MICROSCOPY RESEARCH AND TECHNIQUE 45:313–324 (1999)

Sensilla on the Antenna and Ovipositor of the Parasitic Wasps

Trichogramma galloi Zucchi and T. pretiosum Riley
(Hym., Trichogrammatidae).
FERNANDO LUÍS CÔNSOLI,1* ELLIOT WATANABE KITAJIMA,2 AND JOSÉ ROBERTO POSTALI PARRA1
1Department of Entomology, Escola Superior de Agricultura ‘‘Luiz de Queiroz’’ - ESALQ/USP, 13418–900, Piracicaba, SP, Brazil
2NAP-MEPA, Núcleo de Apoio à Pesquisa em Microscopia Eletrônica na Agricultura, ESALQ/USP

KEY WORDS in vitro rearing; host selection; host recognition; host evaluation; host suitability;
biological control; artificial diets; artificial host
ABSTRACT The understanding of the stimuli perceived by these parasitoids to accept and
exploit a potential host may support the development of artificial hosts for their in vitro rearing. The
most common structures involved in the host selection process in parasitoids are the antenna and
the ovipositor. Sensilla present on these structures are involved in host recognition and acceptance,
and the identification of which kind of stimulus (physical or chemical) is perceived by the sensilla
could provide data about the host characteristics that elicit the parasitism behavior. The antenna
and ovipositor of the wasps Trichogramma galloi and T. pretiosum were studied to identify their
sensillar structure and their possible functions in the host selection process, as a support to the
development of an artificial host for the in vitro rearing of these parasitoids. Seven sensilla and one
setiform structure were present on the antenna of both parasitoid species, with a mixed, chemo-, or
mechanoreception function. The ovipositor has four different sensilla and their role in the host
evaluation is suggested.Microsc. Res. Tech. 45:313–324, 1999. r 1999 Wiley-Liss, Inc.

INTRODUCTION (Schmidt and Smith, 1985, 1987). Sensilla on the tip of

Trichogrammatidae, a widespread family of Chalcidoi-
dea consisting of 620 species in 80 genera, are egg
parasitoids that ranges in length from 0.2 to 1.5 mm.
Trichogramma and Oligosita are the most speciose
genera of this family, representing 40% of its diversity
with approximately 145 and 110 species, respectively.
The large number of species described for Tricho-
gramma is related to its importance as a biological
control agent; and, although it is the most well-known
genus in this family, Trichogramma is not the most
typical nor the most common genus (Pinto and
Stouthamer, 1994).
Trichogramma is a cosmopolitan genus, consisting
exclusively of egg endoparasitoids. They are usually
associated with eggs of moths (Fig. 1), but also have
been recorded on eggs of beetles, flies, bugs, hoppers,
wasps, and lacewings (Pinto and Stouthamer, 1994).
Adults of Trichogramma are 0.2–0.3 mm long, and they
parasitize single or clustered host eggs from 0.3 mm
(0.014 mm3) to 3 mm (14 mm3) in size (Pinto, 1997;
Schmidt, 1994). Host recognition and acceptance are
driven by behavioral mechanisms by using sensilla on
the antenna and ovipositor that assesses the suitability
of a host by examining the host’s surface and internal
contents (Schmidt, 1994). After contacting the host,
females of Trichogramma exhibit intensive antenna-
tion of the host’s surface, drilling the chorion before its
acceptance (Klomp and Teerink, 1962; Salt, 1935). The
antenna is also important in measuring the curvature
of the host, which will be used as an estimate of the host
size. Trichogramma will use this information to decide
the clutch size to be allocated in that particular host
the ovipositor also seem to be used for the evaluation of
the internal quality of the host or at least to recognize
ovipositional stimulants (LeRalec and Wajnberg, 1990;
Nettles et al., 1983; Qin and Wu, 1988; Rajendram,
1978).
Trichogramma is an important biological control
agent and it has been released against many different
pests on a wide variety of agroecosystems in approxi-
mately 32 million hectares/year worldwide (Li, 1994). It
has been successfully mass-produced on eggs of moths
of stored products, especially Sitotroga cerealella (Ol.)
(Lep., Gelechiidae), Anagasta kuehniella (Zeller), and
Corcyra cephalonica (Stainton) (Lep., Pyralidae) since
1930 when Flanders reported the possible use of eggs of
non-natural hosts (factitious hosts) to mass produce
Trichogramma (Flanders, 1930; Li, 1994; Smith, 1996).
The use of biological control through entomophagous
insects is a very old science and has been implemented
with success in many different agroecosystems. It is one
of the best biological methods to control insect pests and
avoid contamination from broad-spectrum toxic pesti-
cides. This biological method is also compatible with
any other method of pest control in an integrated
management program (DeBach 1974; Flint and van den
Bosch 1981; Hoy and Herzog, 1985; Huffaker, 1971;
Rosen et al., 1994; Stehr, 1975).
Contract grant sponsor: Fundação de Amparo à Pesquisa do Estado de São
Paulo (FAPESP).
*Correspondence to: Fernando Luı́s Cônsoli, Texas A&M University, Entomol-
ogy Research Laboratory, College Station 77843–2475, TX.
E-mail: f-consoli@tamu.edu
Received 25 June 1998; accepted in revised form 12 February 1999

r 1999 WILEY-LISS, INC.

314 F.L. CÔNSOLI ET AL.
Fig. 1. Female of the parasitic wasp Trichogramma pretiosum parasitizing an egg of a moth, the corn
earworm, Helicoverpa zea (Lep., Noctuidae) (photo by H. Negri de Oliveira).
Because of its economical importance, new lower cost
and low labor technologies for mass production of these
minute wasps have been developed. The in vitro rearing
system for the mass production of parasitoids has been
intensively studied since the 1970s, with the develop-
ment of artificial diets for 72 species of parasitoid
wasps, of which 18 were species of Trichogramma
(Cônsoli and Parra, 1997). This new technique of insect
rearing allows the production of the natural enemy
without the necessity of rearing a second species (facti-
tious host), therefore, reducing the costs and labor of
the rearing system. However, only Chinese researchers
have been successful in mass producing and releasing
in vitro–reared parasitoids (Cônsoli and Parra, 1997;
Grenier, 1994, 1997; Li, 1992). The development of an
artificial diet and artificial host that allows the parasi-
toids to grow up from egg laying to adult emergence is
essential to establish in vitro rearing techniques.
The development of a suitable artificial diet depends
on basic knowledge in fields such as functional morphol-
ogy, physiology, nutrition, and parasitoid/host interac-
tions. The intensity of research in each of these fields
depends whether an endo- or ectoparasitoid is being
reared (Vinson, 1994). For the development of an
artificial egg, it is necessary to understand the mecha-
nisms involved in the host selection process. Chemical
and/or physical cues are always associated with host
recognition and exploitation by parasitoids (Vinson,
1976, 1985). The knowledge of what kind of stimulus
(physical and/or chemical) is used by parasitoids to
select a host would help the development of attractive
artificial hosts and diets.
After parasitoids locate the habitat of their hosts in
field conditions, they usually locate and recognize their
hosts by stimuli produced by the host (Bin et al., 1989;
Cave and Gaylor, 1987; Isidoro et al., 1996; Noldus and
van Lenteren, 1985; Vinson, 1985; Weseloh, 1972).
Sensillar structures present on the antenna of parasi-
toid females play a basic role in this process, recogniz-
ing chemical and/or physical stimuli produced by the
host (Hays and Vinson, 1971; McIver, 1975; Schneider
and Steinbrecht, 1968; Zacharuk, 1980, 1985). Addition-
ally, the ovipositor also plays an important role in host
recognition and acceptance by responding to chemicals
associated with the host, besides being used only as a
puncturing or a poison injection organ (Hawke et al.,
1973; Le Ralec and Wajnberg, 1990, 1996; van Veen and
Wijk, 1985; Vinson, 1975ab). Also, the ovipositor can
work to externally mark the host after parasitism
(Hokyo et al., 1966; Rabb and Bradley, 1970; Strand
and Vinson, 1983).
In the present work, we describe the external mor-
phology of the sensilla on the antenna, with especial
reference to the ones located on the last antennomere,
and ovipositor of the two most studied Trichogramma
species in Brazil, T. galloi and T. pretiosum. These data
when used in association with behavioral studies will
allow us to better understand their roles during the
host selection process (host location, recognition, and
acceptance) (Klomp and Teerink, 1962; Klomp et al.,
 SENSILLA OF T. GALLOI AND T. PRETIOSUM
Fig. 2. Antennae of Trichogramma galloi showing the general
distribution of the sensilla. The only difference from that of T.
pretiosum is the presence of one sensillum basiconica instead of two at
the fourth flagellomere. A: Dorsal view; B: Lateral view; C: Ventral
1980; Pak, 1988; Schmidt and Smith, 1985 Salt, 1935,
1937). Also, these studies should provide information
for the development of a suitable artificial host for the
development of in vitro rearing techniques of both
species.
MATERIALS AND METHODS
Stock Culture of Parasitoids
Trichogramma galloi and T. pretiosum were reared
on UV-treated eggs of the factitious host, Anagasta
kuehniella (Lep., Pyralidae), and held in incubators
(temperature: 25 ⫹ 1°C; relative humidity: 60 ⫹ 10%
RH; photophase: 14 h). A. kuehniella was bred on
artificial diets composed of wheat flour and yeast or
corn and wheat flour (Parra 1997), following the tech-
niques described by Parra et al. (1989).
Scanning Electron Microscopy
Antenna of newly emerged, ether-anesthetized fe-
males of T. galloi and T. pretiosum were fixed in 3%
glutaraldehyde in a 0.1 M phosphate buffer solution
(pH 7.2) for 2 hours, dehydrated in acetone (30–100%),
critical point dried in a CO2 Balzers Critical Point
Dryer CPD 030, sputter coated (Balzers Union MED
010 evaporator) with gold, and viewed with a Zeiss
315
view. a ⫽ uniporous sensilla trichoidea; b ⫽ multiporous sensilla
trichoidea type C; c ⫽ multiporous sensilla trichoidea; d ⫽ aporous
sensilla trichoidea; e ⫽ multiporous sensilla placoidea; f ⫽ sensilla
basiconica; g ⫽ setiform structure; h ⫽ coeloconicum sensillum.
(Thornwood, NY) DSM940A scanning electron micro-
scope (SEM) at an accelerating voltage of 5.0– 10.0 kV.
The structure of the ovipositor was observed after
dissection of newly-emerged females previously fixed as
described above. The proximal portion of the ovipositor
was glued on a cover slide, mounted on a stub, air-dried,
sputter-coated with gold, and examined in the SEM.
Ovipositors were also observed by exposing their tip
from the genital capsule of fixed females.
RESULTS AND DISCUSSION
Antennae
Seven morphologically distinct sensillar structures
and one setiform structure were found on the antennae
of females of Trichogramma galloi and T. pretiosum.
The overall structures are essentially similar to what
have been previously described for other Tricho-
gramma species (Amornsak et al. 1998; Olson and
Andow, 1993; Voegelé et al., 1975). Antennae of T. galloi
and T. pretiosum are composed of 7 segments: scape,
pedicel, and 5 flagellomeres (Figs. 2, 3A). The first
flagellomere is cylindrical, short, and slightly longer
than wide. The second is smaller, and when viewed
laterally, partially covers the third flagellomere. The
316 F.L. CÔNSOLI ET AL.
Fig. 3. General structure of the antenna of Trichogramma. A:
Scape (S), pedicel (P) and flagellomeres 1 to 5 (F); B: Flagellomere 2 to
4. Arrows indicate the distribution of the coeloconicum sensillum.
third and fourth flagellomeres are alike. They are
cylindrical, being as long as wide. The last flagellomere,
clava, is considerably larger than the others and is
endowed with many more sensillar structures than the
other flagellomeres (Figs. 2, 3).
Antennal Sensilla
Although Amornsak et al. (1998) have recently pro-
posed a new terminology for the sensilla of Tricho-
gramma based only on external morphology, we decided
to follow the nomenclature previously used by Olson
and Andow (1993) that was based on one proposed by
Snodgrass (1935) and Zacharuk (1980). We also decided
on the use of this terminology to avoid proposing new
names for structures without knowing their real func-
tion, because transmission electron microscope were
not used by Amornsak et al. (1998).
The sensilla present on the antennae of females of T.
galloi and T. pretiosum are also found in parasitic
wasps of other families. The aporous sensillum trichoi-
dea is found in 14 other species of parasitoids, the
uniporous sensillum trichoidea in 11 spp., the multi-
porous sensilla trichoidea type C in 5 spp., the multi-
porous sensillum trichoidea in 9 spp., the multiporous
sensillum placodea in 17 spp., the multiporous grooved
basiconica sensillum in 15 spp., and the coeloconica
sensillum in 5 spp. These numbers were adapted from
Olson and Andow (1993) with the inclusion of T. nubi-
lale, T. australicum, T. maltbyi, and T. brasiliensis
(Amornsak et al., 1998; Voegelé et al., 1975). They
represent 19 species from eight different families of
parasitic Hymenoptera (wasps).
Aporous Sensilla Trichoidea (d in Figs. 2, 4A,C,
5B). These sensilla are found on the scape, pedicel,
and flagellomeres. They are long (about 40 µm) with a
ridged surface inserted in a protuberance (Fig. 5B).
They are usually identified as a mechanosensilla, some-
times perceiving airborne vibrations because they con-
tain tubular bodies in which the dendrites terminate, a
feature typical of mechanoreceptors (Barlin and Vin-
son, 1981; McIver, 1975). In Trichogramma, it has been
suggested that they may provide females with informa-
tion regarding the position of their antennae in relation
to the substrate, based on their distribution pattern
along the antennae (Olson and Andow, 1993). Such
information must be useful in host searching and
exploitation, during which Trichogramma females ac-
tively employ their antennae (Schmidt and Smith,
1985).
Uniporous Sensillum Trichoidea (a in Figs. 2,
4A,B, 5E,F). This sensillum, about 10 µm long, slightly
curved, is exclusively ventrally located at the tip of the
clava. It is inserted on a distinct elevated socket and is
provided with longitudinal ridges without any visible
pore. At the tip of the sensillum there is a single pore.
Olson and Andow (1993) reported the presence of this
sensillum on the male antenna, but in different posi-
tions from that of females.
Multiporous Sensilla Trichoidea. Two types of
multiporous sensilla trichoidea were present in the
antenna of T. galloi and T. pretiosum females differing
in their position and type of insertion. Type C is
ventrally located in the clava, being a curved and
laterally flattened structure with oblique grooves along
its surface (b in Figs. 2, 4A,B, 5A). Pores are located
along the outer margin of the sensillum at the point of
groove convergence. Zacharuk (1980) designated this
kind of sensillum as a multiporous pitted sensillum
because the pores are uniformly distributed along the
wall while a multiporous grooved sensillum show these
pores within the grooves present along its length. These
sensilla are found in larger number on T. pretiosum
antenna (x̄ ⫾ sd - 30.7 ⫾ 0.80) than on T. galloi (x̄ ⫾ sd -
26.6 ⫾ 0.63), being statistically different between these
species (n ⫽ 23; F ⫽ 60.6; df ⫽ 1, 45; P ⬍ 0.05, by
Tukey’s test). The number of these sensilla seems to be
species-specific for Trichogramma. Voegelé et al. (1975)
had already noticed such differences for other species of
these egg parasitoids, T. maltbyi, T. brasiliensis, and T.
evanescens, and they suggested the possibility of using
this character to distinguish Trichogramma species
using females instead of males. This would be a solu-
 SENSILLA OF T. GALLOI AND T. PRETIOSUM 317

Fig. 4. Sensilla distribution on the last flagellomere (clava). A: Lateral view; B: Ventral view;
C: Dorsal view. a ⫽ uniporous sensilla trichoidea; b ⫽ multiporous sensilla trichoidea type C; c ⫽
multiporous sensilla trichoidea; d ⫽ aporous sensilla trichoidea; e ⫽ multiporous sensilla placoidea; f ⫽
sensilla basiconica; g ⫽ setiform structure.
 Fig. 5. Antennal sensilla of T. galloi and T. pretiosum. A: Multi-
porous sensilla trichoidea type C. Arrow indicates the type of basal
insertion; B: Aporous sensillum trichoidea. Arrow indicates the type of
basal insertion; C: Multiporous sensillum trichoidea. Arrow indicates
the type of basal insertion; D: Detail of a multiporous sensillum. Arrow
indicates a pore; E: Uniporous sensillum trichoidea (opened arrow)
and the basiconica sensillum (arrow head) at the tip of the last
flagellomere; F: Close-up of the pore at the tip of the uniporous
sensillum, with the arrow indicating the pore.
 SENSILLA OF T. GALLOI AND T. PRETIOSUM 319

Fig. 6. Antennal sensilla of T. galloi and T. pretiosum. A: Multiporous sensillum placoidea, flattened
structures attached to the surface of the antennae; B: Surface of the sensillum placoidea showing the
pores; C: Basiconica sensillum, peglike structures; D: Coeloconicum sensillum, depressed structure with
a single, central pore.

tion to identify those parthenogenetic Trichogramma
strains that produce an exclusive female progeny.
Because the multiporous sensilla trichoidea are al-
ways contacting the host surface during the drumming
behavior, some authors have suggested they may be
related with host exploitation by the parasitoid (Olson
and Andow, 1993). Yet, recent ultrastructural studies
developed by Isidoro et al. (1996) revealed a gustatory
function for these sensilla, implying chemical recogni-
tion. If these sensilla are involved in the host recogni-
tion process, such chemicals should be available on the
host surface to induce host recognition or acceptance.
However, data available on the parasitization of artifi-
cial plastic eggs by T. galloi and T. pretiosum provide
evidence that both species of parasitoids do not need
chemical stimulants on the host surface for their evalu-
ation and acceptance (Cônsoli and Parra, 1999). Never-
theless, the gustatory function attributed to these
sensilla in the host selection process could be associated
with host rejection instead of host acceptance. This can
be true in the case where moths cover their eggs with
scales that have chemicals that disturb the parasitiza-
tion behavior of Trichogramma, like in the association
between Spodoptera litura and T. chilonis (Ohta et al.,
1994). Yet, the identification of chemicals associated
with the gustatory sensilla from Trichogramma differs
from that of other species because there is no secretory
glands associated with them (Isidoro et al., 1996). This
fact brings up the question on how the chemical stimu-
lus from the host surface is transported to the sense
organs if secretions are not produced.
Another type of multiporous sensillum trichoidea
present on the antenna of T. galloi and T. pretiosum has
a distinct shape and insertion. It is latero-dorsally
320 F.L. CÔNSOLI ET AL.

Fig. 7. Thin cross section of the ovipositor

of Trichogramma. A: Ovipositor enclosed by a
sheath of the genital capsule; B: Detail of the
ovipositor (V1 ⫽ valvula 1, V2 ⫽ valvula 2; s ⫽
sheath of the genital capsule; NS ⫽ nervous
system).
 SENSILLA OF T. GALLOI AND T. PRETIOSUM
located on the clava, and its basal insertion is continu-
ous with the tegument (c in Figs. 2 and 4A,C, 5C,D). It
has a reduced number of pores and has been associated
with olfaction (Olson and Andow, 1993).
Multiporous Sensilla Placodea. Both Tricho-
gramma species under study show 5 long (up to 25 µm),
flattened (about. 2 µm wide) structures attached to the
antennal surface throughout their length except for
their tip. They are laterally and dorsally located on the
clava, running almost half of the clava length. Pores are
abundantly located on the sensillum surface (e in Figs.
2, 4A,C, 6A,B). Doutt (1964) suggested parasitoids may
use these sensilla for long-range chemical orientation
allowing the detection of chemicals produced by the
host in its habitat, and such a possibility has been
extended to many other parasitoids species (Schneider
and Steinbrecht, 1968; Slifer, 1969; Weseloh, 1972;
Norton and Vinson, 1974; Barlin and Vinson, 1981).
Specifically for Trichogramma, the multiporous sen-
silla placoidea may be used to orient parasitoid females
to the sex pheromone released by female hosts, which
may be used by parasitoids to locate a habitat with
possible egg hosts. Sex pheromones, accessory gland
secretions, and chemicals present on adult host scales
were identified as the major chemical components
involved in parasitoid foraging behavior (Noldus, 1989).
Multiporous Grooved Basiconica Sensilla. These
are short (2 µm), with a bulbous tip (about 1.5 µm in
diameter), with a short (about 1 µm), cylindrical stalk
with pores longitudinally distributed along the bulb
region, and are found on the 3rd, 4th, and 5th antennal
segments (f in Figs. 2, 4A, 6C). The one found at the tip,
on the ventral surface of the clava, shows a distinct
bulbous shape. It seems to be more slender than all the
other multiporous grooved basiconica sensilla (MGBS)
(Fig. 5E), as described by Amornsak et al. (1998). There
were differences in the number of this type of sensillum
in the 4th segment between T. pretiosum (2) and
T. galloi (1). Multiporous grooved sensilla have been
identified on the antennae of many species and have
been shown to have many different olfactory functions,
with their functional specificity depending upon their
wall thickness. Thick-walled sensilla were suggested to
be more selective to specific source of chemicals, such as
pheromones, than the thin-walled ones. Thermo and/or
hygrosensitive capabilities were also suggested for the
multiporous grooved sensilla in many other species
(Bin et al., 1989; Olson and Andow, 1993; Zacharuk,
1985).
Coeloconic Sensilla (h in Fig. 2; Figs. 3B, 6D).
These sensilla are located on the 2nd and 4th flagello-
meres, being an almost circular (5 µm in diameter),
depressed structure with a very small (approximately
0.3 µm) single pore in the center. Although these
sensilla are similar on T. galloi and T. pretiosum, they
have a variable shape and distribution on other Tricho-
gramma species. Their number and distribution are
similar to that on T. australicum (Amornsak et al.,
1998), but differ from that on T. nubilale, which has
three of these structures distributed on the 3rd, 4th,
and 5th segments (Olson and Andow, 1993) while T.
evanescens has two, but on the 4th and 5th segments
(Voegele et al., 1975). Olson and Andow (1993) de-
scribed these sensilla as campaniform-like structures
but suggested no function for them. However, these
321
sensilla are very similar to those described for worker
honey bees, believed to be sensitive to one or more
stimuli, such as temperature, CO2, and humidity based
on electrophysiological findings (Schneider, 1969; Dietz
and Humphreys, 1971).
Setiform Structures (g in Figs. 2, 4A,B). They are
needle-like (6–7-µm-long) appendages, with indistinct
basal insertion, without pores, and possibly without
receptive function. They appear over the entire anten-
nal surface, but more frequently on the ventral side of
the clava, near the base (Fig. 4). Because they are
present in large numbers and widely distributed, Olson
and Andow (1993) suggested they may have a protec-
tive function.
Ovipositor (Figs. 7, 8)
The overall organization of the female genitalia of
T. galloi and T. pretiosum is similar to that described to
T. maidis (Le Ralec and Wajnberg, 1990), and not
different from the basic genitalia structure of Hymenop-
tera (Smith, 1970). The ovipositor consists of two pairs
of valvifers and three pairs of valvulae derived from the
8th and 9th abdominal segments (Le Ralec and Wajn-
berg, 1990). The ovipositor is cylindrical and needle-
like in shape, and is encased by a sheath. It is composed
by three coupled styli (two valvulae 1 and one valve 2),
with many distally located sensilla (Figs. 7, 8A). Trans-
verse sections showed that valve 2 functions as a
conductor of valvulae 1, holding all styli together and
interdigitated (Fig. 7A,B), while the female drills the
host surface and oviposits. Valve 2, which is the stylus
facing the abdomen, has a depression near to the distal
portion, and has many irregular cuticle projections at
the tip (Fig. 8B). Four needle-like projections are
present laterally on each side of the stylus (Fig. 8C).
These projections are slanted normally downward,
towards the ovipositor base and are probably used to
pierce the chorion of the egg host. Both valvulae 1 have
four types of sensillar structures distributed proxi-
mately (a,b,c,d in Fig. 8D), behind two lateral cuticular
expansions at their tip (Fig. 8D). Two hook-like sensilla
within an ovoid depression (a, Fig. 8D) and a pore at
their base are located distally on the valvulae 1.
Towards the proximal region, ventrally located in rela-
tion to these sensilla, there is another sensillum (b, Fig.
8D), with an irregular and depressed cuticular struc-
ture. This is followed by a tusk-like projection (c, Fig.
8D) evenly distributed along a line on the surface of the
stylus, just after a cuticular constriction (Fig. 8A,E).
After the first tusk-like projection there is a non-
differentiated structure (d, Fig. 8D), indicated by a
circular depression of the cuticle, resembling a campani-
form sensillum. Both valvulae 1 have a great number of
projections arising from the internal surface (Fig. 8E).
Thin section studies under transmission electron
microscope of these sensilla in other parasitoid species,
disclosed a chemo and/or mechanoreceptor function
(Greany et al., 1977; Hawke et al., 1973; van Veen and
van Wilk, 1985). However, Le Ralec and Wajnberg
(1990) suggested a possible mixed function for sensil-
lum a but only mechanoreception for sensilla c and d.
However, no function was suggested for sensillum b (Le
Ralec and Wajnberg, 1990).
Chemoreceptors on the ovipositor of Trichogramma
should be expected because they are used by the
 Fig. 8. Structure of the ovipositor of Trichogramma. A: Ovipositor
partially distended from the genital capsule. The arrow indicates the
position of a tusk-like structure present on the ovipositor surface, a
detail of which is in the insect; B: Cuticle projections on the ventral
side of the valvula 2; C: Cuticle constriction on the upper surface of the
valvulae 1; D: Sensilla on the tip of the ovipositor. a ⫽ sensillum a
(mixed function); b ⫽ sensillum b (no suggested function); c ⫽
sensillum c (mechanoreceptor); d ⫽ sensillum d (proprioreceptive
mechanoreceptor). Arrow indicates a pore (detail: 30,000x); E: hook-
like structures on the tip of the valvifer 2 (* indicates the presence of
cuticle projections on the internal surface of the valvula 1); F: Detail of
cuticle projections on the internal surface of the valvula 1.
 SENSILLA OF T. GALLOI AND T. PRETIOSUM
parasitoid to evaluate the host suitability. Chemical
substances (amino acids and inorganic ions) are per-
ceived by the ovipositor and stimulate the female to
oviposit (Nettles et al., 1982; Wu and Qin, 1982). These
chemical stimulants are probably recognized by sensil-
lum a, the only one provided by pores. Sensillum d may
act as a proprioreceptor to indicate the relative position
of the ovipositor within the host egg while female is
drilling and ovipositing. Such information would be
useful especially when oviposition occurs in host eggs
partially hidden under others, as suggested for the
sense organs found on the legs of T. maidis (Cals-
Usciati et al., 1988). The mechanoreceptor role sug-
gested for sensilla c and a by Le Ralec and Wajnberg
(1990) may be related to the detection of the host’s
embryonic development through the perception of physi-
cal changes inside the host, although these alterations
are also followed by drastic changes in the chemical
composition of the host (Vinson, 1994).
Based on the sensillar structures found on the an-
tenna and ovipositor of T. galloi and T. pretiosum and
their suggested function in the host selection process,
we conclude that both species of parasitoids do not need
contact chemical stimulants for host recognition and
the chemoreceptors found on the antenna are used for
short- and long-range chemicals as those involved in
habitat and host location. Host recognition for both
species relies on the physical cues of the host upon
contact. However, chemical stimuli are involved in host
acceptance since the chemoreceptors of the ovipositor
may only act upon contact with the host contents. These
conclusions suggest that special attention should be
addressed on the physical traits of the host for the
development of artificial host eggs as well as on the
chemical contents of the artificial diet (not only on its
nutritional quality), because both physical and chemi-
cal stimuli must be provided to elicit host recognition
and acceptance of artificial hosts in an in vitro rearing
system.
ACKNOWLEDGMENTS
We thank Dr. F. Bin and Dr. N. Isidoro (University of
Perugia, Italy) and Dr. S. B. Vinson and Dr. S. O’Keefe
(Texas A&M University, College Station, USA) for
reviewing the early version of this manuscript. We also
thank the Fundação de Amparo à Pesquisa do Estado
de São Paulo (FAPESP) for providing finnancial sup-
port.
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