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Gustation Across the Class Insecta: Body Locations

Article in Annals of the Entomological Society of America · February 2023

DOI: 10.1093/aesa/saac027

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Annals of the Entomological Society of America, XX(X), 2023, 1–7
https://doi.org/10.1093/aesa/saac027
Review

Review

Gustation Across the Class Insecta: Body Locations

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Bethia H. King1, and Panchalie B. Gunathunga
Department of Biological Sciences, Northern Illinois University, DeKalb, IL 60115, USA and 1Corresponding author,
e-mail: bking@niu.edu

Subject Editor: Vonnie Shields

Received 5 July 2022; Editorial decision 28 November 2022

Abstract
This review summarizes which body parts have taste function in which insect taxa. Evidence of taste by mouth-
parts, antennae, and tarsi is widespread. Mouthparts that commonly have taste function are the labium,
including the labella and labial palps, the maxillae, including the galeae and maxillary palps, the inner surface
of the labrum or clypeolabrum of chewers, and inside the precibarium/cibarium of hemipterans, which have
piercing-sucking mouthparts. Tasting with mandibles has not been found, and tasting with the hypopharynx
is seldom reported. Use of the antennae appears uncommon among fly species, but common among species
of lepidopterans, hymenopterans, beetles, and bugs. Although tasting with legs, especially tarsi, is reported
mostly for fly and lepidopteran species, there is also evidence of it for multiple species of beetles, grasshop-
pers, and hemipterans, and one species of a roach, an ant, and a bee. Ovipositor taste function has been sup-
ported for some species of flies, lepidopterans, hymenopterans, orthopterans, and odonates. Taste by wings
has been much less studied, but has been documented in a few fly species. Taste remains unstudied for any
species or any body parts of Archaeognatha, Dermaptera, Mantodea, Mecoptera, Phasmatodea, Megaloptera,
Neuroptera, Phthiraptera, Psocoptera, Siphonaptera, as well as Raphidioptera, Strepsiptera, Embioptera,
Notoptera, and Zoraptera. Across holometabolous insects, larvae have not often been examined, the excep-
tion being some species of lepidopterans, flies, and beetles. Taste studies of antenna and legs are uncommon
for even lepidopteran and beetle larvae.

Key words: antennae, gustatory, mouthpart, ovipositor, tarsi

In humans, gustation versus olfaction is defined not only by the re- tests show that the antennae of the beetle Chrysomela populi L.
sponse being to nonvolatile versus volatile compounds, but also by (Coleoptera: Chrysomelidae) have taste function (Pentzold et al.
the body part that detects the chemical, tongue versus nose. However, 2019). The labium of the mosquito Anopheles gambiae (Diptera:
some animals taste with parts of their body other than just their Culicidae) has both gustatory (Supp Table 1 [online only]) and ol-
mouthparts (Caprio et al. 1993). For example, some insects taste factory function (Kwon et al. 2006). To date, olfactory receptor
with their tarsi as they walk on their food source prior to feeding proteins appear to be restricted to olfactory function, yet they are ex-
(Yarmolinsky et al. 2009). This may allow insects to more quickly pressed not just on the antennae, but also on the nectar-sucking pro-
locate appetitive compounds and avoid aversive compounds (Scott boscis and the legs of adults of the butterfly Heliconius melpomene
2018). In insects, the rough equivalent of human taste buds are taste L. (Lepidoptera: Nymphalidae) (Briscoe et al. 2013).
sensilla, which are often hair shaped. We tend to associate human Most recent reviews of insect taste focus primarily, or even ex-
eyes, nose, mouth, and ear each with a different sensory function. In clusively, on taste in one species, the common fruit fly, Drosophila
insects, a given body part may have more than one of these sensory melanogaster Meigen (Diptera: Drosophilidae) (e.g., Scott 2018,
functions. Insect maxillary palps and antennae are often referred to Toshima and Schleyer 2019, Chen and Dahanukar 2020, Montell
as olfactory organs (Li et al. 2018, Oh et al. 2021), but their func- 2021, Puri and Lee 2021, Komarov and Sprecher 2022). This is not
tion varies among insect species. In larvae of the beetle Melolontha surprising because this species is by far the best studied. However,
melolontha L. (Coleoptera: Scarabaeidae), there appear to be not there are more than a million species of insects, from about 23 other
only olfactory sensilla but also taste sensilla on the antennae, maxil- orders, not just Diptera, and some are economically important.
lary palps, and labial palps (Eilers et al. 2012). Electrophysiological Understanding insect taste is relevant to control of pest insects, e.g.,

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2 Annals of the Entomological Society of America, 2023, Vol. XX, No. XX

when the control involves the insect eating the pesticide. Thus, nar- on the presence of only a single dendrite and that dendrite having
ratives about insect taste need to incorporate what has been learned a tubular body (Chu-Wang and Axtell 1972, Singh and Singh 1984,
about other insect species in the last 20+ years. For example, there Rist and Thum 2017). Some seem to have hygro and/or thermal func-
is information about which body parts D. melanogaster tastes with; tion, based on features such as location in a pit or the presence of
but what about other species? At least some flies and lepidopterans lamellated dendrites (Altner et al. 1978, Steinbrecht 1984, Schneider
taste with their feet, but is it all, and is it just these taxa? This review et al. 2018). Thus, we suggest that gustatory function should not be
compiles what body parts are used to taste by which insects. Thus, it concluded until more evidence than a terminal pore is available, e.g.,
addresses questions such as: which parts of the mouth and digestive transmission electron microscope (TEM) images that show internal
tract are used? Do all insects use their antennae and legs? Are wings features of the sensilla, behavioral tests, or electrophysiological tests.
commonly used? Do abdominal structures such as ovipositors and Many gustatory sensilla have both taste and touch func-

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male external genitalia ever have taste function? Do internal organs tion (Shanbhag et al. 2001, Sparks and Dickens 2017). However,
other than the digestive tract taste? they are nevertheless called gustatory or taste sensilla or con-
This review also identifies areas that need further research. With tact chemosensilla or contact chemoreceptors (Isidoro et al. 1998,
the recent emphasis on course-based undergraduate research experi- Komendant 2011, Rebora et al. 2019). The published literature that
ences (CURES) (Dolan 2016), it is worth noting that many taste we read is consistent in assigning gustatory function when there is
experiments are fun and feasible for undergraduates to do with ac- a tubular body on one dendrite, which is indicative of that neuron
curacy after fairly minimal training, e.g., projects on specific body being mechanosensory (tactile), and at least one other neuron
parts or specific taste hairs with a variety of tastants, e.g., different without a tubular body. When that was not the case, we looked for
sweeteners and bitter compounds like caffeine and quinine. Many the sensilla being uniporous and no evidence that the function was
taste experiments are also cheap and safe, and they lend themselves thermo- or hydro-sensory, olfactory, or strictly mechanosensory.
to getting undergraduates to really focus on and understand the Evidence of temperature or humidity function includes being set in
most important aspects of experimental design, e.g., controls and a depression, lamellated dendrites, and/or the space at the tip of the
testing blind (i.e., without knowledge of which test subject got which sensillum sometimes so filled with dendrites that no lymph is seen
treatment, to avoid inadvertently biasing the results). (Steinbrecht 1984, Tichy and Kallina 2010, Schneider et al. 2018).
Olfactory function is usually concluded when the sensillum wall is
multiporous (Stocker 1994, Schoonhoven et al. 2005, Rebora et al.
Methods 2019, Sevarika et al. 2021, but see Ruschioni et al. 2012); there is
References on insect taste were located by searching with Google also often, but not always, extensive branching of dendrites (Sutcliffe
Scholar and the databases Web of Science and Scopus, until we were 1994, Shanbhag et al. 1999). Evidence of strictly mechanosensory
no longer finding additional articles, by which point we had found function is one, rarely two, dendrites, each with a tubular body
hundreds of articles. For each body part, we report which species (Dumpert 1972, Keil 1997).
appear to have taste function, as well as the type of evidence: mor- Pharyngeal chemosensilla were assumed not to be olfactory based
phological and neuroanatomical, behavioral, electrophysiological, on their location within the body where very little air movement is
or molecular. Although some studies suggesting the absence of taste expected. Pharyngeal chemosensilla were also assumed not to re-
in a body part have been included, it remains possible that taste func- spond to changes in humidity or temperature, because such function
tion may later be demonstrated for those species. For example, elec- seems like it would be more useful on the outside of an insect. Thus,
trophysiological taste tests with the ovipositor of migratory locusts the presence of a single terminal pore, sometimes with evidence that
were initially unsuccessful (Woodrow 1965) but have since been suc- cobalt solution will pass through it, was considered sufficient to sug-
cessful (Gaaboub and Tousson 2010). Also, a body part may respond gest a taste function in pharyngeal chemosensilla (Zacharuk 1980).
to tastants that were not tested, or techniques of recording neural Further evidence justifying these assumptions is needed.
responses may improve. There are also likely experiments in which
evidence of taste was not found, and the result was not published
Results and Discussion
because the authors were aware of biases against negative results
(Csada et al. 1996, Chouvenc et al. 2011). Best-Studied Larvae
For behavioral and electrophysiological evidence we required Generally, taste has been less well-studied for larvae than for adults
that another sense, e.g., strictly tactile or olfactory, had been ruled (Supp Tables 1–10 [online only]). However, taste has been very well-
out as an alternative explanation. For example, imagine that a body studied in certain mouthparts of Lepidoptera larvae. Among cater-
part is touched to different tastants or concentrations, and they elicit pillars, evidence of taste function of the maxillary galeae’s sensilla,
different responses. All involve touch, so the difference in response is i.e., of the lateral and medial styloconica, has been reported for
likely not just mechanosensory. Differences in viscosity among solu- many species and from more than a dozen families (Supp Table 2
tions are assumed to be inconsequential. If the chemical being tested [online only], Schoonhoven and van Loon 2002, Shields and Martin
is not volatile that indicates gustation rather than olfaction. 2012). These sensilla are especially well-studied because electro-
For morphological evidence, some authors suggest gustatory physiological recordings from them are relatively easy (Schoonhoven
function based on a sensillum having a single terminal (apical or and van Loon 2002). Some caterpillars also seem to taste with their
subapical) pore (e.g., Zacharuk and Shields 1991, He et al. 2019, maxillary palps and their epipharynx (the underside of the labrum)
Taszakowski et al. 2019). However, we required more evidence, be- (Supp Table 2 and 4 [online only]).
cause a single terminal pore might be strictly a molting (ecdysial) Taste has also been studied in fly larvae, although not as ex-
pore, rather than the entry for tastants (Zacharuk et al. 1977), and tensively as in caterpillars. The head region of the larvae of D.
recognizing whether or not a pore is strictly used in molting is not melanogaster, house flies, and some other flies, contains three pairs
straightforward (Zacharuk et al. 1977; Faucheux 2012, 2017). of external chemosensory structures: the dorsal organ (DO), which
Also, some sensilla with a single terminal pore appear to have a may be derived from the antenna segment; the terminal organ (TO),
nongustatory function. Some appear to be mechanosensory, based which is also called the maxillary palp complex; and the ventral
Annals of the Entomological Society of America, 2023, Vol. XX, No. XX 3

organ (VO), which may be derived from the mandibular segment by the larva’s terminal organs is present in all four of the species that
(Fig. 1 in Apostolopoulou et al. 2015, Hartenstein et al. 2018). have been examined. Taste function also appears to be absent in the
In some flies these organs are known to have gustatory neurons maxillary lacinias of adult green head horse flies and in the maxillary
(along with other types of sensory neurons) (Supp Tables 2, 3, and stylets of the one hemipteran in which taste function of the maxillae
5 [online only]). However, taste remains unexplored in most flies has been examined (Peregrine 1972).
with these three pairs of organs, e.g., Piophila casei (Linnaeus) A taste function is infrequently discussed for mandibles, but
(Diptera: Piophilidae) (Sukontason et al. 2005), Parasarcophaga appears to be absent in three species of hemipterans, a caterpillar
dux (Thomson) (Diptera: Sarcophagidae), Chrysomya albiceps species, and adult mosquitoes (Supp Table 3 [online only]). We hy-
Wiedemann (Diptera: Calliphoridae), and Cochliomyia macellaria pothesize that taste sensilla are rare on mandibles because such
(Fabricius) (Diptera: Calliphoridae) (Mendonca et al. 2010, 2014). sensilla would have difficulty withstanding the considerable mech-

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anical force that mandibles are subject to. Mandibles of insets with
Mouthparts and Digestive Tract chewing mouthparts are sometimes subject to considerable stress
For an insect with chewing mouthparts, the labrum is positioned (Weihmann et al. 2015, Rühr and Blanke 2022). The mandibles of
like an upper lip, and its inner side is called the epipharynx Figure. hemipterans are modified into long thin stylets and also may experi-
The labium is positioned like a lower lip or chin. Laterally are a ence more force than more interior mouthparts. In kissing bugs, the
pair of mandibles (jaws) and then ventral to them is a pair of max- mandibular stylets are exposed to force as they pierce through the
illae. A hypopharynx is in the center, and is sometimes described as host’s outer surface, whereas the maxillary stylets enter a blood vessel
tongue-like. There are two pairs of palps, the maxillary palps, and (Escandón-Vargas et al. 2017). Likewise, cotton strainers pierce the
the labial palps. In some insects, mouthparts have evolved from the seed coat with their mandibular stylets (Wang and Dai 2017). Larvae
ancestral chewing parts into new forms, such as sucking, as in mos- of certain fly species have sensory organs called the ventral organs,
quitoes and hemipterans; sponging, as in house flies; and siphoning, which have been hypothesized to develop from the mandibular seg-
as in lepidopterans Figure. Given current evidence, it would be very ment (Hartenstein 1988, Cobb et al. 2009). A taste function has been
surprising if any insects are found to lack taste function in at least seen in the ventral organs of two of four fly species that have been
some mouthparts in life stages that feed. The mouthparts for which examined, in D. melanogaster and house fly larvae, but not in two
evidence of tasting has been reported most often are the labium, species of root fly maggots (Supp Table 3 [online only]).
including the labellum or labial palps, and the maxillae, including Evidence of taste with other parts of the digestive tract also
the maxillary galeae and palps (Supp Tables 1 and 2 [online only]). exists. In sap sucking insects like aphids and other homopterans,
Taste function in labial parts has been found broadly taxonom- the hypopharynx and epipharynx join to make a precibarium and/
ically (Supp Table 1 [online only]). Labial palps with taste func- or cibarial pump, which are located in the base of the proboscis
tion have been documented in species of: a mayfly naiad, a locust, and are described as being part of the pharynx or just anterior to
two roaches, two beetle larvae and an adult, a honey bee, an ant, the pharynx (Ruschioni et al. 2019). Similarly, in flies with spon-
three moths, and three caddisflies. Taste function in other parts of ging mouthparts, just anterior to the pharynx are the cibarial sense
the labium, including the labellum, has been documented in even organs and labral sense organs (Fig. 1 in Stocker and Schorderet
more species: a roach, five true bugs/hemipterans, a thrips, a honey 1981). Evidence of taste function in the precibarium or cibarium has
bee, and almost 28 flies from 11 different families. In adult D. been shown in four families of Hemiptera; in two families of Diptera
melanogaster, the labellum is the only external mouthpart with taste (tsetse flies and a mosquito); and in a thrips species (Supp Table 4
function, although there are taste sensilla within the proboscis (Chen [online only]). Whether the cibarial cavity of adult lepidopterans has
and Dahanukar 2017). a taste function appears to be unexplored (Inoue et al. 2009, Krenn
Tasting with parts of the maxillae also seems to be widespread 2010), although, as noted, the maxillae that form a siphoning tube,
among insects (Supp Table 2 [online only]). In adult Lepidoptera a proboscis, have taste function (Figure, Supp Table 2 [online only]).
that feed, the maxillary galeae form their most obvious mouthpart, Gustatory sensilla on the hypopharynx appear absent in mosquitoes
the sucking, often coiled, proboscis. There is evidence of taste func- and in a horse fly species, both of which take liquid meals. However,
tion of this proboscis in at least seven lepidopteran species from five gustatory sensilla have been reported for a species of roach, which
families. Taste sensilla can be found on the inside and outside of has chewing mouthparts. Tasting with the labrum, epipharynx (the
this proboscis (Inoue et al. 2009, Faucheux 2013). As noted above, internal surface of the labrum), or inner surface of the clypeolabrum
evidence of taste function on the maxillary galeae has been demon- has been reported for a variety of insect species: two locusts, a ter-
strated in many species of caterpillars, with the caterpillars coming mite, a roach, the cotton stainer bug, the larval stage of a beetle,
from more than a dozen different families (Supp Table 2 [online adults of four mosquitoes, and some, but not all, caterpillars (Supp
only], Schoonhoven and van Loon 2002, Shields and Martin 2012). Table 4 [online only]).
Evidence of taste function of the maxillary galeae has also been Pharyngeal taste sensilla have been well-documented in adult D.
documented for five species of beetles from three different families melanogaster (Chen et al. 2021). These sensilla lack a cuticular ex-
and in five bee species from Apidae. tension such as a hair; instead, there is just a pore in the cuticle (Fig.
Evidence of taste function in the maxillary palps has been seen 1h in Chen and Dahanukar 2020). These sensilla respond to sweet,
in six caterpillar species from five different families, as well as in bitter, acid, salt, and amino acids (Chen et al. 2019, 2021). Pharyngeal
species of: two damselfly naiads; three grasshoppers; one termite; taste sensilla also occur in the larval stage of D. melanogaster, where
two roaches; two adult and two larval beetles; a honey bee; three they have been shown to respond to caffeine (Apostolopoulou et al.
caddisflies from three different families; and some flies (Supp Table 2016).
2 [online only]). In D. melanogaster, taste appears to be absent in Pharyngeal taste input in D. melanogaster helps them avoid
adult maxillary palps (Singh and Nayak 1985), but present in the many aversive compounds (Chen et al. 2019). In D. melanogaster
larva’s terminal organs, which appear to develop from the maxillary adults, pharyngeal taste neurons respond to sucrose by inhibiting
segment. Across all of the studies with Diptera, taste with the adult feeding (Joseph et al. 2017), and nutritive-sugar sensors in the brain
maxillary palps is absent in four species and present in one; and taste also help limit overconsumption (Dus et al. 2015). Another benefit
4 Annals of the Entomological Society of America, 2023, Vol. XX, No. XX

to taste receptors on body parts that are not initially exposed to food fly family. Technical challenges in making electrophysiological re-
sources is that not all aversive compounds are present in food before cordings from wing sensilla at least partly explain why taste func-
ingestion. Some are breakdown products of digestion and bind to tion in wings has not been well-studied (Agnel et al. 2017). In D.
receptor(s) on deterrent gustatory receptor neurons only after diges- melanogaster; the anterior wing margins respond to sweet and bitter
tion has begun (Wada-Katsumata and Schal 2021).. molecules (Raad et al. 2016), bits of dead E. coli (Yanagawa et al.
2014), salts, and lipopolysaccharides (Yanagawa et al. 2019). The
Antennae, Legs, Wings, and Abdominal Structures desert locust, Schistocerca gregaria Forskål (Orthoptera: Acrididae),
Apart from mouth parts, the body parts in which taste has been may have chemoreceptors in its wings (Page and Matheson 2004),
most often documented are the antennae and tarsi (Supp Table 5 but whether the responses to sucrose solution and to salt solution
and 6 [online only]). Insect antennae are perhaps best known for were to the chemical per se versus to the contact that presentation of

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functioning in olfaction and touch. However, antennae also are used the solutions entails is difficult to distinguish.
to taste in a wide range of insect taxa, including naiads in three fam- Some evidence of taste has been reported for ovipositors in
ilies of Ephemeroptera, three families of Blattodea, eleven families of six families of flies, five of Lepidoptera, five of Hymenoptera, two
Hemiptera, eleven of Coleoptera (adults in eleven and larvae in one), of Odonata, and one of Orthoptera (Supp Table 8 [online only]).
eight of Hymenoptera, and eight of Lepidoptera, as well as in one Evidence of taste by the ovipositor is not strong for D. melanogaster
family each of Zygentoma, Orthoptera, Thysanoptera, Plecoptera, (Stocker 1994, Dweck et al. 2021), but has been documented with
and Isoptera (Supp Table 5 [online only]). electrophysiological and behavioral tests for D. suzukii (Matsumura)
Bernays and Chapman (1994) note that taste sensilla have not (Biolchini 2016).
been reported for caterpillars on body parts other than the mouth- Taste by the male intromittent (copulatory) organ has been pro-
parts. However, since then; taste function has been demonstrated for posed, but support is limited. An apparent single pore is seen in scan-
larval antennae of the maize stalk borer, Busseola fusca (Lepidoptera: ning electron microscope (SEM) images of the ground beetle Carabus
Noctuidae) (Juma et al. 2008). In Diptera, some larvae have clearly fiduciarius Csiki (Coleoptera: Carabidae) (Kim and Yamasaki 1996)
identifiable antennae that are slender protrusions, and some have less and of the southern sickle bush cricket, Phaneroptera nana Fieber
obtrusive paired structures called dorsal organs (DOs), which seem (Orthoptera: Tettigoniidae), although uniporous sensilla appear
to develop from the same segment of the body as typical antennae to be absent on the male intromittent organ of some other orth-
do. Among larval flies, there is evidence of taste function of antennae opterans (Faucheux 2012). Internal vaginal sensilla have been seen
or DOs for 17 families (Supp Table 5 [online only]), but there is in some damselflies, but are reported as mechanosensory (Córdoba-
evidence for a lack of taste function on the DOs in D. melanogaster Aguilar 2003, Cordero-Rivera 2017), with no vaginal taste sensilla
(Oppliger et al. 2000). Among fly adults, taste sensilla seem to be yet reported.
absent in the antennae of most species that have been examined, in The cerci of a cricket species has taste function (Supp Table 8
yellow fever mosquitoes, a biting midge species, a tephritid species, [online only]), as do the cerci (anal/lateral leaflets/lobes) that are part
house flies, and one of two Tabanus (Diptera: Tabanidae) horse fly of at least some fly species’ ovipositors (Hooper et al. 1972, Rice
species, as well as in at least parts of the antennae of D. melanogaster 1976). Taste function has been documented on the upper surface of
(Supp Table 5 [online only]). However, taste sensilla have been docu- the head of migratory locusts (Supp Table 9 [online only]), and mo-
mented in the antennae of some adult flies: a horse fly species in lecular evidence suggests that it may occur on the thoracic surface of
Tabanus and two species of biting midges in Culicoides (Diptera: D. melanogaster larvae (Cobb et al. 2009).
Culicides). An absence of taste sensilla in antennae has been reported
not only for these flies, but also for a true bug nymph and for adult Well-Studied Species
males of a scale insect species (Supp Table 5 [online only]). For most orders for which there is some information on the location
Tasting with the legs, especially the tarsi (Supp Table 6 [online of taste, only one or two body parts have been examined, although
only]), is best known in Diptera, where it has been demonstrated in there are about fourteen species with evidence of taste in four or
ten families. Evidence for leg taste function has also been reported more body parts (Supp Table 10 [online only]). These well-studied
in eight families of Lepidoptera, three families of Hemiptera, two species are mostly lepidopterans and flies, but also two species of
of Hymenoptera, and one of Orthoptera, Blattodea, and Coleoptera orthopterans, a beetle, and adult honey bees. Drosophila seems to
(Supp Table 6 [online only]). In D. melanogaster adults, the tarsi be the most-studied insect with regards to which body parts have
have more taste sensilla than do the better-studied mouthparts taste function. For adult Drosophila there is evidence of taste func-
(Vosshall and Stocker 2007, Ling et al. 2014). Male D. melanogaster tion for the labellum, labrum, pharynx, tarsi, wings, and ovipositors,
have more taste sensilla on their forelegs than do females. In and evidence of a lack of taste function for the maxillary palps and
D. melanogaster (Joseph and Carlson 2015) and the butterfly antennae. For Drosophila larvae there is evidence of taste function
Heliconius erato (Linnaeus, 1758) (Lepidoptera: Nymphalidae) for the terminal organ (TO), ventral organ (VO), epipharynx, and
(Silva et al. 2017), the three different pairs of insect legs do not re- abdomen.
spond to the same tastants. An apparent lack of tarsal taste has been
reported for the harlequin bug and for a species of ladybird beetle Conclusions
(Supp Table 6 [online only]). In the case of the ladybird beetle, the Tasting with feet is not restricted to species of flies and lepidop-
tarsal sensilla appear to lack dendrites, and tarsal contact with su- terans, but has also been reported for species of orthopterans, some
crose does not elicit a behavioral response, despite sucrose doing so hemipterans, some adult beetles, a roach, a bee, and an ant. Most
upon mouthpart contact (Yosano et al. 2020). We found no evidence insect species that have been tested appear to taste with their feet,
for or against the presence of taste function on legs of caterpillars. the exception being some, but not all, tsetse flies. Antennae have a
Whether there are any entire insect orders without taste function in taste function in most species examined, the exception being some
the legs remains to be seen. flies. Taste function of wings is understudied but has been demon-
Tasting with wings has been reported for only three species of strated in a few flies. For each mouthpart except adult mandibles,
insects (Supp Table 7 [online only]), each of which is in a different there is evidence of taste function in at least one insect species, but
Annals of the Entomological Society of America, 2023, Vol. XX, No. XX 5

taste function has most consistently been reported for the labia, e.g., Cobb, M., K. Scott, and M. Pankratz. 2009. Gustation in Drosophila
labellum and labial palps, and for maxillae. There is evidence of taste melanogaster, pp. 1–38. In P. L. Newland, M. Cobb and F. Marion-Poll
function for the ovipositor in insects from several orders, although (eds.), Insect taste. SEB Experimental Biology Series, vol. 63, 2009/01/30
ed. Taylor & Francis, New York, NY.
surprisingly little such evidence for D. melanogaster. Similarly, evi-
Córdoba-Aguilar, A. 2003. A description of male and female genitalia and
dence of a taste function of male external genitalia is weak, and evi-
a reconstruction of copulatory and fertilisation events in Calopteryx
dence of a vaginal taste function is nonexistent.
haemorrhoidalis (Vander Linden) (Odonata: Calopterygidae).
Odonatologica. 32: 205–214.
Cordero-Rivera, A. 2017. Sexual conflict and the evolution of genitalia: male
Supplementary Data damselflies remove more sperm when mating with a heterospecific female.
Sci. Rep. 7: 7844.

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Supplementary data are available at Annals of the Entomological Society of America
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