Micron 35 (2004) 671–679

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Antennal sensilla of some forensically important flies in families

Calliphoridae, Sarcophagidae and Muscidae
Kom Sukontasona,*, Kabkaew L. Sukontasona, Somsak Piangjaia, Noppawan Boonchua,
Tarinee Chaiwonga, Radchadawan Ngern-kluna, Duanghatai Sripakdeea,
Roy C. Vogtsbergerb, Jimmy K. Olsonc
a
Department of Parasitology, Faculty of Medicine, Chiang Mai University, Chiang Mai 50200, Thailand
b
Department of Biology, Hardin-Simmons University, Abilene, TX 79698-6165, USA
c
Department of Entomology, Texas A&M University, College Station, TX 77843-2475, USA
Received 1 March 2004; revised 14 April 2004; accepted 20 May 2004

Abstract
Antennal sensilla of some forensically important fly species in the families Calliphoridae (Chrysomya megacephala, Chrysomya rufifacies,
Chrysomya nigripes and Lucilia cuprina), Sarcophagidae (Parasarcophaga dux) and Muscidae (Musca domestica) were studied using
scanning electron microscopy. Five types of sensilla were observed: trichoid, basiconic, coeloconic, styloconic and sensory pit. Only trichoid
sensilla are found on the scape of the antenna, while both trichoid and styloconic sensilla are located on the antennal pedicels of all species
studied. Basiconic sensilla are the most numerous of the sensilla found on the antennae of both sexes of all fly species studied and are
comprised of two subtypes: large and small basiconic sensilla. Coeloconic sensilla are characterized by short pegs, with either grooved or
smooth surfaces, that are sunken into deep depressions. No marked difference was observed in the number, morphological structure or
distributional pattern of any of the sensilla among the species studied, with the exception of there being more numerous sensory pits detected
in female P. dux compared to the other species. The suggested function of each antennal sensillum was based on comparison with results of
other investigations on similar sensilla.
q 2004 Elsevier Ltd. All rights reserved.
Keywords: Antenna sensilla; Flies; Receptor; Scanning electron microscopy; Forensic entomology

1. Introduction arthropod succession that inhabit decomposing human

The life cycle of fly species in the families Calliphoridae
(blow flies), Sarcophagidae (flesh flies) and Muscidae
(house flies and relatives) are routinely involved in forensic
investigations. The presence of eggs, larvae and puparia of
fly species belonging to these families collected from human
corpses at death scenes provides entomological information
that is not only useful in estimating the postmortem interval,
but may also be used for analyzing toxic substances that
could have led to death (Smith, 1986; Lord, 1990; Introna
et al., 1998; Goff, 2000; Byrd and Castner, 2001; Greenberg
and Kunich, 2002). According to Smith (1986), flies
belonging to these families are frequently recorded as
among the first insects to arrive at a body in the waves of
* Corresponding author. Tel.: þ 66-53-945342; fax: þ 66-53-217144.
E-mail address: ksukonta@mail.med.cmu.ac.th (K. Sukontason).
0968-4328/$ - see front matter q 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.micron.2004.05.005
cadavers. In particular, species of blow flies have been
shown to arrive at carcasses and lay eggs within a few
minutes postmortem in experimental studies (Goff, 2000;
Byrd and Castner, 2001; Greenberg and Kunich, 2002).
In order to locate the remains, often from long distances,
flies must be equipped with excellent olfactory organs to
detect chemical cues such as ammonia, ammonium carbon-
ate, and sulfur-rich volatiles being produced by corpse
decomposition (Greenberg and Kunich, 2002). Antennal
sensilla in many insects, including flies, have been
extensively recorded as bearing several olfactory receptors
(DeVaney et al., 1970; Greenberg, 1970; Shanbhag et al.,
1999; Gullan and Cranston, 2000; Ochieng et al., 2000;
Merivee et al., 2002). Several investigations of antennal
sensilla in various insect groups have been performed to date,
but relatively little information involving fly species of
importance in forensic entomology exists. Due to this
672 K. Sukontason et al. / Micron 35 (2004) 671–679

deficiency, the purpose of this study was to investigate the

sensilla found on all antennal segments of six species of flies
in the three fly families of greatest forensic importance.

2. Materials and methods

Six species of adult flies were obtained from laboratory

colonies maintained at the Department of Parasitology,
Faculty of Medicine, Chiang Mai University, Thailand.
Four species of adult flies in the family Calliphoridae
(Chrysomya megacephala, Chrysomya rufifacies, Chryso-
mya nigripes and Lucilia cuprina), one species of
Sarcophagidae (Parasarcophaga dux) and one species of
Muscidae (Musca domestica) were used for the investi-
gation. For each species, 15 flies of each sex were processed
for scanning electron microscopy.
Adult flies were first transferred into transparent plastic
bags using a glass tube and were then killed by placing the
bag in a freezer set at 4 8C for 20 min. Each killed fly was
placed on a glass slide and cut under a dissecting
microscope using a scalpel blade in order to obtain only
the head portion for further SEM processing, as described
by Sukontason et al. (2003).
3. Results
All calliphorid, sarcophagid and muscid flies bear
aristate antenna frontally and centrally on the head and
are situated between the large compound eyes as can be seen
in the male C. megacephala in Fig. 1. The antennae contain
three segments: a proximal scape, pedicel and distal
flagellum (Fig. 2). All antennal segments contain sensilla;
however, the most numerous sensilla are concentrated on
the flagellum. For the purpose of classifying sensilla in this
study, the terminology of Zacharuk (1985) was adopted.
Fig. 2. SEM micrograph of features on antennal pedicels of flies. Antennal
morphology of male M. domestica comprised of three antennal segments:
scape (I), pedicel (II) and flagellum (III) bearing a dorso–laterally located
arista (AR). Arrowhead indicates position of styloconic sensilla.
Bar ¼ 100 mm.
3.1. Scape
Unlike in most insects, the scape is the shortest segment
of the antennae in the flies that were studied (Figs. 2,6 and 7).
Only one type of sensillum, known as a trichoid sensillum,
was seen on the scape. Trichoid sensilla are found in a
curved single row on the scape, but their number varied
among the fly species examined (Table 1). Trichoid sensilla
are characterized as long hair-like sensilla with incised
longitudinal grooves and acute tips (Fig. 3). The base of
each sensillum is fitted tightly in a slightly elevated, smooth
socket. The length of this type of sensillum was more or less
comparable in each specimen studied. The cuticular surface
of the scape is otherwise, densely covered with non-
innervated microtrichia or spinules, which are small hair-
like structures with sharp-pointed tips.
Table 1
Mean number ^ SD of sensilla on the antennal scape and pedicel of adult
flies

Fly species Sex No. Scape Pedicel

Trichoid Trichoid Styloconic

C. megacephala M 10 3.9 ^ 0.9 30.2 ^ 2.9 2.0 ^ 3.7

F 10 4.9 ^ 0.9 28.2 ^ 3.7 7.0 ^ 2.0
C. rufifacies M 10 3.6 ^ 0.9 22.9 ^ 2.7 5.5 ^ 0.7
F 10 3.9 ^ 0.9 33.1 ^ 3.5 6.1 ^ 1.1
C. nigripes M 9 5.6 ^ 1.1 25.0 ^ 3.2 3.3 ^ 1.4
F 10 5.5 ^ 1.6 22.7 ^ 3.4 3.1 ^ 0.7
L. cuprina M 10 6.7 ^ 0.9 22.1 ^ 3.5 8.2 ^ 1.7
F 10 6.0 ^ 0.7 24.2 ^ 3.1 9.6 ^ 2.1

P. dux M 9 6.2 ^ 1.3 37.9 ^ 4.7 5.0 ^ 1.0

F 7 7.3 ^ 1.5 40.4 ^ 6.6 6.1 ^ 0.9
Fig. 1. SEM micrograph of features on antennal pedicels of flies. Head of M. domestica M 10 6.1 ^ 1.3 23.0 ^ 2.9 2.1 ^ 0.7
male C. megacephala showing antennae (A) located centrally between F 10 6.8 ^ 1.0 23.5 ^ 2.9 1.6 ^ 0.8
large compound eyes (E), dorsal to the maxillary palps (MP) and proboscis
(P). Bar ¼ 1 mm. M ¼ Male, F ¼ Female.
 K. Sukontason et al. / Micron 35 (2004) 671–679
Fig. 3. SEM micrograph of features on antennal pedicels of flies. Cuticular
surface of pedicel of female M. domestica showing dense covering of non-
innervated microtrichia or spinules (SP) interspersed with large trichoid
sensilla (T) emerging from flexible sockets. The trichoid sensilla (T) are
acute and have a longitudinally grooved cuticular surface. Bar ¼ 10 mm.
3.2. Pedicel
The pedicel has a characteristic longitudinal suture
extending along its outer lateral edge and the cuticular
surface is also densely covered with microtrichia (Figs. 3
and 4). Two types of sensilla were seen on the pedicel of all
specimens examined: trichoid and styloconic sensilla. The
length of the trichoid sensilla on the pedicel varied, in
contrast to those located on the scape (Fig. 2). Styloconic
sensilla were arranged in a frontal group on the pedicel
(arrowhead in Fig. 2) and varied in number among fly
species (Table 1). These sensilla appeared morphologically
similar among the fly species examined. As can be seen in a
male L. cuprina (Fig. 4), each styloconic sensillum contains
a seta that is bulbous at the base and then tapers distally to an
acute tip. The base of each seta is received in a socket with
an elevated rim. A cluster of fine hairs is situated posterior to
the raised rim of the setal socket and is oriented anteriorly
into the air stream (arrowhead in Fig. 4). In the female of
P. dux (Fig. 5), the terminal ends of the bulbous setae are
obtuse compared to those of L. cuprina (Fig. 4). In some
specimens examined, some of the bulbous setae were
Fig. 4. SEM micrograph of features on antennal pedicels of flies. Cuticular
surface of pedicel of male L. cuprina showing a group of seven styloconic
sensilla, each bearing longitudinal grooves on the bulbous seta that tapers to
an acute tip distally. A tuft of fine hair (arrowhead) is present posterior to
the raised rim and seta of each setal socket. Bar ¼ 10 mm.
673
Fig. 5. SEM micrograph of features on antennal pedicels of flies. Styloconic
sensilla of pedicel of female P. dux showing presence of bulbous seta
(right) and lost seta (arrow). Arrowhead indicates posterior hair tuft.
Bar ¼ 10 mm.
missing (arrow in Fig. 5). Minute variations in styloconic
sensilla were seen throughout this study, with some
specimens of M. domestica and C. rufifacies even lacking
the cluster of fine hairs near the setal socket.
3.3. Flagellum
The flagellum is the most prominent antennal segment of
the flies studied, with its length being ca. 65 –70% of the
antenna’s longest axis (Figs. 1,2,6 and 7). A bristle-like
appendage or ‘hair’, called the arista, is located proximally
near the base of flagellum and is positioned along the
dorso – lateral margin in line with the longitudinal suture of
the pedicel (Figs. 1,2 and 6). The entire surface of the
flagellum is densely covered with microtrichia (Fig. 8).
Antennal sensilla observed on this segment include
basiconic sensilla, coeloconic sensilla and sensory pits. In
all fly specimens examined, basiconic sensilla were the most
numerous type and consisted of two subtypes: large and
small basiconic sensilla (Fig. 8). The large basiconic sensilla
are characterized by setae with a smooth surface that are
Fig. 6. SEM micrograph of features on antennal flagella of flies. Antenna of
female P. dux showing multiple sensory pits (arrows) on flagellum.
Bar ¼ 100 mm.
674 K. Sukontason et al. / Micron 35 (2004) 671–679
Fig. 7. SEM micrograph of features on antennal flagella of flies. Sensory pit
(arrow) on flagellum of male L. cuprina. Bar ¼ 100 mm.
thickened proximally and have a slight curvature that
gradually tapers to an acute tip distally; whereas, the small
basiconic sensilla bear shorter, blunt-tipped setae that are
interspersed amongst the other types of sensilla and
microtrichia (Figs. 8 and 9). Basiconic sensilla are oriented
in a direction that is roughly parallel to the length of the
flagellum giving the flagellum a velvety appearance.
Coeloconic sensilla could occasionally be found on the
flagellum, with those occurring proximally on the segment
being more scattered than in other areas. This type of
sensillum, which is characterized by being located in a
deeply sunken depression of the integument called a
sacculus, was present in different forms (Figs. 10– 13).
Sometimes multiple forms could be found in the same
specimen. Without the use of SEM, these particular sensilla
often remain unseen by their being obscured by a dense
covering of microtrichia and/or basiconic sensilla. Detec-
tion of these coeloconic sensilla is sometimes facilitated
using SEM; therefore, they were addressed in this study.
The first form of coeloconic sensillum observed in this study
consisted of a short, blunt-tipped peg with smooth surface,
Fig. 8. SEM micrograph of features on antennal flagella of flies. Cuticular
surface of flagellum of male M. domestica densely covered with thin, acute
microtrichia or spinules (SP). Large basiconic sensilla (LB) that are
characterized by being slightly curved setae with thicker bases and pointed
tips and small basiconic sensilla (SB) that are much shorter with blunt tips
are interspersed over the surface. Bar ¼ 10 mm.
Fig. 9. SEM micrograph of features on antennal flagella of flies. Higher
magnification of a small basiconic sensillum emphasizing its blunt tip.
Bar ¼ 1 mm.
that was slightly curved distally and located centrally within
a deep sacculus. This type of sensillum was observed in
female C. rufifacies (Fig. 10). In some specimens, such as of
those L. cuprina, coeloconic sensilla were located either
singly or in a group of a few sensilla. The second form of
coeloconic sensillum observed was a short, cone-shaped peg
with longitudinal grooves and relatively pointed tip situated
centrally in a deep sacculus. These sensilla appear singly or
in a group, such as seen in a group of three sensilla in a male
L. cuprina in Fig. 11. The third form was rarely seen in the
species examined, but was observed in female L. cuprina
situated lateral to the arista (Fig. 12). This sensillum is
shaped more or less like an inverted triangle centrally
located in a sunken depression surrounded by a doughnut-
shaped base (Fig. 13).
A variety of sensory pits were also observed in the
different fly species examined in this study. Sensory pit
morphology typically appears as a relatively large pit
containing several sensory pegs within the sunken cuticular
surface and occupies an area ca. 3 – 14 mm in width and ca.
6 – 18 mm in length. Some slight variations in this
morphology were noticed among the fly species examined
Fig. 10. SEM micrograph of coeloconic sensilla observed on flagella of
flies. Coeloconic sensilla of female C. rufifacies, each containing a slightly
curved, blunt seta located centrally within a deep sacculus. Bar ¼ 1 mm.
 K. Sukontason et al. / Micron 35 (2004) 671–679
Fig. 11. SEM micrograph of coeloconic sensilla observed on flagella of
flies. Group of three coeloconic sensilla of male L. cuprina arranged in a
linear fashion on the cuticular surface. Each sensillum bears a cone-shaped
projection with longitudinal grooves centrally located within each sacculus.
Bar ¼ 1 mm.
and occasionally multiple forms could be found in the same
specimen. The variety in the sensory pit morphologies can
be seen among female M. domestica, female L. cuprina,
male P. dux and female P. dux (Figs. 14 –17). The sensory
pit in female M. domestica is a single pit (Fig. 14), while that
in female L. cuprina has a partition (arrow in Fig. 15) that
divides it into compartments. Pits in the antennal flagellum
of male P. dux contain separated sensory pegs (Fig. 16);
whereas, fusion of these pegs was observed in female P. dux
(Fig. 17).
The number of sensory pits on the flagellum varies
according to species. In this study, the sarcophagid, P. dux,
exhibited a higher number of pits than any of the other fly
species. The most obvious sensory pits are found along the
dorso – medial aspect of the flagellum. These pits were most
numerous in female P. dux and ranged from 8 to 15 pits
ðn ¼ 9Þ (arrows in Fig. 6), but in the male of this species the
pits were less numerous, ranging from 0 to 6 pits ðn ¼ 9Þ:
Numbers of pits could conceivably be higher due to unseen
or smaller-sized pits that could not be clearly perceived.
Moreover, only the dorsal and lateral views of the flagellum
Fig. 12. SEM micrograph of coeloconic sensilla observed on flagella of
flies. Position of coeloconic sensillum (arrow) near arista (AR) of female L.
cuprina. Bar ¼ 10 mm.
675
Fig. 13. SEM micrograph of coeloconic sensilla observed on flagella of
flies. Higher magnification of coeloconic sensillum of female L. cuprina
situated in a sunken depression surrounded by doughnut-shaped base.
Bar ¼ 1 mm.
could be viewed under SEM. Thus, the number of ventral
pits remains unknown. Lucilia cuprina bears a lower
number of sensory pits that are easily discernable (arrow
in Fig. 7), but counting of the exact number under extreme
scrutiny has not been performed. As for the other species
examined, the sensory pits that are obvious are few in
number, but they may still have more of this type of
sensillum than was recorded in this study.
4. Discussion
Similar to what is seen in other insects, the antennae of
flies in the families Calliphoridae, Sarcophagidae and
Muscidae bear several types of sensilla for chemo- and/or
mechanoreception that are essential for detecting food,
searching for prey, finding oviposition sites, and recogniz-
ing the opposite sex of their species for mating. Adult flies
of C. megacephala, C. rufifacies, C. nigripes, L. cuprina,
P. dux and M. domestica have their antennae densely
endowed with sensory receptors. Evidence presented herein
indicates that these flies bear morphologically similar
Fig. 14. SEM micrograph showing a variety of sensory pit morphologies on
fly flagella. Female M. domestica displaying a large centrally located peg
encircled by smaller pegs deep within a sensory pit. Bar ¼ 1 mm.
676 K. Sukontason et al. / Micron 35 (2004) 671–679
Fig. 15. SEM micrograph showing a variety of sensory pit morphologies on
fly flagella. Female L. cuprina with two compartments of sensory pits
divided by a partition (arrow). Each compartment has a centrally located
blunt peg encircled by similar sized pegs. Bar ¼ 10 mm.
antennal sensilla with no marked difference in the
morphology of each sensillum type being found to indicate
sexual dimorphism among the species examined. In like
manner, Microplitis croceipes (Hymenoptera: Braconidae),
a parasitoid which depends largely on chemical cues to
locate its host, no distinct morphological differences were
found in the antennal sensilla between the sexes (Ochieng
et al., 2000).
Also, for the most part, there seems to be no distinct
difference in the number of each type of sensillum between
sexes of the fly species that were included in the current
study (Table 1). An exception to this trend was the higher
number of sensory pits detected in P. dux compared to the
other species in the study. L. cuprina ranked second in
number of easily discernable pits and the remaining four
species had pits that were either minimal in size or hard to
observe. Based on the observations of P. dux and L. cuprina
in this study, the number of sensory pits in female flies is
greater than in males of the same species. This finding is
consistent with Chapman (1982) who found that there are
over 250 sensory pits in female Sarcophaga, but ca. 50 on
each antenna in males. The females of Sarcophaga
Fig. 16. SEM micrograph showing a variety of sensory pit morphologies on
fly flagella. Male P. dux showing separated sensory pegs with bulbous bases
that taper to acute tips within a sensory pit. Bar ¼ 1 mm.
Fig. 17. SEM micrograph showing a variety of sensory pit morphologies on
fly flagella. Female P. dux indicating fusion of sensory pegs. Spinules (SP),
large basiconic sensilla (LB) and small basiconic sensilla (SB) can be seen
surrounding the sensory pit. Bar ¼ 10 mm.
argyrostoma were found to have 261 pits per funiculus or
third antennal segment by Slifer and Sekhon (1964).
Likewise, the number of pits in the calliphorid, Phormia
regina, were discovered to range from 11– 16 in females,
but only 9– 11 in males (Chapman, 1982). In comparison,
muscid flies have been reported to have many fewer sensory
pits on their antennae. For example, three pits were found in
the flagellum of Musca autumnalis by Bay and Pitts (1976),
two pits were found in Stomoxys calcitrans by Lewis
(1971), and four pits were seen in Hydrotaea viritans by
Been et al. (1988). In addition, Dethier (1971) noted that the
dung-feeding dipterans, which depend largely upon odor for
locating their food, have more olfactory pits than the flower-
visiting species that depend on vision, indicating the
importance of these pits in odor detection.
Many authors (e.g. Chu-Wang et al., 1975; Chapman,
1982; Wasserman and Itagaki, 2003) have concluded that
the most likely function of the sensory pits is for olfactory
reception. Wasserman and Itagaki (2003) based their
conclusions according to the staining pattern of these
antennal cells conducted by using NADPH-diaphorase
histochemistry. This information supports the hypothesis
that the abundant sensory pits seen in sarcophagids would
help the female flies be more sensitive in olfactory
reception. This would allow them to detect even the faintest
odors of decomposition to locate a corpse for larviposition.
Moreover, flesh flies are usually present over a wide range
of the decay process of a corpse. Not only are they present
during the initial fresh stage of decay, but they are also
present during the later stages of corpse decomposition
and/or mummification stage when there is little or no strong
odor to attract females to larviposit (e.g. Smith, 1986; Goff
and Flynn, 1991; Sukontason et al., 2001).
Regarding the fine ultrastructure of the sensory pits
observed in the present study, it appears that some minor
differences in the external morphology occur in the fly
species examined. Not much detail of the shape of each
sensory peg inside the sensory pits of M. domestica
 K. Sukontason et al. / Micron 35 (2004) 671–679
and L. cuprina could be seen, since only top-view SEM
photomicrographs could be produced. In this study, only the
pegs of male P. dux could be seen with much detail and each
could be described as having a bulbous base that gradually
tapers to an acute tip. In general, the overall structure of the
sensory pits of the fly species examined in this study appear
to be similar to those on the third maxillary palp of biting
midges of Culicoides spp. (McKeever et al., 1994), and to
those on the funiculus of female warble fly Hypoderma
bovis (Hunter and Adserballe, 1996).
Basiconic sensilla were the most numerous type of
antennal sensillum in both sexes of all the fly species
examined in this study. The large and small basiconic
sensilla observed in the flies of the present study resemble
those on the antennae of other dipteran species such as the
fruit fly, Drosophila melanogaster (Diptera: Drosophilidae)
(Shanbhag et al., 1999), human bot fly, Dermatobia hominis
(Diptera: Cuterebridae) (Fernandes et al., 2002), Pseudo-
perichaeta nigrolineata (Diptera: Tachinidae) (Rahal et al.,
1996), and type 1 and type 2 of the female warble fly,
H. bovis (Diptera: Oestridae) (Hunter and Adserballe,
1996). Some similarity in morphological appearance of
basiconic sensilla has been noted in other non-dipteran
insects such as in the sphinx moth, Manduca sexta
(Lepidoptera: Sphingidae) (Shields and Hildebrand, 1999)
and the eucalyptus woodborer, Phoracantha semipunctata
(Coleoptera: Cerambycidae) (Lopes et al., 2002).
No sexual dimorphism was detected regarding the fine
ultrastructure of both types of basiconic sensilla in all the fly
species examine in the present study. Lack of sexual
dimorphism in basiconic sensilla was also observed in
P. semipunctata by Lopes et al. (2002). No clear functional
distinction has been made between the two morphological
types of basiconic sensilla, but it has been suggested by
many authors that they both likely play an olfactory role
(Zacharuk, 1985; Hunter and Adserballe, 1996; Rahal et al.,
1996; Shields and Hildebrand, 1999; Lopes et al., 2002;
Broeckling and Salom, 2003).
The fine ultrastructure of trichoid sensilla on antennal
segments of flies that was seen in the present study utilizing
SEM appears similar upon comparison with that of several
other insect species previously reported. Eisenbeis and
Wichard (1987) suggested that the trichoid sensilla function
as contact chemoreceptors in a related arthropod, the
centipede Geophilus longicornis (Geophilidae). However,
a mechanoreceptive function for trichoid sensilla has been
proposed in numerous insects that were studied including
the human bot fly, D. hominis (Fernandes et al., 2002), the
ground beetle, Bembidion properans (Coleoptera: Carabi-
dae) (Merivee et al., 2002), the red imported fire ant,
Solenopsis invicta (Renthal et al., 2003), and the parasitoid
wasp Microplitis croceipes (Hymenoptera: Braconidae)
which was based on an investigation using transmission
electron microscopy (TEM) (Ochieng et al., 2000). Still
another hypothesis, the dual function as both mechano- and
chemoreceptors, has been suggested for the trichoid sensilla
677
in the alfalfa weevil, Hypera postica by Bland (1981) based
on the ultrastructure, including the outward projection of
these sensilla. Therefore, the function of the trichoid sensilla
in the fly species that were studied will require further
investigation and experimentation before their specific role
is known.
Upon comparison of the coeloconic sensilla of this study
with those in previous studies, the grooved and smooth pegs
of these sensilla in the flies morphologically resemble the
‘sensilla coeloconica type 1 and type 2,’ respectively, of the
rhinoceros beetle, Oryctes rhinoceros (Coleoptera: Dynas-
tidae) (Renou et al., 1998). The grooved pegs in the flies of
this study were also found to be similar to those reported in
the antennae of the human bot fly, D. hominis (Diptera:
Cuterebridae) (Fernandes et al., 2002) and the biting
midge, Culicoides impunctatus (Diptera: Ceratopogonidae)
(Blackwell et al., 1992). In female L. cuprina, the coeloconic
sensillum that is situated in a sunken depression surrounded
by a doughnut-shaped base looks very similar to that seen in
the terminal flagellar sub-segment of the female parasitoid
wasp Microplitis croceipes (Hymenoptera: Braconidae). In
accordance with what was seen in the fly species, this type of
sensillum occurs in very low numbers in this wasp species,
having one (or at the most two) per flagellomere (Ochieng
et al., 2000). The probability of these types of sensilla being
involved with thermo- or hygroreception (McIver, 1969;
Altner et al., 1983; Zacharuk, 1985; Ochieng et al., 2000), or
possibly olfactory reception (Olson and Andow, 1993;
Renthal et al., 2003), have all been proposed.
The styloconic sensilla of flies in this study morphologi-
cally correspond with those found in the arista of male
D. hominis by Fernandes et al. (2002). Numbers of these
varied from 6 – 16 per antenna between different specimens.
Among the five families of flies examined by Greenberg
(1970), this type of sensillum was found more in
calliphorids and sarcophagids than in muscids. Styloconic
sensilla in the antennae of the leaf beetle, Psylliodes
chrysocephala (Coleoptera: Chrysomelidae) appear as
cone-shaped pegs emerging from raised areas of cuticle
(Bartlet et al., 1999). Short, stout cones with terminal
nipples describe the styloconic sensilla in the click beetle,
Melanotus villosus (Coleoptera: Elateridae), where the
female has seven sensilla but the male only has three
(Merivee et al., 1999). Mechano- and/or chemoreception
have been suggested by previous authors (Zacharuk, 1985;
Mitchell et al., 1999) as the probable function of styloconic
sensilla, but more research is needed to help clarify this
issue.
Antennae are considered to be the nostrils of insects
(Zacharuk, 1985). The sensitivity of chemoreceptive
sensory neurons to odor varies and those that respond to a
wide range of different chemicals are considered generalist
cells (Chapman, 1982). Different cells, even within the same
sensillum, can exhibit different profiles of response to a
range of chemicals. As for flies, in the electrophysiological
analysis of the antennae of female L. cuprina it was
678 K. Sukontason et al. / Micron 35 (2004) 671–679
demonstrated that the olfactory receptor neurons responded
best to exposure to either 1-octen-3-ol, dimethyldisulphide
or 2-phenylethanol (Park and Cork, 1999). Likewise,
Kelling et al. (2002) reported a large variability in response
profiles to several odor stimuli by electrophysiological
characterization of the third antennal segment of
M. domestica. The odors of 1-octen-3-ol, amyl acetate,
3-methylphenol, 2-pentanone and R(þ ) limonene elicited
the largest responses in house fly antennal cells. Kelling
et al. (2002) reiterated that receptor cells with similar or
different odor response profiles can reside in the same
sensillum. Behavioral studies by Pappenberger et al. (1996)
demonstrated that human body odors that the female yellow
fever mosquito, Aedes aegypti, finds attractive actually exert
their effects on the mosquito as a complex mixture of
synergistically acting components. In regard to fly species
of forensic importance, females are rapidly attracted to
corpses emanating odors in the form of chemical cues of
animal decomposition, including blood and/or wounds
(Greenberg and Kunich, 2002). Wasserman and Itagaki
(2003) pointed out the fact that there is a wide variety of
odors from animal-related compounds (e.g. dried blood,
liver, meat, carrion, cheese, CO2, cattle feces, human breath,
acetone, aliphatic alcohols, aldehydes, and mammalian
secretory and excretory compounds) that will attract blow
flies or flesh flies. These compounds represent a huge range
of chemical composition.
Results of this present study clearly show that the
antennae of adult flies in the families Calliphoridae,
Sarcophagidae and Muscidae are equipped with antennal
sensilla types that are morphologically similar. The
probable function of each type of sensillum was suggested
on the basis of comparison with other previous investi-
gations. Fly species of Calliphoridae and Sarcophagidae
could probably be considered more important in regard to
forensic entomology since they are typically more closely
associated with human corpses than species of Muscidae.
Since this is typically the case, more research involving
electrophysiological investigations of each type of olfactory
cell in the antenna would help to clarify the specific role that
each olfactory receptor plays in each species.
Acknowledgements
This work received support from Dr Natth Bhamarapra-
vati’s Fund for Science Research at Chiang Mai University.
We thank Chiang Mai University for funding the cost of
reprints.
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