diversity

Editorial
An Overview of “Insect Biodiversity”
Giorgia Sollai * and Paolo Solari *

Department of Biomedical Sciences, Section of Physiology, University of Cagliari, University Campus, S.P. 8,
09042 Monserrato, Italy
* Correspondence: [email protected] (G.S.); [email protected] (P.S.)

Insects comprise more than half of all described species in the animal kingdom and
account for a considerable proportion of all biodiversity on the planet [1,2]. This great
variability is due to the specificity of the genetic, morphological, and functional aspects that
different insect species have developed to successfully cope with the complex and dynamic
habitats in which they live.
Insects are referred to as pests or disease carriers that influence agriculture, human
health, and natural resources. Many of them are also beneficial for humans, as they pollinate
plants, produce useful substances, control pest insects, act as scavengers, and serve as food
for other animals and, in the near future, possibly for humans too [3]. Furthermore, given
their great biodiversity, insects are valuable objects of study in biology, evolution, and
ecology. In fact, a large amount of scientific knowledge in genetics has been obtained from
fruit fly experiments, as well as population biology in flour beetle studies. Insects are often
used in investigations regarding hormonal action, nerve and sense organ functions, and
many other physiological processes, and also as environmental quality indicators. Even if
the causes of their remarkable diversity remain poorly understood, it has been suggested
that herbivory may have accelerated diversification in many insect clades [4,5].
In this respect, this Special Issue aims to highlight new research and significant ad-
vances in order to better understand, from different perspectives and methodological
approaches, the genetic and the morpho-functional aspects characterizing the great level of





 biodiversity in insects.
Citation: Sollai, G.; Solari, P. An One key but controversial aspect in assessing insect biodiversity is that taxonomy
Overview of “Insect Biodiversity”. sometimes appears inadequate and/or incomplete, and in a case where a species falls
Diversity 2022, 14, 134. https:// outside of the intraspecific variability range of closely related species, many taxonomists
doi.org/10.3390/d14020134 tend to list it as a new species [6]. An article by Deng et al. [7] in this Special Issue deals
Received: 7 February 2022
with this concern. These authors collected 1261 articles containing 4811 new insect species
Accepted: 10 February 2022
between 2009 and 2017 and reported that, despite the increased taxonomic efforts for the
Published: 14 February 2022
discovery of more species and their geographical distribution information, more than 21%
of these new species were described from only one specimen and/or one locality, and
Publisher’s Note: MDPI stays neutral
half of all new species were reported based on fewer than five specimens. On the other
with regard to jurisdictional claims in
hand, these authors encourage taxonomists to adopt better practices, such as increasing the
published maps and institutional affil-
number of specimens and geographical coverage of sampling, including DNA data, and
iations.
improving international collaboration in the description of new species.
Alternatively, the possibility exists that global species numbers might be underesti-
mated because of cryptic diversity [2,8]. Therefore, the use of good morphological methods
Copyright: © 2022 by the authors.
and intensive studies with large specimen numbers from many localities would help sepa-
Licensee MDPI, Basel, Switzerland. rate most species previously found to be cryptic. Advances in revealing cryptic diversity
This article is an open access article may also come from the use of DNA methods. This topic is supported in this Special
distributed under the terms and Issue by de Moya et al. [9], who explored the Bemisia tabaci complex of whiteflies, which
conditions of the Creative Commons are considered pests worldwide and are thought to contain cryptic species corresponding
Attribution (CC BY) license (https:// to geographically structured phylogenetic clades. Based on their automatic barcode gap
creativecommons.org/licenses/by/ discovery (ABGD) analyses, these authors reported the existence of at least five species
4.0/). from both the analyses of nuclear orthologs and cytochrome oxidase I.

Diversity 2022, 14, 134. https://doi.org/10.3390/d14020134 https://www.mdpi.com/journal/diversity

In a different paper of this Special Issue, Han et al. [10] revisited the phylogenetic
position of the genus Yaeprimus within Chironomini on the basis of both morphological
and molecular evidence. Their molecular results strongly support Yaeprimus as a sister
to Imparipecten Freeman, 1961, rather than to the Microtendipes group, thus countering a
previously reported systematical position exclusively based on morphological analyses.
An important, emerging threat for insect biodiversity is posed by the widespread
contamination of ecosystems with plant protection chemicals, such as fertilizers, pesticides,
and herbicides, which ultimately cause a rapid decline in both insect biomass and diver-
sity [11,12]. In this Special Issue, an article by Giglio et al. [13] examined this aspect. By
way of combined field and laboratory trials, these authors tested the effects of exposure to
realistic doses of pendimethalin-based herbicides on the constitutive immunity of Harpalus
(Pseudoophonus) rufipes, a beneficial carabid species that inhabits croplands. They reported
that exposure to herbicides can have sublethal effects, as herbicides interfere with some key
components of the immune response in insects. These effects depend on both the different
field conditions from which the insect population comes and on the cumulative effects of
repeated applications over time and suggest that this highly lipophilic herbicide, applied
in early spring when adults start foraging in the field, may be quickly absorbed through
the cuticle or ingested through the direct consumption of contaminated food.
Insect biodiversity is associated with the quantity and type of host plants available,
environmental factors, and their physiological state [14–17]. The availability of host plants
and an insect’s ability to find them are key factors for the survival of a species because
they represent both suitable oviposition sites for adult females and potential food sources
for the offspring [18–20]. An important role in the choice and recognition of a host plant
is played by the information that the olfactory and gustatory systems send to the central
nervous system (CNS) on the chemical composition of the plant [21–23]. In particular,
insects show a great peripheral plasticity that allows them to adapt to the environment
in which they live [24]. In this Special Issue, Sollai et al. [25] show that sex, physiological
state, and experience can modulate the olfactory sensitivity of the medfly Ceratitis capitata,
a highly invasive species of economic interest. The results show that: (a) lab-reared mated
males are more sensitive to host plant headspace than females, while the opposite is true for
wild insects; (b) wild virgin males are more sensitive than mated ones, while no difference
was observed among lab-reared medflies; (c) lab-reared virgin females are more sensitive
than mated ones, while few differences were found within wild medflies; (d) lab-reared
mated males are more sensitive to host plant extracts than wild ones, while the opposite
was found for females. Taken together, these findings highlight that the physiological
state and habitat contribute to the peripheral plasticity of insects of both sexes, modulating
their olfactory sensitivity to ensure the most appropriate adaptations for the survival of
the species.
Remaining in the context of environmental conditions, in this Special Issue,
Viterbi et al. [26] published an interesting study on the effects of temperature and its
changes on the biodiversity of insects in mountains. In particular, the authors observed
significant differences between groups of species and along the altitudinal gradient, al-
though only small changes emerged in the overall biodiversity patterns. The effects of
temperature increase could be more pronounced for spiders and butterflies and could
be particularly detrimental for high-altitude species. They observed significant changes
in community composition and species richness, especially in the alpine belt, but a clear
separation between vegetation levels was also retained in the warming scenarios. This
conservative approach suggests that even a moderate temperature increase (of about 1 ◦ C)
could influence animal biodiversity in mountain ecosystems.
Insects’ biodiversity may represent a threat for rare plant species, which are reliable
indicators of environmental changes but also are a resource in various economic sectors,
such as pollination and human health. The loss of biodiversity is related to several key
factors, such as human activity (fragmentation and loss of habitat, pollution, etc.), climatic
events, and geological processes [27]. This course can be reversed through a reduction in
Diversity 2022, 14, 134 3 of 5

the use of environmental pollutants and pesticides and the increase in favorable habitats
for the species [28]. To this end, it could be of particular importance to learn the factors
that regulate the relationships between plants, the environment, and herbivores [29]. For
example, the effects of herbivorous species capable of influencing the viability of a host
plant and the development of its reproductive structures should be considered [30]. In this
Special Issue, Bonsignore et al. [31] showed the effects that several species of herbivorous
insects have on a rare plant species, Salvia ceratophylloides, endemic of southern Italy. They
found bottom-up and top-down effects on plant health and reproduction associated with
herbivorous action. Among the herbivores, mainly Squamapion elongatum affected this rare
species of sage: the acquired data indicate that the density of the herbivore in the area of
diffusion of sage does not represent a quantitative regulating factor of flowering, but it can
rather condition the survival of the species.
In this Special Issue, Enkhtur et al. [32] showed that moths are creatures with an
important role in the ecosystem and have the potential to serve as environmental indicators.
In particular, Geometrid moths (Geometridae), constituting one of the biggest families of
Lepidoptera, are a species-rich and easily recognizable family and have served as indicators
for environmental changes [33]. By analyzing the distribution pattern, species richness,
and biodiversity in the Mongolian ecoregions and correlating them with environmental
variables, the authors concluded that annual precipitation and the maximum temperature
of the warmest month were the most important environmental variables that correlated in
an analysis of geometrid assemblages of different ecoregions in Mongolia.
Finally, Hristov et al. [34] published an interesting review describing how the reduction
in honey bee populations affects various economic sectors, as well as human health. Despite
the important role played by these insects, a progressive decline in bee colonies is being
observed due to the effect of the excessive use of pesticides in agricultural production,
genetically modified plants, electromagnetic radiation, inadequate honey bee nutrition,
crops growing in monoculture, and biodiversity loss. Honey bees are the most economically
valuable pollinator in the world: 9.5% of the total economic value of agricultural production
comes from insect pollination, which totals an amount of just under USD 200 billion globally.
They pollinate not only a large number of commercial crops (cereals, vegetables, fruits,
edible oil crops, stimulants, and nuts and spices), but also many wild plants, some of which
are threatened with extinction, playing a significant role in every aspect of ecosystem,
facilitating the growth of trees, flowers, and other plants that serve as food and shelter for
many creatures large and small. Honey bee products, such as honey, pollen, royal jelly,
propolis, bee venom, wax, and bee bread, are important resources with regard to human
nutrition and the production of pharmaceuticals and food additives.
In conclusion, even if this Special Issue does not entirely cover the vast range of the
possible topics related to insect biodiversity, it provides insight into the multiple directions
with which biodiversity intersects. Unfortunately, we are currently experiencing a biodiver-
sity crisis: many species, among which are insects, are becoming extinct, probably before
we can even detect their existence and describe them.This loss in insect biodiversity, which
is mainly a consequence of anthropogenic pressure, will ultimately lead to a subsequent
decline in ecosystem stability and functioning. In this respect, better knowledge of the
genetic, morphological, and functional aspects characterizing the great level of biodiversity
in insects will be decisive for the safeguarding of ecosystems.

Author Contributions: G.S. and P.S., writing—original draft preparation, review and editing. All
authors have read and agreed to the published version of the manuscript.
Funding: This research received no external funding.
Acknowledgments: We would like to thank all the authors and referees for their remarkable contri-
bution to this SI.
Conflicts of Interest: The authors declare no conflict of interest.
Diversity 2022, 14, 134 4 of 5

References
1. Tihelka, E.; Cai, C.; Giacomelli, M.; Lozano-Fernandez, J.; Rota-Stabelli, O.; Huang, D.; Engel, M.S.; Donoghue, P.C.J.; Pisani, D.
The evolution of insect biodiversity. Curr. Biol. 2021, 31, R1299–R1311. [CrossRef] [PubMed]
2. Stork, N.E. How Many Species of Insects and Other Terrestrial Arthropods Are There on Earth? Ann. Rev. Entomol. 2018, 63,
31–45. [CrossRef] [PubMed]
3. Manno, N.; Estraver, W.Z.; Tafur, C.M.; Torres, C.L.; Schwarzinger, C.; List, M.; Schoefberger, W.; Coico, F.R.M.; Leon, J.M.; Battisti,
A.; et al. Edible Insects and Other Chitin-Bearing Foods in Ethnic Peru: Accessibility, Nutritional Acceptance, and Food-Security
Implications. J. Ethnobiol. 2018, 38, 424–447. [CrossRef]
4. Sollai, G.; Tomassini Barbarossa, I.; Masala, C.; Solari, P.; Crnjar, R. Gustatory sensitivity and food acceptance in two phylogeneti-
cally closely related papilionid species: Papilio hospiton and Papilio machaon. PloS ONE 2014, 9, e100675. [CrossRef]
5. Wiens, J.J.; Lapoint, R.T.; Whiteman, N.K. Herbivory increases diversification across insect clades. Nat. Comm. 2015, 6, 8370.
[CrossRef]
6. Lim, G.S.; Balke, M.; Meier, R. Determining species boundaries in a world full of rarity: Singletons, species delimitation methods.
Syst. Biol. 2012, 61, 165–169. [CrossRef]
7. Deng, J.; Guo, Y.; Cheng, Z.; Lu, C.; Huang, X. The Prevalence of Single-Specimen/Locality Species in Insect Taxonomy: An
Empirical Analysis. Diversity 2019, 11, 106. [CrossRef]
8. Adis, J. Thirty Million Arthropod Species-too Many or too Few? J. Trop. Ecol. 1990, 6, 115–118. [CrossRef]
9. De Moya, R.S.; Brown, J.K.; Sweet, A.D.; Walden, K.K.O.; Paredes-Montero, J.R.; Waterhouse, R.M.; Johnson, K.P. Nuclear
Orthologs Derived from Whole Genome Sequencing Indicate Cryptic Diversity in the Bemisia tabaci (Insecta: Aleyrodidae)
Complex of Whiteflies. Diversity 2019, 11, 151. [CrossRef]
10. Han, W.; Wei, J.; Lin, X.; Tang, H. The Afro–Oriental Genus Yaeprimus Sasa et Suzuki (Diptera: Chironomidae: Chironomini):
Phylogeny, New Species and Expanded Diagnoses. Diversity 2020, 12, 31. [CrossRef]
11. Brühl, C.A.; Zaller, J.G. Biodiversity Decline as a Consequence of an Inappropriate Environmental Risk Assessment of Pesticides.
Front. Environ. Sci. 2019, 7, 177. [CrossRef]
12. Raven, P.H.; Wagner, D.L. Agricultural intensification and climate change are rapidly decreasing insect biodiversity. Proc. Natl.
Acad. Sci. USA 2021, 118. [CrossRef] [PubMed]
13. Giglio, A.; Cavaliere, F.; Giulianini, P.G.; Kurtz, J.; Vommaro, M.L.; Brandmayr, P. Continuous Agrochemical Treatments in
Agroecosystems Can Modify the Effects of Pendimethalin-Based Herbicide Exposure on Immunocompetence of a Beneficial
Ground Beetle. Diversity 2019, 11, 241. [CrossRef]
14. Dangles, O.; Irschick, D.; Chittka, L.; Casas, J. Variability in Sensory Ecology: Expanding the Bridge Between Physiology and
Evolutionary Biology. Q. Rev.Biol. 2009, 84, 51–74. [CrossRef] [PubMed]
15. Reside, A.E.; Butt, N.; Adams, V.M. Adapting systematic conservation planning for climate change. Biodivers. Conserv. 2018, 27,
1–29. [CrossRef]
16. Sollai, G.; Biolchini, M.; Solari, P.; Crnjar, R. Chemosensory basis of larval performance of Papilio hospiton on different host
plants. J. Insect Physiol. 2017, 99, 47–57. [CrossRef]
17. Thompson, J.N.; Pellmyr, O. Evolution of Oviposition Behavior and Host Preference in Lepidoptera. Ann. Rev. Entomol. 1991, 36,
65–89. [CrossRef]
18. Sollai, G.; Biolchini, M.; Crnjar, R. Taste sensitivity and divergence in host plant acceptance between adult females and larvae of
Papilio hospiton. Insect Sci. 2018, 25, 809–822. [CrossRef]
19. Sollai, G.; Biolchini, M.; Loy, F.; Solari, P.; Crnjar, R. Taste input from tarsal sensilla is related to egg-laying behavior in Papilio
hospiton. Entomol. Exp. Appl. 2017, 165, 38–49. [CrossRef]
20. Sollai, G.; Solari, P.; Crnjar, R. Olfactory sensitivity to major, intermediate and trace components of sex pheromone in Ceratitis
capitata is related to mating and circadian rhythm. J. Insect Physiol. 2018, 110, 23–33. [CrossRef]
21. Feeny, P.; Städler, E.; Åhman, I.; Carter, M. Effects of plant odor on oviposition by the black swallowtail butterfly, Papilio polyxenes
(Lepidoptera: Papilionidae). J. Insect Behav. 1989, 2, 803–827. [CrossRef]
22. Nishida, R. Chemosensory basis of host recognition in butterflies–multi-component system of oviposition stimulants and
deterrents. Chem. Sens. 2005, 30 (Suppl. 1), i293–i294. [CrossRef] [PubMed]
23. Solari, P.; Corda, V.; Sollai, G.; Kreissl, S.; Galizia, C.G.; Crnjar, R. Morphological characterization of the antennal lobes in the
Mediterranean fruit fly Ceratitis capitata. J. Comp. Physiol. A Neuroethol. Sens. Neur. Behav. Physiol. 2016, 202, 131–146. [CrossRef]
[PubMed]
24. Sollai, G.; Biolchini, M.; Crnjar, R. Taste receptor plasticity in relation to feeding history in two congeneric species of Papilionidae
(Lepidoptera). J. Insect Physiol. 2018, 107, 41–56. [CrossRef] [PubMed]
25. Sollai, G.; Solari, P.; Crnjar, R. Differences in the Olfactory Sensitivity of Ceratitis capitata to Headspace of Some Host Plants in
Relation to Sex, Mating Condition and Population. Diversity 2020, 12, 207. [CrossRef]
26. Viterbi, R.; Cerrato, C.; Bionda, R.; Provenzale, A. Effects of Temperature Rise on Multi-Taxa Distributions in Mountain Ecosystems.
Diversity 2020, 12, 210. [CrossRef]
27. Corlett, R.T. Plant diversity in a changing world: Status, trends, and conservation needs. Plant Divers. 2016, 38, 10–16. [CrossRef]
28. Wagner, D.L.; Van Driesche, R.G. Threats posed to rare or endangered insects by invasions of nonnative species. Ann. Rev.
Entomol. 2010, 55, 547–568. [CrossRef]
Diversity 2022, 14, 134 5 of 5

29. Souza, L.; Zelikova, T.J.; Sanders, N.J. Bottom–up and top–down effects on plant communities: Nutrients limit productivity, but
insects determine diversity and composition. Oikos 2016, 125, 566–575. [CrossRef]
30. Ancheta, J.; Heard, S.B. Impacts of insect herbivores on rare plant populations. Biol. Conserv. 2011, 144, 2395–2402. [CrossRef]
31. Bonsignore, C.P.; Laface, V.L.A.; Vono, G.; Marullo, R.; Musarella, C.M.; Spampinato, G. Threats Posed to the Rediscovered and
Rare Salvia ceratophylloides Ard. (Lamiaceae) by Borer and Seed Feeder Insect Species. Diversity 2021, 13, 33.
32. Enkhtur, K.; Boldgiv, B.; Pfeiffer, M. Diversity and Distribution Patterns of Geometrid Moths (Geometridae, Lepidoptera) in
Mongolia. Diversity 2020, 12, 186. [CrossRef]
33. Ashton, L.; Maunsell, S.; Bito, D.; Putland, D. Macrolepidopteran assemblages along an altitudinal gradient in subtropical
rainforest—Exploring indicators of climate change. Mem. Queensl. Mus. 2011, 55, 375–389.
34. Hristov, P.; Neov, B.; Shumkova, R.; Palova, N. Significance of Apoidea as Main Pollinators. Ecol. Econ. Impact Implic. Hum.
Nutrition. Divers. 2020, 12, 280.