Title: Florida Entomologist
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Permanent Link: http://ufdc.ufl.edu/UF00098813/00114
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Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1979
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
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Volume ID: VID00114
Source Institution: University of Florida
Holding Location: University of Florida
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(ISSN 0015-4040)

Volume 62, Number 1 March, 1979


Symposium: Sociobiology of Sex
LEPPLA, N. C.-Sociobiology of Sex ~.1....--.------. ...... ............ 1
in Male Parasitic Wasps of the Genus Megarhyssa: I. Sexual
Discrimination, Tergal Stroking Behavior, and Description of
Associated Anal Structures .............--- -..... .. ......-............ ........ 3
MORRIS, G. K.-Mating Systems, Paternal Investment, and Aggressive
Behavior of Acoustic Orthoptera ... .... ~. ~.. ......... .. 9
LLOYD, J. E.-Mating Behavior and Natural Selection ......................... 17
-- -- -- --^ ^--- -^-
Sex Pheromone Biology of the Adult Tomato Pinworm, Keiferia
lycopersicella (W alsingham) -. -- .... -------...........__ 35
BROWER, J. H.-Mating Competitiveness in the Laboratory of Irradi-
ated Males and Females of Ephestia cautella .... .......- ......__ 41
SHIH, C. I., S. L. POE, AND H. L. CROMROY-Biology, and Predation of
Phytoseiulus macropilis on Tetranychus urticae -....-----........... 48
NIGG, H. N., R. F. BROOKS, AND R. C. BULLOCK-Chlordane Residues in
Florida Citrus Soils .......---............ .. ...................... .. 54
Ru, N., AND R. I. SAILER-Colonization of a Citrus Whitefly Parasite,
Prospaltella lahorensis, in Gainesville, Florida -... ............-.... 59
WANI, R. L., S. L. POE, AND G. L. GREENE-Emergence Pattern of the
Sorghum Midge, Contarinia sorghicola, and its Parasite, Apro-
stocetus diplosidis ------- -......... ................ ........ 65

Scientific Notes
RU, NGUYEN AND R. B. WORKMAN-Seasonal Abundance and
Parasites of the Imported Cabbageworm, Diamondback
Moth, and Cabbage Webworm in Northeast Florida .... 68

Continued on Back Cover

Published by The Florida Entomological Society



President .....
Vice-President .
Secretary .......
Treasurer --------.

Other Members of Executive Committee

-- R. F. Brooks
..... N. C. Leppla
...--. F. W. Mead
E. S. Del Fosse

J. B. Taylor
R. E. Brown
C. A. Musgrave
R. C. Bullock
A. K. Burditt, Jr.
W. L. Peters


Associate Editors

C. A. Musgrave
.... A. B. Hamon
J. E. Lloyd
J. R. McLaughlin
C. W. McCoy
H. V. Weems, Jr.
E. S. Del Fosse

Business Manager

THE FLORIDA ENTOMOLOGIST is issued quarterly-March, June, Septem-
ber, and December. Subscription price to non-members is $15.00 per year in
advance, $3.75 per copy. Membership in the Florida Entomological Society,
including subscription to The Florida Entomologist, is $10 per year for
regular membership and $2 per year for students. Inquiries regarding
membership and subscriptions should be addressed to the Business Manager,
P. O. Box 12425, University Station, Gainesville, FL 32611. The Florida
Entomologist is entered as second class matter at the Post Office in DeLeon
Springs and Gainesville, FL.
Manuscripts and other editorial matter should be sent to the Editor,
Department of Entomology and Nematology, Archer Road Lab-Building 339,
University of Florida, Gainesville, 32611. Business matters for other Society
officers can be sent to that individual at the University Station address above.
When preparing manuscripts, authors should consult "Instructions to
Authors" on the inside cover of all recent issues.

This issue mailed March 7, 1979

Symposium: Sociobiology of Sex




Insect Attractants, Behavior and Basic Biology Research Laboratory
Federal Research, Science and Education Administration, USDA
P.O. Box 14565, Gainesville, FL 32604

This intriguing topic, developed by combining the unifying concepts of
sociobiology with particularly illustrative examples of mating behavior in
arthropods, was presented originally as an exceptionally popular sym-
posium. Sociobiology, an emerging and therefore somewhat controversial
field, provides a framework for exploring the genetic bases, ultimate causes,
and evolutionary consequences of animal behavior. Sexual reproduction, the
other component of our topic, is an extremely costly yet pervasive form of
propagation-one that commits an apparently inordinate amount of time and
physical energy to a seemingly overcomplicated activity. We observe differ-
ential parental investment and almost automatically question its relation-
ship to associated mating strategies and sexual selection. We ponder, for
example, the adaptive significance of complex and prolonged courtship dis-
plays that include the use of elaborate chemical and physical cues. How can
the fitness of insects be maximized by devoting an entire developmental stage
almost exclusively to this singular activity? We intended that this sym-
posium would stimulate these kinds of questions by proffering relevant ob-
servations and preliminary interpretations. Our goal was to create an aware-
ness and appreciation for the subtleties of insect reproduction among
entomologists who might otherwise consider the subject esoteric.
The conceptual framework for our discussion may be gleaned from E. O.
Wilson's prodigious monograph, Sociobiology, the New Synthesis, and the
equally intriguing Sociobiology and Behavior by David Barash. Wilson
loosely defines sociobiology as "the systematic study of the biological bacis of
all social behavior" and emphasizes animal societies, population structure,
communication, and the physiology that underlies these social adaptations.
Both he and Barash based their compendia on the work of R. D. Alexander,
R. A. Fisher, W. D. Hamilton, M. J. Smith, R. L. Trivers, and G. C. Williams,
among other pioneering zoologists, who analyzed animal behavior in evolu-
tionary terms and thereby discovered unifying principles that apply to both
social and solitary sexually reproducing organisms. The quintessence of
sociobiology is examined by Richard Dawkins in The Selfish Gene, which
describes societies and individuals (survival machines) as mere vehicles of
genetic evolution. For our purposes, however, the "Sociobiology of Sex" ap-
plies to all animals that engage in any form of sexual reproduction.

'Adapted from a symposium, "Mating Behavior: Sociobiology of Sex," co-organ:zed by
N. C. Leppla and J. E. Lloyd and presented at the Fifty-Second Annual Meeting of the
Entomological Society of America-Southeastern Branch, held in Gainesville, FL, on 26 Jan-
uary 1978.

The Florida Entomologist 62(1)

March, 1979

Since comparative animal behavior is fundamental to sociobiology, we
must understand the historical development and current state of this field
before we can discern the ways in which the two are linked. Animal behavior
has been studied from the time that man first experienced an interdependence
with other animals. Indeed, the classical works of Aristotle, Pliny, Descartes,
Darwin, Fabre, and their respective contemporaries abound with behav-
ioral anecdotes. However, it was not until 1894 that Lloyd Morgan founded
the so-called "science of animal behavior" by advocating quantitative ob-
servation rather than anthropomorphism and teleology. In 1910, Oskar
Heinroth established ethology as the study of species-specific, instinctive
behavior patterns and the discipline subsequently proliferated in Europe
primarily through the efforts of Konrad Lorenz, Niko Tinbergen, and their
associates. Specificity is determined by the comparative method, and in-
stinctive behavior includes that which is genetically programmed and
environmentally mediated. Both aspects emphasize the adaptive significance
of behavior and therefore necessitate an evolutionary approach. Now
ethology generally is known as the study of animal behavior in nature.
The stereotyped character of insect behavior is ideally suited to intensive
ethological analysis and this has prompted such prominent zoologists as Karl
von Frisch, T. C. Schneirla, Howard Evans, Charles Michner, G. P.
Baerends, and Erik Nielsen to select insects as their experimental animals.
ITheir dissections of complex behavioral sequences ultimately yielded
"ethograms" composed of discrete elements. However, these kinds of subtle
elements often are inaccessible, being integrated into complex hierarchies
with built-in contingencies and alternatives. Others may operate by means
of simple switches, with the triggering stimuli either present or absent. In
any case, behavioral repertories must be described before we can understand
the reproductive strategies of insect species.
Our distinguished authors represent a new generation of ethological in-
quiry. Robert Matthews is well-known for his detailed studies of social wasps
and he just coauthored Insect Behavior, the first comprehensive text on this
subject. His paper introduces the topic by describing ongoing studies of
sexual behavior associated with aggregation in male Megarhyssa wasps. His
approach demonstrates the importance of relating observed behavior to the
subject's morphological capabilities. Glen Morris has concentrated his sci-
entific efforts on the aggressive behavior of acoustic Orthoptera. His article,
therefore, focuses on specific mating systems, relates them to paternal in-
vestment, and culminates by enumerating the conditions under which ag-
gression escalates. James Lloyd, internationally recognized for his research
on the photic communication of fireflies, concludes the symposium by em-
phasizing relationships between systematics and behavior. He uses fireflies
and bed bugs to illustrate the use of a one gene analysis model, describes
and speculates on several unusual mating systems, and incorporates Smith's
concept of evolutionarily stable strategies. These papers are enhanced con-
siderably by the incorporation of many previously unpublished ideas and
much new information.

We thank C. O. Calkins (Insect Attractants Laboratory) and T. J.
Walker (University of Florida) for their assistance in developing the
symposium; A. N. Sparks and H. R. Gross, Jr. (Southern Grain Insects

Symposium: Sociobiology of Sex 3

Laboratory), and L. C. Kuitert (University of Florida) for including us in
an already crowded program and personally handling our local arrange-
ments; and the Florida Entomological Society for their gracious offer to
publish these proceedings.


Department of Entomology
University of Georgia
Athens, GA 30602

Parasitic wasps comprise a species-rich group for which a number of
casual and fragmentary observations on mating and courtship exist, but
few detailed studies have been published (Matthews 1975). A singular
phenomenon among them is provided by the pre-mating behavior of
Megarhyssa, a genus of large, long-tailed ichneumon wasps parasitizing
Tremex horntail larvae in dead trees. Males form conspicuous mixed-species
aggregations on the bark of trees from which new adults are actively
emerging (Fig. 1). All facing inward around a circular area of about 50
mm diameter, they jostle about and jockey for position, but show little overt
aggression toward one another. When a female wasp finally chews through
the bark, a conspecific male quickly mates with her, and the other males at
least temporarily disperse. The most detailed previous studies of Megarhyssa
behavior are those of Heatwole et al. (1963, 1964) and Heatwole and Davis
(1965) in which most references to earlier observations may be found.
Most previous reports of Megarhyssa male aggregation have asserted
that species identification and sex discrimination are determined as the
emerging adult penetrates the outer bark surface. Several workers have
noted that the waiting males often probe bark crevices with the tips of their
abdomens, including penetrating the partially chewed out emergence hole.
Nuttall (1973) and others have suggested that this may result in fertiliza-
tion of the female within its tunnel. However, this idea has usually been
discounted because of the relatively greater length of the female abdomen
as compared to the male.
During the summers of 1977 and 1978 we studied male aggregation in 3
species of Megarhyssa found in beech-hemlock forests of an upstate New
York biological station, the E.N. Huyck Preserve in Rensselaerville. Our
study sites consisted of 8 isolated, dead but still standing beech trees. The
maximum distance between any 2 trees was less than 100 m. Among the
results described in this paper are a previously unappreciated behavior, a
possible new gland, and the shedding of some new light on previous interpre-

'Hymenoptera: Ichneumonidae.

The Florida, Entomologist 62(1)

March, 1979


Fig. 1. Typical aggregation of several male Megarhyssa wasps at an emer-
gence site on a dead beech. Two species are present and several individuals
are marked with spots of paint to permit recognition.
stations of the nature and function of male aggregation in Megarhyssa.
In contrast to searching or patrolling males, those in aggregations are
unusually tenacious; if disturbed they may fly away but normally will re-
group within minutes at the same precise spot on the tree. On a longer time
scale, males also show a remarkably high fidelity to particular host trees,
returning to patrol them day after day. Because of this tenacity, once an
aggregation is found, it is relatively easy to follow it through time and, by
.distinctively marking males with spots of Testor's@ enamel, to monitor in-
dividual interactions. For example, 'in July 1977, 45 males were marked and
the host trees checked at least once daily during the following month.2 Of
these, 15 were later seen again, and 64% of these recaptured individuals
were found patrolling the same tree from which they had originally been
recorded. This tendency to return to a particular tree was previously noted
by Abbott (1934) and Heatwole and Davis (1965) and applies to both sexes.

2One male was sighted 24 days after being marked, a longevity which concurs with
Heatwole and Davis' (1965) record of 27 days for 1 male.

r~ 9

Symposium: Sociobiology of Sex

The latter authors obtained an even higher fidelity to particular trees (81%
and 100% for 24 and 15 males, respectively, marked in 2 consecutive years).
Within a given aggregation, the behaviors which occur seem at first
quite random. In our study, these commonly included mutual antennation,
antennal lashing by 1 male upon the body of another, bouts of wing vibra-
tion, abdomen flexing, probing crevices with the abdominal tip, and head
butting, all of which confirmed the previous observations of other re-
searchers. Both before and after the emerging adult chewed through the
bark, some males exhibited a very distinctive behavior which we called
tergall stroking". Males performing this behavior adopt a stereotyped
posture (Fig. 2), with the abdomen thrust well forward between the legs
until its dorsal surface touches the substrate, its tip lying between the tips
of the antennae. Only the dorsum of the ultimate abdominal segment
touches the bark; the genitalia at the tip of the abdomen do not contact the
substrate (contrary to observations of similar behavior mentioned by Heat-
wole et al. 1963), nor are the claspers extended. In this position, the male
wasp repeatedly thrusts its abdomen forward in short, anteriorly-directed
strokes along the substrate.
On males of all 3 species of Megarhyssa, there is a large membranous
and partially eversible sac (Fig. 3, inset) opening at the apex of the last
(8th) tergite, adjacent to the base of the genital capsule. The occurrence of
this possible anal gland does not seem to have been reported previously. On
its dorsal surface, scanning electron microscopy reveals the presence of a
dense tuft of setae resembling an applicator brush (Fig. 3). Although close
examination of this anal sac after treatment with 10% potassium hydroxide
solution failed to reveal a cuticle-lined glandular structure, it is possible
that tergal marking materials are derived from cells of the hind gut, which

Fig. 2. Sketch of a male Megarhyssa macrurus (L.) performing the
tergal marking behavior. (Drawing by Joan W. Krispyn)

The Florida Entomologist 62(1)

Fig. 3. Scanning electron micrograph of partially everted "anal gland" of
Megarhyssa greenei Viereck showing the dense tufts of setae which form a
brush-like applicator (dorsal aspect, 200x). Abdomen of a male Megarhyssa
macrurus showing the large membranous area on the posterior dorsal sur-
also terminates at this point. Hand-held males will readily excrete a droplet
or 2 of fluid, possibly liquid waste products, from this opening. No odor is
discernable from the male anal area. Although a possible protective odor
has been noted for Megarhyssa (Townes 1939), our observations indicate
that this odor appears to emanate from glands in the head. Whatever further
study clarifies about the histology of this possible gland, analysis of films of
individuals performing tergal stroking make it quite apparent that the setal
brush actively strokes the substrate. A closely similar behavior is that
which Heatwole et al. (1963) term "dipping," in which males bend the
abdominal tip downwards and drag it along the substrate for several centi-
meters, in an action reminiscent of trail-laying in ants. We have also ob-
served this behavior on a few occasions, but it differs from tergal stroking in
that the abdomen is arched so that the tip of the genitalia contact the bark.
Also dipping tends to be performed while walking over the bark, rather than
at aggregation sites.

March, 1979

Symposium: Sociobiology of Sex 7

In 1978, 16 aggregations involving marked individuals were studied for
extensive time periods. Thirteen of these aggregations were of mixed species
composition, the other 3 consisting of a single species. In each case, 1 or 2
individuals performed the majority of the tergal stroking bouts and crevice
insertions. When the aggregation was artificially dispersed, those returning
appeared to maintain the same relative status, and these dominantss" re-
sumed their stroking bout with the same frequency as they had shown previ-
ously. Such "dominance" may be simply a reflection of internal milieu. It
may also be related in some manner to conspecificity with the emerging
adult, although in 3 of the 13 mixed species aggregations the emergent was
of a different species from the "dominant" stroking/inserting male.
Tergal stroking does not appear to be related to the sex of the emerging
individual. Males in aggregations are much less discriminating in this re-
gard than has previously been thought. It appears that little sex discrimina-
tion is taking place at all at the moment the new adult first penetrates the
bark, as previous workers have asserted (Heatwole et al. 1964). Even when
males subsequently emerged, tergal stroking and insertion proceeded, un-
abated, up to the time of actual mounting attempts. On several occasions, as
the new male emerged it was mounted almost immediately, and repeated
copulatory attempts were made by the aggregated males.
By affixing small screen cages over emergence sites where males were
congregating, we were able to record the sex and species of the emerging
adults for 35 aggregations. Of these, 12 yielded males and 23 produced fe-
males. Males aggregated with equal intensity at sites from which either sex
emerged, and no behavioral differences correlated with the future emergent's
sex were demonstrable. Although Heatwole et al. (1964) observed 31 ag-
gregations no data are recorded for the outcome of 20 of these except that 3
yielded wood-boring insects other than Megarhyssa. The remaining 11 all
yielded females.
The functions) of the anal gland and of tergal stroking are unknown.
Were a clear-cut sexual bias evident, classic possible roles might include
aphrodisiac effects or other involvement in female receptivity alteration,
territorial marking, or an olfactory "display" function. We suspect that it
may be related instead to the behavior in which males repeatedly attempt to
insert their abdomens into bark crevices and emergence holes. Frequently
they penetrated the emergence hole, thrusting their abdomen deeply (often
all the way to the thorax) and snugly along the side of the emerging adult.
The positional relationship of the inserting male to the emerging adult is
such that the setal brush rubs along the body of the emerging adult. It
seems unlikely that such behavior is attempted copulation, given the fre-
quency with which such insertion behavior occurs at sites from which males
subsequently emerge.
If the rate at which receptive females are encountered determines a
male's overall reproductive success, selection should favor extreme male
mobility or the guarding of areas where receptive females are most likely to
be found (Parker 1970, 1973). It should also favor any other process which
increases the rate at which females are encountered. Tergal stroking and
insertion might both contribute to such an increase. For example, the
propensity to perform tergal stroking at all emergence sites irrespective of
sex or species of the emerging individual may serve to reinforce the male's
tendency to return to particular trees. This would be adaptive from the

The Florida Entomologist 62(1)

March, 1979

male's standpoint, since if Megarhyssa are emerging from the tree, there is
a strong likelihood that more adults will issue from the same tree in sub-
sequent days and that at least some of them will be conspecific females.
Concurrently, time invested with a slowly emerging male might cost a
female emergence to be missed on the other side of the tree. Selection would
be expected to favor any male behavior which minimized the time invested in
unfruitful aggregations, as well as behaviors increasing the rate of success-
ful encounters. Pre-mating discrimination, although potentially advanta-
geous, apparently does not occur. If the insertion behavior hastens the
emergence of the new adult which is being stroked, it would be advantageous
for a male to perform this behavior with any emerging wasp regardless of
sex or species. It would decrease the length of time he would need to invest
at a particular emergence site. Further study of the sexual behavior of male
Megarhyssa wasps will clearly be revealing in the context of sexual aggrega-
tion and discrimination.


This research was supported by grants from the Edmund Niles Huyck
Preserve, Rensselaerville, New York. Our special appreciation is extended to
Joan Krispyn who executed the ink drawing and to Karl Scott who took the
SEM photograph. Kathy Jablonski, Wayne Gardener and Kenneth Suarez,
members of an EARTHWATCH team assigned to the E. N. Huyck Preserve,
were enthusiastic and helpful assistants during 1977.


ABBOTT, C. E. 1934. Notes on Megarhyssa lunator. Psyche 41:238-40.
HEATWOLE, H., D. M. DAVIS, AND A. M. WENNER. 1963. The behavior of
Megarhyssa, a genus of parasitic hymenopterans (Ichneumonidae:
Ephialtinae). Zeit. Tierpsychol. 19:652-64.
AND 1964. Detection of mates and hosts by parasitic
insects of the genus Megarhyssa (Hymenoptera: Ichneumonidae).
Amer. Midi. Nat. 71(2) :374-81.
AND 1965. Ecology of three sympatric species of parasitic
insects of the genus Megarhyssa (Hymenoptera: Ichneumonidae).
Ecology 46:140-50.
MATTHEWS, R. W. 1975. Courtship in parasitic wasps. Pp. 66-86 In P.W.
Price, ed., Evolutionary Strategies of Parasitic Insects and Mites,
Plenum Publ. Co., N.Y.
NUTTALL, M. J. 1973. Pre-emergence fertilization of Megarhyssa nortoni
(Hymenoptera: Ichneumonidae). N. Z. Ent. 5:112-7.
PAEKER, G. A. 1970. Sperm competition and its evolutionary consequences in
the insects. Biol. Rev. 46:528-68.
1973. Courtship persistence and female-guarding as a male time
investment strategy. Behaviour 48:157-82.
TOWNES, H. K. 1939. Protective odors among the Ichneumonidae (Hy-
menoptera). Bull. Brooklyn Ent. Soc. 34:29-30.



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Symposium: Sociobiology of Sex


Department of Zoology and Erindale College
University of Toronto, Mississauga

The term mating system has been defined as a "behavioral strategy em-
ployed in obtaining mates" (Emlen and Oring 1977). Such a definition makes
differences in the mechanisms of pair formation the logical basis of dis-
tinguishing and classifying system types. For crickets, katydids, and many
grasshoppers, sound signals have a special importance to mating. Among
these acoustic Orthoptera 2 broadly distinct mating systems recur fre-
quently: male call and male search.
The males in a call system remain geographically localized for long
periods and by repetition of an acoustic display, the calling song, elicit the
approach of non-singing females. The calling song functions as a naviga-
tional aid, the basis of phonotactic response. By contrast, the males of a
search system are not sedentary. They actively explore their environment
for females. Instead of functioning as a fixed beacon, sounds call attention
to the momentary location of an exploring male or they occur in conjunction
with visual and chemical signals exchanged at close range with encountered
A meadow katydid, Orchelimum gladiator Bruner (Tettigoniidae), and a
band-winged oedipodine grasshopper, Dissosteira carolina (L.) (Acrididae),
provide a contrasting illustration of male call and male search types, re-
Orchelimum gladiator live in visually occluded swale habitats where they
form aggregations of relatively sedentary, stridulating males. Within these
aggregations, likened by Alexander (1975) to the leks of certain vertebrate
species, males repeat through daylight hours, a calling song consisting of a
series of ticks alternating with a buzz. Each song is only a few seconds in
duration, but 1 follows another in rapid succession and the result is an
acoustic display remarkable for its continuity. At peak densities, singers
within the aggregation are separated from their nearest stridulating
neighbor by an average distance of 1.7 m (Morris 1967). Experiments with
speaker broadcasts of recorded calling song have shown that the song
evokes and guides the approach of both sexually receptive females (Morris
et al. 1975a) and of nearby male competitors (Morris 1972).
Dissosteira carolina occurs in areas with little or no vegetation. During
the heat of the day males walk and make short flights (1 to 3 m) with no
apparent goal (Kerr 1974). Over the last part of these flights their yellow-
bordered black wings move in a visual display accompanied by a crackling
sound known as crepitation. As they detect other D. carolina, the males ap-
proach by flying and/or walking. After arriving next to a conspecific male or
female, a male invokes a diverse repertoire of signals involving movement of
his jumping legs. Certain of these signals, e.g. femur-tipping with wing
striking (Kerr 1974) generate sound. Dissosteira carolina also produces a

The Florida Entomologist 62 (1)

March, 1979

hovering crepitation display in which a male flies up 0.5 to 2 m, crepitates
without lateral displacement, and then returns to approximately its original
ground location. Only other males have been observed to approach these
crepitating flight displays (Otte 1970).
The strategy of generating a beacon by which the female will navigate
requires that the male provide her, for at least many minutes, with a rela-
tively fixed goal. Orchelimum gladiator males therefore show strong site
fidelity. This in turn means that a higher proportion of their competitive
interactions will involve a rather small group of nearby males. The system
tends to be static with reduced social contact. Conversely, for D. carolina
crepitating flight draws conspecifics to a site only momentarily occupied by
the calling male. Crepitating displays are ephemeral in both time and space.
Such a system is more dynamic and the incidence of contact with strangers
very high. Individuals do not show fidelity to display sites.
The site fidelity of call systems is most marked where the singer oc-
cupies a burrow. Field crickets (Gryllus spp.) have a burrow-based call
system. They defend geographically fixed territories for which the burrow is
the focal point (Alexander 1961). Orchelimum gladiator males do not con-
struct or occupy burrows. They apparently shelter in the lower reaches of
the grass and sedge. Consequently, within limits of habitat suitability, they
are indifferent to their exact geographical location; the area of their activity
(one might better say the "volume" of their activity since they move
vertically as well as horizontally within the vegetation) drifts many meters
over the course of several days (Morris 1967).
The signalling in D. carolina's search system is intermittent and op-
portunistic. It is given in response to an encountered conspecific. What fol-
lows initial contact is a rapid alternation of the roles of sender and recipient
as the 2 (or more) insects exchange leg signals. By contrast the signalling in
a call system is sustained and endogenous. No evoking stimulus, such as
another conspecific, is required. Information is broadcast rhetorically, with-
out anticipation of imminent female response. For hours on end the sender
remains the sender and the recipient, if even present, remains the recipient.
The mating systems of most Tettigoniidae including 0. gladiator, in
contrast to those of most crickets and grasshoppers, seem to involve sub-
stantial nutritional investment by the male in his future offspring. This in-
vestment takes the form of a remarkably large gelatinous mass, the
spermatophylax, which accompanies the sperm ampullae and is consumed by
the female after copulation.
The food value of tettigoniid spermatophores (sperm ampullae and
spermatophylax) is practically unknown. But in the meadow katydid
Conocephalus brevipennis (Scudder) about 20% by weight of the spermato-
phore is protein, the remainder being largely water (personal communica-
tion, Gordon E. Kerr). The spermatophore with its spermatophylax is
such a substantial fraction of a male's body weight, as much as 25% in
Ephippiger bitterensis Finot (Busnel et al. 1956), that taken together with
its probable high protein content, it is best viewed (Thornhill 1976) as
paternal investment.
In addition to the spermatophore and various glandular products (Fulton
1915), nutritional investment by males of acoustic Orthoptera can extend to
female consumption of male body parts. For example, males of the
tettigoniid Cyphoderris buckelli Hebard possess fleshy, cream-colored wings

Symposium: Sociobiology of Sex

beneath their tegmina upon which the female feeds during copulation, con-
suming both wing substance and accompanying haemolymph.
Cyphoderris buckelli is also an instructive example of the potential for
female exploitation of the male, which is inevitably linked to the evolution of
a sizable paternal investment. This species, occurring in northwestern North
America, is one of only 4 survivors of an ancient and large insect group, the
Haglidae (Prophalangopsidae) considered ancestral to modern-day katydids
and crickets (Zeuner 1939). Males produce their calling songs at night from
low understory shrubs and the lower trunks of trees. On reaching a male,
the female mounts over his back, the primitive mating posture of ensiferan
Orthoptera (Alexander and Otte 1967) and begins to chew on his wings
(Fig. 1A). A substantial spermatophore, suggesting tettigonioid rather than
grylloid affinity, is passed to the female.
Typically, female katydids can only obtain the spermatophore food
package by submitting to insemination. But in C. buckelli it would be possible
for a female to begin copulation, obtain nourishment via a male's wings and
then withdraw before the male succeeds in attaching his spermatophore.
Mated females might pursue coitus interruptus with a succession of partners
as a foraging technique. The presence of what Hinton (1946) terms a gin
trap (pinching organ) on the male's abdominal tergites testifies to past
selection that limits this sort of exploitation. Cyphoderris buckelli's gin trap
(Fig. 2) consists of recurved hooks, 1 pair directed forward and another
directed backward, on the 10th and 8th tergites, respectively. As the terminal
abdominal segments are telescoped inward, the hooks converge on a portion
of the female's venter and prevent her from pulling away (Fig. 1B).
Katydid males, via their spermatophores, make a large parental invest-
_~ Ci n.q~l a"* i3 PbS -~ aI;L~~

Fig. 1. A male and female of Cyphoderris buckelli Hebard copulating on
the forest floor near Kelowna, B.C. Visible are the spermatophore, the
chewed and bleeding ends of the male's wing, and the strain folds of the
female's abdomen produced as she pulls against the restraint of the male's
gin trap.

The Florida Entomologist 62(1)




Fig. 2. Rear portion of abdomen of a Cyphoderris buckelli Hebard male in
dorsal view showing the gin trap.
ment with each copulation. In oedipodines and other Acrididae, the spermato-
phore has been reduced in the course of evolution to a temporary sac-like
extension of the penis, penetrating to the spermatheca (Davey 1960).
Oedipodines thus invest little in mating beyond their sperm. Low parental
investment by a sex is linked to the practice of multiple mating by that sex
and to increased variance in reproductive success (Trivers 1972, Wilson
1975). Oedipodine males will tend to be polygynous and compete with other
males of their species to inseminate as many females as possible. Katydids
on the other hand will tend toward monogyny; having apparently evolved
with a lower expectation of contacting mates, they invest heavily in the 1 or
2 matings that they do secure.
There is abundant experimental evidence that the calling song in male
call mating systems governs pair formation and that it evokes and guides
the approach of receptive females (Regen 1913, Duijm and van Oyen 1948,
Busnel et al. 1955, Walker 1957, Morris et al. 1975b). Use of this same
calling song to simultaneously mediate male competitive interactions, though
probably next to universal in Tettigoniidae, has been neglected as a subject
for experimentation and understated in the literature.
Most tettigoniids never or only rarely fight. But among males of
Orchelimum vulgare Harris and 0. gladiator, overt intraspecific aggression
is common (Morris 1971). Such fighting is ultimately a manifestation of

March, 1979

Symposium: Sociobiology of Sex

male competition. Somehow the dominant male must be improving his access
to females.
An Orchelimum singer will periodically 'track' a neighbor's song se-
quence by interposing his own songs in a fixed time relationship (personal
communication, Marianne Feaver). Then suddenly he may move from the
relative permanence of his singing post to walk and leap a meter or more in
the direction of the other male. A grappling fight often ensues, involving
vigorous biting and kicking. So far 6 species of Orchelimum are known to
engage in grappling aggression.
The role of Orchelimum calling song in aggression also has been demon-
strated experimentally. Orchelimum gladiator males will track song broad-
cast from a distant speaker, then make a phonotactic approach. If the
speaker is repositioned one can evoke further approaches (Morris 1972).
There can be no doubt that Orchelimum calling song releases attack by
nearby males. The tracking leads us further to assign a role for the song in
the threat of attack.
Among Tettigoniidae a high incidence of overt aggression is known only
in Orchelimum, but tracking and other forms of phonoresponse are almost
universal in the family. Some type of male-male mutual calling song influ-
ence (alternation, synchrony) seems to have been uncovered wherever it has
been sought (Jones 1966, Shaw 1968). Given that in Orchelimum phono-
response via calling song is aggressive, then we may suppose that in other
katydid species, even though overt aggression is absent, the occurrence of
phonoresponse reflects the function of calling song in mediating male com-
petitive interactions.
In support of this one can cite also the universal tendency for regular
spacing of singing katydid males (Alexander 1956). Such regularity implies
ongoing knowledge of the location of singing neighbors; at night or in
visually occluded habitats only the calling song seems likely to provide this
information. Almost all katydids, therefore, can be expected to use the calling
song in aggression and Orchelimum differs only in the readiness of competi-
tors to escalate to an extreme expression of competition for mates.
In a search system male-male contesting signals occur independently of
female solicitation. Femur tips are a display separate from crepitation. In
systems employing a calling song, male-male aggressive information is inti-
mately combined with information directed at females. This goes some way
to explain the amplitude modulation complexity of many katydid calling
songs (Morris and Walker 1976). The remarkable diversity of oedipodine
signal repertoires is less a reflection of greater need for information transfer
than a result of the feasibility of employing different information units in
different contexts. In search systems the context, intra or inter-sexual, is
usually known to the signaller. Senders in a call system are, by contrast, true
broadcasters; since information about recipients is often lacking, senders
must allow simultaneously for these 2 contexts.
Game theory has recently been applied to animal contests (Maynard
Smith 1976, Maynard Smith and Parker 1976). It provides some insight into
the readiness of Orchelimum spp. to escalate male aggressive encounters. We
should expect escalation to be associated with the following conditions: (A)
the protagonists are evenly matched, (B) the species has very limited po-
tential for inflicting severe damage, and (C) the pay-off for success in a
particular encounter is substantial.

The Florida Entomologist 62 (1)

March, 1979

Only evenly matched animals need escalate. If an obvious asymmetry in
fighting ability exists (i.e. substantial size or weight difference) it could be
used by the participants early in the interaction to predict the outcome.
Orchelimum males meet on vegetation that moves readily beneath their
weight and being diurnal with well-developed eyes, they have good visual
information about their opponent's size. Selection will favor individuals that
can anticipate the loss of a fight and thereby avoid both risk and wasted
Because of the large spermatophylax, Orchelimum males undergo a sub-
stantial weight loss with the passage of a spermatophore. Thus we might
suspect that recently mated males are at a disadvantage in aggressive en-
counters and yield more readily. Escalation to overt fighting should be
occurring most often between males of comparable size and with well-formed
There are some emphatically predatory katydids (Capnobotes, Rehnia,
Phlugis) but most acoustic Orthoptera, and certainly Orchelimum, are not
adapted strongly for predation. They lack formidable weapons and it seems
unlikely that fighting Orchelimum males could inflict serious damage on each
other. However, one should bear in mind the immense importance of stridula-
tion in securing a mate. A torn tegminal cell will reduce a singer's intensity
(Morris and Pipher 1967) and compromise his chances of attracting females.
The reward for success in aggression is improved access to females.
Ready escalation by Orchelimum males suggests that their pay-off per en-
counter is unusually high. In D. carolina domination in an aggressive en-
counter gains for a male only a short-lived opportunity to search without
the presence of a rival (Kerr 1977). The incessant exploration by males in
search systems ensures a high challenge rate. Thus an oedipodine male's im-
provement in mate access per encounter is comparatively trivial; the pay-off
does not warrant escalation.
In Orchelimum's mating system, with males calling rather than search-
ing, challenges may occur at a much lower rate, contributing to greater per
encounter gain in exclusive calling time. Calling systems may thus have an
inherently greater pay-off per encounter than search systems, but this im-
proved pay-off cannot account for the extent of Orchelimum escalation. Most
other tettigoniids have call systems yet they do not show a comparable in-
cidence of fighting.
Perhaps Orchelimum fighting is linked somehow to a scarcity of sexually
receptive females and to the adaptiveness of paternal investment. There is
evidence that O. gladiator females become unresponsive to the calling of
males after mating once (Morris et al. 1975a). Suppose that most females
mate with only 1 male in their lifetime. Together with tendencies toward
synchronized female maturation, this would cause receptive females to be a
relatively scarce commodity in an Orchelimum population. The operational
sex ratio (Emlen and Oring 1977) would become rapidly skewed in favor of
males in the course of the breeding period. The increasing rarity of potential
mates would give increased importance to success in aggressive encounters
and perhaps justify more frequent escalation.
The existence of the spermatophylax tells us that females have benefited
historically from a large nutritional input on the part of the male. There
must have been strong selection for females to mate with sedentary sustained
singers because this mechanism of male aggression mediated by song ensured

Symposium: Sociobiology of Sex

that a female would obtain an adequate food gift. Females respond prefer-
entially to singers. Steady singing from a fixed location (necessary if the
female is to successfully navigate to the male) can only be achieved by re-
peatedly defeating competing males in aggressive interactions. The presence
of an adequate spermatophore confers a weight advantage which enables a
male to win fights. Recently mated males, males only just become adult, or
males for whatever reason inefficient in foraging, will have a smaller
spermatophylax, weigh less, and tend to lose encounters. Escalation, its cost
of less significance in species lacking formidable weapons, may be seen as the
most effective way to avoid deceit. In a grappling fight it is hard to lie about
one's weight.
The mating systems of acoustic Orthoptera range from active searching
to sedentary calling. Call-answer systems and burrow-centered territoriality
are among several variations in the kinds of mating systems. Many struc-
tural and behavioral peculiarities of males e.g. a large spermatophylax, overt
fighting, a gin trap, can only be properly understood as evolutionary prod-
ucts of a system maximizing individual reproductive success. The different
interests of males and females have shaped these systems. Each sex has
evolved to exploit features of the other's behavior and to defend against such
exploitation. Thus, males vulnerable to 'eat and run' females evolved a
structure that restrains feeding females and ensures paternity. Females
may benefit by preferring males whose ability to sing undisturbed derives
from success in overt fighting; this success is in turn derived from the
presence of a large spermatophylax that confers upon the male a competitive
weight in his aggressive encounters.


This paper was prepared during a research leave at the University of
Florida. I would like to thank the Department of Entomology and Nema-
tology at that institution for their kind hospitality. Appreciation is extended
to Norm Leppla for greatly improving the written manuscript. Jim Lloyd
directed my attention to the term "gin trap". I wish particularly to thank
Tom Walker for his friendship and for many productive and enjoyable
katydid conversations. Funds for the study of Cyphoderris were provided as
part of Operating Grant 4946 of the National Research Council of Canada.

ALEXANDER, R. D. 1956. A Comparative Study of Sound Production in In-
sects, with Special Reference to the Singing Orthoptera and Cicadidae
of the Eastern United States. Ph.D. Thesis, Ohio State Univ. 529 pp.
1961. Aggressiveness, territoriality, and sexual behavior in field
crickets (Orthoptera: Gryllidae). Behaviour 17:130-223.
AND D. OTTE. 1967. The evolution of genitalia and mating behavior
in crickets (Gryllidae) and other Orthoptera. Univ. Mich. Mus. Zool.
Misc. Pub. 133. 62 pp.
1975. Natural selection and specialized chorusing behavior in acous-
tical insects. Pages 35-77 In D. Pimentel ed. Insects, Science and
Society. Academic Press, N.Y.
BUSNEL, R. G., B. DUMORTIER, AND F. PASQUINELLY. 1955. Phonotaxie de
femelle d'Ephippiger (Orthoptere) A des signaux acoustiques syn-
th6tiques. C.R. Soc. Biol. Paris 149:11-3.

The Florida Entomologist 62(1)

- AND M. C. BUSNEL. 1956. Recherches sur le comportement
acoustique des EphippigBres (Orthopteres, Tettigoniidae). Bull. Biol.
Fr. Belg. 90:219-86.
DAVEY, K. G. 1960. The evolution of spermatophores in insects. Proc. Roy.
Ent. Soc. Lond. (A) 35:107-13.
DUIJM, M. AND T. VAN OYEN. 1948. Het sjirpen van de Zadel sprinkhaan. De
Levende Natuur 51:81-7.
EMLEN, S. T. AND L. W. ORING. 1977. Ecology, sexual selection and the evolu-
tion of mating systems. Science 197:215-23.
FULTON, B. B. 1915. The tree crickets of New York: life history and bionom-
ics. N.Y. Agric. Exp. Stn. Tech. Bull. 42. 47 pp.
HINTON, H. E. 1946. The "gin traps" of some beetle pupae; a protective de-
vice which appears to be unknown. Trans. Ent. Soc. Lond. 97:473-96.
JONES, M. D. R. 1966. The acoustic behaviour of the bush cricket Pholidop-
tera griseoaptera I. Alternation, synchronism and rivalry between
males. J. Exp. Biol. 45:15-30.
KERR, G. E. 1974. Visual and acoustical communicative behaviour in Dis-
sosteira carolina (Orthoptera: Acrididae). Can. Ent. 106:263-72.
.1977. Uncertainty Analyses of the Behaviour of Dissosteira carolina
(Orthoptera: Acrididae). Ph.D. Thesis, Univ. of Toronto. 76 pp.
MAYNARD SMITH, J. 1976. Evolution and the theory of games. Am. Scientist
AND G. A. PARKER. 1976. The logic of asymmetric contests. Anim.
Behav. 24:159-75.
MORRIS, G. K. 1967. Song and Aggression in Tettigoniidae. Ph.D. Thesis,
Cornell Univ. 229 pp.
AND R. E. PIPHER. 1967. Tegminal amplifiers and spectrum con-
sistencies in Conocephalus nigropleurum (Bruner), Tettigoniidae. J.
Ins. Physiol. 13:1075-85.
1971. Aggression in male conocephaline grasshoppers (Tettigoni-
idae). Anim. Behav. 19:132-7.
1972. Phonotaxis of male meadow grasshoppers (Orthoptera: Tet-
tigoniidae). J. N. Y. Ent. Soc. 80:5-6.
G. E. KERR AND D. T. GWYNNE. 1975a. Ontogeny of phonotaxis in
Orchelimum gladiator (Orthoptera: Tettigoniidae: Conocephalinae).
Can. J. Zool. 53:1127-30.
- AND -- 1975b. Calling song function in the bog katydid,
Metrioptera sphagnorum (F. Walker) (Orthoptera, Tettigoniidae):
female phonotaxis to normal and altered song. Zeit. fur Tierpsy-
chologie 37:502-14.
AND T. J. WALKER. 1976. Calling songs of Orchelimum meadow katy-
dids (Tettigoniidae). I. Mechanism, terminology, and geographic dis-
tribution. Can. Ent. 108:785-800.
OTTE, D. 1970. A comparative study of communicative behavior in grass-
hoppers. Univ. Mich. Mus. Zool. Misc. Pub. 141. 168 pp.
REGEN, J. 1913. Uber die Anlockung des Weibchens von Gryllus campestris
L. durch telephonisch iibertragene Stridulationslaute des Minnchens.
Pflugers Arch. Ges. Physiol. 155,:193-200.
SHAW, K. C. 1968. An analysis of the phonoresponse of males of the true
katydid, Pterophylla camellifolia (Fabricius) (Orthoptera: Tettigoni-
idae). Behaviour 31:203-60.
THORNHILL, R. 1976. Sexual selection and paternal investment in insects.
Am. Nat. 110:153-63.
TRIVERS, R. L. 1972. Parental investment and sexual selection. Pages 136-79
In B. Campbell ed. Sexual Selection and the Descent of Man, 1871-
1971. Aldine, Chicago.

March, 1979

Symposium: Sociobiology of Sex

WALKER, T. J. 1957. Specificity in the response of female tree crickets
(Orthoptera, Gryllidae, Oecanthinae) to calling songs of the males.
Ann. Ent. Soc. Am. 50:626-36.
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ZEUNER, F. E. 1939. Fossil Orthoptera Ensifera. Brit. Mus. Nat. Hist.,
London. 321 p.


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

Laymen get the impression that biologists have an inordinate preoccupa-
tion with sex. We are immoderate, and it is excusable: sexual behavior is the
key to understanding biological species. Only when he turns his attention to
mating behavior does the biologist begin to use more than inferential evi-
dence for species "boundaries."2 Most basic and applied investigations ulti-
mately depend upon a knowledge of species, whether for acquiring species-
pure samples, or for identifying and manipulating vulnerable points in the
ecology of some competing organism. Questions that are addressed to the
way species found their origin, got to be the way they are, and exist today,
begin and end with a discussion of biological species.
Gene flow is the phenomenon at the center of understanding and defini-
tion of biological species. In the final analysis, when reduced to its smallest
moment of flux, to its irreducible whit of displacement, genes flow down an
aedeagus and into another individual, and the genes of 2 parents flow to-
gether. (Sometimes the actual mechanics are not exactly like this-genes are
handed over in sacks, left on posts, or squirted into the surrounding me-
dium.) Mating behavior arranges for and accomplishes gene flow. It com-
prises the activities and events that take place as the animals seek, identify,
win over and appraise, and finally accept partners in reproduction. Thus
sexual behavior, in all its intimate and diverse detail in the animal kingdom,
becomes a necessitous obsession with biologists. Further, because the biologies
of most organisms are constructed around sexual success, there is more at
stake in understanding mating behavior than "merely" straightening out
species. This knowledge is fundamental to understanding biology at all, and
its most important principles.
Natural selection is the choreographer, ,composer, and lyricist of the en-
tire sexual performance. It brings about change in gene (=allele) fre-
quencies, and this, in a reasonable working definition, is evolution. Natural
selection occurs when certain genetic sorts of individuals in a population
leave a greater number of progeny than do others. As simple and old-hat as
it sounds, this elementary fact can be used with considerable reward and
success when one addresses mating behavior studies. Surprisingly, many

'Fla. Agr. Exp. Sta. Journal Series No. 1199.
2In Shakespeare's The Merchant Of Venice copulation is "the deed of kind" (Shipley 1977).



'' b+
. ~ Ic

W^alf-9 7

LF kw

The Florida Entomologist 62(1)

March, 1979

published studies, speculations, and conclusions, indicate that not all biolo-
gists understand, use, or profit from this simple, old but fresh biological
verity. One practical application or approach-plan is the one-gene-analysis-
model (OGAM) that Richard Dawkins (1976) presented at a popular level
in his book The Selfish Gene, as did David Barash (1977) in his highly
readable Sociobiology and Behavior. This technique has been used by some
biologists for more than a decade (e.g. Williams 1966). The OGAM pits 2
opposing phenotypes against each other in the reproductive game. For pur-
poses of simplification, the competing phenotypes are based on 2 allelic forms
of a gene (or as more commonly expressed, upon the 2 "genes" that are com-
peting for a locus). Simple genetic logic is followed, or rather is pushed
through to its seeming endpoint. The conclusion, as to what should or should
not be, is not final or binding on nature: it merely provides a guide and pre-
vents certain kinds of errors, raises suspicions of certain explanations or
observations, suggests lines of research to be followed, and provides a sound
criterion for recognizing significant observations on natural phenomena. The
OGAM places in proper probability perspective some erroneous explanations
that otherwise seem credible or plausible (such as the heart-warming story
of the mutualistic yucca-moth, that appears in ecology texts).
To begin with a simple example, consider a species of beetle in which the
female emits a pheromone that males smell at a distance and approach.
Males with long antennae (gene L+) are more successful in getting to fe-
males than males with shorter ones (gene L-)-the former detect lower
levels of the pheromone and are able to track it better. It is obvious that un-
less the disadvantages of the longer antennae outweigh the advantages, as
measured in reproductive success, L4 will gradually replace L-. (We really
don't need the OGAM to come to this plain, straightforward conclusion.)
In a more complicated case, males of S.E. Asian Pteroptyx fireflies con-
gregate in great numbers in trees and flash in synchrony. Males and females
are attracted to the pulsing trees, and it has been speculated that by syn-
chronizing their flashes these males are providing a huge beacon-tree to help
(proposed context of selection) other members of their species get to a
gathering place quickly and thereby avoid predation by bats3. In other words,
the synchrony is said to be a group adaptation (biotic as opposed to organic,
Williams 1966), evolved and maintained in the context of group benefit. Let
S+ result in synchronous flashing, and its competitor for that locus on the
chromosome (S-) not produce such behavior. By devoting more of their ac-
tivity (energy, attention) to their own reproductive success, males with S-
will find and inseminate more females, and leave more S- progeny than their
rivals leave S+, in each generation. In fact, we would predict that the syn-
chronizing behavior should be lost; indeed, it should never have evolved. But
we observe that males do flash in synchrony, and, therefore, conclude that
selection producing this behavior must be acting in some context other than
that proposed. Synchrony must be doing something for the competing, S+-
bearing and perpetuating male. The assisted individuals, those using the
beacon-effect to reach the tree, are but cueing in on a conspicuous and highly
relevant marker for locating available eggs to fertilize. As a working hy-
pothesis the benevolent-beacon theory is certainly worse than none at all-

3Entomologists often speak of aggregation pheromones without ascertaining actual func-
tion, or even being aware that such an explanation is not simple and demands special factual

Symposium: Sociobiology of Sex 19

it flies in the face of simple genetics-and we are guided by the OGAM to
seek other explanations before setting out across the Pacific to study beacon-
Bedbugs and kin are reproductively eccentric. Males inject sperm through
the female body wall and into the hemocoel, where evolutionarily new struc-
tures within (paragenital system) conduct as well as store sperm prior to
fertilization. Males of the genus Afrocimex have external paragenital struc-
tures like, in fact in some respects, more developed than those of conspecific
females. Males are found with copulation scars where other males have
jabbed them, and with spermatozoa within. Males of Xylocoris mount
mounted males and inject them with sperm, some of which finds its way into
the sperm ducts of the prime-positioned males and hence into the females
with their ejaculate. These developments were not recognized as belonging to
male competition (Fig. 1)-the latter circumstance might correctly be called
autocuckoldry-but instead it was even imagined that male bedbugs would
evolve helper ducts to assist the alien sperm (uplifting larceny from pilferage
to grand theft!) (Anon. 1974). Let D+ build the helper duct and D- not do
so. D+ cannot gain. Although it may sometimes help other D+, it will on oc-
casion, to one extent or another, contribute to the success of D-, thereby re-
ducing its own proliferation. Because D-, on the other hand, will selfishly
keep all possible fertilizations, it will always deny passage to D+. It is ironic
that such a theory could be preferred when it appears that male competition
probably was a major selective force resulting in the evolution of internal
fertilization (Parker 1970; Fig. 2) and of the extra-genital, traumatic in-
semination of these bugs.
In these examples the underlying assumption that permitted and led to
the originally speculated "adaptations" was that behavior occurs for the
benefit of the population or species. One of the greatest values of the OGAM
is that it will often uncover this unlikely, if not completely erronous, as-
sumption. It forces one to focus on the selfish gene, its fate, and its conse-
quences. Selfish genes instruct their gene machines, to use Dawkin's meta-
phor, to be competitive in every aspect of their mating behavior, from the
beginning search to final fertilization-to yield nothing without a net re-
productive gain. Consider these insectan examples: sperm put into a dung
fly female by a male is largely pushed aside by sperm from the next male
(Parker 1970). A male heliconiid butterfly puts a chemical on the female
that deters other males (Gilbert 1976). After copulating, males of many
insects, including flies and Lepidoptera, put plugs behind their sperm which
prevent the entrance of sperm from subsequent males (Parker 1970; Leo-
pold 1976). Male walkingsticks remain with and ride their females, as living
chastity belts, for days or weeks (Sivinski 1977), and a Parnassius butterfly
glues his genitalia to those of a female (Wigglesworth 1965). In some
Diptera, males inject accessory secretions (matrone) that inhibit the mating
behavior of the female by affecting components in the central nervous system
(Leopold 1976 and refs.). Males of a firefly inject flashes into the coded
patterns of rivals, possibly making them appear to be those of a different
species (Lloyd 1978)4. The external "superfemale" genitalia on Afrocimex

4Competitive mimicry: In a competitive situation, the presentation of false information,
permitting an individual to gain an advantage over a rival: 1) by making another individual
of the interaction (competing male, or female) appear to belong to another class of objects (a
different species), or 2) by emitting signals appropriate to the female (contested resource),
and diverting the rival's approach from the correct object.

The Florida Entomologist 62(1)

March, 1979

males may also perpetrate sexual dirty tricks, directing the copulatory
thrusts of rivals to the wrong spot, more or less masturbating them as they
compete to fertilize the eggs of the contested females. Males of a New
Guinea longhorned grasshopper insert raspberries (i.e. Bronx cheers) into
the rhythmic, "nasal", bleating songs of nearby males (Fig. 3). Male army-
worm moths emit a pheromone that inhibits the responses of rivals to avail-
able females (Hirai et al. 1978)5. Males of the horseshoe crab pry and
shove each other as they gather around and press up against the female of a
tandem-pair at the oviposition site, perhaps stealing fertilizations from the
consort by releasing sperm when the eggs are laid (Fig. 4). Finally, in more
straightforward competition, males of other species, such as scarabs (Fig. 5)
and phengodid beetles, bite, horn, pinch, shove, kick, and maim each other in
the presence of females (Otte and Stayman 1978; Eberhard 1978; Tiemann
If, in the analysis of any behavior, one finds that success in the competi-
tion doesn't seem to correlate with reproductive success, (i.e. that certain
males seem to be fighting while others are doing the mating), understanding
OGAM, he will not conclude that they were simply "naturally aggressive" (a
meaningless expression), but pause in coming to a conclusion, and give the
behavior closer scrutiny. When a particular mating strategy is observed in
males, a rival counter-strategy will be sought, as will ecological imperatives
that make certain other strategies potent. For example, a male firefly that
makes the pattern of a rival appear to be that of a sympatric species has his
strategy safely anchored in reproductive isolation. When it was found that
the male heliconiid put a deterrent chemical on the female, Gilbert (1976)
addressed himself to the question of why males continued to be deterred by
the chemical and did not evolve out of the trap. The ethologist that is trying
to construct an ethogram, or the sociobiologist making a sociogram, should
also notice that unless competitive situations simulating possible natural
events and triggers are arranged, a critical part of the catalogue is being
omitted. Similarly, if mating analyses are always made under unnatural,
crowded conditions in the laboratory, one cannot expect to see the entire
repertory of behavior.
The mating behavior discussed so far has been rather simple. To round
out this overview, I should mention insects with different ecologies and
phylogenetic backgrounds. In insects that require resources that are spatially
limited such as carrion, bracket fungi, dung, stream oviposition sites, or per-
haps even certain insects themselves (Fig. 6), male success may depend upon
the ability to fight, to take over and defend these locations or territories on
them or nearby. When resources are not limiting, and populations dense,
males may easily reach females, presenting each female with many suitors
and the opportunity to exercise choice. If such ecological circumstances exist
for some time, females may force males into leks6 where they must compete
by singing, flashing, strutting, or butting heads for hours (Lloyd 1978).
Under other circumstances, no better understood, males may bring resources
or tokens to females, such as seeds, dead flies, and empty balloons (Thornhill
1976 and refs.) and the nature of the associated male rivalry is different, as

"The discoverers of this suggested that its "biological significance . might be the in-
creased (species) reproductive efficiency that results"! Molecularly marvelous studies in chem-
ical ecology often seem to lack biological sophistication.
leck = mating aggregation or assembly (Alexander 1975, p. 67).

Symposium: Sociobiology of Sex 21

is its theoretical interest and application. Other phylogenetic and ecological
circumstances have resulted in females dispensing with males and sex,
seasonally or completely, as in certain aphids, weevils, and flies, the Surinam
roach in Florida, and an Australian grasshopper, to mention a few. There is
even a coccid that is a self-fertilizing hermaphrodite (Britton et al. 1970).
Sexual parasitism, in which females of a parthenogenic species copulate
with males of another, such as the obligate, sexually parasitic spider beetle
Ptinus mobilis (Woodroffe 1958), may be based on the purloining of nutri-
tional ejaculate. Other explanations, that the egg requires a sperm trigger
for development (easily corrected through evolutionary time), or that the
female is preventing the mating of a potential mother of the ecological com-
petitors of her progeny, are by themselves inadequate and unsatisfying.
To the naturalist and the theorist, and each should have some of the
other in him (Fig. 7), among the more interesting behaviors are those in
which he sees an evolutionary tracking and one-ups-man-ship, or evidence
that this has taken place. In male-male contests it is obvious that for every
new strategy or ploy that evolves, a counter move may be expected sooner or
later. As observers we never know when we have tuned in, in evolutionary
time: is it sooner or later? Going from insect to insect is like being in a time
machine in which one remains stationary as coevolutionary phylogeny
moves past. Theorists and observers alike, but not the animals themselves
though they may reach the point, are pursuing the so-called evolutionarily
stable strategy (ESS) (Maynard Smith 1976). This is the space-time-
genome coordinate at which no new mutant strategy can displace the existing
By adding yet another dimension to the mating behavior hypervolume I
can indicate some behavior that I believe must exist and that may be very
common, and some counter-behavior that may have evolved to subvert it. This
will return to our point of departure, the species problem, and will link the
biological species with all of their subtle sexual competition and complica-
tion, to those reminders of a simpler day, the morphological cabinet species of
the museum taxonomist. In many animals, probably most insects, the ener-
getic contributions made by the 2 parents to each individual offspring are
quantitatively different, with females giving more. This fact has stimulated
a great flurry of theoretical activity and animal inspection over what the
consequences for mating behavior ought to be. Presently it is pretty generally
concluded that this fundamental and ancient asymmetry should result in
hot-to-trot males which have been selected to drop their sperm and dash to
the next female, and choosy (coy) females that are very particular and se-
lective with respect to their mates. The generalization seems legitimate, and
to go a long way in explaining, for example, male coyness observed in species
in which there are sex role reversals and the males have taken on rearing
chores generally found in females (Fig. 8) (Trivers 1972 and refs.; Williams
1975 and refs.; Alexander and Borgia 1978 and refs.). With the occurrence
of cannibalism in mantids and some few others (Fig. 9), these males may
sometimes be coy also (Thornhill 1976 and refs.).
Selection for haste in males, and coyness in females, results in what
amounts to competition between the sexes. Males may be selected to bypass
any choice that the females attempt to exercise, and then females selected to
maintain their options, to not be misled or to have their choices subverted.
If males subdue and seduce females with true aphrodisiacs, females may be

The Florida Entomologist 62(1)

March, 1979

expected to escape sooner or later in evolutionary time. And after sperm has
been placed in a female, she should manipulate it: store, transfer (from
chamber to chamber), use, eat, or dissolve it, as she makes additional obser-
vations on males. Females may accept and store sperm from a male for in-
surance that they will get a mate, and then become choosy. And if the male
ejaculate contains nutritional elements which he contributes to his zygotes,
females should evolve to get this from him for free (recall sexual para-
sitism). Then males should be selected to prevent it, and to make sure that
their genes are used with their expensive, nutritional contributions. It is
possible for sperm to be manipulated in the female (e.g. sex determination in
Hymenoptera). Female reproductive morphology often includes sacs, valves,
and tubes that could have evolved in this context. In fact, it is possible that
some reported examples of sperm competition are actually cases of sperm
Given that females, to one extent or another, subvert male interests by
the internal manipulation of ejaculate, it is not inconceivable that males will
have evolved little openers, snippers, levers and syringes that put sperm in
the places females have evolved ("intended") for sperm with priority usage
-collectively, a veritable Swiss Army Knife of gadgetry! Remember copula-
tion in the bedbug and the male blade? Males of some scutellerid Hemiptera
have large, bizarre genitalia, half the size of the female abdomen. In Hotea
they are spiky and heavily sclerotized, and apparently tear their way through
the vagina and body cavity to reach the spermatheca (Leston, in Hinton
1961). Also recall the diverse shapes of male genitalia that taxonomists have
exploited for decades. These variations and elaborations may in many in-
stances have evolved to bypass female resistance and sperm manipulation,
and represent present or past sexual success strategies. Carrying this line
of reasoning a bit farther, it may be possible to make certain inferences and
predictions about sexual selection and courtship, and even ecology, on the
basis of the diversity (adaptive radiation) of male and female genitalia
within a group. This suggests that intraspecific variation, perhaps poly-
morphism, is to be expected and in many instances another source of
taxonomic confusion.
Recognition that sexual conflict-coevolution occurs, and the consequent
revolutionary perspective of mating and courtship behavior, followed from
an appreciation of natural selection: the genteel view that assumed the indi-
vidual got mated to serve the good of the species and to prevent its extinc-
tion was a conceptual desert, and a researcher's dead end.


I thank Thomas J. Walker and John M. Sivinski for helpful discussion
and reading the manuscript; Frank J. Maturo, Jr. for use of Sea Horse Key
Marine Station; J. C. Webb for technical assistance in the analysis of
Hexacentrus acoustical interactions; and the following colleagues for calling
my attention to pertinent papers in the literature, providing helpful informa-
tion on the natural history of several species, identifying insects, and assist-
ing in the field: F. C. Johnson, Robert S. Lloyd, James L. Nation, Reece I.
Sailer, John M. Sivinski, George Steyskal, Robert T. Sullivan, Thomas J.
Walker, Minter J. Westfall, Howard V. Weems, Jr., F. Willemse, and Robert
E. Woodruff. Personal research reported here was supported by Depart-

Symposium: Sociobiology of Sex

mental, N.S.F., and family funds; Hexacentrus research was carried out
during the 1969 Alpha Helix Expedition to New Guinea, sponsored by the
Scripps Institution of Oceanography and funded by N.S.F.


ALEXANDER, R. D. 1975. Natural selection and specialized chorusing behavior.
Pp. 35-77 In D. Pimintel, ed. Insects, science and society. Academic
Press, New York.
-- AND G. BORGIA. (In press). On the origin and basis of the male-female
phenomenon. In M. Blum and A. Blum, eds. Reproductive competition,
mate choice and sexual selection. Academic Press, New York.
ANON. 1974. The oversexed bedbug. Newsweek 84:48.
BARASH, D. P. 1977. Sociobiology and behavior. Elsevier North-Holland.
N.Y., 378 p.
BRITTON, E. B., W. L. BROWN, JR., ET AL. 1970. The insects of Australia.
(CSIRO). Melbourne Univ. Press. 1029 p.
DAWKINS, R. 1976. The selfish gene. Oxford Univ., N.Y. 224 p.
EBERHARD, W. (In press). The function of horns in Podischnus agenor
(Dynastinae) and other beetles. In M. Blum and A. Blum, eds. Re-
productive competition, mate choice and sexual selection. Academic
Press, New York.
FLEMING, W. E. 1972. Biology of the Japanese beetle. A.R.S. Tech. Bull
1449, U.S.D.A. Washington, 129 p.
GILBERT, L. E. 1976. Postmating female odor in Heliconius butterflies: a male
contributed antiaphrodisiac? Science. 193:419-20.
HINTON, H. E. 1961. Sperm transfer in insects and the evolution of
haemocoelic insemination. Pp. 95-107 In K. C. Highnam, ed. Insect re-
production. Roy. Ent. Soc., London.
HIRAI, K., H. H. SHOREY, AND L. K. GASTON. 1978. Competition among court-
ing male moths: male-to-male inhibitory pheromone. Science 202:
LADD, T. L., JR. 1966. Egg viability and longevity of Japanese beetles treated
with Tepa, apholate, and metepa. J. Econ. Ent. 59:422-5.
LEOPOLD, R. A. 1976. The role of male accessory glands in insect reproduc-
tion. Ann. Rev. Ent. 21:199-221.
LLOYD, J. E. (In press). Sexual selection in luminescent beetles. In M. Blum
and A. Blum, eds. Reproductive competition, mate choice and sexual
selection. Academic Press, New York.
MAYNARD SMITH, J. 1976. Evolution and the theory of games. Amer. Sci. 64:
OTTE, D. 1977. Communication in Orthoptera. Pp. 334-61 In T. Sebeok, ed.
How animals communicate. Indiana, Bloomington.
-- AND K. STAYMAN. (In press). Beetle horns: some patterns in func-
tional morphology. In M. Blum and A. Blum, eds. Reproductive com-
petition, mate choice and sexual selection. Academic Press, New York.
PARKER, G. A. 1970. Sperm competition and its evolutionary consequences in
the insects. Biol. Rev. 45:525-67.
SHIPLEY, J. T. 1977. In praise of English. Times, New York. 310 p.
SIVINSKI, J. 1977. Factors affecting mating duration in the stick insect
Diapheromera velii Walsh (Phasmatodea, Heteronemiidae). M.S.
Thesis, Univ. of New Mexico. 169 p.
SMITH, R. L. 1976. Male brooding behavior of the water bug Abedus herberti
(Hemiptera: Belostomatidae). Ann. Ent. Soc. Amer. 69:740-7.
THORNHILL, R. 1976. Sexual selection and paternal investment in insects.
Amer. Nat. 110:153-63.

The Florida Entomologist 62(1)

March, 1979

Fig. 1. Wheel bugs (Reduviidae: Arilus cristatus (L.)) stacked in a
sexual "encounter," with 2 males (top) competing to fertilize the eggs of the
female. Probably similar interactions were significant in the evolutionary de-
velopment of traumatic insemination in bed bugs. Males are touching anten-
nae, and the upper one has his wings spread (to maintain his precarious
perch?). The female may be evaluating them, or simply awaiting the victor
of whatever contest is being waged. How much subtle data-processing is
possible in the CNS of these animals is unknown, but mate choice could in-
volve fine-tuning to certain elements of male conduct. Little entomological
attention has been paid this critical aspect of insect biology in spite of its
significance for many insect management programs.

Fig. 2. Enigmatic copulation wheel in the damselfly Ischnura ramburii
(Hagen) (male at top). In odonates the male grasps the female behind the
head with terminal appendages and the female then places her terminal
gonopore over the sperm, which has been placed on "accessory" genitalia be-
neath the basal segments of the male's abdomen. An evolution of this wheel
sec lis as impossible as the evolution of a mechanical wheel, such as rotary
wings, and for the same reason-the seeming impossibility of adaptive inter-
mec'iate stages. Sexual competition or selection may have been a major factor
in its evolution. Assuming an ancestral gonopore-gonopore connection
(though a spermatophore placed on the ground cannot be ruled out as a
sterling point) the male could have pushed this connection against his
ver'ral surface to prevent rivals from prying him loose or slipping in be-
tween, to keep the female from escaping, or to force sperm into her. Males of
some species have ventro-lateral flaps (oreillets) on the second segment,
which may function in these regards. Seizure of the female behind the head
would have evolved later, perhaps from restraining moves by males against
the delicate cervical membranes, or more plausibly, near the center of the
locomotor forces females would use in escaping.

Symposium: Sociobiology of Sex 25

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The Florida Entomologist 62(1)

Fig. 3. The calling song of males of a New Guinea longhorned grass-
hopper (Hexacentrus mundus (Walker); Listroscelinae) consists of a buzz
of several seconds duration followed by a series (75) of short "quanks".
S shows the end of a buzz and 7 quanks: time line at bottom, 1 sec/div.
Spectrogram at upper left shows the structure of a quank, each vertical
band a wing-rub: carrier frequency in kHz at left; upper time line, 0.25
sec/div. Males inject short sounds (raspberries) into the rhythmic quanking
of other males. Spectrogram at upper right shows structure of a raspberry;
note the slower wing-rub rate. D shows 18 sec of a sonic duel. Raspberries
seem to be injected to occur in coincidence with quanks, but sometimes the
quanker, seemingly detecting the raspberry, delays his quank 300-500 msec.
-Several competitive strategies occur among Orthoptera, but one of inter-
fering with a neighbor's song to prevent females from being attracted has
been suggested, discussed, discounted, and reaffirmed (Alexander 1975; Otte
1977).-The heckler's strategy may be to break the quanker's rhythm, to
hide communicatively important elements at the beginning of each quank, or,
like dueling banjos, to out-finesse and outscore him, while attending females
keep track of each by his individually distinctive song. This dueling may
have evolved from the synchrony of identical songs, and the acoustical and
temporal characteristics of the raspberry may provide clues to the nature of
competition in other synchronizers. Perhaps of significance to this behavior
is the fact that H. mundus is a predaceous grasshopper.

Fig. 4. Horseshoe Crabs on a beach at Sea Horse Key, Florida. At spring
high tides during March-May females (large individual at lower left-center)
approach the water's edge, acquire a consort, dig in and lay their eggs. Males
outnumber females at the beach, and fertilization is external. Competing
males pry and push each other as they press up against the female, pre-
sumably stealing fertilizations from the consort (top center). The consort
position, the best place to be, is taken first. Males have hooks for hanging
onto the carapace. One wonders if the mating strategies of this living fossil
(from the Paleozoic, more than 2 million centuries ago) are as unchanged as
its external morphology.

March, 1979

Symposium: Sociobiology of Sex 27

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March, 1979


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Symposium: Sociobiology of Sex


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The Florida Entomologist 62 (1)

March, 1979

Fig. 6. The small dung flies (Sphaeroceridae:Copromyza (Borborillus)
sp.) on the back of this ball-forming dung beetle (Scarabaeidae:Canthon
pilularius ?) are cleptoparasites, and develop in the dung balls. A fly, ap-
parently a mated female, will ride the beetle, appearing like the supervisor
on a piece of heavy machinery, as the dung ball is rolled some distance from
the cow pie. Apparently the egg is placed with the ball after its storage
underground. Tumblebugs, "robot" territories that package, transport, and
bury larval food, are a limited resource, and seemingly sites for the forma-
tion of resource-based leks (see Alexander 1975 for discussion) where males
compete for fertilizations. During a short period of observation I found up
to 11 flies on the back of a single, ball-building beetle.

Fig. 7. A pair of dung beetles (Scarabaeidae:Boreocanthon depressipen-
nis) with a dung ball enroute from cow pie to burying place. In the pairs I
observed males did the pushing (lower right) and females rode, tumbling
over and over as they clutched the balls. Moments after this photo was made
the female was away on whirring wings, while the male remained to push
and then bury the ball. The female of another pair departed after reaching
the burial site, immediately after the male had dug his way out of sight be-
side the ball. Upon viewing the desertion an observer wonders if the female
left a permanent mate, which she will meet and recognize again at the pie, to
begin another ball; or, recalling the theoretical considerations of Robert
Trivers (1972) on parental investment and mate exploitation, could the
female have helped and played the male along to a point where a single
parent could succeed, then have run off to repeat the performance with a new
male? A foresaken partner has no viable choice but to attend the ball.
Might a male abscond first? What are the relative investments and their
scheduling in such pairs?

Symposium: Sociobiology of Sex

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The Florida Entomologist 62 (1)

Fig. 8. A male belostomatid bug (Belostomatidae:Belostoma lutarium
Stal.) burdened with eggs cemented on his back by the female. His invest-
ment in each offspring goes considerably beyond a dab of sperm. Unlike
males of many other belostomatids, he is unable to further his total repro-
duction by pursuing and copulating with many females. Theory predicts that
males of species such as this should be more discriminating in their choice
of sexual partners, past selection having favored males that responded ap-
propriately to certain cues emanating from females during courtship. The
nature of this information is a complete mystery, as it is in virtually all
other insects. One thing is obvious: males must have a way of assuring their
paternity of the eggs they tend. Smith (1976) found that males keep the
eggs wet, expose them frequently to atmospheric air, maintain water flow
over them, and select microhabitats that promote their development. He
suggested that among probable costs to males were reduced hunting ef-
ficiency, increased exposure and susceptibility to predation, reduced dispersal
options, and the already-mentioned limit on total reproduction.

Fig. 9. Firefly femme fatale (female Photuris versicolor complex) eating
a male she has attracted with false mating signals. Various parts of him
are scattered in the foreground, as she stands over and chews on his thorax.
A flight wing protrudes from between her mandibles. Males of prey species
are in a bind: competition is keen and they must hurry to gain mates, but
they must avoid mimics. Selection should strongly favor discrimination! Al-
though cannibalism of conspecifics has not been observed, this mimicry prob-
ably originated with predation on mates and/or conspecific males that were
attracted after insemination. Should a female fertilize her eggs with sperm
from a male she has been able to catch? Getting caught is poor recommenda-
tion for a sire-his sons will presumably have a tendency to be like him, and
being eaten ends prospects for gaining additional fertilizations. Food
abundance, mate availability, and age should have a bearing on this be-
havior. In any event, in carnivorous species the last "question" on the mate
examination may be the tough.

March, 1979

Symposium: Sociobiology of Sex

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34 The Florida Entomologist 62(1) March, 1979

TIEMANN, D. L. 1967. Observations on the natural history of the western
banded glowworm, Zarhipis integripennis (Lec). Proc. Cal. Acad. Sci.
TRIVERS, R. L. 1972. Parental investment and sexual selection. Pp. 135-79. In
B. Cambell, ed. Sexual selection and the descent of man: 1871-1971.
Aldine, Chicago.
WIGGLESWORTH, V. B. 1965. The principles of insect physiology. Methuen,
London. 741 p.
WILLIAMS, G. C. 1966. Adaptation and natural selection. Princeton Univ.,
Princeton. 307 p.
--- 1975. Sex and evolution. Princeton Univ., Princeton. 200 p.
WOODROFFE, G. E. 1958. The mode of reproduction of Ptinus clavipes Panzer
form mobilis Moore (=P. Latro act.) (Coleoptera:Ptinidae). Proc.
R. Ent. Soc. Lond. 33:25-30.


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McLaughlin et al.: Keiferia Sex Pheromone Biology




Laboratory and field tests established that the adult tomato pinworm,
Keiferia lycopersicella (Walsingham), is crepuscular (evening) in its mat-
ing habits. Approximately 75% of males captured in traps baited with fe-
male pheromone extracts and placed in tomato fields in February were
caught during the twilight period (5-8 PM EST). Females exposed to an
abrupt-transition photoperiod in the laboratory exhibited maximum calling
activity 0.5 h after lights-out, but males were responsive to female extracts
throughout the night. Virgin pairs mated only during hours 1-3 of the lab-
oratory scotophase.
Females called most and males were most responsive to the pheromone
in a laboratory olfactometer during the third night after eclosion. Males
were more responsive when bioassayed with dim light from both above and
below the olfactometer than when there was light only from below.

The tomato pinworm (TPW), Keiferia lycopersicella (Walsingham), has
been a serious pest of tomato in southern California for many years (Oat-
man 1970), and since 1970 has become economically important in Florida
(Wolfenbarger 1974, Wolfenbarger et al. 1975). Poe et al. (1975) suggested
that several changes in cultural practices and insecticide use are probably
responsible for its present status in Florida.
Generally TPW infestations are combated with multiple applications of
insecticides; however, Oatman (1970) has demonstrated that, in the absence
of insecticides, parasites can suppress larval pinworm populations. He
further suggested that parasites must be integrated in some manner with
other measures to achieve economic control. Exploitation of lepidopteran sex
pheromones may provide a means of suppressing moth species without im-
pairing the effectiveness of their natural enemies.


The research colony originated from insects collected on tomato at Im-
mokalee, FL. The insects were reared to pupation on tomato plants in a
greenhouse at 27+4C, 60-90% RH and the prevailing natural photoperiod.
Soon after pupation they were placed in environmental cabinets at 2810C,
60-65% RH and 14L:10D photoperiod. The pupae were segregated by sex,
and the resulting moths were collected daily, held in age-segregated groups
in 2.3-liter clear plastic boxes, and fed a 10% sucrose solution.

ILepidoptera: Gelechiidae.
'Mention of a commercial or proprietary product in this paper does not constitute an en-
dorsement of that product by the USDA or University of Florida.
'Insect Attractants, Behavior and Basic Biology Research Laboratory, Agricultural Re-
search, Science and Education Administration, USDA, P.O. Box 14565, Gainesville, FL 32604.
4Present address: Cereal Genetics Research, Agricultural Research, Science and Education
Administration, USDA, Columbia, MO 65201.
'Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611.

36 The Florida Entomologist 62(1) March, 1979


The pheromone-releasing behavior (calling) of female moths was ob-
served in environmental cabinets maintained as described, but with a scoto-
phase background illumination of ca. 1.3 lux (Spectra Illumination Meter
Model F 200-TV-B). The virgin females emerged individually in clear plastic
7-dram snap cap vials and were provided with 10% sucrose in water through-
out the experiment.
Preliminary study had indicated that females exhibited calling behavior
only during the scotophase. Therefore, we observed and recorded the be-
havior of each female at 1-h intervals from 1 h before to 1 h after the scoto-
phase. The study was conducted for 6 nights following the day of eclosion
(which occurs during the morning hours). From 38 to 105 females were ob-
served each hour each night. The hourly percentages of 3-night-old females
that exhibited pheromone-release behavior (wings slightly spread, abdomen
elevated, pheromone gland everted) are diagrammed in Fig. 1.
Only 26% of the females were calling at 1 h into the 1st night. This in-
creased to 39% on the 2nd night, and 58% on the 3rd night and decreased to
ca. 45% from the 4th to 6th nights. The mean percent (for all hours each
night) of females exhibiting calling behavior was 13.2, 15.0, 25.6, 23.2, 20.8,
and 14.8 for nights 1 to 6, respectively. The maximum number of calling
females was observed during the 1st h of the scotophase of all nights; a
secondary increase occurred ca. 4-6 h later. Females did not call in advance


70 1 \ S Females Calling
I \ ---A Mated Pairs
60 \

I \
50 1

J 1 I
0 40 I

30 -

20 I

10 -

0 2 3 4 5 6 7 8 9 10

Fig. 1.

Periodicity of female calling behavior and mating in the tomato

McLaughlin et al.: Keiferia Sex Pheromone Biology 37

of lights-of, but did exhibit some persistent activity during the hour after
lights-on (11% of the 3-night-old moths). Moths ceased to call by the 2nd h
of the photophase.
The duration of individual calling bouts was not recorded, but individual
females frequently remained in the calling posture for 2-3 h during the first
4 h of darkness. This was especially true of females during the 3rd night.


Mating behavior was determined in the environmental cabinets described.
Hourly observations were made of 10 individual male-female pairs of virgin
moths in 7-dram vials or of 10 virgin pairs in a 12.6-liter plexiglass cage. At
each observation the mating pairs were removed from the experiment and
replaced with a virgin pair, each individual of which had been held to that
point under test conditions. The moths were in their 2nd to 5th night follow-
ing eclosion. Four replications were conducted of each of the single-pair and
multiple-pair matings.
The copulatory periodicity of the moths is diagrammed in Fig. 1. Sexual
activity, like female calling, was greatest during the 1st h of darkness, and
very little copulation occurred after the 3rd h. Males ran or walked in their
approach to calling females. Approach was generally from behind or at ca.
900 to the female and was accompanied by rapid wing fanning. Females were
not observed to fan their wings when males approached. The copulatory
strikes of the males were made laterally with males beside the females.
Moths remained in copula from 30 min to 2 h.


One hundred 2-night-old virgin female moths were aspirated into a flask
containing 10 ml of anhydrous ether and held for 24 h. The liquor was
filtered and adjusted with ether to 10 ml. Six graded concentrations of the
extract expressed in female equivalents (FE) were prepared by using ace-
tone (less volatile and easier to use for bioassay) as the diluent. These prep-
arations were stored at 0C until tested.
Bioassays of the extracts were conducted by placing groups of 10 virgin
males of known age in 1.8 x 44-cm plexiglass tubes connected in groups of 15
to a common manifold (Sower et al. 1973). Compressed air filtered through
activated charcoal and a molecular sieve was metered through each tube at
1.0 liter/min via the manifold. A pheromone sample was applied to the sur-
face of a stainless steel applicator that was then inserted through a 0.5-cm
diam hole at the upwind end of each tube. Upwind movement to within 4 cm
of the applicator and behavior associated with sexual excitation were observed
and timed. Only those males more than 4 cm from the applicator at the time
of sample introduction were used in computing the percent responding. Re-
sponse was recorded at 15 and 30 sec after introduction of the sample. Males
were allowed to acclimate to test conditions for at least 15 min before each
The tests in the olfactometer were conducted to determine three param-
eters of male biology: the responsiveness of various ages; the daily periodic-
ity of the response; and the dose-response characteristics in the olfactometer.
The olfactometer was illuminated (1.3 lux) from below by rheostat-dimmed

38 The Florida Entomologist 62(1) March, 1979

incandescent bulbs shielded by a diffuser of white bond paper supported by
0.64-cm plexiglass.
Six age groups of males (1st to 6th nights after eclosion) were tested at
10-1 FE. Each age group was assayed beginning 30 min before lights-out and
every hour until lights-on. Fifty males (5 tubes) were assayed per age group
per night. Each age X time was tested 5 times (250 males, 25 assays).
Response curves for 2-, 3-, and 4-night-old males are shown in Fig. 2.
Newly emerged males in their first night showed little sexual excitation or
upwind movement when exposed to the female extract. On their 2nd night,
the males were attracted to the pheromone source, but excited wing fanning
and copulatory attempts were absent. Males in their 3rd night exhibited the
highest level of excitation, attraction, and numbers of copulatory strikes.
Responses declined on the 4th and each subsequent night.
Males became responsive within 0.5 h of lights-out. The 3-night-old males
exhibited a high level of response throughout the scotophase with peak ac-
tivity at 0.5 and 7.5 h. This tendency toward bimodal activity was present in
all age groups tested. Males did not seem to exhibit decreased or inhibited
response to the pheromone due to multiple exposure.
A dose-response curve was constructed from data obtained by exposing
randomly mixed populations of 3- to 6-night-old males to 6 concentrations
(10-1 to 10-6 FE) of female extracts at 30 min into the scotophase. Each
dose was assayed 15 times. In our olfactometer, 10-2 FE produced the great-
est upwind response (Fig. 3, average of 15 and 30 sec counts). The number
of males attracted to within the 4-cm area decreased at 10-1 FE, but the level

2 nd Night -
3 rd Night A--A
70- 4th Night ---.
S60- S

Z \5 \

"' \ '',
L/f 40- \


LL 20

.250 0.5 1.5 25 3.5 4.5 5.5 6.5 7.5 85 9.5

Fig. 2. Response of male tomato pinworm to a 10-1 FE extract of the
female sex pheromone at hourly intervals from 0.5 h before lights-out until
0.5 h before lights-on. Males were in their 2nd, 3rd, or 4th night after

McLaughlin et al.: Keiferia Sex Pheromone Biology

5 50-
Li 40

.J 30



10' 10-2 10-3 104 105- 10-6

Fig. 3. Attraction response in an olfactometer of male tomato pinworm to
6 concentrations of the female sex pheromone (expressed in female equiva-
lents = FE).
of excitation, wing fanning, and copulatory attempts increased. At concen-
trations below 10-2 FE attraction rapidly decreased as did the other be-
During these preliminary bioassays, we began to suspect that TPW
males were more responsive when the olfactometer was illuminated from
above and below than when it was illuminated only from below (necessary to
provide backlight for observation). We therefore constructed 2 plywood
boxes (15 x 56 x 66 cm) each equipped with five 40-watt showcase light
bulbs. These bulbs were covered by a diffuser consisting of a sheet of 0.13
mm teflon sandwiched between a sheet of clear prismatic polystyrene and a
sheet of 0.64-mm plexiglass. Light intensity from each box was regulated by
a rheostat. The bioassay device rested on the lower box and the upper unit
was 40 cm above. Tests (25 replicates with 10 males/replicate) were then
conducted at 2 light levels (0.2 and 1.3 lux) with light from below only or
from both above and below using a dose of 10-1 FE at 0.5 h of the scoto-
phase. At 0.2 lux, the mean (SE) response with light below was 63.5-
3.4%; the response with light above and below was 83.1+3.3% (significant,
p<0.01, F = 28.3). The respective means at 1.3 lux were 71.8+3.7% and
87.13.9% (significant, p<0.01, F = 12.09). TPW males are more re-
sponsive in the olfactometer to the female sex pheromone when illuminated
from below and above.


In February 1976 a field test was conducted at Homestead, FL, to de-
termine whether males could be captured in sticky traps (Pherocon@ 1C)

40 The Florida Entomologist 62(1) March, 1979

baited with female extracts. Two traps each were baited with 10-1 or 10 FE
placed on a piece of filter paper suspended within the trap. These traps,
alternating doses, were placed at plant level (ca. 45 cm) and 10 m apart
along a row in a tomato field. The experiment began at 6:00 AM and con-
tinued for 24 h. The traps were checked hourly and replaced at that time
with new, newly baited traps. Two unbaited traps were also deployed.
The average number of males captured per trap each hour is shown in
Table 1. A total of 2999 males were captured in the four traps, 66% of
those by the traps baited with 10 FE. Unbaited traps captured 5 males.
Males began to respond to the female extracts as the sun began to set (5:00-
6:00 PM). The maximum response occurred after sunset, from 6:00 to 7:00,
i.e., during twilight. Captures diminished rapidly after full dark. The
bimodal response of males in the laboratory was not evident in the field

The tomato pinworm is crepuscular in its mating habits and may use a
decline in light level as an environmental cue to synchronize mating activity.
Males and females exposed to abrupt changes in light intensity in the lab-
oratory displayed precopulatory and copulatory behaviors shortly after
lights-out. Male response to pheromone in the field and our unquantified
field observations of general moth activity revealed that this activity occurs
primarily during the period immediately preceding and following sunset.
The crepuscular activity of TPW moths may be exploited by applying
pesticides during or just before twilight since a major impact of pesticide
sprays on this species is the death of the adults. Such a multiple-application
of sprays obviously is not desirable, but they would probably be more effec-
tive if applied late in the day.
An interesting contrast exists between the calling behavior of females
and male response in the laboratory (Figs. 1 and 2) and the mating curve
(Fig. 1). Even though females call and the males are quite responsive
throughout the night, virgin pairs do not mate after 3-4 h into the scoto-


Mean Percent Mean temp.
Time capture/trap of total (0C)

6:00-7:00 AM 16.3 2.0 15.2
7:00-12:00 Noon 0 0 23.9
4:00-5:00 PM 0 0 24.4
5:00-6:00 PM 98.3 12.1 22.8
6:00-7:00 PM 413.0 50.8 21.7
7:00-8:00 PM 95.0 11.7 20.6
8:00-9:00 PM 64.3 7.9 19.7
9:00-10:00 PM 59.0 7.3 19.2
10:00-11:00 PM 38.5 4.7 19.2
11:00-6:00 AM 28.5 3.5 18.6

McLaughlin et al.: Keiferia Sex Pheromone Biology

phase. Possibly the females do not release pheromone during the latter hours
of the scotophase even though they do adopt the calling posture. Female
Plodia interpunctella (Hiibner) that call during the scotophase release ca.
13 times more sex pheromone than those that call during the photophase
(Nordlund and Brady 1974). There may also be a female receptivity rhythm
as in Ephestia cautella (Walker) (Barrer and Hill 1977).
The increased response of male TPW in the olfactometer when overhead
illumination was provided is apparently caused by the overhead orientation
of the stimulus rather than the increase in intensity. It is very possible that
this phenomenon would be observed in other species since the dorsal light
response is a well-known orientation mechanism in flying and swimming
animals (Fraenkel and Gunn 1961).

BARRER, P. M., AND R. J. HILL. 1977. Some relationships between the calling
posture and sexual receptivity in unmated females of the moth,
Ephestia cautella. Physiol. Ent. 2:255-60.
FRAENKEL, G. S., AND D. L. GUNN. 1961. The orientation of animals. Dover
Publications, Inc., New York. 376 pp.
NORDLUND, D. A., AND U. E. BRADY. 1974. Factors affecting release rate and
production of sex pheromone by female Plodia interpunctella (Hiib-
ner) (Lepidoptera: Pyralidae). Environ. Ent. 3:797-802.
OATMAN, E. R. 1970. Ecological studies of the tomato pinworm on tomato in
southern California. J. Econ. Ent. 63:1531-4.
POE, S. L., J. P. CRILL, AND P. H. EVERETT. 1975. Tomato pinworm population
management in semitropical agriculture. Proc. Fla. St. Hort. Soc.
SOWER, L. L., K. W. VICK, AND J. S. LONG. 1973. Isolation and preliminary
biological studies on the female-produced sex pheromone of Sitotroga
cerealella. Ann. Ent. Soc. Am. 184-7.
WOLFENBARGER, D. 0. 1974. Small pest causes big trouble for tomato growers.
Sunshine State Agric. Res. Rep. 19:14-5.
BARGER. 1975. Control and sequential sampling for damage by the
tomato pinworm. J. Econ. Ent. 68:458-60.


Stored-Product Insects Research and Development Laboratory
Agricultural Research, SEA, USDA
Savannah, GA 31403

Males or females of the almond moth, Ephestia cautella (Walker), a
serious pest of stored commodities, were irradiated (I) with either 35 krad
(a partially sterilizing dose) or 50 krad (a sterilizing dose) and combined
with pairs of untreated (U) adults at I:U ratios of 1, 5, 10, 15, or 25. Doses
'Lepidoptera: Pyralidae.

The Florida Entomologist 62(1)

March, 1979

of 35 and 50 krad reduced egg hatch to 9.0 and 0%, respectively, when the
ratio was 25 I males per U pair; egg hatch was 1.6 (35 krad) and 2.6% (50
krad) when the ratio was 25 I females per U pair. Both males and females
were slightly less competitive after treatment with 35 krad than after treat-
ment with 50 krad (based on percentage egg hatch). Treated females were
more competitive at both doses than treated males; treated females were
competitive at all release ratios. The I males were competitive with U males
except at the lowest release ratio. Thus, irradiated adults were judged suf-
ficiently competitive for field trials to be justified.

Ephestia cautella (Walker), the almond moth, is a cosmopolitan pest of
stored food commodities. It severely damages many different products and is
probably the most destructive stored-product pyralid moth. In many situa-
tions it is the only species of moth present, and the relatively isolated popu-
lations within commodity storage structures make the species amenable to
control with the sterile insect release technique (SIRT). Amuh (1971) ad-
vocated the use of the SIRT for control of the almond moth, and the sugges-
tion has been repeated by various investigators. In spite of the fact that
radiosensitivity and sterilization doses have now been well defined (Calderon
and Gonen 1971, Cogburn et al. 1973, Gonen 1975, Brower 1979), no serious
effort has been made to apply the technique for population control. One
possible reason is the lack of any information concerning the mating com-
petitiveness of irradiation sterilized adults of this species. Reduced mating
competitiveness is considered a serious obstacle to field application of the
SIRT (North and Holt 1968). The present study was designed to determine
mating competitiveness of irradiated males and females of the almond moth.

Moths were obtained from laboratory stock cultures reared in 3.8-liter
jars on the complete diet described by Silhacek and Miller (1972). Through-
out the experiments, cultures and test insects were maintained at 271C
and 605% RH with a 12-h photophase (0600-1800). Upon emergence, un-
mated moths were collected in No. 000 gelatin capsules, segregated by sex,
and aged 24 h before irradiation. Moths were treated in a 60Co irradiator
with a source strength of ca. 700 Ci and a dose rate of ca. 722 rad/min.
Doses utilized were 35 krad (a partially sterilizing dose) and 50 krad (es-
sentially a fully sterilizing dose) (Brower 1979). All controls were unir-
radiated moths at the same age.
Immediately after treatment, irradiated (I) moths of 1 sex were placed
with untreated (U) moths of the same sex in 1.9-liter jars; then U moths of
the opposite sex were added. Screen tops were attached, and the jars were
inverted over open petri dishes. Irradiated males and I females were tested
separately for sexual competitiveness. Moth density was kept similar by
using the following numbers and ratios (I:U:U): 10:10:10 (1:1:1), 20:4:4
(5:1:1), 20:2:2 (10:1:1), 30:2:2 (15:1:1), and 25:1:1. Three days after the
insects were placed in jars, eggs that had fallen through the wire mesh into
the petri dishes were removed. One hundred eggs from each collection were
placed in petri dishes on black construction paper disks surrounded by rear-
ing medium for hatched larvae. Egg hatch was determined after 10 days (an
egg was scored as hatched only if the larva was successful in emerging com-
pletely from the chorion). Medium containing the hatched larvae from each

Brower: Ephestia cautella Mating Competitiveness

petri dish was added to ca. 200 g of medium in a 0.47-liter jar with the lid
sealed with a double layer of filter paper. As soon as emergence of F, adults
started, jars were examined 3 times/week, and adults were removed and
counted until emergence was completed. There were 10 replications of each
sex, dose, and I:U:U ratio.
Natural egg infertility of the untreated (U) population and egg in-
fertility of U females paired with only I males and I females were also de-
termined. The expected egg infertility calculated from the I:U release ratios
was corrected for control egg infertility and for egg fertility in the I samples
because of the residual fertility of the substerilized males. Competitiveness
values (indicated by C.V.) for the I males were then calculated by dividing
the percentage actual infertility by the percentage corrected expected in-
fertility using the same reasoning as Fried (1971). Because I females laid
infertile eggs and the presence of these eggs would bias the observed per-
centage egg hatch, the following equation was used to estimate a corrected
expected percentage infertility for females:
x (No.I 9) (v) + y (No.U 9) (w)
Corr. exp. % infertility = x (No. I 9 ) + y (No. U 9 ) x 100%

x = No. of eggs of I 9 x U S,
y = No. of eggs of U 9 x U 8,
v = Fraction of infertile eggs laid by I 9, and
w = Fraction of infertile eggs laid by U 9.
Competitiveness values (C.V.) for I females were calculated by dividing the
percentage observed egg infertility by the percentage corrected expected in-
fertility. Total progeny per U 9 was calculated by multiplying the number
of progeny per 100 eggs by the mean number of eggs per female.

MALES-35 KRAD. The mean number of eggs laid per female when confined
with various ratios of males irradiated with 35 krad was not significantly
different from the mean number of the control except at 1 ratio (Table 1).
When the numbers of eggs laid by females at the different male ratios were
compared, there were significant differences between some of them. Only
8.2% of the control eggs were infertile, but the addition of I males increased
infertility to 39.7% at the lowest ratio (1 I & : 1 U & : 1 U 9). As the ratio
of I males to U males increased, the percentage of infertile eggs also in-
creased to 95.0% at a ratio of 15 I 8 : 1 U S : 1 U 9 (Table 1). Because 35
krad is not a fully sterilizing dose for adult males, some eggs produced by U
females paired with I males were fertile (1.9%). Thus, sexual competitive-
ness of adult males irradiated with 35 krad was good at all ratios (0.75-1.0
was considered competitive), though it was generally better at the higher
ratios of I males (Table 1). The mean number of progeny/100 eggs was 80.9
for the controls; this decreased to 5.0 for the 25 I S : 1 U : 1 U 9 ratio.
The number of replicates that produced no progeny also increased markedly
as the I:U ratio increased.
MALES-50 KRAD. There were no significant differences between the various
ratios in the number of eggs per female when males were irradiated with 50
krad, except the 10 I $ : 1 U 4 : 1 U 9 ratio which was less (Table 1). Also,

The Florida Entomologist 62(1)



0 |2





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Brower: Ephestia cautella Mating Competitiveness

there was no difference in the number of eggs when I males and U males
were confined separately with U females. At this dose, egg infertility in-
creased from 37.1% at a 1 I : 1 U 8 : 1 U 9 ratio to 99.5% at a 10 I
S: 1 U S: 1 U 9 ratio to 100% at the 25 I S: 1 U : 1 U 9 ratio (Table
1). Competitiveness values for the various ratios ranged from 0.69 to 1.09.
The average number of progeny per 100 eggs decreased from 82.5 in the U +
U controls to 0 in the 10 I 8 : 1 U 8 : 1 U 9 and 25 I : 1 U : 1 U 9
FEMALES-35 KRAD. Females irradiated with 35 krad and paired with U
males did not produce significantly (P <0.05) fewer eggs than mated U
females (Table 2). Virgin I females produced significantly fewer eggs than
mated females; many of the females at the higher I female ratios probably
did not mate since females greatly outnumbered males. The mean number of
eggs per female at higher ratios was much less than that for mated U 9
though significantly greater than the mean number produced by virgin fe-
males. Females irradiated with 35 krad and paired with U males had an egg
fertility of 0.6%. Egg infertility increased from 54.9% at the 1 I 9 : 1 U 9 :
1 U 3 ratio to a high of 98.4% at the 25 I 9 : 1 U 9 : 1 U $ ratio (Table 2).
Sexual competitiveness of I females was better than that of U females at a
1 I 9: 1 U 9 : 1 U & ratio and slightly better at the higher ratios (Table
2). The number of progeny produced decreased concurrently with an in-
crease in the ratio, and only 0.8 progeny/100 eggs were produced at a 25 I 9 :
1 U 9 : 1 U 3 ratio (Table 2).
FEMALES-50 KRAD. Females irradiated with 50 krad and paired with U
males laid fewer eggs (139.4) than mated U females (235.7, Table 1) but
significantly more eggs than virgin I females (26.3) (Table 2). Except for
the 1 I 9 : 1 I : 1 U & ratio, all ratios produced numbers of eggs between
these 2 values, and these numbers did not differ significantly from either
extreme. Mean percentage egg infertility increased from 54.9% at the 1 I 9 :
1 U 9 : 1 U S ratio to 99.5% at the 10 I 9 : 1 U 9 : 1 U 8 ratio but de-
clined to 97.4% at the 25 I 9 : 1 U : 1 U & ratio (Table 2). The calculated
competitiveness values for all ratios averaged 1.1; that is, the females were
fully competitive. The number of progeny produced depended on the release
ratio and ranged from a high of 41.4 to a low of 0.7 (Table 2). Most of the
cages with the 3 higher ratios produced no progeny.


Control of the almond moth by using the SIRT is feasible according to
models by Brower and Tilton (1975). Mating competitiveness of irradiated
adults had not been determined previously, and this study provides the
needed data. Both males and females irradiated as adults have an adequate
level of sexual competitiveness to be effective in a sterile insect release pro-
gram. Males could be used in one warehouse and females in another, thus
doubling the efficiency of the program without increasing the number of
insects that must be reared. Alternatively, males and females could be ir-
radiated and released together. Ahmed et al. (1976) showed that such an
alternative was feasible for the closely related Indian meal moth, Plodia
interpunctella (Hiibner). Moreover, Brower (1978) summarized the work
on competitiveness of irradiated adults of the Indian meal moth and con-
cluded that irradiated females were fully compentive but irradiated males

The Florida Entomologist 62(1)






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Brower: Ephestia cautella Mating Competitiveness

were slightly less competitive at 35 krad and 10-20% less competitive at 50
krad. Overall, results with the Indian meal moth were very similar to these
reported here for the almond moth though irradiated almond moth males, if
anything, were more nearly competitive. Stress factors encountered in the
field could reduce the actual competitiveness of sterile insects. However, ir-
radiated almond moths were judged sufficiently competitive for field testing.


The help of James E. Edenfield, Biological Technician at this laboratory,
in setup and data collection is gratefully acknowledged.

AHMED, M. Y. Y., E. W. TILTON, AND J. H. BROWER. 1976. Competitiveness of
irradiated adults of the Indian meal moth. J. Econ. Ent. 69:349-52.
AMUH, I. K. A. 1971. Potentialities for application of the sterile-male tech-
nique to the control of the cocoa moth, Cadra cautella Walk. Pages
7-11 In Application of Induced Sterility for Control of Lepidopterous
Populations (Proc. Panel Vienna, 1970). IAEA, Vienna. 169 pp.
BROWER, J. H. 1978. Mating competitiveness of irradiated males and females
of the Indian meal moth (Lepidoptera: Pyralidae). Can. Ent. 110:
BROWER, J. H. 1979. Radiosensitivity of adults of Ephestia cautella (Walker)
(Lepidoptera: Pyralidae). J. Econ. Ent. 72: (In press).
BROWER, J. H., AND E. W. TILTON. 1975. Potential for control of Cadra
cautella (Walker) by release of fully or partially sterile males. Int. J.
Appl. Radiat. Isot. 26:720-5.
CALDERON, M., AND M. GONEN. 1971. Effects of gamma radiation on Ephestia
cautella (Wlk.) (Lepidoptera, Phycitidae)-I. Effects on adults. J.
Stored Prod. Res. 7:85-90.
COGBURN, R. R., E. W. TILTON, AND J. H. BROWER. 1973. Almond moth:
Gamma radiation effects on the life stages. J. Econ. Ent. 66:745-51.
FRIED, M. 1971. Determination of sterile-insect competitiveness. J. Econ. Ent.
GONEN, M. 1975. Effects of gamma radiation on Ephestia cautella (Wlk.)
(Lepidoptera, Phycitidae)-IV. Sensitivity of maturing sperm in the
adult to a sterilizing dose. J. Stored Prod. Res. 11:97-101.
NORTH, D. T., AND G. HOLT. 1968. Inherited sterility in progeny of ir-
radiated male cabbage loopers. J. Econ. Ent. 61:928-31.
SILHACEK, D. L., AND G. L. MILLER. 1972. Growth and development of the
Indian meal moth, Plodia interpunctella (Lepidoptera: Phycitidae),
under laboratory mass-rearing conditions. Ann. Ent. Soc. Am. 65:

The Florida Entomologist 62(1)

March, 1979


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

Phytoseiulus macropilis (Banks) was studied in the laboratory to de-
termine its potential rate of population increase when fed on eggs of
Tetranychus urticae Koch. The shortest developmental period was 4.7 days
for males, and 5.8 days for females reared at 27 20C and 90% 5% RH
under 12L:12D. Mean generation duration was 14.2 days with an average
reproductive rate of 1.89 eggs per female per day. Predation of P. macropilis,
expressed as the number of prey eggs consumed per individual, was 5.9 eggs
for the immature stages and 194.1 eggs for the adult or 7.9 eggs consumed
per adult per day. Mean adult longevity was 25.3 days. For each predator
egg produced, 4.3 prey eggs were consumed. The intrinsic rate of predator
population increase (rm) was calculated as 0.27 individuals per female per

Phytoseiulus macropilis (Banks) is the most common phytoseiid pred-
ator of spider mites in south Florida (Saba 1974). Its life history is de-
scribed by Smith and Summers (1949) and its life cycle duration by
Prasad (1967). No details of the reproductive rate or predation of P.
macropilis at specific prey densities are given in these studies; data are not
adequate to determine the intrinsic rate of increase, sex ratio, or influence
of prey density on individual life history studies for the predator or its
host, Tetranychus urticae Koch (Watson 1964, Laing 1969, Shih et al. 1976).
The objective of this paper is to provide data on the biology, potential rate
of population increase, and predation of P. macropilis fed on T. urticae.


The life history of P. macropilis was studied at the University of Florida
Agricultural Research and Education Center, Bradenton, Florida. Environ-
mental chamber conditions were 27 20C, 90 5% RH and 12L:12D. The
initial colony of P. macropilis was collected from among twospotted spider
mite populations on celery, statice (Limonium sp.), chrysanthemum, and
blackberry in a greenhouse at Bradenton, in August 1973. Predators were
reared continuously on spider mites fed on 'Henderson Bush' lima beans
(Phaseolus) grown from seed in 6 in pots. Each pot was set in a 6 in diam-
eter (1 in deep) pot saucer, which served to supply water for the plants and
to confine the mites to the unit. Eggs from the collected predators were used
to start 23 individual colonies. Eight days after establishing each stock
colony, all second generation (Fl) P. macropilis adults were transferred

'Acarina, Phytoseiidae.
2Acarina, Tetranychidae.
'Contribution from Florida Agricultural Experiment Station Journal Series No. 5743.

Shih et al.: Phytoseiulus Biology and Predation

to 14 petri dishes containing excised leaf cultures of spider mites. Each
culture contained at least 3 females and 2 males of P. macropilis. A leaf
culture was prepared as follows: twospotted spider mites were transferred
onto dwarf 'Henderson Bush' lima beans 2 days after primary leaves had
expanded. After 24 hours a spider mite infested leaf was excised and placed
upside down on a 70 x 70 mm piece of cheese cloth in a 100 mm diameter
petri dish. Water was added to soak the cloth but not flood the leaves.
Lanolin applied to the leaf periphery helped confine mites to the leaves. A
No. 00 brush was used to manipulate mites.
After 24 hours on the 14 leaf cultures, surviving P. macropilis adults
were transferred to 9 fresh leaf cultures for another 24 hours before being
returned to the stock colony. The 23 (14 and 9) excised-leaf cultures were
kept until the P. macropilis eggs had hatched, after which the young
predator mites were transferred to fresh leaf cultures. Thereafter, the
predators were transferred to fresh leaf cultures every 24 hours until the
colony died. To ensure mating, male predators of the same generation were
added to cultures where no males developed or where males died during the
first 14 days.
Mites were transferred to separate leaf cultures after each molt. All
mites in a subculture were thus in the same developmental stage. Daily prey
consumption, durations of developmental stages, preoviposition period and
daily egg production were recorded every 12 hours (0.5 day). When a devel-
opmental stage was not observed between 2 successive observations, the dura-
tion of the missed state was assumed to be 0.25 day.
Due to the overlap of developmental stages and the active nature of
spider mites, the predatory capability of P. macropilis was evaluated on the
basis of spider mite egg consumption rather than on predation of all prey
stages. The number of spider mite eggs on each leaf culture always greatly
exceeded actual daily egg consumption of P. macropilis. With food supply
ample and laboratory microenvironment uniform, the influence of these fac-
tors was not considered in this study. The sex ratio of P. macropilis was
determined in both the laboratory study and in samples taken twice weekly
from 2 field plots of strawberries at Bradenton, Florida from January
through March 1974.
Phytoseiulus macropilis prey consumption was established as the number
of prey consumed per predator in each age class. The intrinsic rate of nat-
ural increase (rm) or the number of offspring produced per female per day
was determined as a fundamental statistic to explain the predator mite
capacity for numerical increase under laboratory conditions. The increase of
P. macropilis was estimated using the method proposed by Birch (1948).


macropilis IMMATURE STAGES. Appearance and behavioral characteristics de-
scribed for P. persimilis by Laing (1968) were found to be very similar to
P. macropilis. Phytoseiulus macropilis egg mortality was 2.9%, but no
losses were recorded for the other juvenile stages. The average duration of
the juvenile period was 2.2 days for eggs, 0.7 days for larvae, 1.0 days for
protonymphs and 0.6 and 0.1 days for female and male deutonymphs, re-
spectively (Table 1).

The Florida Entomologist 62(1)

March, 1979

Tetranychus urticae EGGS AT 27 20C, 90 5% RH, AND

Stage of Number Days duration cumulative
development observed Mean SD IMin.-Max. days*

Egg 174 2.2 -
Larva 169 0.7 0.5 2-5 2.2
Protonymph 169 1.0 0.5 3-6 2.9
Active deutonymph
Female 122 0.6 0.6 3-7 3.9
Male 47 0.1 0.1
Quiescent deutonymph
Female 122 1.3 0.7 4-10 4.5
Male 47 0.7 0.3 3-7 4.0
Female 122 0.9 4-10 3.8
Male 47 0.8 3-7 4.7

*Duration of each active stage is recorded at the beginning of each stage and then added
to the cumulative duration of all proceeding stages.
Adult stage. The deutonymphal quiescent period was 1.3 days for females
and 0.7 days for male deutonymphs. Development time from egg through
the quiescent deutonymph was 4-10 days for females and 3-7 days for males
(Table 1). Males lived approximately 23 days; females lived an average of
27 days (Table 2). The mean oviposition period was nearly 25 days with an
average total production of 49.1 eggs per female (Table 2). Fifty percent
natural mortality was reached at 26.7 days for females and 22.6 days for
males. Male to female ratios determined from field samples and the lab-
oratory study were 1:5 and 2:5, respectively.
Prey Consumption. The number of T. urticae eggs consumed per individual
P. macropilis is shown in Tables 2 and 3. Each immature stage required
more prey than the preceding one: 0.4, 1.5, and 4.0 eggs per mite larva,
protonymph, and deutonymph, respectively. The daily predation potential
of adults was 7.9 eggs per individual per day (Table 2).

II. BIOLOGY OF P. macropilis.
Influence of predator age on predation. Daily predation by adult P. macro-
pilis increased rapidly to a peak the 4th day after ecdysis, then decreased
gradually until death (Fig. 1). The only exception to this general trend was
an abrupt increase near the end of the life span observed in the few mites
(6) surviving on days 33 and 34. The mean daily consumption rate for all
individuals used in this experiment (44 females and 23 males) was 7.9 eggs
per adult per day.
Mating and daily oviposition rate. Copulation occurred immediately after
female ecdysis. The first eggs were laid 2 days after mating. Unmated fe-
males did not oviposit. Female P. macropilis reached their peak oviposition
rate of 3.18 eggs per female per day on the 7th day after ecdysis. The rate

Shih et al.: Phytoseiulus Biology and Predation

macropilis FED Tetranychus urticae EGGS AT 27 20C, 90 5%
RH, AND 12L:12D:

Characteristic observed Mean SD Min.-Max.

Longevity (days)
Female 44 26.7 6.4 12-40
Male 23 22.6 5.1 14-32
Oviposition period (days) 44 24.6 5.3 12-36
Total number of eggs
laid per female 44 48.3 9.5 24-68
Total number of eggs
laid per female
per day 44 1.9 0.9 0.4-3
Total number T. urticae
eggs consumed per
predator 67 194.1 43.3 134-228
Total number T. urticae
eggs consumed per
predator per day 67 7.9 3.0 0-16
Total number T. urticae
eggs consumed per
predator egg laid 44 6.1 1.5 3-8

20C, 90 5% RH, AND 12L:12D CHAMBER.

Number of
Stages of Number eggs consumed per mite stage
P. macropilis observed Mean SD Min.-Max.

All immature stages

79 0.4 0.4 0-1
76 1.5 0.9 0.3-3
75 4.0 1.9 1-9
75 5.9 3.2 2-15

Potential Rate of Increase. The intrinsic rate of increase, (rm) was calcu-
lated to be 0.27 individuals per female per day. The predator population was
estimated to increase (Ro) 47 times in a mean generation time (T) of 14.2

remained fairly constant for 3 days, then declined gradually to 0 at 1-2 days
before death. Fluctuations in daily oviposition rate occurred in cycles of 2
to 3 days during the latter half of the oviposition period, usually from days
14 to 28 (Fig. 2).
The relationship between T. urticae eggs consumed and predator eggs
produced is noteworthy. During the first 4 days after ecdysis, the ratio of

The Florida Entomologist 62(1)



5 10 15 20 25 30 35 38

Fig. 1. Predation
macropilis (Banks).

14 -

1 2-


LL 8-

W 6-


of Tetranychus urticae Koch eggs by Phytoseiulus


Af A
1 _____ 1 1 1 1 1

5 10 15 20 25 30 35 38

Fig. 2. Daily egg production by Phytoseiulus macropilis and its consump-
tion of Tetranychus urticae eggs.


() 10-
0r 6-


March, 1979

Shih et al.: Phytoseiulus Biology and Predation

prey egg consumption to predator eggs laid was much higher than that of
the subsequent 7 days. Feeding cycles of 3 to 5 days appeared on days 13 to
26 (Fig. 2). Females older than 26 days consumed slightly greater numbers
of prey but laid fewer eggs. Initial fluctuations might be attributed during
the first 5 days to the need for building nutritional reserves and completing
ovarian development during a period of high food intake and low oviposition
rate. A highly active female at this period may also need greater quantities
of food to meet energy requirements; later (days 5-25) the cycling may be
due to the periodic build-up and depletion of nutritional reserves as eggs are
laid; after day 27, the differences may be due to senility or biased data since
fewer than 8 females were alive at 28 days.
The data reported here for Phytoseiulus macropilis differ only slightly
from those provided by Laing (1969) for P. persimilis. The life cycle of
P. macropilis appears to be more rapid at each stage of development, a dif-
ference which might be attributed to the low and fluctuating temperature
used in the P. persimilis study. P. persimilis oviposited an average of 2.4
eggs per day compared to 1.8 eggs per day for P. macropilis. The intrinsic
rate of increase for P. persimilis was 0.219 individuals per female per day
and the population multiplied 44.4 times in a mean generation time of 17.32
days (Laing 1969). In contrast, P. macropilis had an intrinsic rate of in-
crease of 0.271 individuals per female per day and the population multiplied
47 times in a mean generation time of 14.2 days. Since both species are ap-
parently of tropical origin and frequent similar habitats it is not surprising
that their life histories are similar.

BIRCH, L. C. 1948. The intrinsic rate of natural increase of an insect popula-
tion. J. Anim. Ecol. 17:15-26.
LAING, J. E. 1969. Life history and life table of Tetranychus urticae Koch.
Acarologia 11:32-42.
PRASAD, V. 1967. Biology of the predatory mite, Phytoseiulus macropilis, in
Hawaii (Acarina:Phytoseiidae). Ann. Ent. Soc. Amer. 60:905-8.
SABA, F. 1974. Life history and population dynamics of Tetranychus tumidus
in Florida (Acarina:Tetranychidae). Fla. Ent. 57:47-63.
SHIH, C. I., S. L. POE, AND H. L. CROMROY. 1976. Biology, life table, and
intrinsic rate of increase of Tetranychus urticae. Ann. Ent. Soc.
Amer. 69:362-4.
SMITH, L. M., AND F. M. SUMMERS. 1949. The structure and biology of the
red spider predator, "Hypoaspis macropilis" (Banks). Proc. Ent. Soc.
Wash. 51:209-18.
WATSON, T. F. 1964. Influence of host plant condition on population increase
of Tetranychus telarius (Linnaeus) (Acarina: Tetranychidae). Hil-
gardia 35:273-320.

The Florida, Entomologist 62 (1)

March, 1979



Chlordane 10 G applied to the soil surface at 5.6, 8.4, and 11.2 kg AI/ha
remained in the 0 to 2.5 cm soil horizon and reached ppb levels after approxi-
mately 1 year. Chlordane did not leach below approximately 2.5 cm nor was
chlordane detected in irrigation ditch water at or above 1.0 ppb. Chlordane
was not found in 'Valencia' and 'Hamlin' orange leaves or fruit at or above
5.0 ppb. Chlordane disappeared in a pseudo-first order fashion with a half-
life of 27 to 72 days.

Chlordane was substituted for the control of Fuller rose beetle,
Pantomorus cervinus (Boh.), in Florida citrus when aldrin and dieldrin were
banned by the United States Environmental Protection Agency in 1975.
Although chlordane has been used for 25 years to control termites in very
young citrus groves, no data are available under Florida conditions regard-
ing its environmental behavior. Studies in other parts of the United States
have shown that 15-40% of the applied chlordane may persist up to 14 years
in soil (Fleming and Maines 1954, Lichtenstein and Polivka 1959, Nash and
Woolson 1967). Chlordane may also penetrate root crops (Stewart 1975) and
it has been found in a wide variety of agricultural soils where chlordane had
not been used for at least 5 years (Carey et al. 1973). Technical chlordane is
a complex mixture of at least 18 chlorinated components (Brooks 1974). It
has an ecological magnification of about 100,000 as it moves through the
food-web and has a water solubility of 0.056 ppm (Sanborn et al. 1976). In
view of the documented environmental persistence and food-web magnifica-
tion of chlordane, the present study was undertaken to determine the per-
sistence of chlordane and the possible agroecosystem contamination by
that chemical after an agricultural application to Florida citrus soils.


Manual applications of granular 10 G chlordane were made with a
Seymour seeder at rates of 5.6, 8.4, and 11.2 kg AI/ha (5.0, 7.0, and 10.0 lb
AI/acre). Treatments were replicated 4 times in 0.91 ha plots in a Latin
square design. Four no-treatment plots were included as controls.
Two experiments were conducted in 20-year-old 'Hamlin' orange groves
near Lake Alfred, Florida (White Clay Pit Road and Shinn Groves) and 1
experiment was conducted in a 'Valencia' orange grove near Ft. Pierce,
Florida (Fountain Grove). The Ft. Pierce grove was double-bedded (2 rows
between ditches) with 1 m ditches between beds, 2 large feeder ditches at
each end of the grove with an additional large feeder irrigation ditch along
the road. The Lake Alfred groves were not irrigated.
Soil samples were taken by vacuuming five, 20 cm2 x 2.5 cm deep volumes

1Florida Agricultural Experiment Stations Journal Series No. 1372. This study supported
in part by special funds from the Center for Environmental Programs, Univ. of Fla., Gaines-
ville and by a commercial grant from Velsicol Corporation.
'University of Florida, Agricultural Research and Education Center, Lake Alfred, 33850.
3University of Florida, Agricultural Research Center, Ft. Pierce, 33450.
4Use of a specific product does not constitute an endorsement by the University of Florida
to the exclusion of similar products.

Nigg et al.: Chlordane Residues

in each of the plots. A 2.5-10 cm, 10-17.8 cm, 17.8-33 cm and 33-61 cm sample
was taken from the center of each vacuumed area with a standard soil
sampling tube. The 5 samples from each soil horizon were bulked to obtain 1
sample per plot per soil horizon.
Water samples were taken from the Ft. Pierce grove ditches and lateral
ditches in hexane-washed Mason jars. Four water samples from each of 7
irrigation ditches and the 3 supply ditches were taken each sampling day.
Eight fruit and 10 leaves constituted a fruit leaf or sample. Samples not
extracted immediately were stored a maximum of 15 days at -200C.
Chlordane was extracted from soil samples by weighing 10 g of air dried
soil from each bulked sample into 100-ml beakers and wetting with 2.0 ml
of distilled water. Beakers were covered with aluminum foil and held at 40C
for 16 h. Fifty ml of 20:80 benzene:acetone was added to each beaker and
the soil was sonicated at 15,000 Hz for 30 sec. The beaker was recovered with
foil, the soil allowed to settle and a 5.0 ml aliquot was removed. This aliquot
was evaporated almost to dryness in a 50.0-ml volumetric flask under N2 at
500C. After cooling, 5.0 ml of benzene was added to the 50.0-ml flask fol-
lowed by 40.0 ml of 2% sodium sulfate. The flask was capped, shaken, and
the layers were allowed to separate. The benzene was moved to the neck of
the flask with distilled water and removed to brown glass bottles over a few
grams of sodium sulfate for GLC analysis.
Fruit were separated into rind and pulp. The rind was diced, a 10 g sub-
sample was homogenized in 30.0 ml of 1:1 dichloromethane:acetone and a 5.0
ml aliquot taken. Fruit pulp was homogenized and a 10 g subsample ex-
tracted as for rind. Leaves were weighed and homogenized in 50.0 ml of 1:1
dichloromethane:acetone and a 5.0 ml aliquot taken. Leaf and fruit extrac-
tion aliquots were evaporated to dryness under N,, taken-up in 5.0 ml of
benzene and column chromatographed on a 10 g silica gel column. Elution
was with 50.0 ml of benzene.
A 100.0 ml aliquot of each water sample was extracted twice with 20.0
ml of dichloromethane and once with 20.0 ml of benzene. These extractions
were combined, taken to dryness, and transferred to 10.0 ml of benzene over
a few grams of sodium sulfate for gas-liquid chromatographic (GLC)
analysis. All extractions were stored at -20oC prior to analyses.
At the 1 ppm level, recoveries from 8 replications of each fortified sub-
strate were: soil-100.0 + 4.0%; rind-98.5 3.5%; pulp-98.7 2.0%;
and leaves-100.0 3.0%. Chlordane was recovered from water at the 50
ppb level at 95.0 5.5%.
The GLC analysis was done on a 0.8 m x 2 mm glass column packed with
5% SP 2100 on 100/120 mesh Gas Chrom. Q. Operating conditions were:
column 190C, inlet 2200C, Ni6e EC detector 3000C, N2 60 ml/min. All GLC
injections were 5 pl. Burdick and Jackson solvents, used for GLC and extrac-
tions, were assessed for contaminants by evaporating 500.0 ml to 2.0 ml for
Quantification was similar to Cochrane et al. (1975) with the quantifica-
tion of heptachlor, heptachlor epoxide, y-chlordane and y chlordane/
nonachlor (single peak) in technical chlordane by peak height used for
estimation of technical chlordane. Since samples 140 days postapplication
generally contained only the y-chlordane and y-chlordane/nonachlor GLC
peaks, only these GLC peaks were used to estimate technical chlordane for
140 day + samples.

The Florida Entomologist 62(1)

March, 1979

Moisture capacity, pH, and organic matter were measured in composite
samples from each grove and soil horizon according to Black (1965) as
modified by Anderson et al. (1968). Rainfall was measured with standard
fencepost gauges (Weather Measure Corp.4). Technical chlordane and indi-
vidual chlordane component standards were provided by Velsicol Corp.
Statistical analyses were performed on a Tektronix 4051 graphic computing
system with an interactive digital plotter (Tektronix, Inc.; Beaverton,
Oregon 970774).


Chlordane was apparently confined only to the upper 2.5 cm soil horizon
in all 3 experiments for up to 445 days (Table 1). Only inconsistent and un-
interpretable traces of chlordane were detected below 2.5 cm. Consequently
only the 2.5 cm results are reported here. No apparent 'disappearance' of
chlordane occurred until ca. 150 days postapplication. Tafuri et al. (1977)
observed no disappearance of chlordane for ca. 169 days postapplication and
confinement of chlordane to the 0-10 cm soil horizon in a clay loam soil.
The data in Table 1 are variable and the levels of chlordane observed on
day 139 are consistently higher than those observed on day 48 and in some
cases on day 13 postapplication. This variability was probably related to
disking and mowing operations which occurred in all 3 experiments just
prior to the 3rd sampling. Similarly, Talekar et al. (1977) have attributed
variation in dieldrin recovery to rototilling.


Grove name (kg AI/ha) 0-2.5 cm residues (mean S.D.)

Fountain 13 days* 48 days 139 days 395 days

5.6 39.5 43.4 2.6 1.9 10.2 12.0 0.49 0.22
8.4 28.7 25.7 10.9 + 13.7 19.7 18.9 0.10 0.19
11.2 21.5 29.2 13.8 6.3 86.1 53.5 0.42 0.25
No treatment ND** ND ND ND

White Clay
Pit Road 14 days 44 days 139 days 445 days

5.6 4.8 1.8 3.2 1.3 4.0 2.6 ND
8.4 52.2 + 50.1 4.0 2.4 6.8 3.8 0.40 0
11.2 44.9 44.4 7.1 7.2 38.6 40.0 ND
No treatment ND ND ND ND

Shinn 30 days 65 days 153 days 410 days

5.6 6.9 1.8 1.8 0.99 11.3 10.9 0.08 0.09
8.4 3.9 2.6 2.0 1.1 31.6 40.3 0.23 + 0.27
11.2 23.8 20.3 10.2 11.4 61.0 32.1 ND
No treatment ND ND ND ND

*Days postapplication.
**ND = Not Detected.

Nigg et al.: Chlordane Residues

The data fit to the standard 1st order kinetic pesticide decay model
averaged R2 = 0.74 0.14 (Fountain Grove); R2 = 0.82 0.13 (White
Clay Pit Road) ; R2 = 0.64 0.24 (Shinn Grove) (Table 2). Half-lives in
the 3 experiments varied from 23-100 days. The average half-life in each
experiment appeared to be related to the moisture capacity of the 0-2.5 cm
soil horizon with the higher the moisture capacity, the shorter the half-life
(Table 3). Rainfall was about equal among groves: Ft. Pierce 131.06 cm,
White Clay Pit Road 150.88 cm, and Shinn Grove 139.45 cm. The half-life
for chlordane observed here was about 1/3 that of p,p'-DDT and dieldrin
under fall and winter subtropical conditions in Taiwan (Talekar et al. 1977).
The high temperature and soil moisture conditions in Florida probably ac-
count for this non-persistence of chlordane (Talekar et al. 1977, Williams
1975). Irreversible binding of chlordane to these soils is probably not a factor
in either the half-life or 'disappearance' behavior of chlordane (Lichtenstein
et al. 1977). The short half-life and rapid disappearance of chlordane in
Florida, however, may compromise its use as a residual insecticide for weevil


Treatment Half-life
Grove name (kg AI/ha) \ (days) t (R2 x 100)

Fountain 5.6 -0.009 77.0 67**
8.4 -0.01 69.3 90**
11.2 -0.01 69.3 64*
Average 71.9

White Clay
Pit Road 5.6 -0.03 23.1 92**
8.4 -0.02 34.7 90**
11.2 -0.03 23.1 86**
Average 26.9

Shinn 5.6 -0.01 69.3 70**
8.4 -0.007 99.0 38
11.2 -0.03 23.1 85**
Average 63.8

*P 0.05.
**P 7 0.01.
tHalf-life = 0.693/X.

THE 0-2.5 cm HORIZON.

Moisture Organic
Grove capacity (%) pH matter (%)

Fountain 1.25 6.14 1.7 0.2
White Clay Pit Road 4.90 6.22 2.3 0.6
Shinn 1.31 6.32 2.5 0.8

The Florida, Entomologist 62 (1)

March, 1979

Chlordane residues were not detected in fruit, leaf, or water samples and
consequently chlordane residues, if present, must have been below the de-
tectable level of 5.0 ppb in fruit and leaves and 1.0 ppb in water. Sanborn
et al. (1976) found 104 ppm of chlordane in algae when water contained 1.06
ppb of chlordane and Mattraw (1975) detected no chlordane in southern
Florida surface waters, yet 33% of southern Florida sediment samples con-
tained chlordane. Biomagnification and low water solubility of chlordane
shown in these studies suggest additional experiments designed to determine
life-form levels of chlordane after a normal agricultural application of
chlordane would be appropriate.


The technical assistance of Ms. Roberta Woodruff, Mr. Paul Keene, and
Mr. Neil Berger is gratefully acknowledged.

REESE. 1968. Methods of analysis. Fla. Agric. Exp. Sta., Univ. of Fla.,
IFAS, AREC, Lake Alfred, Fla. 33850 #1398. 61 pp.
BLACK, C. A. 1965. Methods of soil analysis: Agronomy 9 (part 2) : 771-1572.
Amer. Soc. Agron. Madison, Wis. 1572 pp.
BROOKS, G. T. 1974. Chlorinated insecticides. Technology and application.
CRC Press 1:113-56.
CAREY, A. E., G. B. WIERSONA, H. TAI, AND W. G. MITCHELL. 1973. Organo-
chlorine pesticide residues in soils and crops of the cornbelt region,
United States-1970. Pestic. Monit. J. 6:369-76.
WILSON. 1975. Gas-liquid chromatographic determination of tech-
nical chlordane residues in food crops: Interpretation of analytical
data. J. Assoc. Off. Anal. Chem. 58:1051-61.
FLEMING, W. E., AND W. W. MAINES. 1954. Persistence of chlordane in soils
of the area infested by the Japanese beetle. J. Econ. Ent. 47:165-9.
LICHTENSTEIN, E. P., AND J. B. POLIVKA. 1959. Persistence of some chlo-
rinated hydrocarbon insecticides in turf soils. J. Econ. Ent. 52:289-93.
"persistent" and "nonpersistent" 14C-labeled insecticides in an agri-
cultural soil. J. Agric. Food Chem. 25:43-7.
MATTRAW, H. C. 1975. Occurrence of chlorinated hydrocarbon insecticides,
Southern Florida-1968-72. Pestic. Monit. J. 9:106-14.
NASH, R. G., AND E. A. WOOLSON. 1967. Persistence of chlorinated hydro-
carbon insecticides in soils. Science 157:924-6.
SANBORN, J. R., R. L. METCALF, W. N. BRUCE, AND P. Y. Lu. 1976. The fate
of chlordane and toxaphene in a terrestrial-aquatic model ecosystem.
Environ. Ent. 5:533-8.
STEWART, D. K. R. 1975. Chlordane uptake from soil by root crops. Environ.
Ent. 4:254-6.
and movement of AG chlordane in soil and its residues in alfalfa. J.
Agric. Food Chem. 25:353-6.
TALEKAR, N. S., L. SUN, E. LEE, AND J. CHEN. 1977. Persistence of some
insecticides in subtropical soil. J. Agric. Food Chem. 25:348-52.
WILLIAMS, J. H. 1975. Persistence of chlorfenvinphos in soils. Pestic. Sci.

Ru and Sailer: Prospaltella in Florida 59


Dept. of Entomology and Nematology
University of Florida
Gainesville, FL 32611

Prospaltella lahorensis Howard, native to Pakistan and established in
California in 1968, was obtained from California in June 1977. A total of 57
females and 130 males were released in sleeve cages placed on small branches
well infested with 2nd, 3rd and 4th nymphal stages of the citrus whitefly,
Dialeurodes citri (Ashmead). Releases were made and colonies were estab-
lished at 1 site in Winter Haven, FL on Citrus sp. and 3 sites in Gainesville,
FL on Citrus spp., Viburnum odoratissimum Ker-Gawl and Ligustrum
lucidum Aiton. At 1 Gainesville site P. lahorensis increased dramatically and
by the end of October 1977 had dispersed into 29 nearby citrus trees and 1
viburnum. At the end of March 1978 viable populations of P. lahorensis were
present at all 4 release sites.

The citrus whitefly, Dialeurodes citri (Ashmead), a pest of citrus and
ornamental shrubs, is native to China and countries of Southeast Asia
(Cockerell 1903, Morrill and Back 1911). It also is found in many countries
of the Mediterranean region and North and South America. In the United
States, its range includes Alabama, California, Florida, Georgia, Louisiana,
Mississippi, North Carolina, Texas, Virginia, and Washington, D.C. (Anony-
mous 1974).
While traveling in British India in 1911, R. S. Woglum found Prospaltella
lahorensis Howard parasitizing the citrus whitefly. He subsequently collected
the species from the region now known as Pakistan, and shipped stock to
Florida. The parasites arrived during the winter at a time when no hosts
were available and thus no culture was established (Woglum 1913).
The purpose of our present project was to introduce and establish P.
lahorensis for the biological control of the citrus whitefly in Florida utilizing
stock obtained from California. In California P. lahorensis is established in
Orange Co. and the city of Sacramento as a result of importation from
Pakistan in 1968 by Dr. Paul DeBach of the University of California, Di-
vision of Biological Control at Riverside.


Prospaltella lahorensis was collected from citrus leaves in Orange County,
California and shipped to the Biological Control Laboratory, Division of
Plant Industry, Gainesville, FL in June 1977. The material, including para-
sitized and unparasitized citrus whitefly nymphs, was held in quarantine
until adult parasites emerged. Within 2 days following their emergence the

SHymenoptera: Aphelinidae.
2Florida Agr. Exp. Sta. Journal Series No. 999.

The Florida Entomologist 62(1)

March, 1979

adults were released on Citrus sp., Viburnum odoratissimum Ker-Gawl and
Ligustrum lucidum Aiton in Gainesville. The parasite was subcolonized on
citrus in Winter Haven, FL in October 1977.
In order to reduce premature dispersal, the adult P. lahorensis were re-
leased in sleeve cages (1 on each host plant) placed over small branches
(Fig. 1). These branches were well infested with 2nd, 3rd, and 4th nymphal
stage citrus whitefly (Fig. 2). The sleeve- were of nylon organdy (70 cm
long, 35 cm diameter; or 40 cm long, 20 cm diameter), reinforced with 2
wire loops to prevent collapse, and with drawstrings at each end. From 2 to
21 virgin or mated females of P. lahorensis were released from a stoppered
vial into each sleeve. A narrow band of the sleeve was smeared lightly with
honey to provide supplemental food for adult parasites.
In the laboratory, P. lahorensis were reared in 50 dram plastic shell vials
(Fig. 3). One or 2 leaves (b) including the petioles and a piece of either
citrus or viburnum stem with ca. 20-30 third stage citrus whitefly nymphs
were placed in each inverted vial. For aeration, a circular opening (a) 2.5
mm in diameter, was cut from the bottom of the vial and covered with 54-
mesh nylon organdy. A single hole was punched in the plastic lid through
which the plant stem, wrapped in a thin layer of cotton, was passed. This
vial was then placed over another vial, filled with water (c), into which the
stem was placed. Water was changed daily. For studying the life cycle, 1
female and 1 male adult P. lahorensis, emerging on the same day, were intro-
duced into the upper vial. They were removed 2 days later. In other trials,
virgin or mated females were placed singly in the vials and allowed to re-
main until they died, usually within 7 days. After 10 days exposure, the
citrus or viburnum leaves were checked daily for parasitized nymphs.
Whitefly nymphs containing an immature parasite were removed 1 or 2 days

Fig. 1. Original release site of Prospaltella lahorensis on citrus at the
University of Florida campus, Gainesville. (a) organdy sleeve cage.

Ru and Sailer: Prospaltella in Florida 61


Fig. 2. (a) Healthy 4th nymphal stage of citrus whitefly, Dialeurodes
citri (Ashmead). (b) Larva and (c) pupa of Prospaltella lahorensis Howard
inside citrus whitefly nymph. (d) Empty whitefly nymphal case after P.
lahovensis emergence.
before emergence of the adult parasite. After removal the parasitized
nymphs were placed individually in No. 1 gelatine capsules to prevent

The number of parasites and dates of each release are in Table 1. Twenty
days after the 21-22 June release, larvae and pupae of P. lahorensis (Fig.
2b and c) were observed at the release sites on both Citrus sp. and Ligustrum
odoratissimum. Adults emerged 2 days later (Fig. 2d) and all were females.
In order to strengthen the parasite colonies and insure simultaneous and
continuing presence of both adult males and females, 5 additional releases
totaling ca. 100 males from California stock and involving all 3 Gainesville
release sites were made during the period 14-19 July 1977. By the end of
August, a second generation had appeared and hosts containing P. lahorensis
pupae were present throughout the citrus tree as well as the V. odoratis-
simum and L. lucidum hedges where the first releases were made. By the end
of October 1977, P. lahorensis was found on 29 citrus trees and 1 viburnum
tree on the University of Florida campus near McCarty Hall. One tree is
200 m from the site of the original release site.
The populations of P. lahorensis present at all release sites were mon-
itored 7 times from 30 January to 13 March 1978. Table 2 shows the results

The Florida Entomologist 62 (1)



Fig. 3. Cage for rearing Prospaltella lahorensis on citrus whitefly in the
laboratory; (a) circular vent in cage; (b) citrus leaf; (c) vial of water.
of 10 leaf samples taken on each sampling date from a citrus tree immedi-
ately adjacent to the tree on which the species first was released. Despite
evidence of considerable mortality during the winter, the number of larvae
and pupae present on 13 March 1978 indicated survival of a viable popula-
tion. In early April both sexes of P. lahorensis were observed at all 3 Gaines-
ville sites and at the site in Winter Haven, FL where the species was sub-
colonized on citrus in October 1977.
Laboratory studies showed that P. lahorensis was autoparasitic (= adel-

March, 1979

Ru and Sailer: Prospaltella in Florida

TABLE 1. NUMBER OF Prospaltella lahorensis RELEASED AND NUMBER OF F1

Host plant

Citrus sp.
Citrus sp.
Citrus sp.
Citrus sp.
Ligustrum lucidum
Viburnum odoratissium

Release date

22 June 1977
25 June 1977
28 June 1977
8 July 1977
21 June 1977
24 June 1977

No. P. lahorensis
Females Males


Prospaltella lahorensis

Larva Pupa Adult

Larva Pupa


30 Jan 1978 6 8 0 6 4 2
10 Feb 1978 7 16 0 1 6 4
20 Feb 1978 4 11 1 5 6 6
25 Feb 1978 4 1 0 10 13 4
27 Feb 1978 4 3 0 4 11 6
7 Mar 1978 14 19 2 8 17 7
13 Mar 1978 10 12 0 5 12 5

phoparasitic, Flanders 1937; Zinna 1962) with females produced as endo-
parasites of the citrus whitefly, but with males produced as hyperparasitic
ectoparasites on immature females of their own species (Fig. 4). The devel-
opment from egg deposition to adult at 24C required 12 to 15 days for the
adelphoparasitic males and 24.5 days for females (N=6 males and 4 fe-
males). In 6 tests involving single, mated females caged on whitefly infested
leaves, 30 females and 1 male progeny resulted. When 18 virgin females were
similarly tested, no progeny were produced.
In a small grove of 50 citrus trees located near Red Hill in Orange
County, California, ca. 30-40% of the citrus whitefly were parasitized by P.
lahorensis. The sex ratio of material collected and shipped to Gainesville, FL
varied in 1977 from 1 female:6 males in June to 1.5 females:100 males in
July and 1 female:2 males in August. This variation in sex ratio may have
been due to season-related changes in the citrus whitefly population levels
and subsequent response of the adelphoparasitic P. lahorensis.
According to Rose (1977, personal communication), the dispersal of P.
lahorensis has been very slow in California, averaging only about 6 m a year
in Sacramento. In Orange County where it was introduced in 1968 it has

F, progeny


The Florida Entomologist 62(1)

BplHB^ C1

^ LA

Cd 4^


-~ r-


Fig. 4. Male larva (b) on female pupa (a) of Prospaltella lahorensis.
dispersed less than 1609 m from the site of the original release and remains
uncommon in the citrus groves of that area. This slow rate of dispersal may
be characteristic of the species under conditions peculiar to California. Our
evidence of movement over a distance of 200 m in 3 months suggests a much
greater dispersal capability in Florida.

March, 1979



Ru and Sailer: Prospaltella in Florida


Thanks are due to Dr. Paul DeBach, Mr. M. Rose, Ms. M. Reynolds, Div.
of Biological Control, University of California, Riverside, California for
sending Prospaltella lahorensis to Gainesville and for the technical advice
and assistance generously provided the senior author while he was in Cali-
fornia to observe and collect the citrus whitefly parasite during June 1977.

ANONYMOUS. 1974. Distribution maps of pests. Map No. 111. Commonw. Inst.
Ent. London.
COCKERELL, T. D. A. 1903. Whitefly and its allies. Fla. Agr. Exp. Sta. Bull.
FLANDERS, S. E. 1937. Ovipositional instincts and developmental sex differ-
ences in the genus Coccophagus. Univ. Calif. Berkeley. Publ. Ent. 6:
MORRILL, A. W., AND E. A. BACK. 1911. Whiteflies injurious to citrus in
Florida. USDA. Bur. Ent. Bull. No. 92:1-109.
WOGLUM, R. S. 1913. Report of a trip to India and the Orient in search of the
natural enemies of the citrus whitefly. USDA. Bur. Ent. Bull. 120:
ZINNA, G. 1962. Richerche sigli insetti entomofagi. III. Specializzazione
entomoparassitica negli Aphelinidae: Interdipendenze biocenotiche
tra due specie associate. Studio morfologico, etologico e fisiologico del
Coccophagoides similis (Masi) e Azotus matritensis Mercet. Boll. Lab.
Ent. Agr. (Fillippo Silvestri) Portici. 20:73-184.



Seed heads of 'DeKalb E-57' sorghum, planted on 26 April, 16 May, and
1 June 1976, were harvested in the soft dough stage and placed in emergence
cages to determine the temporal pattern of emergence for the sorghum
midge, Contarinia sorghicola (Coquillet), and its parasite, Aprostocetus
diplosidis Crawford.
Midge emergence began at the end of the milky stage, peaked 5-6 days
later, and was completed in 10-11 days. Parasite emergence began 7-8 days
after the end of the milky stage, peaked at 12-14 days, and was completed in
18-19 days. The emergence of the 2 insects overlapped from the 7th to 11th
day after the end of the soft dough stage. Toxic residues of insecticides, ap-
plied to kill emerging adult midges, could be present for 7 days after the end
of the soft dough stage without harming the adult parasites.

2Eulophidae: Hymenoptera.
WFlorida Agricultural Experiment Station Journal Series No. 1080.
4Department of Entomology and Nematology; University of Florida; Gainesville 32611.
5University of Florida Agricultural Research and Education Center; Quincy, FL 32351.
Present address: Kansas State University; Garden City, KS 67846.

The Florida Entomologist 62(1)

March, 1979

The sorghum midge, Contarinia sorghicola (Coquillet), an important in-
sect pest of grain sorghum in many countries around the world has inflicted
losses totalling millions of dollars per year (Geering 1953, Harris 1961, 1969,
Thomas and Cate 1971, Huddleston et al. 1972). The midge is most difficult
to observe and control because of its concealment within the seeds of the
sorghum spikelets. Field observations (Dean 1910) have indicated that the
endoparasite Aprostocetus diplosidis Crawford produces mortality in late
season midge populations. These observations were confirmed by Dean (1911)
and Walter (1941) who referred to A. diplosidis as the most prominent and
aggressive parasite of the sorghum midge. Increasing midge parasitization
by A. diplosidis was observed as the sorghum growing season progressed
during 1976 at Quincy, Florida. Adults of the parasite A. diplosidis and
adults of the sorghum midge emerged from infested sorghum heads in an
overlapping pattern, an observation which agreed with that of Dean (1911)
who recorded a 6:1 ratio of parasite/midge from samples taken in late
Knowledge of the temporal pattern of emergence for the 2 insects could
help to avoid possible exposure of the parasite adult to chemicals intended
for the adult midge. This would prevent substantial mortality of the parasite
population and aid in biological control of the midge in subsequent plant-
ings. Hence, in 1976, studies were initiated in Florida to determine the
emergence periods for both pest and parasite.

'DeKalb E-57' sorghum was planted on 26 April, 16 May, and 1 June
1976 at the University of Florida Agricultural Research and Education
Center, Quincy, in 4-row plots arranged in a randomized complete block de-
sign with 4 replications. The plots were treated with Atrazine@ at 2.1 kg
Ai/ha for weed suppression and with 560.4 kg/ha of 5:10:15 fertilizer before
planting; plots were 18.5 m long, with 91.4 cm between rows. The sorghum,
grown under rainy conditions, was planted by machine at a seed rate of
about 9.66 kg/ha.
At the milky (soft dough) stage of development, 25 sorghum heads per
replicate were removed from the 2 middle rows of each plot on 26 July, 4
August, and 12 August 1976. The sorghum heads were placed into 25.4 X 34.3
X 50.8 cm emergence cages. Emerging sorghum midges and parasites were
collected in 2.5 cm diam by 4.8 cm plastic vials inserted into the side of each
cage. The vials were coated with petroleum jelly on the inside walls to en-
trap the emerging insects. Samples were held for 20 days emergence under
laboratory conditions of 24 + 20C and 80% R. H., and monitored daily by
inspecting the vials, recording numbers of midges and parasites and removing
all insects that emerged in each 24 h period. Monitoring and counting were
terminated when midge and parasite emergence stopped. Data collected were
averaged for the 4 replicates and converted into percent emergence of the
total population per day.

Sorghum midge emergence from the heads began at the end of the milky
stage, peaked 5 to 7 days later, and was completed in 10-11 days (Fig. 1).

Ru and Sailer: Prospaltella in Florida 67

Parasite emergence began 6-8 days after the milky stage, increased at the
tenth day, and peaked at 12 to 14 days. Emergence of the midge and para-
site overlapped from the 7th to 11th day with small numbers of parasites
emerging before the 11th day regardless of planting dates.

40 o Midges & Parasites from
sorghum planted 4-26-76
M S Midges & Parasites from
O sorghum planted 5-16-76
30 0 Midges & Parasites from
0. sorghum planted 6-1-76


10 Overlap

Midge Parasit e

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19
Emergence(Days After Milky Stage)

Fig. 1. Emergence of the sorghum midge, Contarinia sorghicola (Coquil-
lett), and its parasite Aprostocetus diplosidis Crawford from sorghum heads
at Quincy, Florida.
In programs where parasite-induced midge mortality is expected to com-
plement chemical midge control, a knowledge of emergence patterns provides
a means for optimization of chemical and biological mortality. Insecticide
residues could be present on sorghum heads until the 7th or 8th day after the
milky stage sufficient to kill emerging midge adults. Residues present after
the 8th day, however, must not be lethal to the midge parasite if populations
are to be maintained to aid in controlling subsequent midge generations.

DEAN, H. W. 1910. Some notes upon the life history of the sorghum midge.
J. Econ. Ent. 3:205-7.
1911. The sorghum midge. USDA Bur. Ent. Bull. 85, part IV (Rev.)
58 p.
GEERING, Q. A. 1953. The sorghum midge, Contarinia sorghicola (Coquillett),
in East Africa. Bull. Ent. Res. 44:363-6.
HARRIS, K. M. 1961. The sorghum midge, Contarinia sorghicola (Coquillett),
in Nigeria. Bull. Ent. Res. 52:129-46.
1969. The sorghum midge. World Crops. 21:176-9.
C. E. FOREHAND. 1972. Biology and control of the sorghum midge I.
Chemical and cultural control studies in West Texas. J. Econ. Ent.
THOMAS, J. G. AND J. R. CATE, JR. 1971. The sorghum midge and its control.
Texas Agr. Exp. Sta. PR-2863.
WALTER, E. V. 1941. The biology and control of the sorghum midge. USDA
Tech. Bull. 778, 26 p.

The Florida Entomologist 62(1)

March, 1979


WORM IN NORTHEAST FLORIDA'--(Note). Four successive plantings
of cabbage were grown near Hastings, FL during 1974-75 to follow cabbage
webworm, Hellula rogatalis (Hulst), diamondback moth, Plutella xylostella
(L.), and imported cabbageworm, Pieris rapae (L.), populations and para-
sites as part of an integrated pest management survey. Cabbage webworm
numbers (Fig. 1) and damage were high in 1974 and low in 1975. Two larvae
could damage the bud so that a head was not formed. No parasites were
reared from 513 larvae collected during the period. Diamondback larvae were
common from ca. Feb.-June with 4 population peaks noted each year (Fig.
1). Larvae fed on the cabbage buds producing many small holes which en-
larged greatly as the leaves grew out. Parasites reared from 1415 diamond-

mm 4
< uJ 2
0. o L=-T-

----- Cabbage Webworm Larvae

I' \

I I f

:' I':':' . I

- Diamondback Moth Larvae
----- Diamondback Moth Pupae
Diamondback Moth Parasites

--- Imported Cobbogeworm Lorvae
- Imported Cabbogeworm Eggs

.. Feb. i Ma. Ap. May J _' c. J.' Fb. Mor. Apr. May
I---st planting--& -2nd planting- --- 3rd planting--- -- 4th planting--
1974 1975

Fig. 1. Average number of cabbage webworms (top), diamondback moth
larvae, pupae, and their parasites (center), and imported cabbageworms
(bottom) per cabbage plant at Hastings, FL during 1974-75.

IIFAS Journal Series No. 1353.

Scientific Notes

back larvae and 147 pupae included Diadegma insularis (Cresson), an
ichneumonid, which was reared from 32% of the larvae and pupae examined.
The adults were common all season with highest numbers, 30/100 plants,
observed in May. An eulophid, Tetrastichus sokolowskii Kurdjumov, was
reared from 10% of the diamondback pupae. High numbers of imported
cabbageworms occurred in 1974 with over 6/plant observed in June (Fig. 1).
Numbers were low in 1975. No parasites were found in 3059 eggs taken dur-
ing the study. Parasites reared from 1391 larvae and pupae included a
braconid, Apanteles glomeratus (L.), which was reared from ca. 0.015% of
the larvae. The parasites developed within the larvae and pupated in yellow-
ish cocoons on the leaves. Two larvae were parasitized by a tachinid,
Lespesia aletiae (Riley). Pteromalus puparum (L.), Pteromalidae, was
found throughout the season and was reared from 4% of the pupae. Overall
biotic control of all 3 pest insects in northeast Florida in 1974-75 was lower
compared to some other areas of the United States although the principal
parasites are similar. Limiting factors may include the yearly fluctuation of
pest populations, climatic differences and the use of pesticides.-NGUYEN Ru
AND R. B. WORKMAN, Dept. of Ent. and Nem., Univ. of Florida, Gainesville
and Agr. Res. Center, Hastings, FL, respectively.

-L -C- -- --^_-- -^- -^- -*--- I -L

(Note). Scientists often must determine the length, width, and/or area of
insect wings to conduct studies of aerodynamics and kinematics of insect
flight, genetic divergence, growth and development, sexual dimorphism,
correlations with body weight and wingbeat frequency, and velocity of flight.
In the past, wing dimensions have been derived by using light microscopy or
photography. Although sophisticated equipment is available that rapidly and
accurately determines surface area, the instrument costs several thousand
dollars. This note describes a simple method that enables the user to obtain
the actual measurements of wing length, width, and area by using a
Trisimplex microprojector to enlarge the wings and a compensating polar
planimeter to determine area. The current average cost of the 2 tools are
$300 and $150, respectively.
The procedure is demonstrated below using 6 species of lab-reared
Tephritidae (Table 1). Flies are killed in ethylacetate vapor. The right
wings are excised as near the wing base as possible and dry-mounted on a
microscope slide under a glass coverslip. The slide is mounted on the stage
of the microprojector, and the magnified wing image is projected downward
onto tracing paper where the focused outline is pencil traced. The length of
the wing tracing is measured from the point of articulation in the center of
the axillary region (Ax) to the farthest point of the remigium (Rm). The
width (w) is measured from the widest part of the vannus region (Vn)
through the remigium to the costal margin along a line transecting the

'Trade names are used in this article solely for the purpose of providing specific informa-
tion. Mention of a trade name does not constitute a guarantee or warranty of the product by
the U.S. Department of Agriculture or an endorsement by the Department over other products
not mentioned.

The Florida Entomologist 62(1)


Wing dimensions X+SE
Length Width Area
(mm) (mm) (mm2)

Dacus dorsalis (Hendel)
$ 6.3 .04 2.4 2.02 10.8 .10
9 6.4-.03 2.5.02 11.3.11
Dacus cucurbitae Coquillett
8 6.4-.05 2.5.02 11.3.13
9 6.8 .05 2.7.02 13.3.18
Dacus oleae (Gmelin)
$ 4.1.04 1.6.02 4.8.09
9 4.3+.04 1.7.01 5.2 .09
Ceratitis capitata (Wiedemann)
S 4.6.04 2.3.02 7.7.12
9 4.6.04 2.2 .02 7.2.12
Anastrepha suspense (Loew)
$ 6.0 .04 2.4 .02 10.9 .18
9 6.2.04 2.5 .02 11.6.17
Rhagoletis pomonella (Walsh)
S 4.0.05 1.8.05 5.5.01
9 4.7.06 2.2.05 7.5.02

*Flies were 8 days old except D. oleae males and females were 2 days
were taken from 20-30 flies/sex per species.

old. Measurements

widest points and extending perpendicularly through the line designating
length (1) (Fig. 1) (terminology from R. E. Snodgrass, 1935. The Wings,
Ch. X, p. 226, In: Principles of Insect Morphology, McGraw-Hill Book Co.,
Inc., New York and London. A micrometer is fastened to the stage of the
microprojector to determine the magnification power. Actual length and

Costal margin



Fig. 1. The right wing of a male Ceratitis capitata showing the ap-
propriate location of length (1) width (w) measurements. Ax, axillary; Rm,
remigium; Vn, vannus.

March, 1979

Scientific Notes

width is obtained by dividing the projected measurements by 20 because the
1-mm scale of the micrometer projected onto paper is 20-mm long. For ex-
ample, the enlarged length of a wing of Ceratitis capitata (Wiedemann)
was measured as 95.0 mm and the width, 46.5 mm; thus, the actual length,
(95 -20) was 4.75 mm, and actual width (46.5-20) was 2.33 mm. The area
of the wing was determined by moving the tracer arm of the planimeter
clockwise along the entire wing outline (Keuffel and Esser Co., 1963. Com-
pensating Polar Planimeter Instruction Manual, 32 pp.). Again, the reading
on the dial, (i.e., 0322 venier units or 32.2 cm2) is divided by 202 to obtain the
actual area of the wing (0.0805 cm2 or 8.05 mm2).-J. L. SHARP. Insect At-
tractants, Behavior and Basic Biology Research Laboratory, Agricultural
Research, Science and Education Administration, USDA, P.O. Box 14565,
Gainesville, FL 32604.

- -* -^- *- -- -^- -^--- -- --^

PINE GROWTH-(Note). Heavy infestations of a native pine tortoise scale,
Toumeyella parvicornis (Ckll.), (det. G. W. Dekle, Fla. Div. Plant Industry,
Gainesville) consistently attended by workers of the red imported fire ant,

I -7 N I**
m 7



1 2 3 4 5
Fig. 1. Mean heights of 2 groups (5 plots/group) of 3-year-old slash pines
in a Clay Co., FL plantation in July 1972. Pines heavily infested (=I; 5
trees/plot) with pine tortoise scale, Toumeyella parvicornis (Ckll.) tended
by red imported fire ants, Solenopsis invicta Buren, were only ca. 60% as tall
as insect-free pines (= NI, 5 trees/plot).

72 The Florida Entomologist 62(1) March, 1979

Solenopsis invicta Buren, (det. W. F. Buren, Univ. of Fla., Gainesville) were
found in a 3-year-old, bedded, typical slash pine (Pinus elliottii Engelm. var.
elliottii) plantation located at the junction of state hwys. 16 and 21 in
Clay County, FL during July 1972. At least 50% of the trees in a 40 ha area
were infested, resulting in pockets of chlorotic, stunted trees, often covered
with sooty mold. Measurements taken in 5 plots each with 5 infested and 5
non-infested trees showed that mean height growth was significantly (p -
.01) reduced by 40% in scale-infested trees (Fig. 1). A survey of 37 forested
counties in north Florida was conducted during the fall of 1972 by personnel
of the Florida Division of Forestry, but failed to detect similar damaging
scale infestations in young slash pine plantations. Live T. parvicornis scales
were not found in the Clay County plantation 1 year after the outbreak was
first detected and no similar outbreaks have been reported to date.
Toumeyella parvicornis commonly infests slash pine seedlings in 2-year-
old plantations in Florida, but seldom persists into the 3rd growing season
(unpubl.). Reasons for the above T. parvicornis-S. invicta outbreak are un-
known. The potential for damage to young pine plantations would appear to
be great if such scale-ant infestations should develop and persist in the
future.-R. C. WILKINSON, Univ. of Fla., Dept. of Entomology and Nema-
tology, Gainesville, and C. W. CHELLMAN, Fla. Div. Forestry, Tallahassee.

(Note). Six-spined pine engraver beetle adults sometimes make brood tun-
nels in the inner bark of living Pinus spp. (Felt 1906, N.Y. State Mus. Mem.
8: 349; Swaine 1918, Can. Dept. Agr. Bull. 14: 113; Blackman 1922, Miss.
Agr. Exp. Sta. Tech. Bull. 11: 114; Beal and Massey 1945, Duke Univ. Sch.
For. Bull. 10: 145). Facultative tunneling behavior observed during a host
resistance study (unpubl.) might explain the partial success of this species
in attacking living trees.
Adult Ips calligraphus attacks were induced on the scionwood portion of
32 (8 clones x 4 ramets) grafted slash pines by attaching attractive pine
bolts artificially infested with I. calligraphus males to the trunks (cf. Wilkin-
son 1964, Fla. Ent. 47: 57). Some ramets of 1 clone were characterized by
relatively low growth rate and large numbers of induced Ips attacks. Atyp-
ical tunnel patterns were most common in a tree which had a maximum
oleoresin exudation pressure (OEP) of only 2.6 atoms, and which was killed
by repeated attacks over a 4-week period. Atypical attacks were also found
in some of the remaining 31 trees (OEP : 8 atmos.), but none of these trees
Brood gallery systems of the kind usually present in non-resistant host
material were found in the inner baik of the attractive bolts (Fig. 1A). In
this case a male was found in the central nuptial chamber and 1 female in
each of the 3 adjoined egg galleries. In 1 apparently resistant tree (10
atmos. OEP), 6 live females were found side-by-side, tunneling vertically
upward in a broad tunnel in the inner bark (B). Females were almost en-
gulfed with oleoresin, which drained out of the bottom of the tunnel through
the entry hole. This same pattern was repeated in some of the other ap-
parently resistant trees. In the 1 tree which eventually died, adults had suc-

Scientific Notes

cessfully established 2 egg galleries in the bark, but they and their brood
were subsequently killed by infiltrating oleoresin (C).
The facultative tunneling behavior of I. calligraphus adults appears to
be related to the ability of this species to colonize living pine trees. The
broad, vertical tunnels made by I. calligraphus females in slash pine char-
acterized by relatively high OEP (B) are very similar to brood tunnels made
by the black turpentine beetle, Dendroctonus terebrans (Olivier), which also
attacks living pines.-R. C. WILKINSON, Dept. of Entomology and Nema-
tology, Univ. of Fla., Gainesville 32611.











Fig. 1. Typical (A) and atypical ( C) galleries produced by Ips cal-
ligraphus (Germ.) in slash pines.


74 The Florida Entomologist 62(1) March, 1979

Prof. Joseph R. Watson was the first editor of The Florida Entomologist
(nee Florida Buggist) and continued in that office for the first 28 volumes of
publication. It was only through his devotion and even personal financial ex-
penditure that the journal survived its early difficulties.
Prof. Watson was a charter member of the Florida Entomological So-
ciety and was its first president in 1916. He served again as president in
While Florida had several earlier Experiment Station entomologists, be-
ginning with a brief term by Dr. W. H. Ashmead in 1888, Prof. Watson
became the first chairman of an entomology department in Florida. His ap-
pointment was in the Experiment Station, where he served from 1911 until
his death in 1946.
He came to Florida from a professorship at the University of New Mexico
and earlier positions at Rochester College in Indiana, Berea College in
Kentucky, and Adelbert College of Western Reserve. He was awarded the
B.S. degree by Baldwin College and the A.M. by Western Reserve.
I thank Dr. A. N. Tissot for providing background material.-S. H. KERR,
Dept. of Entomology and Nematology, Univ. of Florida, Gainesville 32611.


! Cc~~ tp.I

Book Review

ICA. R. W. Merritt and K. W. Cummins, eds. 1978. Kendall/Hunt Publishing
Company, Dubuque, Iowa. xiii + 441 p., profusely illustrated. $19.95.
For the past several years aquatic biologists have looked forward to the
publication of a volume to replace the late Dr. Robert L. Usinger's Aquatic
Insects of California. At last Drs. Merritt and Cummins have brought to
completion the formidable task of assembling such a book with the assistance
of 21 specialists in the various insect orders.
The book will be of greatest use to those students taking courses in
aquatic entomology. Others, such as general biologists or entomologists con-
ducting surveys of the aquatic environment, will also find the book to be
most helpful to them in identifying their insects.
The editors, with others, have written introductory chapters, including 1
on general morphology, another on collecting and rearing methods, 1 con-
cerned with the ecology and distribution of aquatic insects, another treating
phylogeny and evolutionary adaptations, and 1 on general classification in-
cluding a key to orders. The bulk of the book is devoted to keys to the orders,
with each chapter written by a specialist or specialists. Except for the im-
portant aquatic families of Diptera, the keys stop at the family level. The
dipterous families Tipulidae, Culicidae and Simuliidae are keyed to genus
but the Chironomidae stop with the subfamily. The book is certain to be
criticized for not extending the keys of all orders to the generic level; how-
ever, were the authors to have done this, the size of the book would have been
substantially larger, publication delayed, and the cost prohibitive for a text-
book in courses in aquatic entomology.
The text drawings are clear, large, well labeled and excellently printed.
Near the end of each chapter there are additional taxonomic references to
help the user go beyond family level of identification if he needs to do so.
Each chapter ends with a summary table giving taxa, habitats, habits, trophic
relationships, North American distribution, and ecological references. The
book is printed in a large, easily read type and is well organized generally.
The extensive bibliography lists 1712 references. All in all, the editors,
authors, and publishers are to be congratulated on producing this excellently
assembled work.-LEWIS BERNER, Dept. of Zoology, Univ of Florida, Gaines-
ville 32611.

The Florida Entomologist 62 (1)

March, 1979


The first issue of The Florida Entomologist, Vol. 62, finds the Florida
Entomological Society in a period of transition and growth. In addition to
new officers and a new Editor (see inside cover), the Society has a new
printer, a slightly revised format for the journal and a few new instructions
to authors for preparation and submission of manuscripts.
The new printer is the E. O. Painter Printing Company of DeLeon
Springs, Florida. This organization is no stranger to technical publications;
they have printed the Journal of the Florida Soil, Crop and Science Society
since 1956, the Proceedings of the Florida State Horticultural Society since
1962, The Journal of Nematology since 1976, and other volumes in engineer-
ing, education and the social sciences. The Executive Committee feels that it
has made a good choice of printers in terms of economy, efficiency, and the
quality of printing, photographic reproduction, and binding. As an added
benefit, page charges for many but not all items in Vol. 62 of The Florida
Entomologist will be comparable with those charged for Vol. 61.
A glance at the March issue will show 2 changes in format. First, all
full-length papers are "placed continuously" in the first and longest section
of this and future issues. This means that when there is adequate space
following the end of 1 paper, another paper will be started on the same page.
Also, new papers may begin on either a left- or a right-hand page. Neither
practice will affect the appearance of reprints since extraneous printing will
be deleted before reprints are run. Set type can be manipulated so that all
reprints will begin on right-hand pages as before, thus conserving reprint
paper and placing all essential bibliographic information immediately before
the reader.
Shorter items including scientific notes, book reviews, photo stories, and
similar material are grouped toward the end of each issue. Like full-length
papers, these shorter items will be placed continuously, and the set type will
be manipulated so that reprints will include only the author's contribution.
The advantages for grouping items and continuous placement of manu-
scripts are obscure to those not acquainted with printing procedures for The
Florida Entomologist. In the past, the Editor was concerned with the ap-
pearance of not only the journal but also the reprints since the latter were
made from 300 unbound copies of the journal. Short items were inserted into
available blank spaces and unused, even-numbered pages after the galley
proofs were printed. The Editor proofed all shorter pieces at the page proof
stage (this follows the galley proof and precedes the actual printing of the
journal). With the new system and some additional procedures recommended
by the printer, all manuscripts, scientific notes, photo stories and other short
items are sorted, numbered and submitted to the printer in 1 package; this
saves time, handling and postage since the Editor and Printer are in different
towns. Every author proofs his contribution. Reprints now will be sold only
in multiples of 100; the minimum order will be 100 and the maximum will be
"unlimited," theoretically.
"Instructions to Authors" will appear in every issue. There are 2 changes
in these instructions that should be noted. When submitting a paper to the
Editor, please send the original manuscript plus original figures and tables
and 3 copies of the entire paper. The original will remain unmarked during
the review process as a reference copy for the author, Editor and Associate

Message from the Editor

Editors. Xerox copies are used for peer review. Associate Editors will make
every effort to recruit peer reviewers not employed by the same agency or
institution as the authors. Reviews from individuals working out-of-state or
in nearby countries (e.g., Canada) will be used where possible. This policy
is in keeping with national and international recognition of The Florida
Entomologist in entomology and acarology research circles.
The second change in "Instructions to Authors" requires the ribbon plus
one extra copy of the revised, reviewed manuscript. The extra copy is for the
Editor who retains it for safe-keeping until the manuscript finally appears
in print; the printer works with the original copy.
Authors still may use recent issues of The Florida Entomologist for ex-
amples of format. The name and year system for references is recommended
for the Literature Cited section of all full length manuscripts. This is the
style currently used in The Florida Entomologist; details and examples are
given in the CBE Style Manual, 3rd edition. Authors of scientific notes and
other short items will continue to insert abbreviated citations directly into
the text; this style is informative and also space-saving.

This article is the first in a series of messages from the Editor, other
members of the Executive Committee, and spokesmen of special committees
serving the Florida Entomological Society. Some of these messages will call
attention to changes in journal procedures while others will inform readers
of significant events in Society business. An interpretation of results from
the recent questionnaire to Florida Entomological Society members will be in
a later issue.

The Florida Entomologist 62 (1)

March, 1979


The Florida Entomological Society will hold its 62nd Annual Meeting on
4-7 September at the Ramada Inn West, 2121 West Tennessee Street, Talla-
hassee 32304 (904-576-6121, 800-228-2828). Room rates will be $26.00 double
and $19.00 single occupancy. The Inn is located on US Route 90 (Tennessee
St.) between US 27 and State Route 263 (Capital Circle). If arriving from
the south on SR 19 & 37, enter Tallahassee on SR 27, turn right on Monroe
Street and left on Tennessee. The registration fee will be $10.00 ($3.00
Student). Questions concerning local arrangement should be directed to:
R. Wills Flowers
Local Arrangements Chairman
Florida A&M University
P.O. Box 111
Tallahassee, FL 32307
If you plan to present a paper, the tear out sheet must be completed and
postmarked no later than 15 July 1979 and sent to:
N. C. Leppla, Program Chairman
P.O. Box 14565
Gainesville, FL 32604
A maximum of 8 minutes will be allotted for presentation of submitted
papers, with an additional 2 minutes for pertinent discussion. The best 3
student papers will be awarded monetary prizes based on content and de-
livery. Projection equipment for 2 x 2 slides only will be available. Authors
should keep slides simple, concise, and uncluttered with no more than 7 lines
of type on a rectangle 2 units high by 3 units wide. Abstracts will be pro-
vided at the meeting.

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