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Received: 24 January 2022
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Revised: 2 May 2022
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Accepted: 5 May 2022
DOI: 10.1002/arch.21917
SHORT COMMUNICATION
Report on antennal sensilla of Aphelocheirus
ellipsoideus (Hemiptera, Heteroptera,
Aphelocheiridae)
Tong‐Yin Xie
1
|Shi‐Yan Ma
1
|Fu‐Xia He
2
1
College of Agriculture, Northeast
Agricultural University, Harbin, China
2
College of Life Sciences, Northeast
Agricultural University, Harbin, China
Correspondence
Fu‐Xia He, College of Life Sciences, NE
Agricultural University, 150030 Harbin,
China.
Email: hefuxia969@163.com
Funding information
the “Academic Backbones”Project of
Northeast Agricultural University,
Grant/Award Number: No. 19XG03
Abstract
The comparative morphology study on antennal sensilla of
Aphelocheirus ellipsoideus from the family Aphelocheiridae,
carriedoutwiththeuseofascanning electron microscope, is
provided. Five main types of mechano‐,chemo‐, and thermo‐
hygroreceptive sensilla with two subtypes of sensilla basico-
nica were found and described on their surface, including
sensilla trichodea, campaniformia, basiconica, ampullacea, and
plate‐like. Antennal sensilla of A. ellipsoideus on macropterous
and brachypterous forms were different.
KEYWORDS
antennal sensilla, Aphelocheiridae, Aphelocheirus ellipsoideus,
Hemiptera
Key points
•Report on antennal sensilla of Aphelocheirus ellipsoideus.
•Antennal sensilla of A. ellipsoideus on macropterous and
brachypterous forms were different.
1|INTRODUCTION
The family Aphelocheiridae is distributed exclusively in the Old World and reaches its greatest species richness in
the Oriental Region (Sites et al., 2011). It is represented by a single genus Aphelocheirus, with 104 species currently
described in the world (Basu & Subramanian, 2016; Basu et al., 2013; Liu & Ding, 2005a,2005b; Liu & Zheng, 1994,
2000; Nieser & Chen, 1991; Nieser & Millán, 1989; Nieser et al., 2004; Polhemus, 1994; Polhemus & Polhemus,
1989,2013; Sites, 2005; Sites et al., 2011; Thirumalai, 2007,2008; Tran & Nguyen, 2016; Xie & Liu, 2014,2015;
Zettel et al., 2008; Zettel, 1993,1998,1999a,1999b,2000,2001,2003,2012; Zettel & Pangantihon, 2010; Zettel
& Tran, 2009). Aphelocheiridae have four‐segmented antennae, the antennae in Aphelocheiridae are easily
Arch Insect Biochem Physiol. 2022;e21917. wileyonlinelibrary.com/journal/arch © 2022 Wiley Periodicals LLC.
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https://doi.org/10.1002/arch.21917
visible since they are not concealed. Species of Aphelocheirus are known for their ability to stay submerged
underwater (Andersen & Weir, 2004), which is possible due to their plastron respiration system that also allows
them to complete their entire life cycle underwater (Polhemus & Polhemus, 1989).
They feed on immature forms of various aquatic insects such as Ephemeroptera and Trichoptera. Biological
knowledge of Aphelocheiridae is based mostly on the European species Aphelocheirus aestivalis, which feeds on the
larvae of Chironomidae (Diptera), Hydropsychidae (Trichoptera), and various molluscs belonging to the genera
Cyclas,Pisidium, and Viviparus (Thorpe, 1965). By contrast, knowledge of the biology and ecology of tropical and
subtropical species of Aphelocheirus is very scarce (Chen et al., 2005; Polhemus & Polhemus, 1989; Sites, 2005).
Up to now, 27 species of the genus Aphelocheirus (Supporting Information: Figure S1) were known from China
(Liu & Ding, 2005a,2005b; Liu & Zheng, 1994,2000; Xie & Liu, 2014,2015). Many species of Aphelocheiridae have
both macropterous and brachypterous forms of individuals. The uncommon macropterous forms, which are often
quite different structurally from the more common brachypterous morphs, are now known for the majority of the
species (Polhemus & Polhemus, 1989). Aphelocheirus ellipsoideus (Liu & Ding, 2005a,2005b) macropterous and
brachypterous forms were reviewed in Xie and Liu (2015).
Sensillar systems are especially important for aquatic insects that live in dense, dark, highly complex habitat
conditions or species with poor vision capabilities (Brönmark & Hansson, 2000; Wisenden, 2000). Moreover, in
aquatic insects, the distinction between taste and olfaction is vague, but is still used based on the structure and
particular location of the sensilla or behavioral response (Crespo, 2011; Zacharuk, 1980). Insects' antennae are an
important location for the structures responsible for sensorial perception. These structures have different
morphologies and respond to different mechanical and chemical stimuli (Nowińska et al., 2020). Although some
papers have shed light on the antennal structures and reception of environmental stimuli in nepomorphan species,
the information on morphology and physiology and the understanding of the olfactory system in water bugs are still
poor (Garza et al., 2021; Nowińska & Brożek, 2019,2020,2021; Nowińska et al., 2020).
2|MATERIALS AND METHODS
The material was obtained from the Institute of Entomology, Nankai University (NKU), Tianjin, China. All specimens
have been cleaned in an ultrasound cleaner, the antennae have been dissected, dried in ethanol, mounted,
sputtered with gold or chromium, and observed using the scanning electron microscope (SEM) Hitachi S‐3400 in
the large‐scale instruments and equipment sharing service platform of the Northeast Agricultural University in
Harbin. We follow the terminology and classification reported in other papers on antennal sensilla of insects
(Altner & Prillinger, 1980; Nowińska & Brożek, 2019,2020,2021).
3|RESULTS AND DISCUSSION
In A. ellipsoideus we observe around 1‐mm‐long and thin four‐segmented antennae. The length of scapus and pedicel is
around 140 and 190 μm, and antennomers of the flagellum are visibly longer, about 240 and 340 μm. The antennae are
rather smooth, with only some groups of thick sensilla visible on the surface of the flagellum (Figures 1–4).
3.1 |Antennal sensilla
The antennae of insects include many elementary sensilla, and they are treated as a complex multimodal
insect organ (Chapman, 1998). The morphological analysis of antennal sensory organs of the studied
Aphelocheiridae shows the presence of different sets of sensilla, forming three basal sensory systems:
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olfactory, mechanoreception, and thermo‐hygroreceptive systems. Their different polymorphic shapes
constitute different subtypes, which may be assigned to different functions depending on their morphology
and the structure of receptors (Nowińska & Brożek, 2020).
The aquatic environment is well suited for the transmission of chemical information. Aquatic animals have
evolved highly sensitive receptors for detecting these cues (Wisenden, 2000). The comparative morphology
study on antennal sensilla of A. ellipsoideus from the family Aphelocheiridae of Nepomorpha was performed
using an SEM. This study is important for the continuation of other morphological studies on the nepomorphan
taxa and suggests possible functions of sensilla based ontheirmorphologyanddistributionontheantennae.
FIGURE 1 Aphelocheirus ellipsoideus (brachypterous, male). SB, sensilla basiconica; SCa, sensilla campaniformia;
SPl, sensilla plate‐like; ST, sensilla trichodea.
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Similar studies have already been performed in most of Nepomorpha families—Belostomatidae and Nepidae
(Nowińska & Brożek, 2019), Ochteridae, Gelastocoridae, and Aphelocheiridae (Nowińska & Brożek, 2020),
Corixidae and Micronectidae (Nowińska et al., 2020), Notonectidae, Helotrephidae, and Pleidae (Nowińska &
Brożek, 2021), and Pleidae (Garza et al., 2021). During these studies, 12 morphological types of sensilla have
been observed and their probable functions have been indicated.
In this study, based on morphological characteristics, five main types of sensilla have been found
(ST, sensilla trichodea; SCa, sensilla campaniformia; SB, sensilla basiconica; SA, sensilla ampullacea; SPl, sensilla
plate‐like) (Figures 1–4). These types have been described in heteropteran insects (Nowińska & Brożek, 2020).
FIGURE 2 Aphelocheirus ellipsoideus (brachypterous, female). SA, sensilla ampullacea; SB, sensilla basiconica;
SPl, sensilla plate‐like; ST, sensilla trichodea.
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STandSCa,becauseoftheirflexiblesockets,arebelieved to play a mechanoreceptive function. SB (subtypes 2
and 3) function as chemoreceptive sensilla given their porous surface and inflexible socket. Pores on the
surface of these sensilla indicate an olfactory function. Another function is thermo‐hygroreception and it is
performed by SA and sensilla coeloconica. Morphological features and distribution of antennal sensilla are
summarized (Supporting Infomation: Table S1) and the distribution of sensilla on the antennae is reported in
specific figures (Figures 1–4).
FIGURE 3 Aphelocheirus ellipsoideus (macropterous, male). SB, sensilla basiconica; SPl, sensilla plate‐like; ST,
sensilla trichodea.
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3.2 |Mechanoreceptive sensilla
The mechanoreception system of the antennae in the studied aquatic bug species is represented by trichoid sensilla of
different shapes. Many trichoid sensilla are fine enough to detect air vibrations (Shields, 2010). In A. ellipsoideus,STand
SCa belong to mechanoreceptive sensilla. ST occur as a single hair. They are embedded into flexible sockets (Figures 1–4).
These are usually long sensilla resembling hair, tapered from the base to the tip and curved toward the antennal shaft or
they are nearly straight hairs. SCa are round plates with a singlepore,embeddedinflexiblesockets.Theyhavebeen
documented only in A. ellipsoideus (brachypterous, male) (Figure 1). They were observed singularly and the types of
FIGURE 4 Aphelocheirus ellipsoideus (macropterous, female). SB, sensilla basiconica; SPl, sensilla plate‐like; ST,
sensilla trichodea.
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mechanoreceptive sensilla were very limited on the antenommeres in comparison with the other nepomorphans (Nepidae
and Belostomatidae), which also prefer the underwater lifestyle (Nowińska & Brożek, 2019,2020). Probably, the absence
of long mechanoreceptive sensilla on the antennal surface in Aphelocheirus might be an adaptation to the type of
environment. The antennae are situated on the head without any physical protection from environmental
stressors and therefore are not concealed in grooves beneath the eyes as observed in Gelastocoridae or Nepoidea
(Nowińska & Brożek, 2019; Popov, 1971). Mechanoreceptive sensilla can detect vibrations from sound or movement,
which are carried further than in the atmosphere, due to the water's density. Moreover, considering that the water
medium is not ideal when it comes to visibility, the role of mechanosensilla in orientation becomes even more important
(Nowińska et al., 2020;Reboraetal.,2019).
3.3 |Hygro‐/thermoreceptive sensilla
Temperature and humidity are major determinants of distribution patterns for species. Thermo‐/hygroreceptive sensilla
play a role in finding new water bodies (Nowińska et al., 2020). Generally, water bugs fly from one body of water to
another during mating season or to find new water habitats. They can also disperse in other ways (Wróblewski, 1980).
SA is found only in the middle of IV antennomere of A. ellipsoideus (brachypterous female) (Figure 2). These are
pegs in pit sensilla with an external opening, which is cuticular and round, and in which no peg is visible because it is
set internally at the bottom of a tube. SA rise from inflexible sockets.
3.4 |Chemoreceptive sensilla
Chemoreceptive sensilla help with the detection of small chemicals dissolved in water. They help in intra‐and interspecific
relationships, such as predator avoidance or foraging (Brönmark & Hansson, 2000;Wisenden,2000). At this stage of
research, it is difficult to clearly indicate the extent of olfactory perception in these bugs, as they are mainly associated with
the aquatic environment. Such differences in sensilla sets may imply the specific reception of chemical compounds. All
sensilla categorized as belonging to the olfactory system meet the standards regarding olfactory sensilla previously
morphologically documented in other insectsbymanyauthors(Chapman,1998;Nowińska & Brożek, 2017,2019,
2020,2021).
3.5 |SB
SB2 and SB3 are long structures that are thick and curved or tapered at the tip. They are inserted in inflexible
sockets. Code of SB2 and SB3 are consistent with Nowińska and Brożek (2020). In A. ellipsoideus, only two types of
SB (SB2 and SB3) were observed, albeit in fewer numbers, as well as several plate‐like sensillum (SP1).
Analyzing the distribution and number of olfactory sensilla on antennae in the particular taxa examined, SB2
and SB3 were found on the penultimate and last antennomers. Several SB2 occurred in groups on the penultimate
and last antennomere most of the time (Figures 1,2, and 4); however, we also found sensilla occurring in groups
or single in the macropterous male of A. ellipsoideus (Figure 3). This SB had the same width along the whole surface,
but was slightly rounded at the tip. They bear pores along the whole surface. The presence of SB2 in all tested
species can point to a monofunctional sensillum, probably responsible for the diagnosis of water chemistry.
SB3 was singularly located at the end of the last antennomere (Figures 1–4). These are porous SB that are long
and flattened along the whole length.
SPl occurs on the distant half of the III and IV antennomere of A. ellipsoideus (male) and is more or less
distributed (Figures 1and 3). They occur only on the distant half of the IV antennomere of A. ellipsoideus (female)
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and are less evenly distributed (Figures 2and 4). These are round sensilla, with uneven edges. Probably, these
sensilla are highly specific for the mentioned taxa and are responsible for a specific behavior. Taking into account
the external morphological features (shape and build) of SPL and multilobated, Nowińska & Brożek (2020) assume
that it performs a chemoreceptive function.
3.5.1 |Comparison to the other heteropteran groups
Sets of sensilla were compared within the studied families, as well as with other nepomorphan families, semiaquatic
bugs (Gerromorpha), and other heteropteran species. The studied families show variations in their adaptation to
aquatic habitats, as well as in antennae's shapes. The basal set of sensilla (ST, SCa, SB2, and SA) is similar in all of
them and is not different from the sensilla present in Nepoidea.
In Aphelocheiridae, the antennae are long and extend over the head, making them very similar to the long and thin
antennae in Gerromorpha and other heteropterans (Andersen, 1982; Schuch & Slater, 1995). This type of antennae also
exists in Ochteridae of Nepomorpha. While in the other families of Nepomorpha, the antennae are short and concealed
beneath the head (Nowińska & Brożek, 2020; Zettel & Lane, 2010). Nonetheless, both types of antennae involve
complex sensilla of different modalities and appearances, despite a common underlying architecture.
Differences in structure/types of sensilla in particular taxa show how sensory systems have adapted to new
lifestyles. There are six sensilla types in Belostomatidae, seven sensilla types in Nepidae (Nowińska & Brożek, 2019);
five sensilla types in Aphelocheiridae and Ochteridae, seven sensilla types in Gelastocoridae (Nowińska & Brożek,
2020); five sensilla types in Corixidae and two sensilla types in Micronectidae (Nowińska et al., 2020); six sensilla
types in Notonectidae, three sensilla types in Helotrephidae, and four sensilla types in Pleidae (Nowińska & Brożek,
2021); and 12 sensilla types in Pleidae (Garza et al., 2021). In the two studied due to the different views of species
and the counting type, the set of sensilla are quite different.
The shapes of antennae and sets of sensilla in the studied families of Nepomorpha are different compared
to semiaquatic bugs (Gerromorpha). In Gerromorpha, there are more sensilla subtypes (18) and their antennae
are very long and thin. In Nepomorpha, the antennae bear fewer types of sensilla (2–12) and their antennae are
muchshorter.TheantennaeofAphelocheiridae bear sparsely distributed sensilla, which are mostly SB. It is
possible that the number of sensilla has been reduced because of their preference for running water habitats.
TheantennaeofOchteridaeandGelastocoridearemostly covered by ST, with lone SB and sensilla coeloconica
sparsely distributed on the surface. There was one new type of chemoreceptive sensilla (SPL) present in
Aphelocheiridae.
AUTHOR CONTRIBUTIONS
Tongyin Xie: Conceptualization (lead); data curation (lead); formal analysis (lead); funding acquisition (lead);
investigation (lead); methodology (lead); project administration (lead); resources (lead); software (lead); supervision
(lead); validation (lead); visualization (lead); writing—original draft (lead); writing—review and editing (lead). Shiyan
Ma: Data curation (equal); formal analysis (equal); methodology (equal); resources (equal); visualization (equal). Fuxia
He: Conceptualization (supporting); funding acquisition (supporting); investigation (supporting); methodology
(equal); project administration (equal); supervision (supporting); visualization (supporting).
ACKNOWLEDGMENT
This study is supported by the “Academic Backbones”Project of Northeast Agricultural University (No. 19XG03).
CONFLICT OF INTEREST
The authors declare that there is no conflict of interest.
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ORCID
Tong‐Yin Xie http://orcid.org/0000-0001-7059-9141
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SUPPORTING INFORMATION
Additional supporting information can be found online in the Supporting Information section at the end of this
article.
How to cite this article: Xie, T.‐Y., Ma, S.‐Y. & He, F.‐X. (2022) Report on antennal sensilla of Aphelocheirus
ellipsoideus (Hemiptera, Heteroptera, Aphelocheiridae). Archives of Insect Biochemistry and Physiology,
e21917. https://doi.org/10.1002/arch.21917
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