Neotrop Entomol (2020) 49:783–794

https://doi.org/10.1007/s13744-020-00781-y

FORUM

Ecosystem Services Provided by Insects in Brazil: What Do

We Really Know?

DL RAMOS1, WL CUNHA1, J EVANGELISTA2, LA LIRA2, MVC ROCHA1, PA GOMES1, MR FRIZZAS3 ,

PHB TOGNI4
1
Programa de Pós-Graduação em Ecologia, Univ de Brasília – UnB, Brasília, DF, Brasil
2
Programa de Pós-Graduação em Zoologia, Univ de Brasília – UnB, Brasília, DF, Brasil
3
Depto de Zoologia, Instituto de Ciências Biológicas, Univ de Brasília - UnB, Brasília, DF, Brasil
4
Depto de Ecologia, Univ de Brasília – UnB, Brasília, DF, Brasil

Keywords
Diversity, biological control, decomposition,
ecological services, Insecta, pollination
Correspondence
M.R. Frizzas, Depto de Zoologia, Instituto de
Ciências Biológicas, Univ de Brasília - UnB,
Campus Universitário Darcy Ribeiro,
Brasília, DF, 70910-900 Brasil; frizzas@unb.
br
Edited by Lessando Moreira Gontijo – UFV
Received 19 December 2019 and accepted
28 April 2020
Published online: 28 May 2020
* Sociedade Entomológica do Brasil 2020
Abstract
Insects are the most abundant and diverse organisms on Earth and pro-
vide essential ecosystem services. However, Brazilian society rarely con-
sider the importance of insects in their diverse country. Therefore, in this
review, we provide an overview of ecosystem services provided by insects
in Brazil. A database search returned 136 articles, published in English or
Portuguese, on ecosystem services provided by insects in Brazil. The first
article was published in 1982, and majority of the studies were conducted
in the Atlantic Forest or the Cerrado biomes. The most frequently studied
insect-provided ecosystem services were pollination, decomposition, and
biological control of pests. The studies focused primarily on natural and
anthropic ecosystems, and most followed an experimental approach. We
noted that the term “ecosystem services” was not used frequently in
studies on insects in Brazil. The information available was mostly taxon-
biased. We discuss the implications of these findings in relation to recon-
ciling economic interests and the need for insect conservation for contin-
ued provision of ecosystem services in a broader perspective. In conclu-
sion, we argue that the scientific community should focus on understand-
ing the ecosystem services provided by insects other than those strictly
related to economic activities, and on improving communication with
policymakers and citizens. As a tropical and megadiverse country, Brazil
has the potential to become a protagonist in conserving and using the
ecosystem services provided by insects, both locally and internationally,
by providing scientific information to policymakers and citizens.

Introduction services), the regulation of ecosystem processes (e.g., soil

The concept of ecosystem services has advanced our under-
standing of how conservation and biodiversity at all levels of
organization are critical to human well-being (Daily et al 1991,
CDB 2000, Costanza et al 2014, IPBES 2018). Ecosystem serv-
ices are the direct and indirect benefits provided by nature to
the society (CDB 2000). These benefits are related to the
products obtained directly from nature (provisioning
fertility and pollination) (regulating services), the basis for
production of all ecosystem services (i.e., supporting serv-
ices) (e.g., maintenance of genetic diversity), and non-
material and esthetic services to the society (cultural serv-
ices) (MEA 2005, IPBES 2018). The value of global ecosystem
services in 2011 was estimated at $125–145 trillion/year,
which represents a loss of $4.3–20.2 trillion/year from 1997
to 2011 due to land-use change in this period (Costanza et al
784
2014). Land-use change is one of the main drivers of biodi-
versity loss worldwide, and the rapid decline in biodiversity
results in the economic loss of global ecosystem services
(IPBES 2018).
Insects are the most abundant and diverse organisms on
Earth, with approximately 1 million described species, and at
least 5 million species yet to be discovered (Stork 2018). They
provide important ecosystem services essential for humans
(Schowalter et al 2018), such as pollination (e.g., Garibaldi
et al 2016), biological control of pests (e.g., Togni et al
2019a), decomposition (e.g., Nichols et al 2008), herbivory
(e.g., Schowalter 2012), and seed dispersal (e.g., Braga et al
2013, Milotić et al 2018). These services are not restricted to
natural environments, as they also operate in and benefit
anthropic landscapes and semi-natural habitats. However,
the benefits of insects are seldom appreciated by the society
(Prather and Laws 2018). Recent evidence indicates that at
the current insect extinction rate, 40% of the world’s insect
species would be lost within a few decades (Sánchez-Bayo
and Wyckhuys 2019). It is expected that the ecosystem serv-
ices provided by these organisms would also be negatively
impacted. Therefore, it is important to recognize the insects
that provide key ecosystem services, and the services that
are in demand by the society and policymakers.
The loss of ecosystem services is of particular concern in
the tropical regions, where most of the global biodiversity is
concentrated (Myers et al 2000, Sala et al 2000, Lewinsohn
and Prado 2005). The highest biodiversity of insects in the
world can be found in the megadiverse country of Brazil
(Rafael et al 2009), which accommodates six biomes that
vary in size, geomorphology, climatic patterns, species rich-
ness, and endemism (IBGE 2004). The forest biomes com-
prise of the internationally known Amazon Rainforest, which
is the country’s largest biome (IBGE 2004), and the Atlantic
Forest, which is one of Brazil’s biodiversity hotspots and is
located mainly along the Brazilian coast (Morellato & Haddad
2000, Myers et al 2000). Another biodiversity hotspot is the
Cerrado, which is a tropical savannah and the second largest
biome that spans approximately one quarter of the country
(Ribeiro & Walter 2008). The Cerrado is considered to be
Brazil’s last agricultural frontier (Sano et al 2019). The
Pantanal biome is the largest continuous wetland in the
world, with flood regimes influenced primarily by the
Paraguay River (Harris et al 2005). The semiarid Caatinga,
which is the only biome exclusive to Brazil, is the largest
tropical dry forest in South America, and is sensitive to de-
sertification in some areas (Overbeck et al 2009, Silva et al
2017). Finally, the Brazilian Pampa is a predominantly
grassland-like biome, with both subtropical and temperate
climates (Roesch et al 2009).
In recent decades, drastic changes have occurred in Brazil,
such as the increase in agricultural activities, livestock, defor-
estation, burning, and urbanization, which have resulted in a
Ramos et al
decrease in the biodiversity and provision of related ecosys-
tem services. Considering the species and ecological diversi-
ties of Brazil, it is reasonable to assume that the country plays
a pivotal role in the maintenance and provisioning of impor-
tant ecosystem services worldwide (IPBES 2018). Brazil also
has continental dimensions, which when allied to its ecolog-
ical importance in the global scenario, makes it a good model
to understand the provisioning of ecosystem services in trop-
ical regions. However, most research on insect-provided eco-
system services has been conducted in temperate regions,
particularly in North America and Europe, with little informa-
tion available about the tropics (Noriega et al 2018). Thus,
our understanding of insect-provided ecosystem services is
limited in a country that should be a leader on the subject.
The lack of scientific evidence also hampers the ability of
policymakers to make effective decisions for biodiversity
conservation as stated by the Convention on Biological
Diversity (CBD), to which Brazil is a signatory (CDB 2000).
To address these knowledge gaps, we surveyed the scien-
tific literature and reviewed the information available on
ecosystem services provided by insects in Brazil, in order to
evaluate research trends, identify opportunities, and better
understand the demands of studying the most diverse taxa
among the life kingdoms in a megadiverse country. Our aim
is to stimulate debate among researchers, society, and poli-
cymakers, for incorporating insect-provided ecosystem serv-
ices in decision-making, especially in the design of
biodiversity-friendly landscapes. We believe that to accom-
plish this objective, the scientific information available on
ecosystem services provided by insects in Brazil, and how
such studies are being conducted in Brazil, should first be
addressed. Therefore, answering basic questions on the eco-
system services provided by insects in Brazil could be the
starting point to promote the interplay among researchers,
society, and decision-makers.
A Brief Overview of Insects and Ecosystem Services
Considering the high diversity of insects in all ecosystems, it is
believed that they play a central role in providing ecosystem
services belonging to all categories defined by the
Millennium Ecosystem Assessment—MEA (2005). It is impor-
tant to note that these categories are not mutually exclusive.
Here, we focused on an overview of the most extensively
studied ecosystem services related to insects. The most ob-
vious provisioning service catered by insects is their use as a
food source by humans. Using insects such as grasshoppers
as food can cut down the gas emissions by half, reducing the
land used for agriculture and the pollution caused by insecti-
cides, in addition to providing a rich source of protein for the
human diet (Wegier et al 2018). Silk and honey have been
used by humans for manufacturing and industrial purposes
Ecosystem Services and Insects in Brazil
for a long time (Zhang et al 2008). Maggot therapy has been
used clinically for the treatment of non-healing wounds and
other diseases with promising results (Stadler 2019). In this
sense, there are several provisioning services that can direct-
ly or indirectly benefit anthropic activities.
Several studies have been conducted on the regulating
services provided by insects. Services that are directly asso-
ciated with socio-economic activities, such as biological con-
trol and pollination, are the most widely studied of these
regulating ecosystem services (Noriega et al 2018). The ben-
eficial effects of biological control and pollination on crop
yield are directly related to the richness of pollinators and
natural enemies (Dainese et al 2019). Herbivory by non-pest
insects is essential for regulating primary production in for-
ests worldwide, consequently affecting carbon and nutrient
cycling (Schowalter 2012, Metcalfe et al 2014). By removing
dung from natural and anthropic environments, dung beetles
indirectly suppress pathogens (Ryan et al 2011), act as gas-
trointestinal parasite controls (Sands and Wall 2017), and
reduce the populations of horn flies in cattle (Nichols et al
2008). Soil-dwelling insects, such as termites and leaf-cutting
ants, are directly linked to the carbon and nitrogen cycles,
due to their effects on soil biological and physical properties
and organic matter inputs from their nests (Jouquet et al
2011, Swanson et al 2019). These services have a high eco-
nomic value associated with them (Costanza et al 2014).
Biodiversity conservation is central to supporting several
other ecosystem services provided by insects. Biodiversity
conservation studies are often used as proxies of ecosystem
service provision (Noriega et al 2018, Cardoso et al 2020). In
fact, knowledge of insect biodiversity conservation and their
biodiversity patterns is primarily required to understand how
insects deliver ecosystem services. However, it must be not-
ed that supporting services also include nutrient accumula-
tion in the soil (Swanson et al 2019), recycling of organic
matter (Ulyshen et al 2014), soil structure and aeration
(Jouquet et al 2011), provision of resources to the food chain
(Evans et al 2011), and modification of the succession and
distribution of plants in natural environments (Schowalter
2012).
Insects are also a part of the cultural heritage in many
human societies. They are used as religious symbols and in
folklore (Lenko and Papavero 1997, Cardoso et al 2020). The
Tucandeira ritual is a part of the ethnic identity of the Sateré-
mawé Indians (Botelho and Weigel 2011). Ancient Egyptians
were fascinated by the sacred scarab beetles; they have been
found in religious rites, tomb hieroglyphics, and drawings,
and are believed to have been used as amulets, commemo-
ratives, good luck charms, and hand-carved scarabs, and
were even buried with the dead (Cherry and Kritsky 1985).
Massive migration of insects, such as that of the Monarch
butterfly in North America (Tucker 2004) and the nightly
aggregation of fireflies in the mangroves in Malaysia, attract
785
several tourists to these regions (Nallakumar 2003). In addi-
tion, many insect species are used as bioindicators in terres-
trial and aquatic ecosystems (McGeoch 2007, Maleque et al
2009).
A large amount of information is available on the ecosys-
tem services provided by insects worldwide. During the prep-
aration of an Insect Ecology course, we conducted a prelim-
inary survey of studies on this subject undertaken in Brazil.
This preliminary survey returned a very low number of stud-
ies, which drew our attention. Several studies in this research
area, including those on pollination, biological control, de-
composition, herbivory, insect conservation, and many other
services, are known to us (including some of our previous
studies) that were not found in this basic search. We also
noticed that our current knowledge on ecosystem services
provided by insects is strongly biased towards studies con-
ducted in temperate regions. For example, Noriega et al
(2018) studied the research trends of ecosystem services
provided by insects globally, but only included a few studies
from tropical regions such as Brazil. This is a cause of con-
cern, because although studies are being conducted on eco-
system services provided by insects in tropical regions, this
information is not easily available. This highlights the fact
that if such information is not easily retrievable by research-
ers themselves, it would be considerably more difficult for
the society and policymakers to use or retrieve the informa-
tion available. In order to conduct a more in-depth investi-
gation of studies and research trends related to ecosystem
services provided by insects in Brazil, we systematically
reviewed the available scientific literature, using a similar
approach as that of Noriega et al (2018).
Literature Survey for Ecosystem Services Provided
by Insects in Brazil
We conducted a broad literature survey, using online plat-
forms to search for peer-reviewed articles on ecosystem
services provided by insects in Brazil. For this study, we de-
fined “ecosystem service” according to the conceptual ap-
proach established by the Millennium Ecosystem
Assessment, which includes provisioning, regulating, sup-
porting, and cultural services (MEA 2005).
We used a two-step approach to select papers for our
analysis. First, we searched the ISI Web of Knowledge and
Scopus using the keywords “(ecosystem* service* AND
Brazil* AND insect*).” These keywords are similar to those
used by Noriega et al (2018), who searched for papers on
ecosystem services provided by insects worldwide. Because
many studies in Brazil are published in Brazilian Portuguese,
we also used the keywords “(serviço* ecossistêmico* AND
Brasil* AND inseto*)” to search the ISI Web of Knowledge,
Scopus, and Google Scholar. We searched for papers with
786
these keywords in the title, abstract, keywords, or topic. The
procedure was the same as adopted by Noriega et al (2018),
which enabled us to compare our results with the global
scenario of studies on ecosystem services provided by
insects, without a sampling bias. However, as was the case
for Noriega et al (2018), this method focused mostly on eco-
system services in terrestrial ecosystems.
Our initial survey returned a total of 2635 publications, of
which 2560 were obtained from Google Scholar, 42 from the
ISI Web of Knowledge, and 33 from Scopus. We carefully
reviewed these articles and eliminated all conference papers,
articles in press, congress papers, theses and dissertations,
technical documents, and papers characterizing taxa other
than insects. Review articles were not included in our analy-
sis, except for those that analyzed secondary data. Majority
of papers returned in this step focused on the ecosystem
services of decomposition, pollination, biological control,
seed dispersal, and nutrient cycling. Therefore, in the next
step of our survey, we included the following keywords in
English and Portuguese: “((decomposition OR decomposição
OR pollination OR polinização OR bio-control OR biocontrol
OR controle biológico OR secondary seed dispersal OR nutri-
ent* cycl*) AND insect* AND (Brazil OR Brasil)).” We applied
the same search exclusion criteria to these papers as in the
first step. By adopting this approach, we included papers that
assessed specific ecosystem services (the most common
services based on the primary literature survey), but did
not include the term “ecosystem services” as a keyword.
We are aware that this procedure could produce a potential
bias in our data; however, we assume that at least the most
cited ecosystem services in the literature were included in
the survey, since most studies did not use the term “ecosys-
tem services” in their texts.
Research Trends in Brazil Over Time
To identify research trends over time, we calculated the
number of publications per year, from 1982 to 2018. We
identified 136 articles on ecosystem services provided by
insects in Brazil. Since the first published article in 1982, there
has been an exponential increase in the number of articles
published on this subject each year (Fig 1), with most articles
published after 2010 (Fig 1).
The first article published using the term “ecosystem ser-
vice” in relation to insects in Brazil dates back to the 1980s.
However, this research topic became more prominent in the
scientific community after the 2000s. During this decade,
important legislations on biodiversity conservation were
implemented, including the National System of Nature
Conservation Units (Brasil 2000), National Biodiversity
Policy (Brasil 2002a), and National Strategic Plan for
Protected Areas (Brasil 2002b). These policies, along with
Ramos et al
results from the Millennium Ecosystem Assessment (MEA
2005), stimulated research and research grants in the coun-
try, resulting in the subsequent production of scientific evi-
dence necessary to support more specific initiatives.
Consequently, the Brazilian Platform on Biodiversity and
Ecosystem Services (BPBES) was established in 2017 (BPBES
2018). The IPBES (2018) has included some ecosystem serv-
ices provided by insects, such as pollination, in their reports.
An increase in the number of studies on insect-provided eco-
system services in Brazil accompanied these events. In addi-
tion, the number of studies on this subject conducted in
Brazil followed the same trend as observed for studies con-
ducted globally (Noriega et al 2018). Thus, the political and
social demands associated with national and international
conventions, treaties, and science funding can be important
drivers of scientific inquiries into the role of ecosystem serv-
ices in human well-being (Hill et al 2013), as well as the role of
insects in these services.
Although studies on insect-provided ecosystem services in
Brazil have been conducted for a long time, the term “eco-
system services” is not always used by the researchers, even
when specific ecosystem services, such as pollination, are
being studied. This can be attributed to the different inter-
pretations of the term, or to the fact that the term is more
familiar in some other research areas, such as ecology and
conservation biology, where it is used more frequently.
It is evident that significant research is conducted on eco-
system services provided by insects in Brazil, generating con-
siderable amounts of information. However, the low use of
the term makes it difficult to retrieve the information avail-
able. We recommend that researchers use this term to clarify
the connection between insects and ecosystem services so
that this information can be easily retrieved when searching
scientific databases. We do not want to trivialize the use of
the term. However, the scientific community involved in
studying ecosystem services provided by insects should stan-
dardize the use of this term and its implication in conserva-
tion, human well-being, economics, and decision-making.
We propose the use of the term “ecosystem services” in light
of the definitions used by the CDB (2000) and MEA (2005).
This is because Brazil is a signatory of the CDB (2000), and
such conventions and treaties define the terms and legal
mechanisms for biodiversity use and conservation in the
country.
Ecosystem Services, Insects, and Brazilian Biomes
To determine if the studies were distributed equally across
the Brazilian biomes, we plotted the study areas on a map
using the ArcGIS software and calculated the total number of
published studies per biome. Most studies were conducted
in the East and Northeast states, with 39.13% in the Atlantic
Ecosystem Services and Insects in Brazil
Fig 1 Cumulative number of articles on ecosystem services provided by insects in Brazil from 1982 to 2018
Forest, 20.29% in the Cerrado, 10.87% in the Caatinga, 6.52%
in the Amazon Rainforest, and 2.90% each in the Pampa and
Pantanal biomes (Fig 2). Five articles did not specify the bi-
ome where the study was conducted, and information re-
quired to determine where it was performed was lacking.
In addition to being the most megadiverse country in the
world, Brazil is a very large country occupied by six distinct
biomes (Brandon et al 2005, Lewinsohn and Prado 2005).
The higher number of studies carried out in the Atlantic
Forest and the Cerrado biomes might be due to two main
reasons. First, the most important economic activities of the
country are concentrated in these biomes. Second, most
qualified research centers can be found, along with densely
populated cities, in both these biomes (Guimarães 2002,
Coutinho 2006, Sano et al 2019). Between 2000 and 2015,
an estimated 236,000 km2 of natural ecosystems in the
Cerrado were converted to anthropic ecosystems. The
Atlantic Forest biome, which had already lost most of its
forested area, lost approximately 290 km2 of its remaining
natural area between 2015 and 2016, mainly due to agricul-
tural expansion (BPBES 2018). Because both biomes are bio-
diversity hotspots (Myers et al 2000), the large number of
studies in these biomes is desirable. Nevertheless, more in-
formation on the supporting, provisioning, and cultural serv-
ices is required.
The low number of studies carried out in other Brazilian
biomes, including the Amazon Rainforest, is a cause of con-
cern, as it indicates that the role of insects in these biomes
has been overlooked. The Amazon Rainforest provides sev-
eral supporting and regulating services to the society, includ-
ing climate regulation. Therefore, more research on the sup-
porting and provisioning services, such as nutrient cycling, of
the natural environments should be undertaken in this re-
gion. Land-use change and disturbance in the Amazon
787
Rainforest reduces the conservation value of the region
(Barlow et al 2016). Studies on how land-use change and
disturbance affect insects and their services should be
promoted. For example, Solar et al (2016) demonstrated that
forest areas support almost double the number of species
than production areas. More importantly, the composition of
ant fauna is drastically affected by land-use change, which
can further affect the ecosystem services provided. Such
well-designed studies have not been realized in all biomes.
On analyzing scientific literature related to the Caatinga,
Pantanal, and Pampa biomes, we encountered a lack of basic
research on insects. It must be noted that this does not mean
that research is not being conducted on this aspect. The
studies that we reviewed mainly focused on regulating serv-
ices, such as pollination, decomposition, and biological con-
trol. This highlights the tendency of scientists to focus on
ecosystem services related to agricultural production in the
country. We argue that studies should also aim to fill the
gaps in biodiversity sampling in terrestrial and freshwater
ecosystems. It is evident that many researchers are working
towards filling these gaps; however, we reiterate that the link
between insects and ecosystem services is not always clear in
majority of the papers. In addition, there is a need to provide
greater scientific financial support to projects related to bio-
diversity conservation, particularly in these biomes.
Services Provided by Insects in Different Brazilian
Ecosystems
The studies we reviewed were classified based on the type of
the ecosystem service evaluated (e.g., pollination, decompo-
sition, seed dispersal). Those reporting the results of more
than one ecosystem service provided by multiple species
788
Fig 2 Number of studies evaluating insect-provided ecosystem services in each Brazilian biome
were classified as “multiple services.” The ecosystems inves-
tigated were classified as natural ecosystems (e.g., natural
protected areas), anthropic ecosystems (cities and rural
areas), semi-natural ecosystems (fallow and regeneration
areas), or natural + anthropic ecosystems (when the study
evaluated either both ecosystems simultaneously or the re-
lationship between them). The most frequently studied
insect-provided ecosystem service in our survey was pollina-
tion (44.30%), followed by decomposition (29.11%), biologi-
cal control (14.56%), multiple ecosystem services (8.86%),
herbivory (1.90%), seed dispersal, and other ecosystem serv-
ices (0.63% each) (Fig 3). These studies were primarily carried
out in natural (39.87%) or anthropic ecosystems (25.32%). In
natural ecosystems, studies on pollination and decomposi-
tion accounted for 86.30% of the analyzed studies. Similarly,
in anthropic ecosystems, studies on pollination and biological
control were found to be more frequent in our analysis
(66.67%) (Fig 3).
Regardless of the ecosystem type, the most extensively
studied ecosystem services provided by insects were pollina-
tion, decomposition, and biological control of pests. These
services are essential to agriculture, and therefore, to the
Brazilian economy. Insect pollination increases the economic
value of agricultural production in Brazil by about 30%, which
corresponds to US $12 billion per year (Giannini et al 2015). At
Ramos et al
the global level, approximately 85% of crops are dependent,
to some degree, on pollination by animals (IPBES 2016). In
conjunction, the global pollination crisis and the importance
of this ecosystem service for crop production explains the
relatively high number of studies on pollination in natural
and anthropic ecosystems. Moreover, it is evident that polli-
nator species richness has a beneficial effect on crop produc-
tion worldwide (Dainese et al 2019). Therefore, in tropical
regions such as Brazil, conservation of pollinator species
requires an interplay between conservation biology and crop
production. The second insect-provided service, i.e., decom-
position, is required for nutrient cycling and soil fertility
(Tilman et al 2002). Although fungi and bacteria are primarily
responsible for decomposition, macroinvertebrates, espe-
cially insects belonging to taxa such as Coleoptera, assist in
this process through bioturbation and fragmentation of or-
ganic matter (detritivory) in the soil (Nichols et al 2008,
Power 2010, Schowalter et al 2018). Coprophagous beetles
of family Scarabaeidae, which feed on animal excreta, ma-
nipulate feces during the feeding process. These dung bee-
tles instigate a series of ecosystem services ranging from
secondary seed dispersal to nutrient cycling and parasite
suppression, and also provide valuable services such as bio-
logical pest control and soil fertilization (Nichols et al 2008).
The biological control of pests is generally studied on
Ecosystem Services and Insects in Brazil
Fig 3 Number of Brazilian studies on insect-provided ecosystem services conducted in different ecosystems. The ecosystems investigated were
classified as natural ecosystems (e.g., natural protected areas), anthropic ecosystems (cities and rural areas), semi natural ecosystems (fallow and
regeneration areas), or natural + anthropic ecosystems (when the study evaluated either both ecosystems simultaneously or the relationship between
them)
agricultural land, including interactions between agricultural
land and natural areas. This ecosystem service is provided
primarily by predatory and parasitoid insects (Bianchi et al
2006). Recent studies have reported that greater biodiversi-
ty of natural enemies in Brazil is associated with improved
biological control of pest insects (Pacheco et al 2017, Dainese
et al 2019, Togni et al 2019a), and successful large-scale bio-
logical control programs involving crops such as sugarcane
have been reported in Brazil (Parra 2014).
It was interesting to note that studies on insect-provided
ecosystem services were mostly conducted in natural and
anthropic ecosystems. A similar trend was observed for stud-
ies undertaken in other countries, mainly in the temperate
region (Noriega et al 2018). Natural ecosystems are the main
source of ecosystem services provided by insects to human
activities such as agriculture. Conversely, the conversion of
land to agricultural use has been accompanied by a decline in
native habitat remnants, protected areas, and native vegeta-
tion on private lands in Brazil (Ferreira et al 2012). This is
reinforced by the fact that majority of the studies reviewed
focused on ecosystem services related to economic activities
relevant to Brazil. There is an opportunity to reconcile the
interests of agribusiness with the need for insect biodiversity
conservation, since agriculture is highly dependent on natu-
ral ecosystems. The provision of scientific basis for support-
ing decision-making demands such integration. However, re-
cent cutbacks in the budget for science in Brazil (Magnusson
et al 2018, Andrade 2019), along with the lack of government
interest in biodiversity patterns, can be major obstructions to
this.
789
The Taxonomical Bias
We identified the insect orders and families used as models
in the reviewed studies to determine if the studies were
taxon-biased. Hymenoptera and Coleoptera were the most
frequently studied insect orders (Fig 4a). In the case of the
insect families studied, many articles described results of
biodiversity surveys; therefore, studies on “multiple families”
were the most common, followed by those on Apidae and
Scarabaeidae (Fig 4b). Families that were found only two
records or less in the revised articles were grouped as
"others" (Supplementary Material).
The most commonly studied taxa belonged to megadi-
verse orders, such as Coleoptera, Diptera, Hymenoptera,
and Lepidoptera. At the family level, many researchers stud-
ied multiple insect families simultaneously, which can be
explained by the predominance of biodiversity monitoring
studies rather than that of more mechanist studies.
However, as shown by Oliveira et al (2016), other arthropods,
angiosperms, and vertebrates in natural areas are generally
sampled near access routes. The taxonomical, spatial, and
temporal biases reported by Oliveira et al (2016), and partial-
ly confirmed in this study, indicate that rare and endemic
species have been neglected in several studies.
Consequently, our estimation of biodiversity patterns, such
as species richness, community composition, and endemism,
could be inaccurate. Unlike the temperate regions, tropical
regions are characterized by insect communities composed
mainly of uncommon and rare species in agricultural and
natural habitats (e.g., Price et al 1995, Togni et al 2019b).
790
Fig 4 Insect orders (a) and insect families (b) included in studies evaluating the ecosystem services provided by insects in Brazil
These rare species could be contributors of many ecosystem
services, which, along with how these species participate in
them, remain unknown.
Globally, natural ecosystems are being lost to agricultural
land at an unprecedented rate. This land use often results in
significant reductions in abundance and diversity of the flora
and fauna as well as alterations in their composition. Despite
this, there is little public awareness about the most
important taxa, in terms of their total biomass, biodiversity,
and the ecosystem services they perform. Such awareness is
important for conservation, as without appreciation of their
value and conservation status, species are unlikely to receive
adequate protection. Snaddon et al (2008) investigated
children’s perceptions of rainforest biodiversity, visiting the
University Museum of Zoology, Cambridge, and found that
children have a sophisticated view of the rainforest, incorpo-
rating many habitat features and a diverse range of animals.
However, some taxa were over-represented (particularly
mammals, birds, and reptiles), while others were under-
represented (particularly insects and annelids). Scientists
and naturalists must continue to emphasize the diversity
and functional importance of lesser-known taxa through
public communication, to aid invertebrate conservation
(Snaddon et al 2008).
Identical taxonomical and functional biases were also
reported by Noriega et al (2018) in studies conducted world-
wide. Globally, the information on ecosystem services
Ramos et al
provided by insects is particularly biased towards studies
conducted in Europe and North America. The insects usually
sampled in these regions are conspicuous and large-bodied
species, species that are easy to find, groups that are the
easiest to study (e.g., insects easily reared in the laboratory),
charismatic fauna, and insects that are readily or easily iden-
tifiable (Noriega et al 2018, Cardoso & Leather 2019). A global
underestimation of insect biodiversity can lead to phyloge-
netic, functional, habitat, spatial, and temporal biases
(Cardoso & Leather 2019, Cardoso et al 2020). We also found
that Brazilian research followed this trend, in case of habitat,
functional, and temporal data on insect-provided ecosystem
services. It appears that we experienced the same method-
ological shortfall as the developed world. We are aware that
sampling these highly diverse insects found in Brazil in dis-
tinct ecosystems (including some hard-to-reach areas across
the biomes) is a difficult task inherent to tropical regions.
However, we need to develop ways to deal with such highly
species-rich assemblages that rarely occur in the developed
temperate regions. The advent of molecular tools, such as
DNA metabarcoding, can greatly contribute to this. Unlike
the traditional taxonomic tools, larger quantities of materials
from different sources can be processed using molecular
methods. Such an approach will allow the assessment of
taxonomical diversity, genetic structure, gene flow, and com-
plex trophic links related to insects in different ecosystems.
These are not definitive suggestions, but just one of the
Ecosystem Services and Insects in Brazil 791

several approaches to recognize the importance of insect-
provided ecosystem services in Brazil, and to deal with the
large number of species present in these ecosystems.
The Experimental Setup of Studies
We classified the studies according to how ecosystem serv-
ices were quantified as (i) non-quantitative (i.e., the authors
assumed a relationship between the taxonomic group and
the provision of specific or nonspecific ecosystem services,
but the ecosystem services were not directly measured); (ii)
proxy (i.e., the authors used a specific measure of biodiver-
sity, such as species richness, abundance, or diversity, as a
proxy for the provision of specific ecosystem services, but the
services were not directly measured); and (iii) experimental
(i.e., a specific ecosystem service was quantified using a spe-
cific or nonspecific experimental setup). These categories
were similar to those used by Noriega et al (2018), which
allowed us to compare our results of insect-provided ecosys-
tem services in Brazil to those of such services worldwide.
For the most representative insect orders, insect-provided
ecosystem services were studied using an experimental ap-
proach (62.32%), followed by proxy studies (18.84%) and
then by non-quantitative studies (18.84%) (Fig 5).
In Brazil, experimental studies that quantify the insect-
provided services are the most common, which differs from
the global trend observed by Noriega et al (2018) (see Fig 5).
These results show that, although we lack definitive data
about the ecosystem services provided by insects, Brazil is
moving in the right direction of improving the understanding
and conservation of these services, and consequently, more
effective application and management. However, it is
important to note that Noriega et al (2018) distinguished
“quantification of ecosystem services” from “experimental
studies.” The same distinction was not possible in our study
due to the low number of “quantification” studies in our
sample. Although experimental setup was more common in
our survey, many studies included a very limited group of
taxa in their analyses. We expect that the number of studies
on ecosystem services provided by insects will increase in the
near future, and if a higher number of such papers are pub-
lished, a new survey addressing more experimental setups
can be conducted.
The shortfalls in the knowledge of insect-provided ecosys-
tem services in the world are related to a variety of method-
ological approaches employed (Noriega et al 2018). Most
studies relied on an observational approach, manipulative
experiments, or use of proxies of ecosystem services. It must
be noted that the relationships between richness and abun-
dance are not always appropriate as proxies for ecosystem
services and function (Solar et al 2016). They fail to link bio-
diversity patterns with species traits and related ecosystem
services (Noriega et al 2018). The experimental design is
more desirable as these links can be directly established.
Our concern is that the experimental setup of many studies
was biased towards the most studied insect-provided eco-
system services. Though experiments were conducted with
sufficient replicates, laboratory studies lacking a clear rela-
tionship with field data and patterns were also common.
However, for some ecosystem services, such as nutrient cy-
cling and seed dispersal, it will be much more challenging to
standardize the experimental setup than for other ecosystem
services, such as pollination and biological control (Moretti
et al 2017).
Irrespective of the experimental approach, certain short-
falls concerning ecosystem services and their relationship
with biodiversity need to be overcome. According to Hortal
et al (2015), studies on biodiversity patterns and function
present seven key shortfalls, which are knowledge gaps in

Fig 5 Approaches used to evaluate insect-provided ecosystem services in Brazil. The studies were classified according to how ecosystem services were
quantified as (i) non-quantitative (i.e., the authors assumed a relationship between the taxonomic group and the provision of specific or nonspecific
ecosystem services, but the ecosystem services were not directly measured); (ii) proxy (i.e., the authors used a specific measure of biodiversity, such as
species richness, abundance, or diversity, as a proxy for the provision of specific ecosystem services, but the services were not directly measured); and
(iii) experimental (i.e., a specific ecosystem service was quantified using a specific or nonspecific experimental setup)
792
species description and taxonomy (Linnean shortfall), geo-
graphic distribution at different spatial scales (Wallacean
shortfall), abundance in time and space (Prestonian short-
fall), evolution of species and their traits (Darwinian short-
fall), link between species’ traits and ecological function
(Raunkiaeron shortfall), responses and tolerances to abiotic
conditions (Hutchinsonian shortfall), and species interactions
(Eltonian shortfall) (Hortal et al 2015). The Linnean and
Wallacean shortfalls are probably related to the taxon and
biome biases reported here. Assuming that there is a rela-
tionship between biodiversity and ecosystem services, the
experimental setup of future studies should primarily focus
on overcoming the Hutchinsonian and Eltonian shortfalls.
Overcoming these two shortfalls will enhance the ability to
predicting how insect-provided ecosystem services will oper-
ate and how they can be managed in a changing world.
Challenges and Perspectives
Although Brazil is a large, megadiverse country encompass-
ing a range of ecosystems and biomes (ecological diversity),
our research into the role of insects in providing ecosystem
services demonstrated that scientific knowledge on this topic
is very scarce in the country. In recent years, Brazil has be-
come the largest consumer of pesticides, with an annual
expenditure of US $17 billion for pest control (Oliveira et al
2014), and has experienced increased deforestation for agri-
cultural intensification, urban expansion, higher rates of dis-
eases such as measles and dengue, and environmental prob-
lems caused by dam disruption. Insect-provided ecosystem
services contribute considerably to the functioning of various
processes. For example, insects can substantially increase
crop production through pollination, serve as natural ene-
mies against pests, and assist in the cycling of nutrients.
Therefore, it is imperative to provide scientific evidence dem-
onstrating the importance of this megadiverse group to the
society. Using the term “ecosystem services” in research
articles can help to make the information easily retrievable
by those who seek scientific evidence to support decision-
making. The spatial, economic, functional, and taxonomic
biases of studies performed in Brazil highlight the need to
expand the research efforts to include (1) biomes other than
the Cerrado and Atlantic Forest, (2) ecosystem services relat-
ed to human well-being other than economic activity, and (3)
rare and endemic insect species. Filling these gaps is the first
step towards fully understanding the role of insects in pro-
viding ecosystem services that are essential to humans, mon-
etizing the value of this megadiverse taxonomic group, and
incorporating scientific evidence of their services in decision-
making.
Presently, in Brazil, there is a clear tension between agri-
business expansion and conservation efforts mediated by
Ramos et al
some politicians and some sections of the society. Such ten-
sions have resulted in cutbacks in scientific efforts and the
amount of budget available, particularly for conservation bi-
ology (Magnusson et al 2018, Andrade 2019). We argue that
this tension, although unnecessary, is an opportunity to con-
duct integrated studies, focusing on both basic and applied
issues. Such integration can increase the number of sites
being sampled both within and between biomes, and im-
prove and standardize the experimental setup. Studies on
insects in Brazil could be at the forefront of the integration
of several areas, such as ecology, evolution, agronomy, and
taxonomy, in light of the ecosystem services provided by
these organisms. However, integrated studies will not be
sufficient to reach the society and policymakers of the coun-
try. We need to improve our ability to communicate with the
society and policymakers, which could be the basis for chang-
ing and valuating the ecosystem services provided by insects.
As researchers in a tropical, megadiverse, and politically po-
larized country, it is our duty to ensure that reliable scientific
information is available to the policymakers, which can aid in
reconciling economic interests with the need for biodiversity
conservation. Entomologists involved in different research
areas will play a pivotal role in improving our understanding
of these ecosystem services, for developing a truly sustain-
able country.
Acknowledgments This study was supported by research grants and
fellowships provided to the authors by the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq) and Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES). MRF is a CNPq
fellow (process no. 313952/2018-3).
Electronic supplementary material The online version of this article
(https://doi.org/10.1007/s13744-020-00781-y) contains supplementary
material, which is available to authorized users.
Author Contributions PHBT and MRF conceived the study. DLR, WLC,
JE, LAL, MVCR, and PAG collected the data and performed the analyses,
under the supervision of MRF and PHBT. DLR and PHBT led the writing
of the manuscript. All authors contributed critically to the drafts and
have approved the final version.
Electronic supplementary material The online version of this article
(https://doi.org/10.1007/s13744-020-00781-y) contains supplementary
material, which is available to authorized users.
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