda, Amphipoda and Ostra-
coda), Nematoda, Oligochaeta, Gastropoda, and
some unidentifiable material. These food items
numerically were only a small part of total gut
contents in most fish examined (data not shown).
To estimate relative selective feeding by blue-
gill on examined chironomid larvae, percent com-
position of chironomid larvae in gut contents of
collected fish was compared to overall percent
composition of chironomid larvae in study lakes
and percent composition of midge larvae in the
nearshore areas with firm sediments representa-
tive of the areas from where the fish were caught,
collected concurrently and reported by Lobinske
(2001) (Fig. 1). In Lake Dora, Tanytarsini were
most common, exhibiting similar percent compo-
sitions in fish gut contents in the entire lake as
well as in nearshore areas. During July, Septem-

June 2002

Scientific Notes


Taxa May 1999 July 2000 September 2000 December 2000

Lake Dora
No. fish collected 10 20 20 20
Chironomus crassicaudatus 0 + 0 (0.0) 0.1 + 0.3 (0.3) 0 + 0 (0.0) 0.8 + 2.6 (16.1)
Glyptotendipes paripes 0.4 + 0.7 (0.1) 1.8 2.5 (6.0) 0.2 0.7 (0.5) 0.3 + 1.1 (5.4)
Cryptochironomus spp. 0 0 (0.0) 2.5 3.4 (8.2) 4.4 + 7.3 (10.4) 0.3 + 1.1 (6.5)
Pseudochironomus spp. 0 0 (0.0) 0 0 (0.0) 0.5 + 1.5 (1.2) 0 + 0 (0.0)
Geoldichironomus spp. 0 0 (0.0) 8.3 + 11.0 (27.5) 10.5 + 13.1 (24.7) 0.7 + 1.7 (14.0)
Tanytarsini 25.1 + 18.2 (57.0) 16.9 + 17.5 (56.3) 26.7 + 22.1 (62.8) 2.6 + 6.3 (55.9)
Tanypodinae 0.7 + 1.3 (1.6) 0.5 + 1.1 (1.5) 0.1 + 0.3 (0.2) 0 + 0 (0.0)
Unidentified/other midges* 17.8 + 7.8 (40.5) 0 0 (0.0) 0.1 + 0.4 (0.2) 0.1 + 0.4 (2.1)

Total Chironomidae 44.0 + 21.0 30.0 + 31.6 42.5 + 34.2 4.7 + 12.0

Lake Yale
No. fish collected 5 7 15 10
Chironomus crassicaudatus 6.2 13.9 (75.6) 0 0 (0.0) 0 0 (0.0) 0 + 0 (0.0)
Glyptotendipes paripes 0.2 0.4 (2.4) 1.7 + 4.1 (9.0) 0 0 (0.0) 0.2 + 0.4 (20.0)
Cryptochironomus spp. 0 0 (0.0) 1.7 + 3.1 (9.0) 0.5 1.3 (18.5) 0.2 + 0.4 (20.0)
Pseudochironomus spp. 0 0 (0.0) 8.6 9.6 (45.5) 0.6 1.6 (22.2) 0.2 + 0.4 (20.0)
Tanytarsini 0.4 0.9 (4.8) 6.0 6.4 (31.7) 1.3 1.3 (48.1) 0.3 + 0.5 (30.0)
Tanypodinae 1.4 + 3.1 (17.1) 0.4 + 1.1 (2.1) 0 + 0 (0.0) 0.1 + 0.3 (10.0)
Unidentified/other midges* 0 + 0 (0.0) 0.5 + 1.1 (2.6) 0.3 + 0.8 (11.1) 0 + 0 (0.0)

Total Chironomidae 8.2 + 13.5 18.9 + 17.2 2.7 + 3.0 1.0 + 0.9

*Larvae could not be identified due to damage to head capsules.

ber and December 2000, Geoldichironomus spp.
larvae in gut contents represented a much larger
percentage of total chironomids when compared
to their percentage composition in the entire lake
and nearshore populations. In Lake Dora, no sig-
nificant differences (t-test) were noted between
percent composition of individual midge taxa in
gut contents and the prevailing populations in
the nearshore or entire lake area. In Lake Yale,
C. crassicaudatus was the most common chirono-
mid in gut contents during May 1999, but was
only a minor component (<20%) of overall midge
community in the lake during that time. During
July 2000, Pseudochironomus spp. and Tanytar-
sini comprised a similar percentage of nearshore
chironomids and in the gut contents of collected
fish. In the remaining periods of the study in Lake
Yale, Tanytarsini were the most common midge
larvae in fish gut contents, as well as in the firm
sediments. Glyptotendipes paripes was the most
common midge in Lake Yale during July and Sep-
tember 2000 and composed a large proportion of
total midge larvae during December 2000, but
comprised a smaller component of the gut con-
tents. This was probably because G. paripes lar-
vae were aggregated in deeper areas of the lake

(Lobinske 2001) and thus were not in the immedi-
ate grazing area of the collected fish. Mean per-
cent composition of G. paripes larvae in all gut
contents and their overall percent composition in
lake total larval population was significantly dif-
ferent (t = 2.938, P = 0.026, n = 4), but there was
no significant difference noted with percent com-
position in firm sediment areas (t = 0.440, P =
0.676, n = 4). No significant differences were
noted among any other chironomid taxa collected
from Lake Yale.
Based on the data, bluegill (L. macrochirus)
seem to be indiscriminate feeders on midge larvae
inhabiting nearshore areas of Lakes Dora and
Yale. However, Gilinsky (1984), Rieradevall et al.
(1995), and Wolfram-Wais et al. (1999) reported
selectivity of fish predation on chironomids. The
significant difference between G. paripes in Lake
Yale benthos and fish gut contents indicates blue-
gill collected during this study were possibly fo-
cusing their feeding efforts in sandy, nearshore
areas, a behavior similar to the preference of
midge predatory fish foraging in restored, sand
bottom areas of Lake Tohopekaliga reported by
Butler et al. (1992). Rieradevall et al. (1995), But-
ler et al. (1992) and Gilinsky (1984) reported that

Florida Entomologist 85(2)

Lake Dora
80 Fish gut contents May 1999
60 Larval composition in firm sediments n 10

40 Larval composition in entire lake
S 60- July 2000
0 Mn 20
- 40 -I
o 20

g 60 September 200
40 |

60 December 2000
40 -n= 20

Cc Gp Cryp Pseu Geol Tant Tanp Oth

Lake Yale

July 2000

1__tIl __L M _J_

Cc Gp Cryp Pseu Geol Tant Tanp Oth

Taxon Collected
Fig. 1. Composite percent composition of total midge larvae of various midge taxa in bluegill (Lepomis macro-
chirus) gut contents and prevailing field populations in Lakes Dora and Yale (Lake County, Florida) May 1999-De-
cember 2000. Chironomus crassicaudatus (Cc), Glyptotendipes paripes (Gp), Cryptochironomus (Cryp),
Pseudochironomus (Pseu), Geoldichironomus (Geol), Tanytarsini (Tant), Tanypodinae (Tanp), and unidentified/
other chironomid taxa (Oth).

fish predation significantly lowered chironomid
larval density in various studied habitats. How-
ever, Batzer et al. (2000) reported that fish preda-
tion in a marsh weedbed did not significantly
reduce chironomid densities when compared to
fish-excluding enclosures because of increased
predation by other macroinvertebrates. Rasmus-
sen (1990) reported midge larvae were a major
part of whitefish diet in Hjarbaek Fjord, Den-
mark, but whitefish only consumed 2-5% of total
chironomid productivity. Further work is needed
to refine these preliminary results, enumerate ac-
tual standing crop of bluegill in Florida lakes to
estimate the total consumption of midge larvae in
relation to the chironomid standing crop and to
determine if bluegill are a major source of midge
Gratitude is expressed to the Florida Fish and
Wildlife Conservation Commission for necessary
permits. This is Florida Agricultural Experiment
Station Journal Series No. R-08369.


Compositions of midge larvae consumed by the
predaceous fish, bluegill (Lepomis macrochirus)
were examined in relation to associated standing
crop of midge larvae in two central Florida lakes.
Overall, these preliminary results indicate that
bluegill may be indiscriminate predators on
midge larvae in shallow, nearshore areas and dis-
play limited predation on larvae aggregated in
deeper areas of the lakes.


ALI, A. 1995. Nuisance, economic impact, and possibili-
ties for control, pp. 339-364. In P. D. Armitage, P. S.
Cranston, and L. C. V. Pinder (eds.), The Chironomi-
dae: the biology and ecology of non-biting midges.
Chapman and Hall, London, UK.
ALI, A. 1996. Pestiferous Chironomidae and their man-
agement, pp. 487-513. In D. Rosen, F. D. Bennett, and
J. L. Capinera (eds.), Pest management in the sub-

June 2002

Scientific Notes

tropics: integrated pest management-A Florida
Perspective. Intercept, UK.
pacts of fish predation on marsh invertebrates: di-
rect and indirect effects. Wetlands 20: 307-312.
BOWEN, S. H. 1996. Quantitative description of the diet,
pp. 513-532. In B. R. Murphy and D. W. Willis (eds.),
Fisheries techniques. 2nd Edition. American Fisher-
ies Society. Bethesda, MD.
WILLIAMS. 1992. Littoral zone invertebrates communi-
ties as affected by a habitat restoration project on Lake
Tohopekaliga, Florida. J. Freshwat. Ecol. 7: 317-328.
CRANSTON, P. S. 1995. Medical significance, pp. 365-384.
In P. D Armitage, P. S. Cranston, and L. C. V. Pinder
(eds.), The Chironomidae: the biology and ecology of
non-biting midges. Chapman and Hall, London, UK.
EPLER, J. H. 1995. Identification manual for the larval
Chironomidae (Diptera) of Florida. Revised Edition.
Florida Department of Environmental Protection.
Tallahassee, FL.
GILINSKY, E. 1984. The role of fish predation and spatial
heterogeneity in determining benthic community
structure. Ecology 65: 455-468.

LOBINSKE, R. J. 2001. Ecological studies of larval Glyp-
totendipes paripes (Chironomidae: Diptera) in se-
lected central Florida lakes for creating an
exploratory temporal and spatial model of nuisance
populations. Ph. D. Dissertation. University of Flor-
ida, Gainesville. 161 pp.
RASMUSSEN, K. 1990. Some positive and negative ef-
fects of stocking whitefish on the ecosystem redevel-
opment of Hjarbaek Fjord, Denmark. Hydrobiologia
200/201: 593-602.
1995. Chironomids in the diet of fish in Lake
Banyoles (Catalonia, Spain), pp. 335-340. In P. S.
Cranston (ed.), Chironomids: from genes to ecosys-
tems. CSIRO. Canberra, Australia.
AND A. HAIN. 1999. Feeding habits of two introduced
fish species (Lepomis gibbosus, Pseudoorasbora
parva) in Neusiedler See (Austria), with special ref-
erence to chironomid larvae (Diptera: Chironomi-
dae). Hydrobiologia 408/409: 123-129.

Florida Entomologist 85(2)


University of Florida, IFAS, Indian River Research and Education Center
2199 South Rock Road, Ft. Pierce, FL 34945

Previous studies with diflubenzuron (AI of
Micromite, Dimilin [Cromton Corporation,
Greenwich, CT 06831] and Thompson-Hayward
TH-6040) revealed inhibition of embryogenesis in
eggs of several citrus root weevils fed on treated
foliage and provided folded wax paper strips as
oviposition sites (Schroeder et al. 1976; Love-
strand & Beavers 1980). In addition, Schroeder et
al. (1976) found that foliar sprays of diflubenzu-
ron applied at rates of 56.6 g and 113.2 g (AI)/378
liters of water with a high-pressure handgun
sprayer reduced egg hatch for 10 days. An aerial
application delivering 283 g (AI) + the extender
Pinolene in 45 1/A was effective for at least 26
days against Diaprepes abbreviatus (L.). Loves-
trand and Beavers (1980) also obtained signifi-
cant reduction of egg hatch of D. abbreviatus,
Pachnaeus litus (Germar) and Artipus floridanus
Horn for 28 days after treatment with handgun-
applied foliar sprays equivalent in rates to those
used by Schroeder et al. in 1976.
By eliminating wax paper strips as oviposition
sites, Schroeder (1996) limited the effect of di-
flubenzuron to residues present on the leaf sur-
faces used for oviposition. This indicated that
direct egg mass contact with diflubenzuron was
an additional mode of action. The two rates eval-
uated by Schroeder (1976, 1996), 142 g and 284 g
(AI)/984 1 of a 1% oil emulsion, were the same
used by Lovestrand and Beavers' 1980 rate,
which were equivalent to lx and 2x the 568 g rate
of Micromite 25W (142 g AI) recommended for
rust mite control (Knapp 2001).
Schroeder (1996) harvested field-sprayed
leaves, paper-clipped them together to form an
oviposition site, and placed treated and untreated
leaf-pairs in cages containing lab-reared oviposit-
ing female adult D. abbreviatus. The lx and 2x
rates resulted in reductions of 77% and 86% in
egg hatch for all time periods with no significant
differences between rates.
Our research, conducted at Ft. Pierce, FL, and
reported here, compares the effect of residues on
egg hatch reduction from aqueous and 0.5% oil
emulsion sprays containing 568 g rate of Micro-
mite 25W
The 1996 Test. A total of twenty shoots bearing
mature flush were selected and tagged on nine 6-
year-old 'Ruby Red' grapefruit trees. On each
twig, 4 distal leaves that could be paired and held
together with a paper clip were selected. These
provided juxtaposed plant surfaces that were
suitable for oviposition. The 2 leaf-pairs were iso-

lated on the twig by removing other leaves to
facilitate caging of the oviposition site. Nine hun-
dred and fifty liters of aqueous and 0.5% oil emul-
sion sprays (FC 435-66) containing 568 g of
Micromite 25W were prepared. These materials
were applied as foliar sprays to "run-off" to 3 trees
per treatment. Three unsprayed trees served as
After spray application, the leaves were al-
lowed to dry. The paired leaves on 4 terminals
were paper-clipped together and each terminal
inserted into a 5.1 cm diameter, 15.2 cm long cy-
lindrical mesh cage on the tree. Two male and 4
female field-collected D. abbreviatus adults were
placed in each cage with a sprig of tender foliage
as food. Fresh, tender foliage was provided as food
every other day. Adults were removed from the
cages on the 7th day post-treatment. Cages were
left on the tree an additional 7 days to protect the
egg masses from predation. Cages and leaf-pairs
were collected on the 14th day and brought to the
lab where egg masses were exposed and exam-
ined under a stereomicroscope to determine their
viability. Eggs with embryos, empty eggs and ne-
onates were counted. This same procedure was
repeated with fresh weevils on the 7th, 14th, 21st
and 28th day after spraying to provide data on the
activity of residues on egg hatch over time. The
test was terminated after 5 weeks. Rainfall was
recorded during the test (Table 1).
There were 116, 1267 and 4054 neonates recov-
ered from leaves harboring eggs treated with Micro-
mite 25W and oil emulsion, aqueous mixture of
Micromite 25W, and untreated control, respec-
tively. This is an average of 98% and 70% reduction
in egg hatch for all time periods with significant
differences between treatments (Table 2).


1996 1997

Treatment applied: 30 May 21 May

21-31 May 0 1.39
31 May 2.21 0
1-8 Jun 2.54 2.57
9-16 Jun 2.18 13.18
17-24 Jun 7.54 3.66

Gauge checked 0800 h daily.

June 2002

Scientific Notes


% Reduction in egg hatch at indicated days post treatment'
Rate Average
Treatment per 950 1 +7 +14 +21 +28 +35 % reduction

Micromite 25W+ 568 g
FC 435-66 Oil 4.751 99 c 100 c 98 c 94 b 99 c 98 c
Micromite 25W 568 g 55 b 74 b 72 b 75 b 71 b 70 b
Untreated 0 2 a 14 a 3 a 6 a 4 a 6 a

'Percent separation within columns by Duncan's Multiple Range Test, 1% level.


% Reduction in egg hatch at indicated days post treatment'
Rate Average
Treatment per 950 1 +7 +14 +21 +28 % reduction

Micromite 25W+ 568 g
FC 435-66 Oil 4.751 70 b 100 b 50 b 100 b 80 b
Micromite 25W 568 g 25 a 39 ab 59 b 78 ab 50 a
Untreated 0 2 a 56 a 2 a 28 a 22 a

'Percent separation within columns by Duncan's Multiple Range Test, 5% level.

Schroeder (1996) found no difference between
two Micromite 25W treatments, viz., 568 and
1136 g of Micromite 25W in 1% oil emulsion
sprays. However, we found that the addition of
0.5% oil emulsion to the Micromite 25W treat-
ment significantly reduced egg hatch compared to
the aqueous spray.
The 1997 Test. Materials and methods in this
experiment differed in 3 respects from the 1996
test: (1) the weevil was P litus, (2) 17-year-old
"Navel" orange trees were used, and (3) the test
was terminated after 28 days.
The results of the test reveal that there were
110, 747 and 1167 neonates found on leaves
treated with Micromite 25W oil emulsion, aque-
ous and no spray, respectively. This is an average
of 80% and 50% reduction for all time periods
with significant differences among treatments
(Table 3).
While Lovestrand and Beavers (1980) compared
568 and 1136 g of Micromite 25W in aqueous
sprays and found that egg hatch was significantly
reduced by both treatments, we found that 568 g
in 0.5% oil emulsion treatment was equivalent in
its impact on the weevils to their low rate of 568
g. Our aqueous spray residue, however, per-
formed poorly compared to their similar aqueous
treatment, perhaps partly due to the rainfall that
occurred during our test (Table 1). However, the
oil component of our emulsion spray apparently
improved residue retention and contributed to its
acceptable performance.
Florida Agricultural Experiment Station Jour-
nal Series No. R-07067.


Residues of the 568 g Micromite 25W spray
rate were less effective in reducing egg hatch of
Pachnaeus litus than the same rate of Micromite
25W fed by Lovestrand and Beavers (1980) to the
same insect, but the addition of oil in our studies
improved the effectiveness of the foliar residues
in the absence of feeding.
Furthermore, Micromite 25W in the 0.5% oil
emulsion was equal in efficacy to Micromite 25W in
1% oil emulsion (Schroeder 1996) in reducing Dia-
prepes abbreviatus egg hatch and would provide
the grower equivalent performance for less cost.
There is no reduction in oviposition attempts
or hatchability because of oil residue (Schroeder
& Green 1983).

KNAPP, J. L. 2001. Florida Citrus Pest Management
Guide: 1999. Fla. Coop. Ext. Service SP-43.
LOVESTRAND, S. A., AND J. B. BEAVERS. 1980. Effect of
diflubenzuron on four species of weevils attacking
citrus in Florida. Florida Entomol. 63(1): 112-115.
A. G. SELHIME. 1976. Ovicidal effect of Thompson-
Hayward TH-6040 in Diaprepes abbreviatus on cit-
rus in Florida. J. Econ. Entomol. 69(6): 780-82.
SCHROEDER, W. J., AND D. S. GREEN. 1983. Diaprepes
abbreviatus (Coleoptera: Curculionidae): oil sprays
as a regulatory treatment, affect on eggs treatment.
J. Econ. Entomol. 76(6):1395-1396.
SCHROEDER, W. J. 1996. Diflubenzuron residue: reduc-
tion of Diaprepes abbreviatus (Coleoptera: Curcu-
lionidae) neonates. Florida Entomol. 79(3): 462-3.

Florida Entomologist 85(2)


'Institut Valencia d'Investigacions Agraries, Ctra. Montcada a Naquera km 5, E-46113-Montcada, Spain
Present address: Universitat Jaume I, Departament de Ciencies Experimentals, Campus del Riu Sec
E-12071-Castello de la Plana, Spain

2University of Florida, Tropical Research and Education Center, Homestead, FL 33031

Phyllocnistis citrella Stainton (Lepidoptera:
Gracillariidae), is an oligophagous leafminer of
Rutaceae, especially of Citrus spp. (Jacas et al.
1997), occurring in all citrus-growing areas
worldwide. Pheromone traps have been used suc-
cessfully in Japan (Ando et al. 1985, Ujiye 1990)
but unsuccessfully in China (Tongyuan et al.
1989), Italy (Ortu 1996), Spain and Florida (pers.
obs.), and Turkey (Uygun, N., Univ. (ukurova,
Adana, Turkey, pers. comm.). Negative results
could be attributed to the use of a non-specific at-
tractant, and to a poor understanding of the be-
havior of the pest. The objective of this study was
to determine patterns of calling of two popula-
tions of P citrella when exposed to three different
photoperiods as a first step toward determination
of a specific sex pheromone.
A first set of experiments was carried out dur-
ing June-July 1998 at the Institut Valencia d'In-
vestigacions Agraries (IVIA) Spain (0.4W long.,
39.6N lat., 33 m alt). A second set was performed
at the Tropical Research and Education Center
(TREC) Florida (80.2W long., 25.3N lat., 1 m
alt.), during October-November 1998. Pupae of
P citrella were gathered from citrus orchards.
Female pupae (Garrido & Jacas 1996) were
placed in 12 cm-diameter petri dishes containing
2% agar and filter paper and held in a cabinet at
25 + 1C, under a photoperiod matching local one
at that moment. Emerging adults were trans-
ferred daily to analogous petri dishes (max. 6
moths per dish) and fed droplets of honey plus
pollen. These were placed in a cabinet at 25 + 1C,
and exposed to one of the three photoperiods:
L12:D12, L16:D8, and L8:D16. At 60 min inter-
vals during scotophase, and using a portable red
light, moth activity (quiescence, locomotion, or
calling) was observed in the cabinet and recorded.
Calling females opened and vibrated their wings
and rhythmically protruded and retracted the
ovipositor. Observations were classified per scoto-
phase hour and sorted by photoperiod, age, and
geographical/seasonal origin. Age was expressed
as the scotophase number (1 to 7), starting with
the first complete scotophase after emergence.
The onset of calling was determined as the time
(hours after lights off) halfway between the first
time calling was observed and the previous obser-
vation. Differences in percentages of female call-

ing were subjected both to G-test (Sokal & Rohlf
1969) and two-way ANOVA for unbalanced de-
signs (SAS Institute 1985). Onset calling times
were also analyzed using a two-way ANOVA.
About 10,000 observations of moth activity were
Table 1 shows the influence of age and photo-
period on the percentage of calling females. Call-
ing patterns for moths subjected to the same
photoperiod regime at the two locations were sig-
nificantly different (G = 24.71; df = 6; P < 0.005).
Nevertheless, the age pattern exhibited by fe-
males exposed to L16:D8 at IVIA did not signifi-
cantly differ from that exhibited by those exposed
to L8:D16 at TREC (G = 7.59; d.f. = 6; P = 0.2245).
Photoperiod (F = 15.56; df = 2, 38; P < 0.0001) and
age (F = 2.67; df = 6, 38; P = 0.0352) affected the
proportion of females calling, whereas the geo-
graphical/seasonal origin did not (F = 0.70; df = 1,
38; P = 0.4102). Calling activity of females ex-
posed to L12:D12 photoperiod was the highest, es-
pecially for age 1 females. Moths aged 4 to 7 called
more than those aged 2 and 3, with 1-day old fe-
males being the less active ones. Females exposed
to a photoperiod opposite to the one occurring
when they were collected in the field barely re-
sponded during the whole experimental period.
Mean onset calling times (MOCT) are shown
in Figure 1. Both geographical/seasonal origin (F
= 781.26; df = 1, 374; P < 0.0001) and photoperiod
(F = 31.06; df = 2, 374; P < 0.0001) affected the
MOCT, but age did not (F = 1.35; df = 6, 374; P =
0.2337). Moths studied during the first set of as-
says called significantly earlier than those tested
during the second set, and, similarly, the longer
the photophase experienced by a moth, the sooner
it called. The MOCT for females collected at IVIA
was 8.62 1.29 h (N = 130; mean SE) when ex-
posed to L12:D12 and 8.11 + 0.93 h (N = 90) when
exposed to L16:D8, whereas for those collected at
TREC, it was 11.67 + 0.58 h (N = 71) when ex-
posed to L8:D16, and 10.65 0.61 h (N = 84) when
exposed to L12:D12. Calling occurred mostly to-
ward the end of the scotophase. Non-calling fe-
males (8L:16D for the experiments at IVIA and
16L:8D at TREC) were usually very active toward
the end of the scotophase.
Phyllocnistis citrella has been considered a
crepuscular and dawn-active moth. Our results

June 2002

Scientific Notes 379

00 .00 00
m2 (0 to^( ^1

> E >(>C
0IC 0

^ ~~ ~ ~ 0 u 0 lCO O OC

ti .4 O^ii o o ^
E- 00ll

0 x

x ,q Cq
EI 00
Sm &i 2 10o'o




rt] --I x -- x 00
x 00 It' COq

Ei- C q x 0

O2 .i
m s m


"I 0 In C<

In 00 Cl 0 0CO
@C g "Co Cl
>6 2 ^' 1(0Io o
S LD^l O (MO t il
[-0 C <; 0(
*.< C0 0 <

0 ~ ( ", M OC O Ol

~I ,3 (0_ oo o ^^i

0 H m -H In

Z 6 S (0

1- << <
-H ^1 (0>

Cl^n0- C 0 l CI(

Florida Entomologist 85(2)



E 14

.E 13



O 9


E1 E - "


- A 1-16L:8D
- O I-12L:12D
0- I-8L:16D
-*- T-16L:8D
-0-- T-12L:12D
p T-8L:16D

1 2 3 4 5 6 7

Fig. 1. Influence of age (1-7) on the onset of calling (in h since start of scotophase) of virgin females of P. citrella
under three different photoperiods (161:8d; 121:12d; 81:16d). The first set of assays was carried out at ivia (i-) and
the second one was at TREC (T-). initial number of females was 30.

show a marked dawn sexual activity. Virgin fe-
males modified their calling patterns in response
to photoperiod with some limitations. Females
could not adapt to the photoperiods completely
opposite to the ones they had experienced in the
field. Furthermore, for a given location, the
MOCT under different photoperiods extraordi-
narily matched current natural scotophases (9 h
at IVIA and 14.5 h at TREC). Therefore, it is pre-
sumed that differences observed at the two loca-
tions were due to seasonal changes rather than to
geographical differences between populations.
The ability of virgin females of many Lepidoptera
to modify the periodicity of their calling in re-
sponse to different environmental changes, in-
cluding day length (Haynes & Birch 1984), has
been interpreted as an adaptation to reduce the
impact of climatic fluctuations on mating success
(Delisle & McNeil 1987).
Onset calling time was not affected by moth
age, although age increased the proportion of fe-
males calling (1 < 2 = 3 < 4 = 5 = 6 = 7 days). At a
given temperature, virgin females of numerous
crepuscular and nocturnal species of Lepidoptera
advance the onset time of calling on successive
nights (Delisle 1992). This has been interpreted
as an adaptation increasing the probability of
older females to attract a mate before younger
moths initiate their calling (Swier et al. 1977).

Age and photoperiod are two of the many fac-
tors affecting the occurrence of calling behavior of
female insects (Howse et al. 1998). Further labo-
ratory experimentation and field testing should
take our results into account.
This research was partially supported by a
grant of the Conselleria de Cultura, Educaci6 i
Ciencia de la Generalitat Valenciana, Spain, to
J. E. P., and a fellowship under the OECD Co-op-
erative Research Program: Biological Resource
Management for Sustainable Agricultural Sys-
tems to J. A. J.
Florida Agricultural Experiment Station Jour-
nal Series No. R-08436.


The effect of age and photoperiod on the calling
behavior of Phyllocnistis citrella was studied.
P citrella is sexually active at dawn. Activation is
presumably caused by the cumulative time
elapsed since sunset. Onset calling time was not
affected by the age of moths, but the proportion of
females calling increased with age.

H. KUROKO. 1985. (7Z-11Z)-7,11-hexadecadienal sex

June 2002

Scientific Notes

attractant of the citrus leafminer moth. Phyllocnis-
tis citrella Stainton (Lepidoptera: Phyllocnistidae).
Agricultural and Biological Chemistry, Tokyo 49:
DELISLE, J., AND J. N. McNEIL. 1987. Calling behaviour
and pheromone titre of the true armyworm Pseuda-
letia unipuncta (Haw.) (Lepidoptera: Noctuidae) under
different temperature and photoperiodic conditions.
J. Insect Physiol. 33: 315-324.
DELISLE, J. 1992. Age related changes in the calling be-
haviour and the attractiveness of obliquebanded lea-
froller virgin females, Choristoneura rosaceana, under
different constant and fluctuating temperatures. En-
tomologia Experimentalis et Applicata 63: 55-62.
GARRIDO, A., AND J. A. JACAS. 1996. Differences in mor-
phology of male and female pupae of Phyllocnistis ci-
trella Stainton (Lepidoptera: Gracillariidae). Florida
Entomol. 79: 603-606.
HAYNES, K. F., AND M. C. BIRCH. 1984. The periodicity
of pheromone release and male responsiveness in
the artichoke plume moth, Platyptilia carduidac-
tyla. Physiol. Entomol. 12: 1597-1600.
HOWSE, P. E., I. D. R. STEVENS, AND 0. T. JONES. 1998.
Insect pheromones and their use in pest manage-
ment. Chapman & Hall, London, UK, 369 pp.
CAIDE, AND J. A. PINA. 1997. Screening of citrus root-

stocks and citrus-related species for resistance to
Phyllocnistis citrella (Lepidoptera: Gracillariidae).
Crop Protection 16: 701-705.
ORTU, S. 1996. Field observations on sexual attraction
of Phyllocnistis citrella Stainton: (7Z, 11Z)-7,11-
hexadecadienal. Proc. XX Intl. Cong. Entomol. p. 496,
Firenze, Italy, Aug. 25-31.
SAS INSTITUTE. 1985. SAS User's guide: Basics. SAS In-
stitute, Cary, NC.
SOKAL, R. R., AND F. J. ROHLF. 1969. Biometry. The
Principles and Practice of Statistics in Biological Re-
search. W.H. Freeman and Company, San Francisco.
776 pp.
SWIER, S. R., R. W. RINGS, AND G. J. MUSIC. 1977. Age-
related calling behavior of the black cutwormAgrotis
ipsilon. Ann. Entomol. Soc. Amer. 70: 919-924.
1989. 7Z, Z11-hexadecadienal: sex attractant of
Phyllocnistis wampella Liu et Zeng. Kunchong Zhi-
shi 3: 147-149 (In Chinese).
UJIYE, T. 1990. Studies on the utilization of a sex at-
tratctant of the citrus leafminer moth, Phyllocnistis
citrella Stainton (Lepidoptera: Phyllocnistidae). I.
Analysis of seasonal population trends and some be-
havioral characteristics of the male moths by the use
of synthetic sex attractant traps in the field. Bulletin
of the Fruit Tree Research Station 18: 19-46.

Florida Entomologist 85(2)


Entomology, Florida A&M University, Tallahassee, Florida 32307, USA

Tshernova (1972) established the genus Viet-
namella for a new species of Asian Ephemerel-
lidae, Vietnamella than. Allen (1980), with little
discussion, placed Vietnamella as a subgenus of
Cincticostella Allen, 1971, in the tribe Ephemer-
ellini. Later, Allen (1984), with somewhat more
discussion of the nymphal characters, in particu-
lar the nymphal gills, admitted that the place-
ment of Vietnamella as a subgenus of Cincticostella
had been incorrect. He established a new subtribe
of the Ephemerellini, Vietnamellae, for Vietna-
Edmunds and Murvosh (1995) transferred the
genus Vietnamella to the related subfamily Telo-
ganodinae and retained the tribe name Vietnam-
ellini for it.
McCafferty and Wang (1997) presented evi-
dence that Vietnamella was closely related toAus-
tremerella Riek, 1963, and, apparently over-
looking the existence of the subfamily name Viet-
namellinae, established a new subfamily, Aus-
tremerellinae, in the family Teloganodidae for the
two genera. Then, McCafferty and Wang (2000)
proceeded to raise the taxon to a family, which
they called Austremerellidae.
According to the International Code of Zoolog-
ical Nomenclature (ICZN 1999: Article 36.1) es-
tablishment of Vietnamellae (Allen 1984) had the
effect of also establishing names for all other
ranks in the family group (coordinate taxa: e.g.,
family, subfamily, tribe). This means that the
name Vietnamellidae became available and valid
at that time.
In determining the valid name of a zoological
taxon, the "Principle of Priority" applies (ICZN
1999: Article 23). It states that "The valid name of
a taxon is the oldest available name applied to
it.. ." Although the Code does make exceptions
for family names in prevailing usage (IC2N 1999:
Article 35.5), Article 23.9.1 describes prevailing
usage to require, among other things, that the se-
nior synonym has not been used as a valid name
after 1899. This is clearly not the case here. Viet-
namellae (and its coordinate taxa) was estab-
lished in 1984 and was an available and valid

name at the time Austremerellidae was estab-
Therefore the valid name for this family of
Ephemerelloidea is Vietnamellidae Allen, and
Austremerellidae McCafferty and Wang is a jun-
ior synonym.


Vietnamellidae Allen is shown to be a senior
synonym of Austremerellidae McCafferty and


ALLEN, R. K. 1971. New Asian Ephemerella with notes
(Ephemeroptera: Ephemerellidae). Canadian Ento-
mologist 103: 512-528.
ALLEN, R. K. 1980. Geographic distribution and reclas-
sification of the subfamily Ephemerellinae (Ephe-
meroptera: Ephemerellidae), pp. 71-91. In J. F.
Flannagan and K. E. Marshall (eds.), Advances in
Ephemeroptera Biology. Plenum, New York.
ALLEN, R. K. 1984. A new classification of the subfamily
Ephemerellinae and the description of a new genus.
Pan-Pacific Entomologist 60: 245-247.
EDMUNDS, G. F., JR., AND C. H. MURVOSH. 1995. Sys-
tematic changes in certain Ephemeroptera studied
by R. K. Allen. Pan-Pacific Entomologist 71: 157-160.
NOMENCLATURE. 1999. International Code of Zoolog-
ical Nomenclature. Fourth Edition. International
Trust for Zoological Nomenclature, London. 306 p.
MCCAFFERTY, W. P., AND T.-Q. WANG. 1997. Phyloge-
netic systematics of the family Teloganodidae (Ephe-
meroptera: Pannota). Ann. Cape Province Museum
(Natural History) 19: 387-437.
MCCAFFERTY, W. P., AND T.-Q. WANG. 2000. Phyloge-
netic systematics of the major lineages of pannote
mayflies (Ephemeroptera: Pannota). Trans. Amer.
Entomol. Soc. 126: 9-101.
RIEK, E. F. 1963. An Australian mayfly of the family
Ephemerellidae (Ephemeroptera). J. Entomol. Soc.
Queensland 2: 48-50.
TSHERNOVA, 0. A. 1972. Some new species of mayflies
from Asia (Ephemeroptera, Heptageniidae, Ephe-
merellidae). Entomologicheskoe Obozrenie 51: 604
614. (In Russian)

June 2002

Scientific Notes


1North Carolina State University, Department of Entomology, Raleigh, NC 27695-7613

2Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, Republic of South Africa

The southern mole cricket, Scapteriscus vicinus
Giglio-Tos, and the tawny mole cricket, S. borellii
Scudder, damage turfgrass in southeastern
United States. The two species are univoltine in
most of their range. They also have similar life cy-
cles and morphology. However, southern mole
cricket is primarily carnivorous, whereas tawny
mole cricket is herbivorous (Taylor 1979, Ulagaraj
1975, Matheny 1981). The African mole cricket,
Gryllotalpa africana Palisot de Beauvois, is a
world-wide pest (Sithole 1986). It damages plants
including wheat, maize, rice, sorghum, millet, bar-
ley, oats, potatoes, cassava, groundnuts, straw-
berries, turnips, tobacco, and vegetables in Africa,
Asia, and Europe. It also causes severe damage to
turfgrass on golf courses in South Africa and Asia
(Brandenburg, unpubl. data). Tsedeke (1979) re-
ported that surface tunneling behavior, which is
partly determined by feeding preference, is differ-
ent between the two species in the U.S. We there-
fore speculate that tunnel architectures of three
species are also different judging from the differ-
ences in their feeding behavior and damage. This
study used fiberglass resins to compare tunnel
architecture of three species of mole crickets in
two locations
Tawny and southern mole cricket tunnel cast-
ings were made on the driving range of Oyster
Bay Golf Course, Brunswick County, NC, during
1998 to 2000. The turfgrass on the driving range
was hybrid bermudagrass, Cynodon dactylon (L.)
Pers. in sandy loam soil. African mole cricket tun-
nel castings were made in typical heavy clay soil
at Silver Lakes Golf and Country Club, Pretoria,
South Africa. The turfgrass on the fairway was
Kikuyu grass, Pennisetum clandestinum Hochst.
ex Chiov.
We located mole cricket tunnel entrances by
hand, and cleaned foreign matter, debris, and soil
from the area around the entrance. We then used
a soapy water flush (Short & Koehler 1979) as an
irritant to flush the mole cricket from the tunnel
for species identification. The soapy water flushing
also helped to find other entrances to the tunnel
and make the soil around the entrance firm. Areas
without turf were avoided because the tunnels are
often blocked by loose soil during the flushing.
We have previously reported that fiberglass
resin is the best material for mole cricket tunnel
casting (Brandenburg et al. 2001). Bondo fiber-
glass resin and hardener (Dynatron/Bondo Corp.,
Atlanta, GA), was used in the U.S. study and a

similar product used in South Africa. This and
other similar products are widely available at
local hardware and automobile repair stores.
Approximately of the recommended amount of
hardener was added to the fiberglass resin (about
1 ml hardener/100 ml resin). The fiberglass resin
hardens quickly after adding hardener, therefore,
the whole procedure must be done quickly. The fi-
berglass resin container was covered and shaken
after adding hardener. The contents were then
poured immediately into the tunnel entrance in a
steady stream. The excavation of the castings
started 1-2 h after pouring. The fiberglass resin in
one can (1 1) usually filled two to three mole
cricket tunnels. We used a large screwdriver to
clear away the grass roots surrounding the tunnel
entrance and to determine the direction of the
casting before starting to dig the cast. Finding
other entrance(s) of the tunnel helps to judge di-
rection the tunnel casting. There are at least two
entrances for tawny and African mole cricket tun-
nels. The soil on tunnel casts was washed away
with water following excavation.
We made over 100 castings and excavations
during 3 years. Tunnels of tawny mole crickets
were almost always (90%) in the shape of '"Y" with
two entrances for each tunnel (Fig. 1A, B, C). Vari-
ations were occasionally observed in the tunnel ar-
chitecture. There might be two parallel '"Y"s
linking together to form a tunnel, or, two entrances
observed at each end of a tunnel. The length of
most tawny mole cricket tunnels ranged from 50 to
70 cm. Tunnels of African mole crickets also typi-
cally showed 'Y" shape (Fig. 1G, H, I). The length of
African mole cricket tunnels ranged from 10 cm to
23 cm. This was much shorter than that observed
in tawny mole crickets. The tunnels of southern
mole crickets were more likely in a reversed "Y"
shape with only one surface entrance (Fig. ID, E,
F). The tunnels often branched within 10 cm deep
of the soil surface. The tunnels were usually much
shorter than those of tawny mole cricket.
The difference in tunnel architecture probably
relates to the behavioral difference of the three
species. Southern mole crickets are carnivorous.
They seek prey throughout the soil. Our observa-
tions and research by Tsedeke (1979) suggested
that southern mole crickets were much more ac-
tive in tunneling than tawny and African mole
crickets. This may be why southern mole cricket
tunnels were almost always branched down into
the soil rather than near the soil surface. In con-


Entrance En cc

ot .

Florida Entomologist 85(2)





a r



Fig. 1. Tunnel Castings of the Tawny (A, B, C), Southern (D, E, F), and African (G, H, I) Mole Cricket.

trast, tawny and African mole crickets feed on
roots near the soil surface and their activity is
around the branches of the "Y". They may face fre-
quent threats from predators (Hudson et al. 1988)
and need more than one entrance in a tunnel. The

harder, clay soil in South Africa may also be re-
lated to the shorter tunnel structure for the Afri-
can mole cricket.
This research was funded in part by the
United States Golf Association, Green Section Re-

June 2002





Scientific Notes

search. We would like to thank Jack Bacheler,
Ken Sorensen, and Mike Waldvogel in Depart-
ment of Entomology, North Carolina State Uni-
versity, for reviewing the draft of this manuscript.


The fiberglass resin used to determine tunnel
architecture of three species of mole crickets was
effective in providing an accurate and durable
record of mole cricket activity. The tunnel castings
of the tawny mole cricket, Scapteriscus vicinus
Scudder, and the African mole crickets, Gryllo-
talpa africana Palisot de Beauvois, almost always
exhibit a "Y" shape upper tunnel structure. The
tunnel castings of the southern mole cricket,
Scapteriscus borellii Giglio-Tos, are typically an
inverted "Y" shape. Tunnels of tawny mole crick-
ets typically go deeper into the soil and are usu-
ally more complex than the other two species.


Determining Tunnel Structure of Mole Crickets

(Orthoptera: Gryllotalpidae) Using Three Materials
with an Emphasis on Fiberglass Resin. J. Kansas
Entomol. Soc. (In press).
Biological control of mole crickets (Orthoptera: Gryl-
lotalpidae) in Florida. Ann. Entomol. Soc. Amer. 34:
MATHENY, E. L., JR. 1981. Contrasting feeding habits of
pest mole cricket species. J. Econ. Entomol. 74: 444-
SHORT, D. E., AND P. G. KOEHLER. 1979. A sampling
technique for mole crickets and other pests in turf-
grass and pasture. Florida Entomol. 1979. 62 (3):
SITHOLE, S. Z. 1986. Mole Cricket. Zimbabwe Agric. J.
83(1): 21-22.
TAYLOR, T. R. 1979. Crop contents of two species of mole
crickets, Scapteriscus acletus and S. vicinus (Ortho-
ptera: Gryllotalpidae). Ibid. 62: 278-279.
TSEDEKE, A. 1979. Plant material consumption and sub-
terranean movement of mole crickets (Orthoptera:
Gryllotalpidae: Scapteriscus) as determined by ra-
dioisotope techniques, with notes on materials for
laboratory feeding. M.S. thesis. University of Flor-
ida, Gainesville. 72 pp.
ULAGARAJ, S. M. 1975. Food habits of mole crickets
(Orthoptera: Gryllotalpidae: Scapteriscus). J. Geor-
gia Entomol. Soc. 10: 229-231.

Florida Entomologist 85(2)


Department of Entomology, University of Massachusetts, Amherst, MA, 01003

Cotesia rubecula (Marshall) is a braconid par-
asitoid of Pieris spp. larvae that is relatively spe-
cific to Pieris rapae (L.) (Lepidoptera: Pieridae), a
pest of cabbage and related cole crops. The estab-
lishment of this parasitoid in the eastern United
States to help suppress this garden pest has been
long sought. Efforts to establish it in North Amer-
ica have a complex history. A self-introduced pop-
ulation of uncertain origin was discovered on
Vancouver Island in British Columbia in 1963
(Wilkinson 1966), and the range of this popula-
tion has extended as far south as Oregon (Biever
1992). This strain was later released in Missouri,
New Jersey, South Carolina, and Ontario (near
Ottawa) (Puttler et al. 1970; Williamson 1971,
1972). This strain appears not to have established
in Missouri (Parker & Pinnell 1972), but may
have established in Ontario (Corrigan 1982). Poor
establishment of this strain was attributed to an
improperly timed diapause induction response
(Nealis 1985).
A second population, from the former Yugo-
slovia, was released in Missouri in the mid 1980s,
and subsequently released in Virginia and On-
tario. In 1988, the Yugoslavian strain was recov-
ered in Virginia, but this population later appeared
to have died out, perhaps due to high level of
hyperparasitism (McDonald & Kok 1991). In
1993, C. rubecula, of uncertain origin, was found
to be the dominant parasitoid in Quebec, in farm-
ing areas near Montreal (about 160 km east of
Ottawa) (Godin & Boivin 1998).
In 1988, a population of C. rubecula was col-
lected by David Reed of the USDA in Shenyang,
China (42 north latitude, 123 east longitude), for
release in the eastern United States. This location
matched the intended release location in Massa-
chusetts in latitude, and both locations have conti-
nental type climates. Parasitized host larvae
(P rapae) were shipped to the USDA quarantine
laboratory in Newark, Delaware. Adult parasitoids
were allowed to emerge and, following confirmation
of species identity, 99 female and 49 male C. rubec-
ula adults from this shipment were shipped to the
senior author in Amherst, Massachusetts in July of
1988 and all were released in field cages in a pesti-
cide-free, 0.1 ha collard plot in Deerfield, Massa-
chusetts (42 n. 1.). That C. rubecula was not present
at this site before the release (through spread, per-
haps from some distant source) is demonstrated by
the absence of C. rubecula in large numbers of
hosts collected at this location and dissected for
parasitism rates in a population dynamics study I
ran in 1985 and 1986 (Van Driesche 1988).

We subsequently reared this strain both in the
laboratory and from field-collected larvae be-
tween 1988 and 1993 and made 12 other releases
in Massachusetts, three in Connecticut, and one
in Rhode Island, for 17 release locations in total
(Fig. 1, two sets of MA sites overlap on map).
Same-year recoveries of the parasitoid were made
at seven of these sites, and recoveries were made
after one or more years at seven other sites.
Among the seven sites at which recoveries were
made in subsequent years, we observed the para-
sitoid at three sites one year after release and at
single sites 2, 3, 5, and 8 years after last release.
Recovery efforts varied in different years and not
all sites were visited yearly.
To assess spread away from release sites, we
periodically collected groups of P rapae larvae
from non-release locations. We have recovered
C. rubecula from 13 non-release sites, from just
north of Hartford, Connecticut to Craftsbury, Ver-
mont (north of St. Johnsbury) (Fig. 1). Recoveries
have been made both along the Connecticut River
Valley and in various locations in the Litchfield
Hills in Connecticut, the Berkshire Hills in Mas-
sachusetts, and the Champlain Valley of Vermont.
Towns in which recoveries have been made either
at non-release sites or, if a release site, one or
more years after the release include Winsor and
Falls Village, Connecticut; Williamstown, Lanes-
boro, Westhampton, Northampton, Amherst, Had-
ley, Deerfield, Northfield, and Barre, Massachu-
setts; and Stamford, Rockingham, Hartland, South
Royalton, Plainfield, Burlington, and Craftsbury,
Vermont (Fig. 1), all of which indicate extensive
range expansion in both agricultural valleys and
adjacent forested hill country. Recoveries through-
out Vermont bring the known range ofC. rubecula
near the Canadian border. Godin and Boivin
(1998)'s report of recovery ofC. rubecula of uncer-
tain origin in southern Quebec, seen in the light
of the data presented here, may be a further
northward extension of the Chinese population,
rather than an eastward extension of releases
from near Ottawa. This is uncertain, as no molec-
ular markers have been identified to separate
these populations.
Because establishment ofC. rubecula has been
associated with declines in density of the other in-
troduced P rapae parasitoid, Cotesia glomerata
(L.), in Oregon and Washington (Biever 1992), we
also counted numbers of P rapae larvae and Cote-
sia parasitoid cocoons (as single cocoons for the
solitary species C. rubecula and as cocoon groups
for the gregarious species C. glomerata) on entire

June 2002

Scientific Notes








New York




/New jersey

Fig. 1. Locations of release and recovery sites for Cotesia rubecula (Chinese strain) in the northeastern United
States. Solid circles with cross hatches are release sites at which recoveries were made one or more years after re-
lease; solid circles without cross hatches are non-release sites where recoveries were made; hollow circles with cross
hatches are release sites at which recoveries were made only in the year of release; hollow circles without cross
hatches are release sites at which no recoveries were made; crosses (in Canada) indicate closest sites with known
releases or recoveries of other strains of C. rubecula.

collard plant in two years (1985, 1986) before the
release of C. rubecula (in 1988) in our Deerfield,
Massachusetts plot and for three years (1990,
1991, 1992) after the parasitoid's establishment.
The size and management of this 0.1 ha collard
plot was maintained in a consistent manner from
1985 to 1992. For each week of the growing sea-
son (May through September) in these years, we
examined 20-100 whole collard plants (187 sam-
ple occasions, with an average of 58 plants per
date). Numbers of groups of C. glomerata cocoons
were greatest in July, August and September. For
all samples in these three months in five years,
we classified each plant sampled into a 2 x 2 ma-
trix. One factor was whether or not the plants had
C. glomerata cocoons on them (+/-). The other fac-

tor was the time period (pre- or post-release of
C. rubecula). Data from two pre-release years
were available (1985 and 1986), as well as three
post-release years (1990, 1991, and 1992). Data
from the year of release (1988) and the following
year (1989) were not included in order to allow for
population interactions to reach a stable end
point before analysis. We then used a X2 test on
these data to determine if the percentage of
plants with C. glomerata cocoons on them varied
between the pre and post release periods. Of 4098
plants examined in these months in 1985 or 1986,
16% (661) bore live C. glomerata cocoons, com-
pared to only 3% (82) of 2708 plants examined in
July-September of 1990, 1991 or 1992, a signifi-
cant difference (X2 = 288.7, df= 1,P < 0.005). Total

Florida Entomologist 85(2)

numbers of larvae on sampled plants in the prer-
elease period (9980) were either the same or
lower (2.45 larvae per plant) than in the post re-
lease period (8683 larvae on 2708 plants, or 3.21
per plant) and therefore the decline in the propor-
tion of plants bearing C. glomerata cocoons in the
post release period cannot be explained as being
due to a decrease in the number of larvae per
plant available for parasitization. Furthermore,
in the 1990-1992 period, C. rubecula accounted
for over half of all Cotesia parasitism of P. rapae
larvae in the Deerfield, MA, release plot (74% [n =
175], 91% [n = 87], and 49% [n = 76], in 1990,
1991, and 1992 respectively, with percentage be-
ing based cocoons of each parasitoid species seen
on sampled plants). These data suggest that
C. glomerata declined in density in the study area
following establishment of C. rubecula. However,
regionally C. glomerata remains common in New
England and we cannot say if it has declined at
that larger spatial scale.
We conclude that C. rubecula has become an
important parasitoid of P. rapae in parts of New
England since its establishment in 1988 and we
suspect that its range is still increasing and
should be examined in other states in the region.
Potential effects of this new parastioid on related
native Pieris butterflies have been examined
(Benson et al. unpubl.).


A population of Cotesia rubecula, collected from
near Beijing, China and released in Massachusetts
in 1988, has established and spread throughout
much of New England. It has become a common
parasitoid of Pieris rapae in agriculture fields and
is also found in meadow habitats. Cotesia glomer-
ata appears to have declined in abundance follow-
ing establishment of C. rubecula.


BIEVER, K. D. 1992. Distribution and occurrence of Cote-
sia rubecula (Hymenoptera: Braconidae), a parasite
of Artogeia rapae in Washington and Oregon. J.
Econ. Entomol. 85: 739-742.
CORRIGAN, J. E. 1982. Cotesia (Apanteles) rubecula [Hy-
menoptera: Braconidae] recovered in Ottawa, On-
tario ten years after its release. Proc. Entomol. Soc.
Ontario 113: 71.
GODIN, C., AND G. BOIVIN. 1998. Occurrence of Cotesia
rubecula (Hymenoptera: Braconidae) in Quebec, 30
years after its introduction in North America. Cana-
dian Entomol. 130: 733-734.
McDONALD, R. C., AND L. T. KOK. 1992. Colonization
and hyperparasitism of Cotesia rubecula (Hymen.:
Braconidae), a newly introduced parasite of Pieris
rapae, in Virginia. Entomophaga 37: 223-228.
NEALIS, V. 1985. Diapause and the seasonal ecology of
the introduced parasite, Cotesia (Apanteles) rubec-
ula (Hymenoptera: Braconidae). Canadian Entomol.
117: 333-342.
PARKER, F. D., AND R. E. PINNELL. 1972. Further stud-
ies of the biological control of Pieris rapae using sup-
plemental host and parasite releases. Environ.
Entomol. 1: 150-157.
THEWKE. 1970. Introduction of Apanteles rubecula
Marsh. and other parasites of Pieris rapae in British
Columbia. J. Econ. Entomol. 63: 304-305.
VAN DRIESCHE, R. G. 1988. Survivorship patterns ofPi-
eris rapae (Lep.: Pieridae) larvae in Massachusetts
kale with special reference to mortality due to Cote-
sia glomerata (Hymen.: Braconidae). Bull. Entomol.
Res. 78: 199-208.
WILKINSON, A. T. S. 1966. Apanteles rubecula Marsh.
and other parasites of Pieris rapae in British Colum-
bia. J. Econ. Entomol. 59: 1012-1013.
WILLIAMSON, G. D. 1971. Insect liberation in Canada.
Parasites and predators 1970. Agric. Canada Liber-
ation Bull. 34.
WILLIAMSON, G. D. 1972. Insect liberation in Canada.
Parasites and predators 1971. Agric. Canada Liber-
ation Bull. 35.

June 2002

Scientific Notes


'Instituto de Ecologia, A.C. Departamento de Entomologia. Km 2.5 antigua carretera a Coatepec, Apdo
Postal 63, 91000, Xalapa, Veracruz, M6xico

2UADY. Facultad de Medicina Veterinaria y Zootecnia. Departamento de Zoologia, Apdo
Postal 4-116 Itzimnd. M6rida, Yucatan, M6xico

Tephritid flies (Diptera: Tephritidae) are
known as "true fruit flies" due to the close relation-
ship between their immature stages and their wild
and domesticated host plants. They are the most
important dipteran pests of agriculture world-
wide (Christenson & Foote 1960) and include 481
genera and 4352 species (Norrbom et al. 1998).
Anastrepha is the most economically important
and diverse genus of fruit flies in the Americas,
with 197 species distributed throughout tropical
and subtropical areas (Norrbom et al. 2000). To
date, 32 species are known to occur in Mexico, and
seven of them have been reported for the Yucatan
Peninsula (YP). This includes the Mexican states
of Campeche, Quintana Roo and Yucatan. Anas-
trepha species reported for each state are:
Campeche (Anastrepha fraterculus, A. hamata,
A. limae, A. obliqua, A. serpentina); Quintana Roo
(A. hamata, A. ludens, A. obliqua, A. serpentina);
and Yucatan (A. fraterculus, A. ludens, A. serpen-
tina, A. striata) (Hernandez-Ortiz 1992).
This work provides new locality records for
Anastrepha species already reported for YP, first
records of three species in YP (A. ampliata, A. pal-
lens and A. spatulata), and the first record of
A. compressa for Mexico. Material examined is
deposited at the Colecci6n de Insectos, Instituto
de Ecologia A.C. Xalapa, Veracruz (IEXA), and
Colecci6n Entomol6gica Regional of the Univer-
sidad Aut6noma de Yucatan (CER).

Anastrepha ampliata Hernandez, 1990
This species has only been reported in the
Mexican state of Chiapas, and Guatemala
(Hernandez-Ortiz 1990, 1992); new record for YP
Material examined. MEXICO. CAMPECHE,
Calkini, Concepci6n, 5-VIII-1997. Huchin (1 Y
IEXA); QUINTANA ROO, Chunhuhub, McPhail-
trap, 12-VIII-1997, Xool-Cetz (8 Y IEXA);
YUCATAN, Dzilam Reserve, Rancho San Salva-
dor, 30-IX-1992, light-trap, Delfin & Manrique (2
Y CER), Ibid. butterfly-trap (1 Y CER), Ibid. 30-
XI-1992, net (2 Y CER).

Anastrepha compressa Stone, 1942
This species was described on the basis of ma-
terial from several localities in Panama (La Cam-

pana, El Cermeho, and Balboa) (Stone 1942), and
updated records include material from Venezuela
(Norrbom et al. 1998). This is the first report from
Mexico which extends its distribution to the north
of Central America.
Material examined. MEXICO. CAMPECHE,
Alfredo Bonfil, 20-1-1997, McPhail-trap (4 2, 1 6

Anastrepha fraterculus (Wiedemann, 1830)
Its known distribution includes the United
States (Texas), South and Central America, and
Trinidad (also introduced to Galapagos Is.) (Stone
1942; Norrbom et al. 1998). Records for Mexico in-
clude Aguascalientes, Campeche, Chiapas, Nuevo
Le6n, Oaxaca, Tamaulipas, Veracruz, Yucatan,
and Zacatecas (Arana et al. 1992; Hernandez-Or-
tiz 1992).Here we report the first record for Quin-
tana Roo state.
Material examined: MEXICO. CAMPECHE,
Tenabo, Tinun, 10-VII-1997, Herrera (1 6 IEXA);
QUINTANA ROO, Felipe Carrillo Puerto, Chunhu-
hub, 5-12-VIII-1997, Xool (11 6, 128 Y IEXA);
YUCATAN, Dzilam Reserve, Rancho San Salvador,
30-XI-1992, light-trap, Delfin & Manrique (1 6
CER); Ibid. 30-XI-1992, butterfly-trap (1 Y CER);
Colonia Yucatan, Kalah Dzonot, 21-22-IX-1993,
butterfly-trap, Delfin & Manrique (2 Y CER).

Anastrepha ludens (Loew, 1873)

This species ranges from southern USA
(Texas) to Central America (Foote 1967). It has
been reported for 25 Mexican states including
Quintana Roo and Yucatan (Huerta et al. 1987;
Arana et al. 1992; Hernandez-Ortiz 1992). Here
we report the first record for Campeche state.
Material examined. MEXICO. CAMPECHE,
La Libertad, 26-VI-1993, net, Manrique (1 Y
CER); QUINTANA ROO, Felipe Carrillo Puerto,
Chunhuhub, 26-XII-1997, Xool (1 2, 1 6 IEXA).

Anastrepha obliqua (Macquart, 1843)
This species has a wide distribution in the
New World, having been recorded from the USA
(Florida, Texas) to South America and the Carib-
bean Islands (Stone 1942). In Mexico its range in-

Florida Entomologist 85(2)

cludes both coasts and other central states (18
states), including Campeche and Quintana Roo
within YP (Hernandez-Ortiz 1992).
Material examined. MEXICO. CAMPECHE,
Palizada, Rancho Santa Isabel, 23-IX-1997, Ca-
brales (1 Y IEXA), Palizada, Rancho Alamilla, 7-
X-1997, Cabrales (1 Y IEXA), Cd. del Carmen,
Matamoros [no date], Dominguez (1 Y IEXA).

Anastrepha pallens Coquillett, 1904
This species has a known distribution in the
Mexican states of Chiapas, Coahuila, Guerrero,
Jalisco, Nayarit, Nuevo Leon, Oaxaca, Sinaloa,
Sonora, Tamaulipas and Veracruz (Hernandez-
Ortiz 1992), with its northern limits in USA
(Texas) and southern limits in El Salvador and
Honduras (Norrbom 1998). Here we report a new
record for YP. The reported hosts for this species
are Sideroxylon celastrinum (Kunth) T. D. Pen-
nington (as Bumelia angustifolia) and S. lanugi-
nosa Michx. Baker et al. (1944) also reported as
host B. spiniflora A.DC. in Mexico, probably a
synonymy of S. celastrinum (see Norrbom 1998).
Material examined. MEXICO. YUCATAN, Ria
Lagartos Reserve, El Cuyo, 8-II-1995, light-trap,
Delfin & Manrique (1 Y CER).

Anastrepha serpentina (Wiedemann, 1830)
This species has a wide distribution in the
New World, from USA (Texas) to South America
and Trinidad (Foote 1967). In Mexico it occurs in
16 states along both coasts and in other central
states, including the YP (Hernandez-Ortiz 1992).
Material examined. MEXICO. QUINTANA
ROO, Vallehermoso, 19-20-VII-93, butterfly-trap,
Delfin & Manrique (1 Y CER); YUCATAN, Dz-
ilam Reserve, Rancho San Salvador, 30-XI-1992,
light-trap, Delfin & Manrique (3 Y CER); Ibid.
butterfly-trap (1 Y CER).

Anastrepha spatulata Stone, 1942
This species has a known distribution in the
Mexican states of Baja California Sur, Chiapas,
Guerrero, Jalisco, Morelos, Nayarit, Oaxaca, Si-
naloa, Sonora, Tamaulipas and Veracruz, with its
northern limit in USA (Texas) and southern limit
in Costa Rica and Panama (Foote 1967; Hernan-
dez-Ortiz 1992). Here we report it for the first
time for YP.
Material examined. MEXICO. CAMPECHE,
Hecelchacan, Blanca Flor, 20-III-1997, Dzul (1 j

Anastrepha striata Schiner, 1868

This species has a known distribution in the
Mexican states of Aguascalientes, Colima, Chia-
pas, Guerrero, Jalisco, Mexico, Morelos, Nayarit,
Oaxaca, Sinaloa, Veracruz and Yucatan, with its

northern limit in USA (Texas) and a southern dis-
tribution in many countries in Central and South
America (Stone 1942; Hernandez-Ortiz & Aluja
1993) Here we report the first record for Quintana
Roo state.
Material examined. MEXICO. QUINTANA
ROO, Felipe Carrillo Puerto, Chunhuhub, 26-XII-
1997, Xool (1 6, 1 2 IEXA), Felipe Carrillo Puerto,
Emiliano Zapata, 15-X-1997, Xool (3 2 Y IEXA).
At this time, a total of 11 Anastrepha species
are known to be present in the states of the
Yucatan Peninsula. The current status of our
knowledge of these species and the most complete
list of Anastrepha in those Mexican states is as
follows: Campeche, 9 species; Quintana Roo, 7
species; Yucatan, 6 species.
The species A. ludens, A. serpentina and
A. obliqua are economically important for fruit
crops in Mexico. At the present time, there are no
reports indicating that they represent an agricul-
tural problem for YP states. However, the pres-
ence of species detrimental to national and
regional agriculture, indicates the need for a per-
manent surveillance campaign to evaluate popu-
lation and damage levels throughout the region.


We make an updated report for theAnastrepha
species that occur in the Yucatan Peninsula. For
the first time A. compressa Stone is recorded from
Mexico, and the occurrence of three other species
for this region is documented:A. ampliata Hernan-
dez,A. pallens Coquillett, andA. spatulata Stone.
Information about localities and collection dates
of voucher specimens are provided.

ARANA, P. J., P. T. VERA, AND J. CACHON. 1992. La
mosca mexicana de la fruta del sur del estado de
Yucatan. Experiencias en desarrollo sostenible, 1-33.
MCPHAIL. 1944. A review of the studies on the Mex-
ican fruitfly and related Mexican species. U.S. Dept.
Agric. Misc. Publ. 531: 1-155.
CHRISTENSON, L. D., AND R. H. Foote. 1960. Biology of
fruit flies. Annu. Rev. Ent. 5: 171-192.
FOOTE, R. H. 1967. Family Tephritidae (Trypetidae,
Trupaneidae), Fascicle 57, pp. 1-91. In N. Papavero
[ed.], A catalogue of the Diptera of the Americas
South of the United States. Museo de Zoologia, Uni-
versidade de Sao Paulo, Brazil.
HERNANDEZ-ORTIZ, V. 1990. Lista preliminary de species
mexicanas del g6nero Anastrepha (Diptera: Tephriti-
dae) con descripci6n de nuevas species, registros y si-
nonimias. Folia Entomol. Mexicana. 80: 227- 244.
HERNANDEZ-ORTIZ, V. 1992. El g6nero Anastrepha Schiner
en M6xico (Diptera: Tephritidae). Taxonomia, dis-
tribuci6n y sus plants hu6spedes. Institute de
Ecologia Publ. 33. Xalapa, Veracruz, Mexico. 162 pp.
HERNANDEZ-ORTIZ, V., AND M. ALUJA. 1993. Listado de
species del g6nero neotropicalAnastrepha (Diptera:

June 2002

Scientific Notes

Tephritidae), con notas sobre su distribuci6n y plan-
tas hospederas. Folia Entomol. Mexicana 88: 89-105.
1987. Distribuci6n geografica de las moscas de la
fruta del g6nero Anastrepha Schiner en M6xico, pp.
128-146. In INIFAP [ed.], Primer Informe sobre Mos-
cas de la Fruta en Mango. INIFAP, Veracruz, M6xico.
NORRBOM, A. L. 1998. A revision of the Anastrepha dac-
iformis species group (Diptera: Tephritidae). Proc.
Entomol. Soc. Washington 100: 160-192.
WHITE, AND A. FREIDBERG. 1998. Systematic data-

bases of names, pp. 65-251. In F. C. Thompson [ed.],
Fruit fly expert identification system and systematic
information database. Backhuys Publ., Leiden,
TIZ. 2000. Phylogeny of the genera Anastrepha and
Toxotrypana (Trypetinae: Toxotrypanini) based on
morphology, pp. 299-342. In M. Aluja and A. L. Norr-
bom [eds.], Fruit flies (Tephritidae): phylogeny and
evolution of behavior. CRC Press, USA.
STONE, A. 1942. The fruit flies of the genus Anastrepha.
U.S. Dept. Agric. Misc. Publ. 439: 1-112.

Florida Entomologist 85(2)


Departamento de Interacciones Planta-Insecto, Centro de Desarrollo de Productos Bi6ticos del Instituto Polit6cnico
Nacional, COFAA, Carretera Yautepec-Jojutla km 8.5, A.P. 24, 62730 San Isidro, Yautepec, Morelos, Mexico

Tuberose, Polianthes tuberosa L. (Liliales:
Agavaceae), is grown as an ornamental plant and
as a source of a fragrant essence for perfumes
(Gonzatti 1981, Watson & Dallwitz 1992). It has
been grown commercially in the state of Morelos,
Mexico, for 60 years. Currently there is grown ap-
proximately 300 cultivated ha, which generates
US $6,300,000 per year (Uribe 2000). However, in
recent years, producers have noticed severe dam-
age caused by an insect that they call black weevil
but unidentified scientifically. The objective of
this work was to identify the pest and to assess
The study was carried out in the central Mexi-
can state of Morelos, between 1822'19" and
1907'10" north and between 99030'8" and
1907'10" west. Mean annual temperature is
20C and the annual rainfall ranges from 900 to
1100 mm (Garcia 1981). Samples were collected
from March to September 2000 in the main areas
of tuberose production to include: the municipali-
ties of(1) Tepalcingo, (2) Emiliano Zapata, and (3)
Coatlan del Rio.
In the first two municipalities, 100 bulbs of
P tuberosa were collected, whereas in the third,
120 plants were collected. A 1-ha plot in each mu-
nicipality was selected, and samples were taken
from the center and four sides of the plot. In the
laboratory, bulbs and whole plants were exam-
ined for insects. The insect larvae and adults were
deposited in 70% ethanol and processed for iden-
tification (see acknowledgments).
The weevil was identified as Scyphophorus acu-
punctatus Gyllenhal (Coleoptera: Curculionidae)
(Muifz 2000, Marfn 2000, Napoles & Equihua
2000). Most damage is caused by weevil larvae
mining the bulb. The samples from Coatlan del Rio

had the highest incidence of damage (69%). Emil-
iano Zapata and Tepalcingo demonstrated 47 and
37% damage, respectively (Table 1).
This paper is the first formal report of S. acu-
punctatus attacking tuberose. Vaurie (1971), in
her revision of Scyphophorus, did not mention
tuberose as a host plant of S. acupunctatus. Dampf
mentioned the weevil as "acapiche del nardo"
(Anonymous 1930). Vaurie (1971) mentions that
this insect is distributed from the southern USA to
the north of South America, the Caribbean (Cuba,
Hispaniola, and Jamaica), East Africa, Hawaii,
Java, and Australia. In Mexico it has been re-
ported attacking several economically important
plants of the family Agavaceae (McGregor &
Gutierrez 1983). In the Yucatan peninsula it has
been mentioned as causing damage up to 50% in
the cultivation of henequen (Agave fourcroydes
Lem.) (Ramirez-Choza 1979). In the states of
Hidalgo, Tlaxcala, and Mexico, it has been re-
ported causing damage of 30% to the cultivated
plant "maguey pulquero" (Agave salmiana Salm-
Dyck ssp. crassispina (Trel.) Gentry and var. culta)
(Ruvalcaba 1983). In the state of Jalisco, it caused
damage of 10% to "agave tequilero" (Agave tequi-
lana Weber var. azul) (Valenzuela 1994). In the
municipalities of Tepalcingo and Coatlan del Rio,
cultivation of A. tequilana began three years ago,
whereas in Emiliano Zapata there are wild agaves,
predominantly A. angustifolia Haw., in the hills
near cultivated P tuberosa. These two agaves may
serve as refuge to S. acupunctatus during chemical
applications or when tuberose is not in the field, so
control is complicated (Uribe 2000). Infestations in
Agave tequilana var. azul have been reported as
from 4 to 24 weevils per plant (Solis et al. 1999).
The present survey showed 6-10 larvae per in-


Bulbs Plants
Locality and date sampled/infested sampled/infested Adults/plant Percent infested

Tepalcingo 6 March 100/36 36
Emiliano Zapata 7 April 100/47 47
Tepalcingo 30 August 45/14 40
Tepalcingo 6 September 120/42 35
Coatlan del Rio 5 September 120/83 4-36 69

June 2002

Scientific Notes

fested bulb and 4-36 adults per infested plant.
S. acupunctatus is a severe pest of tuberose Our
work continue on S. acupunctatus life cycle, popu-
lation dynamics, identification and evaluation of
feeding attractants.
We thank Dr. J. H. Frank (Entomology and
Nematology Department, University of Florida)
for guidance and corrections to a manuscript
draft. This report was supported financially by
Fundaci6n Produce Morelos, A. C. (project num-
ber 4-I/A18/2000). We also thank Uni6n de Pro-
ductores de Nardo del Estado de Morelos, through
Mr. Rodolfo Uribe Landa. We thank R. Mufifz,
A. Marin and Dr. J. Napoles and Dr. A. Equihua
for identification of the insects. The authors of
this paper are scholarship awardees of Comision
de Operaci6n y Fomento de Actividades Academi-
cas del Instituto Politecnico Nacional.


Scypophorus acupunctatus is reported attack-
ing Polianthes tuberosa in Morelos, Mexico. All
growth stages (eggs, larvae, pupae and adults) of
this insect were collected. The average percent in-
festation was 51%.
This paper is in memory of Dr. Mario Camino


ANONYMOUS. 1930. Principales plagas y enfermedades
de los cultivos en la Repuiblica Mexicana, incluyendo
las mas importantes de los Estados Unidos de
Norteam6rica. Oficina Federal para la Defensa Agri-
cola, M6xico. 195, 159, 220.

GARCIA, E. 1981. Modificaciones al sistema de clasifi-
caci6n climatica de Koppen. Institute de Geograffa,
Universidad national Aut6noma del Estado de M6x-
ico. Offset Larios; M6xico, D.F., Mexico.
GONZATTI, C. 1981. Flora Taxon6mica Mexicana II. Ce-
neti; Guadalajara, Mexico (see pp. 87-88).
Mapiv, J. A. 2000. Personal communication. INIFAP-
SAGAR, Celaya, Guanajuato, M6xico.
McGREGOR, R., AND 0. GUTIERREZ. 1983. Gufa de insec-
tos nocivos para la agriculture en M6xico. Alhambra;
M6xico. 166 pp.
MuNIz, R. V. 2000. Personal Communication. Sanidad
Vegetal, INIFAP-SAGAR, M6xico.
NAPOLES, R. J., AND A. EQUIHUA. 2000. Personal Com-
munication. Colegio de Posgraduados, Instituto de
Fitosanidad, Montecillo, Estado de Mexico, Mexico.
RAMIREZ-CHOZA, J. L. 1979. Metodologia para el control
del max del henequ6n Scyphophorus acupunctatus
bajo condiciones de campo como resultados de tres
afos de studio. Fol. Entomol. Mexicana 42: 62-63.
RUVALCABA, J. M. 1983. El maguey manso. Universidad
Aut6noma Chapingo; M6xico. 122 pp.
sectos asociados con Agave tequilana var. azul, en
cinco localidades de Jalisco, M6xico. Memories
XXXIV Congreso Nacional de Entomologia, Aguas-
calientes, M6xico, 23-26 May 1999: 455-457.
URIBE, L. R. 2000. Personal communication. President
de la Uni6n de Productores de nardo del Estado de
Morelos. Coatlan del Rio, Morelos, M6xico.
VALENZUELA, Z. A. G. 1994. El agave tequilero. Litteris;
M6xico, D.F., M6xico (see pp. 121-137).
VAURIE, P. 1971. Review of Scyphophorus (Curculion-
idae: Rhynchophorinae). Coleopts. Bull. 25: 1-8.
WATSON, L., AND M. J. DALLWITZ. 1992. The families of
flowering plants: descriptions, illustrations, identifi-
cation, and information retrieval. [Online] Available:
http://biodiversity.uno.edu/delta/ (4 November 2000).

Florida Entomologist 85(2)


1Rudyard Kipling No. 4800 casa 18, Jardines de la Patria, Zapopan, Jalisco, Mexico, CP 45030
2Departamento de Entomologia Tropical, El Colegio de la Frontera Sur, Ap. Postal 36
Tapachula, Chiapas, M6xico, CP 30700

This paper reports a new record of a moth species, Zamagiria dixolophella Dyar (Pyralidae:
Phycitinae) attacking sapodilla (Manilkara zapota van Royen) in Southern Mexico. This is
the first report of this species in Mexico. The female genitalia and the last instar larva are
described and illustrated.
Key Words: Pyralidae, Zamagiria dixolophella, Sapodilla, Manilkara zapota, Mexico

Un nuevo registro de una especie de palomilla atacando al chicozapote (Manilkara zapota
van Royen) en el sur de Mexico es reportado. La palomilla fue identificada con Zamagiria
dixolophella Dyar de la familiar Pyralidae, subfamilia Phycitinae. Esta especie es reportada
por primera vez para Mexico. La genitalia femenina y el ultimo estadio larvario es descrito
e ilustrado.

The sapodilla (Manilkara zapota van Royen) is
a fruit tree native of the south of Mexico and Cen-
tral America, and is found from the Yucatan pen-
insula to Costa Rica. Sapodilla is grown
commercially in India, the Philippines, Sri
Lanka, Malaysia, Mexico, Venezuela, Guatemala,
and other Central American countries (Mickel-
bart 1996). The entomofauna associated with sa-
podilla trees has been reported from Venezuela,
Puerto Rico and India (Rubio-Espina 1968, Bu-
tani 1975, Sandhu & Sran 1980, Medina-Goud et
al. 1987). For example, in India, over 25 species of
insects have been recorded causing damage to sa-
podilla (Butani 1975, Sandhu & Sran 1980), but
the main insect pests are a galechid caterpillar
(Anarsia achrasella) and the pyralid caterpillar,
Nephopteryx engraphella. The former species is a
bud borer, while the latter feeds on the leaves,
flower buds and young fruits. Unfortunately, the
entomofauna associated with sapodilla in its area
of original distribution has not been studied. In
Mexico and Central America, the main insect pest
of this tree seems to be the fruit flies of the genera
Anastrepha, particularly A. serpentina (Norrbom
& Kim 1988). For several years the sapodilla pro-
ducers of the Soconusco region of Chiapas state,
Mexico have been reporting a bud borer moth,
which has been causing problems in their planta-
tions. This work was undertaken to identify the
moth as a first step towards further studies on the
biology, ecology and control of this species.

The biological material was collected from sev-
eral sapodilla farms located in the municipalities

of Tapachula and Suchiate of the Chiapas state,
Mexico. The pupae from tender young shoots and
fruits were taken to the laboratory and placed in
screen cages (20 x 20 cm) for adult emergence at
25 + 1C, 30-50% RH and a photoperiod 16: 8 (L:
D) h. After adult emergence, the insects were
killed and mounted for identification. Larvae
were killed immediately after collection. They
were fixed in Pampel liquid and preserved in 70%
ethyl alcohol. For morphological studies the lar-
vae were cleared with lactic acid. The terminology
used to describe the setae and other characters is
adopted from Stehr (1987).

The moth species was identified as Zamagiria
dixolophella Dyar (Pyralidae, Subfamily Phyciti-
nae). This species represents the first report in
Mexico. The original description of this species
was based on males collected in Corazal, Panama,
in 1914 (Heinrich 1956). The female has not been
described and is very close to Zamagiria pogery-
thus Dyar (Neunzing & Dow 1993, Heinrich
1956). However, the female genitalia of Z. dixol-
ophella has some differences: the ductus bursae is
short and narrow, while in Z. pogerythus it is
wide, as wide as the bursae. The ductus seminalis
of Z. dixolophella is located in the apical part of
bursae, whereas in Z. pogerythus it is located in
the basal part of the bursae (Figs. 1 A and B).

Description of Adults
In general appearance this insect is a small
gray moth, resting with the wings folded along

June 2002

Scientific Notes


I -T

Fig. 1. A) Female genitalia of Z. Pogerythrus (re-
drawn from Heinrich 1956). B) Female genitalia of Z.

the body. The forewings are dark smoky gray,
with raised scales, colored brown and dark gray.
The hindwings are gray transparent, with the
costal area covered with gray scales. Male and fe-
male wing span is 19 mm. Males may be distin-
guished by a bushy, forward-projecting labial palp
and the presence of an aigrette of the maxillary
palps, which are reddish in color. Male antennae
have a basal sinus covered by strongly developed
gray scale tufts.
Female genitalia (Fig. 1B): signum developed
as a longitudinal, elongate, narrow and sinuous
form, armed with strong spines extending the
length of the bursa. The bursa has a plate armed
with strong spines near the junction with the duc-
tus bursae. The ductus seminalis extends from
the apical part of the bursa and it is slightly scle-
rotized. Ductus bursae are short and narrowed.

Description of Full Grown Larva

The total length of the last instar larva 15-16
mm and it is pinkish in color. The head is notable
for the presence of dark spots located on the dor-
sal and lateral surfaces of the frontal area (Fig. 2).
The stemmatal area is a semicircle and formed by
six stemmata (Fig. 3). Stemmata 3-5 are located
in straight line and higher with respect to the
other stemmata with the border sclerotized. Stem-

Fig. 2. Mature larva of Z. dixolophella, lateral view.

matal seta Sl is situated between stemmata 2
and 3. Stemmatal seta S2 is near the opening of
the stemmatal semicircle and placed in front of st-
emmata 4. The length that divides S2 from the st-
emmata 1 is close to 2/3 of the distance between
this same seta and stemmata 4. S3 is placed out-
side of the stemmatal semicircle. The length that
separates S3 of the stemmata 6 is similar to the
distance between S3 and S2. Pore Sa sited be-
tween stemmata 6 and S3. The length that sepa-
rates the pore Sa from stemmata 6 is the twice
that of the distance from S3. The substemmatal
seta SS1 is situated near to the mandible and sep-
arated from it by a quarter of the length that ex-
ists between this seta and stemmata 5. SS2 is
placed below stemmatas 5 and 6 and with the
same distance separating both stemmata. The
distance that separates SS2 of SS1 is the twice
that of the distance between SS2 and stemmata 5.
SS3 is placed below SS2 and forming a straight
line between them and stemmata 5. The distance
that separates SS2 and SS3 is the same as that
between SS2 and stemmata 5. Pore Ssa is situ-
ated inside of substemmatal area. The length that
divides Ssa from SS1 is the twice of the distance
between the same pore and SS2. The anterior
seta Al is placed in front of stemmata 2 and 3, an-
terior seta A2 is situated directly above Al. The
length that separates Al from A2 is similar to
that which divides Al from stemmata 3.

Fig. 3. Larval head ofZ. dixolophella, lateral view.

i~ Mt


Florida Entomologist 85(2)


Fig. 4. Larval head of Z. dixolophella, frontal view.

Pores Aa are placed above of A2. The length
that divides Aa from the ecdysial line is twice that
which separates Al from the ecdysial line. The
frontal setae Fl is situated near the adfrontal lat-
eral suture. Pores Fa are situated in the middle
part of the front. The distance between adfrontal
setae AF2 with respect to Fl is twice that which
separates AF1 and AF2. Lateral setae LI is
placed anterior to A3 and the separation between
both is the half of the length between LI and st-
emmata 3. The Postdorsal pore Pb is separated by
the same distance that exists between Pbl and
Pb2 setae. The clypeus has a pair of medium setae
C1 and a pair of lateral setae C2, the latter sited
in the intersection of the ecdysial line and the pro-
jection of the adfrontal lateral suture and the di-
viding line between clypeus and the anteclypeus.
The labrum is incised with three pairs of medium
labial setae and two pairs of lateral setae. The
mandibles bear the characteristic tridentate
structure with two basal setae. The intermediate
tooth projects beyond teeth, the oral surface con-
cave. The submentum carries two medium setae
and a bilobed posterior border, recurved towards
the dorsum. A well developed spinneret is
The thoraxic segment T1 with L group is bise-
tose. Segment T2 bears a sclerotized very dark pi-
naculum and evident on the base of the SD1 seta.
Segment T3 has a pinaculum on the SD1 seta but
is less conspicuous than in T2. The thoraxic spir-
acle is semicircular and larger than the abdomi-
nal spiracles, except for the spiracle on segment
A8, which is the same size and form as the tho-
raxic spiracle.
The prolegs are well developed in the abdomi-
nal segments A3-A6 and A10 (caudal). Each pro-
leg carries many crochets. The crochets are
arranged in an uniserial biordinal semicircle


June 2002

* *

from A3-A6. The internal face of prolegs measures
approximately 0.51-0.82 mm. The pinaculum of
SD1 seta of Al-A8 has a circular aspect. Spiracle
A8 is very large in comparison to the other ab-
dominal spiracles and clearly directed towards
the dorsocaudal position. Segment A9 with SV
group bisetose and L group trisetose. In segment
A10, the V setae are separated from each other by
about half of the distance that they are separated
from the V setae of segment A9. The V setae of
A10 are longer or equal to twice the length of the
V setae of A9.


Ten species have been described in the genus
Zamagiria. The related Z. pogerythrus is distrib-
uted from Campeche state, Mexico to Chejel,
Guatemala. The best known species of this genus
is Zamagiria laidion (Zeller), whose larvae also
feed on the leaves and flowers of M. zapota, as
well as on Manilkara emarginata (Sapotaceae)
and Eriobotyra japonica (Rosaceae). This moth
has been collected from the United States (Flor-
ida), Guatemala, Panama, Colombia, Bolivia,
Venezuela and Brazil. Another species of this gen-
era that feeds upon Sapotaceae is Zamagiria fra-
terna, which has been reported in Cuba attacking
Bumelia microcarpa (Heinrich 1956).
Oviposition of Z. dixolophella occurs on the
buds of sapodilla. Larvae feed upon the ovaries
and the petals of the flowers but frequently bore
into tender young shoots. Also, the larvae are
sometimes found inside sapodilla fruit. Prelimi-
nary observations have shown that insect infesta-
tion persists almost throughout the year,
however, the highest populations are found dur-
ing the peak of flowering.


We thank Armando Virgen and Alejandro del Mazo
for their help in collecting the biological material. Also
we thank Rodulfo Munoz Campero (Altamira), Esteban
Moises (Rancho el Nayar) and Abenamar Gonzalez
(Rancho Cazanares) for allowing us to collect material
from their farm.


BUTANI, D. K. 1975. Insect pests of fruit crops and their
control, sapota. Pesticides 9: 37-39.
HEINRICH, C. 1956. American Moths of the subfamily
Phycitinae. United States National Museum Bulle-
tin 207. Smithsonian Institution, Washington, D.C.
NEUNZIG, H. H., AND L. C. Dow. 1993. The Phycitinae of
Belize (Lepidoptera: Pyralidae) Technical Bulletin
304. North Carolina Agricultural Communications.
North Carolina State University. Raleigh, NC.
R. INGLES. 1987. The insects of nispero (Manilkara

Scientific Notes

zapota (L.) P. van Rogen) in Puerto Rico. J. Agric.
Univ. P.R. 71: 129-132.
MICKELBART, M. V. 1996. Sapodilla: a potential crop for
subtropical climates, pp. 439-446. In J. Janick [ed.],
Progress in new crops. ASHS Press, Alexandria, VA.
NORRBOM, A. L., AND K. C. KIM. 1988. A list of the re-
ported host plants of the species of Anastrepha
(Diptera: Tephritidae). U.S. Dept. Agric. APHIS-

RUBIO-ESPINA, E. 1968. Estudio preliminary de los insec-
tos perjudiciales a los arboles de nispero (Achras
zapota Linneus) en el estado Zulia, Venezuela. Revta.
Facul. Agronomia 1: 1-24.
SANDHU, G. S., AND C. S. SRAN. 1980. New records of
Lepidoptera on sapota. FAO Plant Protection Bulle-
tin 28: 43-44.
STEHR, F. W. 1987. Inmature insects. Volume I. Kendall/
Hunt Publishing Co. Dubuque, IA.

Florida Entomologist 85(2)


ALUJA, M., AND A. NORRBOM. 2000. Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior.
CRC Press; Boca Raton. 944 p. ISBN 0-8493-1275-2. Hardback. $169.95.

This book addresses the evolution and behav-
ior of tephritid fruit flies, many of which are of
significant economic importance worldwide. The
volume is the result of a symposium on the evolu-
tion of fruit fly behavior in honor of Drs. Ronald
Prokopy and D. Elmo Hardy, internationally rec-
ognized experts in the field and pioneers of the
study of tephritid fruit fly evolution and behavior.
The volume represents a compilation of sympo-
sium papers arranged into thirty three chapters
and eight sections. The chapters range from broad
topics such the history of the study of tephritid
fruit flies to empirical studies on the use of mito-
chondrial ribosomal DNA in the study of fruit fly
phylogeny. Nevertheless, they are appropriately
organized into Sections that reflect the thought-
fulness and care the editors took to present this
eclectic collection in a reasonably coherent and
logical format. Consequently, the editors achieved
their stated objective that this book, though a
"hybrid" of wide- ranging topics, should serve as a
general reference to specialists in various aspects
of fruit fly evolutionary biology and behavior.
Section I provides reviews of the phylogeny of
the Superfamily Tephritoidea (Chapter 1 by Kor-
neyev), the behavior of fly relatives (tephritoids)
of the Tephritidae (Chapter 2 by Sivinski), and
the history of studies on tephritids including
many of the World's most important Genera of
economic importance such asAnastrepha, Bactro-
cera, Ceratitis, and Rhagoletis (Chapter 3 by
Diaz-Fleischer and Aluja). Together, these three
chapters provide an excellent overview of the
three major themes in the book; phylogeny and
evolution, behavior, and aspects of fruit fly phero-
mones and attractants. Admittedly, Chapter one,
by virtue of the topic, presents a challenge to the
non-taxonomist, especially one unfamiliar with
tephritid systematics. Nevertheless, as the edi-
tors indicate, the book is intended primarily for
specialists. Even to the non-taxonomist however,
a perusal of the chapter allows for a better appre-
ciation of the breadth of the Tephritoidea, far be-
yond our narrow view of the tephritids that are of
agricultural importance. The Glossary will be an
invaluable tool for readers wishing to better un-
derstand the systematics and indeed, other spe-
cialty areas addressed throughout the book. For
specialists and non-specialists alike, the histori-
cal review of fruit fly research (Diaz-Fleischer and
Aluja) and discussion oftephritoid flies (Sivinski)
is refreshing and informative. They provide a
solid background for students and researchers
alike, who wish to study novel tephritid and re-
lated fly species.

Sections III through VI address the behavior
and phylogenetic relationships of the Subfamilies,
Blepharoneurinae and Phytalmiinae, Trypetinae,
Dacinae, and Tephritinae. The chapters within
these sections use various approaches (primarily
cladistics, morphological characters, allozyme vari-
ation, and in at least one case, mitochondrial DNA)
to address the phylogenetic relationships among
members of the respective Subfamilies. As with the
phylogenetic studies of many organisms, several as-
sumptions have to be made regarding convergent
and divergent evolution of a variety of morphologi-
cal (and even allozyme) traits. In some cases con-
trary results are obtained when comparisons are
made among some of the papers. Consequently,
there seems to be a need for the use of existing high
throughput DNA sequencing technologies to begin
to study tephritid fruit fly genomes. Indeed, other
than the few mitochondrial DNA studies presented
in this book, there appears to be no significant dis-
cussion in any of the chapters on the potential use
of DNA sequencing and analysis of specific te-
phritid genes to help resolve some phylogenetic
questions. For example, it would be of tremendous
value to analyze those genes involved in/related to
sexual selection, mating, courtship behavior, and
pheromone production. This book represents the
latest in the field of fruit fly phylogeny and behavior
but provides evidence for the need of complemen-
tary studies using state-of-the-art technology to
help resolve questions on tephritid fruit fly phylog-
eny and the evolution of mating systems.
Section VII contains an interesting and diverse
collection of papers addressing the genetics of
populations, evolution of feeding behavior, sexual
selection and life history traits, and oviposition
behavior. A discussion of Drosophila speciation in
Hawaii raises interesting issues that may also be
addressed in tephritid fruit fly systems. Section
VIII, the Glossary, provides definitions and expla-
nations of various terms used throughout the
book. These are especially valuable in navigating
the papers on tephritid taxonomy/phylogeny.
Overall, I found this book to be an excellent
and comprehensive compilation of research by
experts in the field of tephritid phylogeny and
behavior. I found few errors in the book. Each
chapter is clearly written and with the aid of the
glossary and other outside references, chapters
on taxonomy/phylogeny can be maneuvered to
some degree by non-specialists. The graphics are
satisfactory and the color photographs excellent.
The font size of the nucleotide sequences and cla-
distic tables made reading somewhat difficult.
This is an excellent book for the specialist and for

June 2002

Book Reviews

those seeking new tephritid species for research.
The major gap I found was the absence of nucle-
otide sequences of tephritids (some of which are
available in existing databases) that could help
address phylogenetic questions. The book could
have benefited from a section on the role of cur-
rent molecular technology (such as that used by

Drs. Handler and MacCoombs and colleagues in
Greece) that could help resolve questions on
tephritid fruit fly phylogeny and behavior.
Pauline 0. Lawrence
Department of Entomology and Nematology
University of Florida
Gainesville, FL 32611-0620

Florida Entomologist 85(2)

SCHAEFER, C. W., AND A. R. PANIZZI (eds.). 2000. Heteroptera of Economic Importance. CRC Press;
Boca Raton. (20+) 828 p. ISBN 0-8493-0695-7. Hardback. $94.95.

This book is a comprehensive and up-to-date
review of worldwide literature on Heteroptera
from the point of view of their economic impor-
tance, whether in managed or natural systems.
The book's most important contribution is as a
resource, not just for those involved in the man-
agement of agricultural pests, but for anyone
interested in true bugs. Its organization is so
practical as to invite the reader to simply read it
for edification and enjoyment or to find informa-
tion with immediate applicability. This book rep-
resents a clear guide for future research in
heteropteran biology, and its reviewed references
and the massive bibliography make it a necessary
reference book in entomological libraries.
Currently, Heteroptera, a suborder of the insect
order Hemiptera, include about 37,000 described
species, many of which feed on plants; thousands
more await description or discovery, according to
Schaefer and Panizzi (Introduction, Ch.1). Little is
known about the basic biology and ecology of most
true bugs. This book succeeds in thoroughly sum-
marizing and assessing present knowledge of the
true bugs and presents it in the form of current
reviews and references. The authors reviewed
thousands of articles and books, although not all
references are listed; nonetheless, the number of
references is impressive. For example, Sweet listed
925 references (Lygaeoidea, Ch.6). The references
include information on taxonomy and control of
true bugs, as well information on natural history,
behavior, morphology, embryology, endocrinology,
and ecology, etc. Some of these references guide
the interested reader to identification keys, but
which are not included in this book.
The scope of this book is beyond that of only
North America; for example, research is reviewed
from India, China, Brazil, Thailand, Poland, and
Ukraine. An insect or a plant that is considered a
pest or a beneficial species in the United States
may not be seen in the same way in other parts of
the world. The editors are to be praised for allow-
ing different points of view.
What makes true bugs economically impor-
tant? Many feed on plants, some of which trans-
mit plant-pathogenic viruses. An important
example is Eurygaster integriceps Puton, a seri-
ous pest of wheat (Scutelleridae, Ch.14). Because
heteropterans feed in a unique way, with pierc-
ing-sucking mouthparts, Hori (Ch. 2) explained,
their stylets bypass many of the plants' defenses
against biters and chewers, which also protects
them from many pesticides. Some plant-feeding
bugs are helpful for the control of pest plants; for
example, Zulubius acaciaphagus Schaffner (Aly-
didae, Ch.10) has helped to reduce the seed bank
ofAcacia cyclops, a weed introduced from Austra-
lia to South Africa.

Many true bugs are important predators of in-
sect pests and mites. De Clercq (Pentatomidae:
Asopinae, Ch. 32) stated that the present litera-
ture review has demonstrated the potential value
of several pentatomid predators for the manage-
ment of a wide array of agricultural insect pests.
This information could be extended to other Het-
eroptera families as well, such as Nabidae (Ch.
27), on which there is little information in gen-
eral, especially for the tropics. Berytid predators
(Ch. 31) have also received little attention, but
are potentially important. Mirids (Ch. 28) that
feed on delphacid planthopper eggs have been
used successfully in classical biological control
programs. Other zoophagous heteropterans, mostly
aquatic and semiaquatic, are economically impor-
tant natural enemies because they feed, in part,
on blood-sucking Diptera, especially mosquito
larvae and pupae (Ch. 21 to 25). Biological control
programs have to take into account that some of
these predacious species are cannibalistic, some
may damage crops, or may feed on beneficial
arthropods such as pollinators and spiders.
The authors in this book do not restrict their re-
views to Heteroptera associated with agricultural
crops, but also those associated with ornamentals,
plants not cultivated for major commercial benefit
such as sycamore trees (Tingidae, Ch. 4) and royal
palms (Thaumastocoridae. Ch. 5), and natural sys-
tems, where the intrinsic value of Heteroptera has
not been quantified.
This book is also an excellent resource for be-
ginning entomologists. There is a general descrip-
tion for each of the families included and a review
of their classification and their feeding behavior.
Homeowners will also find it useful because it
contains details about true bugs that occasionally
become nuisances in or around homes, such as
boxelder bugs (Rhopalidae, Ch. 9) which move
towards or into houses in large numbers in the
autumn. Even the general public will find it easy
to read and use because it is devoid of much jar-
gon. Apparently, it has very few errors in spelling;
the only one noticed was in the title of Ch. 8. It is
well organized and indexed. The book would have
benefitted, though, by including some photographs
of the most important pest or beneficial species.
This book is an important resource for health
professionals since it contains information on oc-
casional bites (Ch. 19) by non-blood-feeders,
which do not transmit pathogens, and on Chagas'
disease, of great medical and social importance in
the Western Hemisphere, that is transmitted to
people by infected triatomine bugs (Reduviidae,
Ch. 18). Cimicids (Ch. 17), or bed bugs, also feed
on human blood; studies on whether they can be
vectors for diseases such as hepatitis B, HIV, and
yellow fever are reviewed.

June 2002

Book Reviews

Entomologists and biological control special-
ists will find this book useful, but for heteropter-
ists, it is an essential book. The writers state the
strengths of previous and current research and
recommend research goals for the future. They
emphasize the value that basic biological infor-
mation has with respect to applicable work for the
control of heteropteran pests and the use of bene-
ficial species. Also, the value that good systematic
research has with respect to identification is dis-
cussed by the authors. Not all heteropteran fami-
lies are covered, for which I hope that a second

book of the same style will be written. As a worker
on Lygaeoidea, this book helps me to focus my in-
terests, guides my research and inspires me to
write. I highly recommend this book.
J. Brambila
Florida State Collection
of Arthropods
Division of Plant Industry
Florida Department of Agriculture
and Consumer Services
1911 S.W. 34 Street
Gainesville, FL 32608

Florida Entomologist 85(2)

HOWARD, F. W., D. MOORE, R. M. GIBLIN-DAVIS, AND R. G. ABAD. 2001. Insects on palms. CABI Pub-
lishing; Wallingford, Oxon, UK, xiv + 400 pp. ISBN 0-85199-326-5. Hardback. $120.00. [Sales in the
USA handled by Oxford Univ. Press, New York]

This book seems to be the first on the subject
and thus fills a void. It deals extensively and in-
tensively with the phytophagous insects and mites
that feed on or in palms, and with their natural en-
emies including biological control agents. As such,
it serves as a textbook for anyone wishing to pro-
tect palms from damage by insects and mites, or
from insect-transmitted diseases, or to encourage
pollination by insects. But, it is more than that be-
cause it reviews the behavior and ecology of the in-
sect and mite fauna of palms and makes the
literature available to anyone interested in the ac-
ademic subject of insect/plant relationships.
It is not a multi-authored book with four edi-
tors, but a book written by four authors. It is sub-
divided into only eight chapters; this was possible
only through good integration. F. W. Howard is
the major contributor to it. His opening chapter is
"The animal class Insecta and the plant family
Palmae" which has a useful 29-page condensation
for the uninitiated (e.g., most entomologists)
about the biology, cultivation and use of palms.
There are four pages of introduction for the unini-
tiated about insects. Detailed information about
the insects is, of course, in the other chapters.
The other chapters are: 2, Defoliators of palms;
3, Sap-feeders on palms; 4, Insects of palm flow-
ers and fruits; 5, Borers of palms; 6, Population
regulation of palm pests; 7, Principles of insect
pest control on palms; and 8, Field techniques for
studies of palm insects. The chapters are followed
by a 47-page section of integrated References, and
this by a 20-page integrated Index. Tables in the
text summarize a lot of information usefully. Ta-
ble 3.3 is a list of species of Derbidae (Hemiptera:
Auchenorrhyncha) reported on palms, with host
palms, distributions and literature references.
Table 4.4 lists, by palm genus, arthropods associ-
ated with pollination of palms. It summarizes ar-
thropod behavior, but how can it write "larvae
probably breed in the flowers" when breeding is a
function restricted to adults? Boxes in the text de-
velop items of especial interest that would other-
wise destroy the flow, such as Box 2.2, the story of
classical biological control of a coconut leaf-miner
in Fiji, and Box 5.2, an account of red-ring dis-
ease. The book's numerous black and white photo-
graphs, including electron micrographs, serve
admirably. It even has 16 plates of color photo-
graphs, most of which have been reproduced well.

I have few criticisms. The book (p. 3) claims
that insect biogeography "lacks standardized
place-names." If by this is meant that insect bio-
geography uses names for biogeographical re-
gions that have evolved from some proposed in
the mid-19th century, that is true. But, in making
this claim, the book equates the outmoded expres-
sion "Ethiopian Region" with all of Africa, when
for over two decades sub-Saharan Africa with
Madagascar has been called the "Afrotropical Re-
gion" (Crosskey & White 1977). The northern tier
of African countries has long been attributed to
the Palearctic Region. It asserts (p. 94) that pres-
ence of a mite, Pyemotes ventricosus, detected in
Fiji in 1921, was due to "inadvertent introduc-
tion." This makes three assumptions: 1, that the
mite is not native so must be adventive; 2, that it
was not introduced (deliberately) so it must be an
immigrant; and 3, that it did not arrive by natural
means so must have arrived as a hitchhiker in
some sort of cargo transported by humans. Evi-
dence for those assumptions is not presented. The
weevil Metamasius mosieri, which is stated (p. 279)
to be associated with palms in tropical America,
in fact feeds on bromeliads as an adult and larva
(Larson et al. 2001). Anyone who expects to use
the book as a source of describer (author) names
for the scientific names of palms and their associ-
ated insects and mites will be disappointed-they
are not given. The book's rather high price ($120)
will surely deter some purchasers who are inter-
ested but uncommitted. But, it is carefully edited
and largely free of the typographical and gram-
matical errors, and bloated bureaucratese expres-
sions that are all too common in many technical
books. This makes it a pleasure to read.
J. H. Frank
Entomology & Nematology Dept.
University of Florida
Gainesville, FL 32611-0630


CROSSKEY, R. W., AND G. B. WHITE. 1977. The Afrotropi-
cal Region. A recommended term in zoogeography.
J. Nat. Hist. 11: 541-544.
LARSON, B. C., J. H. FRANK, AND 0. R. CREEL. 2001.
Florida bromeliad weevil. (Available: http://creatures.
ifas.ufl.edu/orn/M mosieri.htm)

June 2002

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

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